Appel A 2 - n ae seas 5g yoy a ; see nnd Te EOS Tate Powe arene a phil TEE Tee OLE Gil NL PEPER oe At wal T Atlee s/s vena AP ee ial, rs POOR 8 EERE S CEES SINE SE BETA LH Bae a ih Lh OPER EH ALE the ge a : - a GNI por Mae ate Sarath Ay re ne: PLL rere Aes Pa te ates RIE SE tne, re stid iv 22h pF eens Sethi ine th Ri a Mell gare i en NATIONAL HERBARIUM : OF VICTORIA 3°14 MAY 1983 LIBRARY ee JOURNAL of the ADELAIDE BOTANIC GARDENS Volume 5 29 September, 1982 Instructions to Authors Topics 4 Papers will be accepted in the following categories: (a) Plant systematics (Australian and horticultural groups); (b) Descriptive plant morphology, anatomy and ecology; (c) Obituaries, biography and history; (d) Bibliographic studies, book reviews; (e) Botanical illustrations; (f) Noteworthy horticultural contributions. Copy Manuscripts must be typed, with double spacing and margins at least 3 cm wide, on one side of the paper only. Three copies must be submitted. Captions must not be italicized, underlined or typed in capitals. All scientific names of generic or lower rank must be underlined. The print area for illustrations is 20 x 13 cm (including captions). Half-tone material should be submitted this size if possible, but will be reduced by the printers, if necessary. Reprints 50 copies of reprints will be provided for each paper. Additional reprints may be purchased at cost. Layout The pattern of the paper should generally be: (i) Title; (ii) Author and Address; (iii) Abstract (except for short papers); (iv) Introduction and subject matter; (v) Acknowledgements; (vi) References. References Text references to publications should be indicated as follows: (Smith, 1959), (Smith, 1959, p. 127), Smith (1959) or Smith (1959, pp. 125-208). The final section of the paper, headed ‘References’, should include only those titles referred to in this way. It should be laid out as follows: Smith, L. L. (1879). The species of Danthonia found in pastures in Victoria. Austral. J. Bot. 65: 28-53. Bentham, G. (1868). “Flora Australiensis”, Vol. 4. (L. Reeve: London). Baker, J. G. (1898). Liliaceae. In Thiselton-Dyer, W. T. (ed.), “Flora of Tropical Africa”, Vol. 7. (L. Reeve: Ashford). Journal abbreviations must be consistent within a paper and authors are recommended to follow “Botanico- Periodicum-Huntianum”. Journals not cited in B-P-H should be abbreviated to conform with this general pattern. The following abbreviations for Australian states should be used: WA, NT, SA, Qld, NSW, ACT, Vic., Tas. Text references to specimens should be italicized, for example Kock 276. Indices When required, follow the pattern on, for example, p. 106 of vol. 1, pt. 2. Recommendations on taxonomic papers Synonymy Authors are requested to include in the synonymy only references to publications containing information additional to that to be published in the paper being submitted. Within this section journal and book titles must be consistently abbreviated. B-P-H journal abbreviations and book titles abbreviated in a similar way are desirable. Authors of references cited in the synonymy should be abbreviated. References may be cited as: Benth., F/. Austral. 4: 111 (1868) OR Benth., F/. Austral. 4: (1868) 111. Citation of specimens 10-30 specimens should be cited for each species (or subspecific taxon), although this may be varied under certain circumstances. The author may decide whether or not to include dates of collections and the sequence, provided a constant pattern is adhered to throughout a paper. Authors wishing to cite all specimens seen may list them all in an index to collectors after the style of the “Flora Malesiana” identification lists. Collections not identifiable by a collection number (assigned by either the collector or herbarium) should cite dates. Correspondence All correspondence concerning the journal should be addressed to: The Director, Adelaide Botanic Gardens, North Terrace, ADELAIDE, South Australia 5000. Editorial Committee B. Morley (Ditector, Adelaide Botanic Gardens) J. P. Jessop (Co-editor) Consultative Editors D. E. Symon (Waite Agricultural Research Institute) H. B. S. Womersley (Botany Department, University of Adelaide) ISSN 0313-4083 J. Adelaide Bot. Gard. 5: 1-304 (1982) TAXONOMIC STUDIES IN EUPHRASIA L. (SCROPHULARIACEAE). A REVISED INFRAGENERIC CLASSIFICATION, AND A REVISION OF THE GENUS IN AUSTRALIA W. R. Barker State Herbarium, Botanic Gardens, North Terrace, Adelaide, South Australia 5000 Abstract _ The infrageneric classification of Euphrasia has been radically revised in the Australasian and Malesian regions. Fourteen sections are proposed. Two sections, Pauciflorae and Euphrasia (Semicalcaratae auctt.), comprise two and four subsections, respectively. The sections Malesianae, Cuneatae, Phragmostomae, Pauciflorae and Lasiantherae, and the subsections Pauciflorae and Humifusae are new and the sections Striatae, Scabrae, Novaezeelandiae, and Anagospermae are given new status. The generic type E. officinalis L. is not a nomen ambiguum in the sense of the ICBN; it should be reinstated as the correct name for E. rostkoviana Hayne. The classification of the genus in Australia has been completely restructured, with 42 taxa formally treated. Eighteen species are recognized, of which E. ramulosa, E. phragmostoma, E. semipicta, E. bowdeniae, E. lasianthera, E. eichleri, E. orthocheilaand E. ciliolataare new and E. caudata, based on E. scabra var. caudata Willis, is given new status. Subspecies are proposed within four polymorphic species. E. gibbsiae has nine subspecies, of which four are new and four have changed status; E. collina, which has caused much confusion in the understanding of the genus in Australia, has 14 subspecies, four new and nine of changed status; E. crassiuscula has three subspecies, one new and one of changed status; and E. orthocheila has two subspecies, one new. In the highly variable species the degree and nature of distinction and amount of intergradation between subspecies is discussed, sometimes as a result of detailed field study. Clines and a limited amount of hybridism occur within species. Interspecific hybrids are rare. In a chapter on morphology, the taxonomic and, where possible, biological significance of morphological attributes in Euphrasia is discussed. Important characters, e.g. the distinctive habit types in the perennial Species, acropetal branch development in the New Zealand annuals, corolla coloration, and the high chromosome numbers of Australian species, have been previously poorly documented or unknown. A number of other characters are redefined to improve their usefulness in classification, while others are shown to be of reduced significance. The origin and diversification of Euphrasia both on a world scale and within Australia are discussed. Cladistic analysis of the genus is difficult at this stage of our knowledge. However, through the postulation of primitive states of a few characters, several relict species are proposed which are remarkable in their scattered distribution in both hemispheres. From the character states present in the proposed relicts, six apparently monophyletic lineages are distinguished. Diversification of the genus seems to have been in at least two phases, one possibly in the early Tertiary or before, the other, from which most extant species are derived, since Neogene times. Apart from the recently derived, highly specialized Sect. Euphrasia, the genus shows no capacity for long-distance dispersal; much evidence points to the need for migration across suitable land. An association of at least two relict species with Nothofagus, which with Fagus has a distribution homologous to Euphrasia, is among the evidence presented for Euphrasia attaining its bihemispheric distribution by the early Tertiary. Deficiencies in the present fossil record allow latitude for this theory. W. R. Barker J. Adelaide Bot. Gard. 5 (1982) CONTENTS Toe Introduction 7s scl ncetet foals ele crscsesnrts teri etercicter croton tient a Eee retncn 49, TRV WE ye sinon ei arts. 3 II. Morphology: measurement, terminology, diagnostic importance and biological relevance ............ 4 AvaMcasurementoficharacterstarrrssiemren mel tatters ieee svar arctan nit arte Pron nTa ies Prva 4 Be Characters j:. tlvyorerscs-2¥eletateps fens stete tt ae aisle eateoe | RE TCY.TeRT Pa OL One, Eo es er ESTE Core Tree pron Paar 5 Me, UE EN eceaonnceroe dr paeta hs daeed Sten Udita deal cata Giandion soko dase keanpanonnon 5 2:@Root'system andjsemi-padrasitism a..- siete el ee eee ee ee tn eres ore eee 5 sho JRE ASCS. ce edhnotdinahacdener sae tcboddaauh co casnapeyn sehr dboseoneng sain is 5 aweHabitiinithe-annualsve se, nets eee Ree en eee en ee en enn, aha ten ar an 6 be aHa bitin thesperennials mene sme ae tl beets onarhare on een eee ment RW el, geese 8 4 pel nd men tum srepelerster eats arta serene coer dey teste ct Oa lela Meare Ne eston Saye ar reSne rece) Peele ie 12 am Glandularnhairsyprmeta 1a -tatttter teen teeny tecten erin Tite iste inte rereietareoe tts iisieisn ie) are 13 bSiEglandularhairs is. otc. Ce Ronee rents aia create astern eta eer cca is 13 cmp Séssile/glands Ryfie eit tcstee, danas ce ae tne ae Ree Meee aE een any 14 SSE sL CAVES ANCIDIACtS tsar wclernerstese hee Te etree oN in Peta a ee We EN oy 14 Gre lnflorescenceregemaeet een tte tats et eee ee eee eae Nee Leen. hee 25 (ee Cal y xtc as attaagh, sree teoirdrs SN ee Apt tule Ren scrapes roe oor rev OW Teeyereie 3 0 tet no i re” 27 8B Ma Corolla ptia are oie sae hale mercy OANA Et IMT. Fy MONS PARTIE de 27 ase Shape and size sth cass ace cer eee necting wee ey eC th ny mont an ENMU. Mme ir Hey a 27 bss Ground colourityet Sti.cr Sine en ertoe te irl Me ee cr ney Nien, Tames Mapes a. Wane 29 C27 Coloration ig tter satelite eee OTe nee tele cheep dn eee Ti Ree sae 29 Gis Corollaiind umentumie rt swrctstetetatestentene ce beer heen or eee tee apne AEE Pe Opera a 31 bd Zero Tin ane EMs creo rote or Anne aahinhbandhnemyronsdrintincpdeloncocohe 31 LOS Gynoecium ©, Spare ste sr aah ne tat a ee nn iar Crean fens Pe TON ee CET Pe eae P|, dee 33 (CET adi nett ditt rinnrdteinh An ddl heroine San db mae ahtdene aetna a4 34 Ae SES otic hod eC oO ic doin Mt omer ie rete naNe la ytd hil dd tty baleen Awl Re eke 35 137 Chromosomenumbergn psc seater ties eee Okabe hac ecininritieerar tie cette 35 alse Method gents. eaaesceiege ees specialty sede oath toll en POON ye diac elas araisa they 37 bh Results ye. A7enrr (0 We Vas HID eee eerie ee ORE VIE, AIRY ETM, aM SN) Teoh PO 37 14sBollenisterility teststaet eee sete te ote ee eT ee een eee. Syne 40 C&Blorallbiolog yates cette 1 eens eet TENT Ae ee POO eR Si NLR ED 40 III. The origin and diversification of Euphrasia .....sse.csccceeeeccsccuvecceeeeccseesccceeeuens 46 AsaPast hypotheses isu. stig meretiemeen recat ote eR erent eee eee eit. baat 46 B. Problems in establishing cladistic relationships of the sections and subsections of Euphrasia ...... 48 C. Relicts of an early phase of world evolutiOn....2..ee.secseeccecceccccccuscesevecevesuceocs 53 BJ WETS A) CUO ETS AME hn sats soon ig sandan do dobubbpudcoapanndacudseieuupondondate 53 E. Timing of dispersal events across barriers to migration ..........0.sececcececcecuccuceeeuces 57 Fa Means olidispersal Finn peceacltur ela eee i ERLE: ne. Pater ennai Rater een 58 G. Diversification of Euphrasia in Australia ....¢.0..+0.e0ccessecssovsceresaceverccetsececess 60 HeEuphrasia—an oldilertiary genus een aera ge ee Ren T nC ee ear et nee ne 66 Tres Conclusions. c' sco: de:afetez sce) sso! lagna olny eee ee TN Te ny Or TO RVR. TN NLT, 68 IV. A revised infrageneric classification of Euphrasia .........0.ceccecuccucuvcuccecucsccuceccucens 69 AvETevious intrapeneric classifications mee tree rtiiermete reese anette are ern ean 69 BIST NE TEVISIONs eon actscteietee cae lscitaret rite Lee tinte etre site Tee TC eT TT ee 72 Wee lntroduction, sitter. othe nave trol tose antes by splt TCL Oy eas, eae es aM rc roy. 72 2 Sima Revision Mey. jcateteic. «t1gee eee be tds atone oni cke Fe Brea SEU eds ion EA HOR ee ae oe 73 V. A taxonomic revision of Euphrasiain Australia .........00..ceeecceeccececceucetanceeaeces mol Fen OIG NSCS Wo odnedsoawe ao Hodcsabobbodddunoamtiatindeond sO Ane Losasnoneousnem noe o: 91 BSTaxonomic'history, seer tne tt cate pear jae ate TN Te ety ne ee wp ED 91 Claxonomic philosophy.and presentations. cee tertecne ea an sen rte emtnineen eet cen 95 DT hesrevision tent tAael kl ere aeieiant. Sere o lee Mee RTOE. UAE rela ne cate Satewl: 98 Key to the sections of Euphrasiain Australias j..00.00s+4s0-c0scescecersisescssacceeoesvis ss 98 Interspecific: hybridsgasMe. oattey. SIMA 4th, Soe 1s Te, CRE AO) SLT SOE ae 287 E. Index to numbered Australian collections ..........0cceseccecceccececcecsceecsccecerces 291 VIeerAcknowledgementsirectaecin meee reece hon tan Ri ie ete tn tee 293 VAL RA Orie nod be cohniapotinadinodnasbebonBenticsughadedbhhooh poAabyaaneabosmoroausnese 294 VITTAIndexito scientific na mes tetetestsree iste eee) ate cern ree ae ee eee ace 299 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) type of E. bowdeniae and the subspecies of E. orthocheila, and the types designated for E. phragmostoma and E. ciliolata should be noted. In addition a formal division of E. caudata into two subspecies has been decided against. Most annotations of relevant specimens seen will not have been modified. Study in the field or of fresh material in 1975 of a few New Guinea species, and field work in early 1979 covering the majority of New Zealand taxa have helped consolidate many ideas on the infrageneric taxonomy and evolution of the genus. A number of essential changes have been made. The remaining observations are left to future revisions of the genus in these countries. Il. MORPHOLOGY: MEASUREMENT, TERMINOLOGY, DIAGNOSTIC IMPORTANCE AND BIOLOGICAL RELEVANCE The following is an enumeration of the attributes which have been described in the present and past taxonomic treatments of Euphrasia. Consideration is given to the nature and significance of the variability of characters and their diagnostic usefulness at the various levels of classification. Floral biology, which involves characters of several organs, is discussed separately. A. MEASUREMENT OF CHARACTERS Much of the taxonomic difficulty in Euphrasia stems from the closeness of the taxa, with the major differences often overlapping to a greater or lesser extent. It has therefore been considered advisable to adhere where practical to a policy of using quantitative terms to describe diagnostically useful characters in cases where the lack of precision, characteristic of many qualitative terms, might obscure the true variability of the characters. Even where precise mathematical definition has been given to some qualitative terms (e.g. the chart of simple symmetrical plane shapes: Stearn 1966), their actual usage is liable to be imprecise. For each taxon the measurements of each character were usually taken from a sample of twenty to thirty specimens selected to cover the extremes of the geographical, ~ ecological and altitudinal range of the taxon. In the more widespread taxa, many more measurements of the variable characters were made. The range of variation in quantitative characters has been portrayed in the descriptions as follows. In presenting the measurements, the extremes of variation are usually placed in brackets at either end of the range. Most commonly, between these bracketed extremes is given the range of values covering the 80% of variation remaining after the 10% smallest and 10% largest values have been removed. If there are insufficient specimens to give an adequate sample only an average of the measurements taken is placed between the bracketed extremes. In a few obviously distinct taxa or where an organ is poorly collected only the overall range of variation has been given. All measurements in centimetres (such as plant height) were taken to the nearest centimetre using a ruler. Small organs requiring millimetre measurements (such as corolla and leaf length and leaf tooth dimensions) were measured under a low-power microscope at 10x magnification with an ocular micrometer and were recorded to the nearest tenth of a millimetre. Measurements of less than one millimetre (such as awn length) were recorded to the nearest twentieth of a millimetre. Some other: measurements are also given to the nearest half of an integer or decimal fraction, depending on the size of the organ involved, and particularly in cases of insufficient material. J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia B. CHARACTERS 1. Life-span Both annual and perennial types of life-span are common in Euphrasia and their distinction is one of the major characters used in the infrageneric classification proposed in this work, and in the previous ones of Wettstein (1896) and Pugsley (1936). The longevity of the perennial species has not been investigated, but it is considered that the sturdy Australian perennials may survive for several years. Generally, in populations of perennials first-year plants are much more scarce than plants of the previous year (detected by weather-worn old infructescences). E. hookeri, however, inhabiting a severe alpine environment, is one species which has the capacity to perennate but may rarely do so (q.v.: note 2). 2. Root system and semi-parasitism The plant is a facultative semi-parasite with European species having the ability to parasitize a range of host species (Yeo 1964). Swellings or “haustoria”, which are located on the finer rootlets, attach to the fine rootlets of the host. Evidence of these haustoria is present in most specimens which have been extracted from the soil with care, but fragments of the host roots still attached to the haustoria are less common. I have observed haustoria in all Australian taxa. The presence of haustoria is widely documented for European species and has been described (Philipson 1959) for E. cockayniana, E. zelandica and E. revoluta of New Zealand. J have also seen haustoria in material (in AD) of E. cuneata (Sect. Cuneatae), all species of Sect. Pauciflorae and E. integrifolia (Sect. Anagospermae) in New Zealand, E. papuana, E. mirabilis and E. lamii (Subsect. Pauciflorae) of New Guinea, and the species of Taiwan (Sect. Malesianae: specimens ex TAI). Material from elsewhere in Malesia and South America was not inspected for haustoria. Yeo (1964) reviewed experimental work in semi-parasitism on the northern hemisphere annuals. Since then Weber (e.g. 1976a,b,c, 1980) has published a continuing series of studies on the haustorial anatomy making comparisons with other genera of Subfam. Rhinanthoideae and Orobanchaceae, while Kuijt (1969) has reviewed physiological aspects of semi-parasitism, including theory that the sessile gland patches on underside of the leaves (see p. 25) produce a high transpirational pull on xylem contents of the host. Simple slender roots may emerge from prostrate axes. This is probably a response to permanently moist conditions. 3. Habit Within both the perennials and annuals of Euphrasia there is a variety of habit types. Differences in habit often appear to be associated with climate. Thus, similar habit types may be found in distantly related species under similar climatic conditions, while closely allied taxa, sometimes within a species or even variants in the one subspecies, may differ markedly in habit in association with corresponding climatic differences. In this work the term stem has been used in the sense of Jackson (1928: “the main axis of a plant, leaf-bearing and flower-bearing as distinguished from the root-bearing axis”), The term stem seems to have been used more broadly by Wettstein (1896), Pugsley (1936) and Du Rietz (1948a,b), as well as the majority of flora writers, in describing the perennials and prostrate annuals. The first-year plants of several Australian species may have a single axis which is erect or ascending, is often profusely W. R. Barker J. Adelaide Bot. Gard. 5 (1982) branched, and produces an inflorescence. On the above definitions this would be the only true stem with any axes arising from this being branches. In subsequent years the stem may die back, often to the region of the upper branches. If the branching is confined to near ground level the plant has a “many-stemmed” appearance. These axes are, however, of a secondary (or greater) nature and should not by the above definitions be called stems. The presence of a single stem is clearer in those perennials which branch high above ground level. In these much of the stem persists throughout the life of the plant. The term main axis(es) refers to the stem, or in perennials after their first year, the robust flowering branches. The term main inflorescence-bearing axis(es) derives from this. Characters useful in defining habit are: the height of the plant or, to exclude variation caused by differences in the stage of development of the branches and inflorescence, the height of the main inflorescence-bearing axis to the base of the inflorescence; the position of the uppermost branches or young shoots onthe maininflorescence-bearing axis (measured both by the number of nodes below the inflorescence, and by the proportion of main inflorescence-bearing axis between the inflorescence and the node bearing the uppermost shoot); the sequence of development and direction of the branches; and the number and length of the internodes on the main inflorescence-bearing axis relative to the length of the leaves. a. Habit in the annuals The habit type consisting of a single erect stem often branched at successive nodes is characteristic of almost all annual species of Euphrasia. In the Australian annuals the branches develop basipetally at consecutive nodes. Elsewhere, however, the order and consistency of branch development is not so strict. In the northern hemisphere species there is also a tendency for branches to develop basipetally but they are often, in the less robust plants, very few or even absent. In the New Zealand annuals of Sect. Novaezeelandiae branching is similarly rather sporadic and tends to be acropetal. The more ordered and consistent development of branching in the Australian species may be related to their apparent capacity for more vigorous growth, enabling stems to bear many internodes and flowers to be produced in large numbers. Two species of Sect. Anagospermae of New Zealand, E. disperma and E. integrifolia, form loose mats with almost completely prostrate branches arising from a reduced stem (see Ashwin 1961). The remaining two species of the section, E. repens and E. dyeri, are intermediate in their habit between these species and Sect. Novaezeelandiae. Until the present revision, habit in the annual species had only been studied in the extensive northern representatives of Sect. Euphrasia, in which the phenomenon of “seasonal (or pseudoseasonal) dimorphism” has been widely documented. Similar habit polymorphism occurs in other genera of the Trib. Rhinantheae e.g. Melampyrum and Rhinanthus (Soo & Webb 1972a,b). The two habit types of Sect. Euphrasia have been characterized as follows by Pugsley (1930). Aestival or early summer-flowering form: Internodes very long. Flowers forming from generally the 4th stem node. Branching consequently limited. Autumnal or late summer-flowering form. Internodes very short in lower parts, [? longer above]. Flowering from the 5th-12th (or more) stem node. Branching copious. Yeo (1964, p. 7) has made slight modifications of these categories. The various habit types have been correlated with climatically based differences in the duration of the growing season. In Euphrasia the aestival form is found in alpine or J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia arctic regions with shorter growing seasons. It is also found in meadows which are frequently mown and where, accordingly, the time between germination and the production of seed is necessarily short. The autumnal or late summer-flowering forms occur in more lowland regions or lower latitudes with correspondingly longer annual periods of growth. Pugsley (1930, 1936) defined a number of series in Sect. Euphrasia (his Sect. Semicalcaratae) partly on the basis of these habit differences. However, he and subsequent workers, such as Juzepcuk (1955), Sell & Yeo (1970) and Yeo (1972), have found the distinctions between the autumnal and aestival types in different complexes too variable to be useful in defining species. They have preferred to describe each taxon by its range of variation in the number of stem nodes, the length of the internodes and the relative abundance of branching. Yeo (1964) has shown that in Sect. Euphrasia habit differences of this kind are retained when different forms are cultivated. together, indicating that they are genetically determined. In addition, he observed precocious flowering (with a consequently reduced stem node number) in some plants of the normally later summer- flowering E. pseudokerneri during periods of unusually high temperatures: “this suggests that temperature is one of the main factors controlling flowering in Euphrasia, and that given sufficiently high temperatures E. pseudokerneri can be made to flower at about as low a node as the high mountain species”. The length of the potential growing season as determined by the climate also appears to be a critical factor in determining the habit type of the Australian annuals (Sect. Scabrae, Sect. Lasiantherae). The longest potential growing season in the regions occupied by the annualsis likely to bein New South Wales, particularly in northern montane habitats where there is both summer and winter rain and winter snows are rare. The two annuals of this region, £. argutaand E. orthocheila, apparently in montane and /or lowland parts possess the tallest habit and the largest number of stem nodes of the Australian annuals (table 1). Elsewhere in Australia the length of the potential growing season is shortened in one of two ways with an accompanying reduction in height and a number of stem nodes (table 1). In montane and lowland temperate Australia, in which E. scabra is widespread, snows are generally absent, but rainfall predominates in the winter months and the summers have irregular rain. The growing season is thus effectively shortened. The reduction in growing season for the remaining annuals which occupy alpine and subalpine zones is caused by the winter snows, with conditions ameliorating earlier with decreasing altitude at a given site, allowing for local differences such as in aspect. There is a progressive reduction in plant height and node number from the predominantly subalpine taxa, such as E. ciliolata and E. caudata through to the predominantly alpine E. alsa. E. eichleri, which occupies both alpine and subalpine localities, is intermediate between these types, while within E. caudata the dwarf variant occurs at a high subalpine site. Table 1. The climatic ranges and variation in selected characters of the Australian annuals of Euphrasia Taxon Height of stem No. of stem No. of flowers No. of ovules Length of Climatic zone to base of nodes in stem seeds _—— inflorescence inflorescence (mm) Alpine Sub- Mon- Low- (cm) alpine tane land E. alsa 1,3-5.0(6.5) (1)3-4(5) ¢.15-30 10-12(15) 1.8-2.5 + (+) E. eichleri (2.5)5-11(14) (5)6-8(12) >15 (20)30(40) 1,5-2.0 ye rs E. caudata dwarf variant —(3.5)6.7(11.0) (5)6-8(9) (10)18(26) (20)37(45) (0.9) 1.2-1.5(1.9) (+) + common variant (7.3)11-24(30) (8)10-14(17) (12)24(32) (30)62(110) (1.0)1.3-1.5(2.0) + (+) E. ciliolata (11)16-23(25) (10) 13-16(19) (10) 16-26(36) c.30-75 (0.9)1.0-1.2(1.4) + + E. scabra (7.5)11-25(45) (6)8-18(24) (10)14-32(54) (57)90(140) (0.4)0.5-0,8(1.0) + + E. arguta (12)22(33) (18)24(30) (30)50-90 .35-80 (0.4)0.6-0.8(1.0) + MH E. orthocheila (15)17.5-37(50) — (10)13-30(36) (15)25-40 (20)45-100 (0.4)0.5-0.8(0.9) oF vee ——— W. R. Barker J. Adelaide Bot. Gard. 5 (1982) b. Habit in the perennials Habit varies greatly in the perennial members of Euphrasia. Some habit types are important in distinguishing species or sections; others may show intraspecific poly- morphism. Asin the annuals, differences in habit type tend to be geographically restricted, but not all variants can be related to climate. The characters used to define the different habits are the portion of the main axis (axes) in which branches develop and the portion which remains simple, the direction of branching, whether the branching is at consecutive or occasional nodes, and the sequence of development of the branches. The various types are illustrated diagrammatically in figs 1 & 2 and are described as follows. The “Malesianae” type (as in E. philippinensis). This habit type is common to all species of Sect. Malesianae, E. papuana and E. culminicola (together with some specimens of E. /amii, which may be conspecific with the latter) of Sect. Pauciflorae Subsect. Pauciflorae and probably to some South American species. First-year plants are very scarce in collections, but may resemble those in the holotype of E. merrillii (Merrill 4720 NY: see Du Rietz 1932b f.5), in which there is apparently a single flower-bearing stem with lateral flower-bearing branches produced from the basal nodes of the plant; subsequently lateral branches probably develop at higher nodes. Most whole-plant material seen is clearly from subsequent years. Arising from their base such plants have several to many main inflorescence- bearing branches along which lateral branches occur sporadically in no fixed sequence. The lack of regular production of aerial branches may be related to the less robust nature of the main axes of these species and the lack of strict seasonal growth in their habitats in the high montane to alpine zones of tropical and subtropical mountains. The “Cuneata” type (as in E. cuneata), fig. 28. This habit type is restricted to Sect. Cuneatae of Australasia and Sect. Atlanticae of the Azores. Plants of this type have branches developing in a more or less strict basipetal sequence at consecutive axils from high on an erect inflorescence-bearing axis just below the inflorescence to ground level, just as in the annuals of the northern hemisphere and Australia. Two variations of this type can be distinguished on the abundance of branches developing from the main inflorescence-bearing axes, with the Azorean perennials tending to produce fewer lateral branches; there is a similar divergence in habit between the annuals of Australia and the northern hemisphere. The exact nature of first-year plants of this habit type is as yet unknown. In E. ramulosa, however, these plants may at least occasionally consist of a single erect or ascending stem in the first year; this dies back completely, producing several erect or ascending stem-like main axes from the perennating base in subsequent years. Alternatively there may be several main branches developing from the base in the first year, with die-back and subsequent development as in the previous case. Rooting may occur in the very proximal part of the axes. The species with this habit type occupy coastal scrub to subalpine habitats. It is proposed that the habit type is primitive in the genus (p. 53). The “Paradoxae” type (as in E. formosissima). This habit type is known only in E. formosissima (Sect. Paradoxae) of the Juan Fernandez Islands. Skottsberg (1921) referred to its unusual habit when he first described the species. The plant consists of a single erect stem with branches developing in consecutive axils and with the stem and lateral branches flowering simultaneously. The number of non-flowering nodes on the branches decreases from the base of the plant, where they have about four nodes, to below the inflorescence, where they lack nodes. The development of the branches is apparently acropetal. This is known in no other perennial of Euphrasia, and reflects a common ancestry with New Zealand annuals which also have acropetal development of branches. E. formosissima is the only J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia First year (pre- sumed) First year Later year “Malesianae” type gH a Later year (portion of plant) “Paradoxae” type “Phragmostoma” type Fig. 1. Habit types in the perennials of Euphrasia (1): the “Malesianae”, “Cuneata”, “Paradoxae” and “Phrag- _ Mostoma” types. (Where individual flowers are not shown, inflorescences are densely flowered and represented by their outline; branch remains of prior years are represented by thick lines, those of the current year by fine lines; nodes are shown as fine horizontal lines on the current year’s branches.) W. R. Barker J. Adelaide Bot. Gard. 5 (1982) species with a means of perennation in which the stem and main branches (possibly after a dormant period) consistently continued to develop leaves and new shoots beyond the inflorescences. The species occurs in montane to alpine zones (Skottsberg 1921). Later year “Humifusa” type “Collina” type Fig. 2. Habit types in the perennials of Euphrasia (2): the “Striata”, “Humifusa” and “Collina” types. (For explanation of diagrams see fig. 1.) 10 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia The “Phragmostoma” type (as in E. phragmostoma), fig. 29. This habit type is confined to a single species in south-east Tasmania. First-year plants are as yet unknown and difficult to envisage. The main inflorescence-bearing axes seen have developed laterally from portions of an older axis from the previous year. These ascending main axes have each developed widely spaced groups of lateral branches, the distal group usually immediately or to 10 nodes below the terminal inflorescence, the next group several to many nodes below the distal one and earlier developed. Development of the branches in each group tends to be strongly basipetal, but occurs over only 2-3(4) consecutive nodes; occasionally, branching is limited to one node only. The more robust of the lateral branches also show development of two widely spaced groups of basipetally produced branches over 2-3 consecutive nodes, and the development of terminal inflorescences. The specimens seen show similar development in the stronger laterals of these branches. Young foliose lateral branches are erect, but the stout axes of previous years growth are procumbent. The remains of the old axes which the main inflorescence-bearing axes had developed and the main axes themselves show die-back at the terminal growing points. Perennation of the plant is therefore usually by means of vegetative lateral branches. There is occasional reversion of the inflorescence rachis to a vegetative axis but it is not consistent as in the “Paradoxae” type. It is probable that E. phragmostoma, by its profuse branching, is one of the largest representatives of the genus. Whether there is any association between climate of the temperate coastal habitat and the initiation of the groups of branches or occasional vegetative growth of the axis after flowering seems worthy of study. The “Striata” type (asin E£. striata), figs 37-41, 43, 44, 46, 47, 66, 68, 75-80, 82-84, 87, 89, 100. The majority of perennials in Australia, east Malesia, New Zealand and Australia have this type of habit, as may some of the South American species. Plants of this type have an erect or ascending stem terminated in the first year by an inflorescence. Vegetative buds are confined to ground level, and branches which are formed are also ascending, or prostrate in the proximal parts (sometimes rooting at the nodes) and distally erect. The habit found in E£. hookeri (q.v.: note 2; figs 46, 47) is clearly a derivative of this type. Plants of this species are single-stemmed, sometimes with shoots forming at the base. The apparently rare plants over one year old have one to several simple, erect branches developed from ground level. The “Striata” type habit mainly occurs in the montane to alpine zones. For further discussion see p. 12. The habit typical of E. collina ssp. muelleri (fig. 68) is somewhat different. The sub- species has been recorded throughout lowland and montane regions. Its habit resembles the “Striata” type in the confinement of the vegetative shoots to near ground level. Although an occasional branch or young shoot may sometimes develop in the lower half of the main branches above ground level, in many plants the great majority of branches arise from a perennating base composed of many densely clustered young shoots. This compact perennating base is not known elsewhere in the genus and may be a response to the prolonged hot dry periods which are characteristic of Australian conditions. The many young shoots may be dormant and develop only after periods of rain. The “Humifusa” type (as in E. humifusa). This is a habit type probably derived from either the “Malesianae” or the “Striata” type of habit. The axes are procumbent and root at the nodes, although van Royen (1972) has stated that the inflorescence-bearing axes may be erect in E. callosa. The nature of first-year plants and hence the stem is not known. The habit type is confined to and typical of Sect. Pauciflorae Subsect. Humifusae of alpine and subalpine grassiauds (at least of New Guinea: van Royen 1972). 11 W.R. Barker J. Adelaide Bot. Gard. 5 (1982) The “Collina” type (as in E. collina ssp. collina), figs 45, 63, 67, 70, 72-74. In this habit type first year plants consist of a single erect stem, terminated by an inflorescence and developing branches above ground level in occasional axils with no fixed sequence. At the end of the first season the stem dies back to the upper branches and in the next season further branches develop from this region and below. There is never the perennation from the base of the plant that often occurs in the “Cuneata” type of habit. The “Collina” type of habit is restricted to the perennials of lowland and montane Australia. The position of the uppermost branches or shoots on the stem relative to its height often differs significantly between taxa having the “Collina” type of branching, as illustrated in fig. 2. For example, the position of branching in the forms of E. collina ssp. speciosa and ssp. paludosa which have the “Collina” type of habit tends to be much lower than in ssp. collina, Within E. collina ssp. tetragona, which has consistently the “Collina” type of habit, there is a geographically based transition in the position of branching, from branching in the upper half of the stem or main branches to branching restricted more to the lower half of the plant (see E. collina ssp. tetragona: note 2; fig. 69). The four habit types of the Australian perennials appear to be genetically determined, for they overlap in their climatic tolerances while generally retaining their integrity. There is no evidence of break-down of the localized “Cuneata” and “Phragmostoma” types, which are geographically sympatric with taxa of the “Collina” type. Many cases of geographical sympatry of taxa with the widespread “Striata” and “Collina” habit types occur in lowland and montane south-eastern Australia, often with no evidence of intergradation. Some taxa, e.g. both ssp. speciosa and ssp. paludosa of E. collina, may have either the “Striata” or “Collina” type of habit. The specimens with the “Collina” type occur only north of Sydney in northern New South Wales. In addition there are several cases of intergradation between taxa with a different habit type. Thus E. collina ssp. collina (of montane and lowland regions) with the “Collina” type of habit and ssp. diemenica (of alpine and subalpine regions) with the “Striata” type, intergrade in the upper limits of forest and woodland (see E. collina: Intra- specific Polymorphism). Similarly, the upper portion of an apparent intergradation between E. gibbsiae ssp. comberi of the alpine zone and of the “Striata” type of habit and ssp. kingii of wet lowland and montane moors with the “Collina” type, has been observed (see E. gibbsiae: Intraspecific Polymorphism). The confinement of the vegetative buds to ground level in the “Striata” type is probably a result of the extreme cold experienced in their natural habitats, and the fact that the ground is somewhat warmer and less variable in temperature than the air or snow above. Plants with this habit are capable of existing at lower altitudes; E. lasiantheraand E. collina ssp. diemenica, ssp. paludosa, ssp. muelleriand ssp. speciosaall occur in montane and lowland regions. In contrast, the “Collina” type of habit is absent from areas seasonally under snow. Vegetative shoots are developed above ground level with a resultant increase in height of the plants which may become even greater in successive years and assist the plant in the competition for light in the forests and dense shrubberies with which it is often associated. 4. Indumentum (excluding that on corolla, stamens and ovary which is discussed under each organ) The distribution, composition, length and density of the indumentum on the axes, mle: J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia leaves, rachis, bracts, pedicels and outer surface of the calyx are of major diagnostic importance at both the specific and infraspecific level throughout the genus. Hairs of both the glandular and eglandular types are common. The length of the different types of hairs varies from organ to organ on a plant and also upon the one organ. For example, the glandular indumentum on the rachis and the axis is generally longer than that on the calyx and leaves, respectively, and the indumentum on the calyx shortens towards the base. Therefore, to compare taxa which differ in indumentum length, the length on each particular organ has been described. Rather than make exact measurements of many specimens, the lengths have generally been described in words covering particular ranges as follows: very short (to 0.05mm), short (0.05-0.1mm), moderately long (0.11-0.29mm), Jong (0.3-c.0.45mm), very long (c.0.45mm upwards). Where the indumentum length has been particularly important in distinguishing taxa an exact statement of the range of variation has been given for one or two organs, as with the glandular hairs on the outer surface of the calyx or on the axis or leaves of the lower parts of the plant. The means by which density has been determined is less exact. If the hairs are almost touching or withing a few hairs’ breadth, they are described as dense. a. Glandular hairs The presence or absence of glandular hairs (more than 0.05mm long when on the leaves, bracts and calyces) over all or some of the axis, leaves, rachis, bracts and calyces is of special diagnostic importance in almost every instance, as it is generally found to correlate with other character differences. The terms non-glandular and glandular have been used to describe these character states. These terms do not encompass the corolla, stamens and ovary, as the indumenta of these vary independently (see relevant sections). The presence of tiny subsessile glandular hairs less than 0.05mm long is of limited diagnostic use. It is likely that these hairs in the Australian species are identical to the very short ones found sporadically in Sect. Euphrasia of the northern hemisphere. Yeo (1972) attributed to these no taxonomic importance. They differ from the sessile glands which are stalkless and occur in indented patches on the underside of leaves and bracts and on the calyx (p. 25). Subsessile glands are common in all Australian taxa at the base of the clefts between the leaf and calyx teeth and along the narrow grooves which are on the upper sides of the leaves and bracts. Morphologically these hairs are similar to the longer diagnostically important glandular hairs; they differ only in the length and the number of cells of the stalk and in E. collina ssp. paludosa and its allied variants a complete transition can be observed. In the Australian Alps this typically non-glandular taxon (q.v.: note 2) has a sparse to dense cover of subsessile glandular hairs. The presence of a number of populations with glandular and non-glandular plants and the lack of correlation with other varying characters have meant that recognition of the plants with subsessile glands as a distinct taxon above the level of forma is not justified. However, in view of the lack of evidence of intergradation, two subspecies of E. orthocheila have been separated on the presence or absence of subsessile to short glandular indumentum. b. Eglandular hairs y The eglandular hairs vary in rigidity and length. Length and density characters for weak eglandular hairs are described as outlined previously. However, in the case of plants which are covered on some or all parts by rigid eglandular hairs, another terminology, more descriptive of the nature and length of the indumentum, has been used. Scabrous has been used for such hairs over c. 0.1mm long (they rarely attain more 13 W.R. Barker J. Adelaide Bot. Gard. 5 (1982) than 0.2mm); these occur in E. scabra and to some extent in E. collina ssp. muelleri. Scaberulous describes a cover of shorter hairs or sharp excresences (c. 0.02-0.1mm long), such as are characteristic of E. ciliolata, E. ramulosa and some individuals of E. collina ssp. muelleri. In related taxa with non-glandular upper parts, the presence or absence of an eglandular indumentum on the external surface of the calyx and sometimes also on the bracts and upper leaves may be a character of diagnostic importance. The length and nature of the eglandular hairs may also be characteristic of taxa. Thus E. crassiuscula is remarkable, at least amongst the southern hemisphere species, in its indumentum which lines the calyces, bracts and leaves; it consists solely of long woolly eglandular hairs. The eglandular hairs on the axis and rachis are rarely of taxonomic value. The indumentum usually becomes much sparser lower down the axis except when dense glandular hairs are present on the lower parts. In Australia eglandular hairs may surround the whole axis, e.g. in E. scabra and E. orthocheila, but more commonly they are confined to two rows or two pairs of lines decurrent from between the leaf bases. In New Guinea EF. humifusa and E. callosa have been separated almost entirely on this character. c. Sessile glands (see p. 25). 5. Leaves and bracts Bract has been used for the leaf subtending the flowers of Euphrasia, following Wettstein (1896), Joergensen (1919: “Deckblatter”), Du Rietz (1932b; 1948a,b) and others. Others, including Pugsley (1930, 1936) and Sell & Yeo (1970) have used instead the term “floral leaf”. Blade has been used in this work in the special sense of that part of leaf or bract excluding the teeth and, if present, the attenuate part of the base (fig. 3). Fig. 3. Usage of the term blade in the leaves and bracts of Euphrasia. In an attempt to give a more precise statement of the shapes of the teeth of the calyces, bracts and leaves, the following method has been found useful (fig. 4). To describe the shape of the tooth overall, ie. from base to apex, the terms obtuse, acuminate, acute, caudate, etc. have been used as defined in Stearn (1966) and Jackson (1928). This seems more descriptive and precise than methods entailing terms such as triangular. The very apex of the tooth is then described by the terms blunt and sharp (which it is realized are the respective English equivalents of obtuse and acute). Although the use of these characters has been avoided in the keys because the terms describing them could be misinterpreted, they are nevertheless often diagnostic, , Leaf and bract characters are of major importance in distinguishing taxa at all levels in Euphrasia. The major characters involved are the overall size and shape, the shape of the base, the number, length and region of distribution of the teeth, the size and shape of 14 J. Adelaide Bot. Gard. 5 (1982) Apical tooth pes, i. lo Vaan 0, oy Fa Os x “, o h ! Fig. 4. Tooth apex terminology for the leaves, bracts and calyx of Euphrasia. bluntly broad-obtuse sharply broad-obtuse bluntly obtuse sharply obtuse bluntly acute sharply acute bluntly broad-acuminate sharply broad-acuminate bluntly acuminate sharply acuminate bluntly narrow-acuminate sharply narrow-acuminate bluntly caudate sharply caudate 15 Studies in Euphrasia Lateral tooth W. R. Barker J. Adelaide Bot. Gard. 5 (1982) the apical tooth, the distribution and pattern of the sessile gland patches on the lower surface, and the indumentum, which has been described in the previous section. The uppermost leaves and lowermost bracts are similar in all characters except sometimes those of indumentum. In the Australian species and probably elsewhere, there tends to be slight narrowing of the blade and broadening of the base (sometimes accompanied by an addition of an extra pair of teeth) from the lowermost bract down to two or three leaves below the inflorescence. Lower down the leaves gradually shorten and sometimes decrease in the number of teeth. Often correlated with these trends are transitions in the nature of the indumentum. From the lowest bracts to the last-formed ones there is a decrease in size and number of teeth. To make consistent comparisons between taxa, the characteristics of the bracts and leaves have been described on the basis of a detailed analysis of the uppermost pair of leaves on the main inflorescence-bearing axis of the plant. The indumentum characters of the bracts and lower leaves are also often described by a comparison with the detailed description of indumentum of the uppermost leaves. A series of figures (figs 5-12) showing the size and shape of uppermost leaves on the main inflorescence-bearing axes of almost all Australian taxa is supplied. Where possible a sample from a population is displayed. If representative collections of a taxon were few, only one or two leaves were removed for display. A number of very distinctive leaf and bract shapes are often of diagnostic importance at a sectional and subsectional level. They involve the size of the blade (defined above) relative to that of the whole leaf, and the shape of the base. Illustrations of extra-Australian types are found in Wettstein (1896: northern hemisphere including Japan and the Azores, New Zealand, South America), Du Rietz (1932b: Malesia), Ashwin (1961: New Zealand) and van Royen (1972: New Guinea). The most widespread, though only sporadic, type is the long attenuate leaf with a prominent blade; it characterizes the sections Cuneatae (fig. 5), Phragmostomae (figs 5, 30) and Paradoxae, some species of Sect. Pauciflorae (E. papuana), Sect. Malesianae and Sect. Euphrasia, and E. bella of Sect. Australes (fig. 6). A prominent blade with a short abruptly attenuate base is typical of Sect. Atl/anticae and the remaining majority of species of Sect. Euphrasia and Sect. Malesianae. Du Rietz’s (1948a) “subdigitate- toothed” and “digitate” leaf types are confined to the southern hemisphere. These have narrow attenuate leaves which comprise largely prominent teeth through the small size of the blade. Such types are characteristic of Sect. Trifidae, Sect. Anagospermae, mainly the New Zealand members of Sect. Pauciflorae and Sect. Striatae. Deeply trifid leaves are typical of the first two sections, but E. integrifolia (Sect. Anagospermae) has lost the lateral pair of teeth. Sect. Striatae (figs 5, 6) and the New Zealand members of Sect. Pauciflorae have a larger blade, and often more teeth. E. hookeri of Tasmania (figs 6, 31, 47) is remarkable for its many digitate teeth and (see below) their greatly recurved margins, which are the basis for retention as a monotypic series in Sect. Striatae. A curious leaf shape is typical of most New Guinea species (Sect. Pauciflorae); the leaf is dominated often for half its length by a hooded apex and the lateral teeth are all but lost. Finally, the only non-attenuated leaf bases are found in Australia and characterize the sections Australes, Lasiantherae and Scabrae (figs 6-12). The leaf is composed mainly of blade through the cuneate, rounded or truncate base. Narrowly cuneate types outside Australia are extremes of variation in species with attenuate leaf bases. The difference in the form of the cuneate to truncate leaf base among the Australian members of Euphrasia seems to be directly related to the nature of vascular system of the leaf at its very base. These non-attenuated types have 3-7 vascular strands arising from the base. Even taxa with narrower leaves, such as FE. collina ssp. collina, may sometimes have 5 vascular strands arising from the base. In other Australian species, i.e. 16 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia Sect. Cuneatae 1. E. ramulosa Grercs v sf Hore-Lacy G y Davis 0 Williams NSW126388 185 NSW22266 11.1963 Sect. Phragmostomae 2. E. phragmostoma Bufton 7 ? Bufton 8 Sect. Striatae 3a. E. gibbsiae ssp. gibbsiae Sve” wee AARAARALLA AA LL 3c. E. gibbsiae ssp. wellingtonensis AR ARR ALAR An Oe A AL Barker 1008 Barker 1123 3d. E. gibbsiae ssp. comberi AAR AALS AALS AALLAAAALA hpeeee 3f. E. gibbsiae ssp. microdonta 0 ] Phillips 87 3g. E. gibbsiae ssp. subglabrifolia Wr eleertvetey tH Barker 1469 . Barker 1470 Fig. 5. Uppermost leaves, actual size dried, of main inflorescence-bearing axes in taxa of Euphrasia in Australia (1).° 17 W.R. Barkers J. Adelaide Bot. Gard. 5 (1982) . (Sect. Striatae: £E. gibbsiae, cont.) | (+ > > — -- - -- ---—- -—--___ - ———. | 3h. E. gibbsiae ssp. discolor a | wey AMAL ALA ARAL 2 Barker 1218A 3i. E. gibbsiae ssp. pulvinestris RAALA ALLALS Ge ¥ ¥ ¥ v DORAASAAAALDA A of Barker ZO | Ss 4. E. striata ae | DOADDA AAA AD EEE OAM Add Aa arre WPT TeNHHV is "Gul 6. E. semipicta Wri bi ' | 7. E. hookeri i ' | | OLE A ARAALAL Le ~ Sect. Aus Australes 8. E. bella Smith | Blake % Blake | BRI 144796 15923 14652 | 10. E. bowdeniae | | ] ) Currie NSW126387 ] Bowden NSW84075 Q Coveny NSW98623 lla. E. collina ssp. collina cc) ae TE ULL LG) 9 es UUM 8 pinirinag ... (arent he Fig. 6. Uppermost leaves, actual size dried, of main inflorescence-bearing axes in taxa of Euphrasia in Australia (2). 18 jo os ME OLB Gothocororsearyeg Coyeveryy Vesertelg Eas Aeeteteunt dortertersevere de t __Eichler 15384 thet: pth igang)... rey W.R. Barker J. Adelaide Bot. Gard. 5 (1982) (Sect. Australes: E. collina, cont.) l1f. E. collina ssp. ‘tasmanica’ 11g. E. collina ssp. gunnii Netty Pec rsrvigy...... (hy Barker 913 11h. E. collina ssp. deflexifolia ¥ SVT FV fiservey ftregorerecge Fouteter gs 11i. E. collina ssp. paludosa Hes HHHA. AEA BAA AA ‘i 1565 Vevey] terse Barker 1505 Barker 1585 Corley fFitererey Bark: 03 arker 16 Barker 1680 q HESS FF. 1li’. E. collina aff. ssp. paludosa HOseesvevsy | Gete VEE Es Cee 11k. E. collina ssp. nandewarensis Johnson & Winterhalder Constable Rupp 22 NE NSW30524 Fig. 8. Uppermost leaves, actual size dried, of main inflorescence-bearing axes in taxa of Euphrasia in Australia (4). 20 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia (Sect. Australes: E. collina, cont.) 111. E. collina ssp. speciosa : & : ¥ Barker 1630 Helms NSW10901 Constable NSW48942 11m. E£. collina ssp. osbornii géhba 066 & & Barker 854 Barker 861 Barker 862 Barker 869 ry eR Ahaenae 7f | Barker 1346 $SGeAteeies HH! Barker 1729 AD9622007 io<«ss tis H Eichler 14452 Hunt 3315 Whibley 1278 Whibley 4155 I1n. E. collina ssp. diversicolor GG 2 OUveGe Ow 8: | VvESSVVOVS € -- OO 1009 Bes ...... V6 SOCHGeSeo4....... WHS CosGoogere ox: eeeHHsE ...... ME hd RL LA LR Fig. 9. Uppermost leaves, actual size dried, of main inflorescence-bearing axes in taxa of Euphrasia in Australia (5). 21 ———EE W. R. Barker J. Adelaide Bot. Gard. 5 (1982) (Sect. Australes: E. collina, cont.) , In’. E. collina aff. ssp. diversicolor Hleeessee us ‘vee llo. E. collina ssp. lapidosa Veovort wootey | seeresy OOH 6 Gray & Totterdell 6525 Gray & T. 6524 Wola Barker 1616 270 —————— llp. E. collina ssp. glacialis : UH MUVESGecres worees | Barker 1685 0900 00Ge8 ...... VHS edertHogo.... 12a. E. crassiuscula ssp. crassiuscula QQrvteses... 000009 000000008 Barker 1546 Fig. 10. Uppermost leaves, actual size dried, of main inflorescence-bearing axes in taxa of Euphrasia in Australia (6). 22 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia (Sect. Australes: E. crassiuscula, cont.) 12b. E. crassiuscula ssp. glandulifera &e 6 : ¥ Barker 1596 12c. E. crassiuscula ssp. eglandulosa GESG0800G08 |. Pooergoogy Sect. Lasiantherae Hassees ae Féiese vi Hlinttiod. 15. E. alsa LAE EY Barker 1696 Vey Ve cetite ¥IVTV SE Barker 1707 Sect. Lasiantherae x Sect. Scabrae Barker 1593 13. E. lasianthera he Barker 1498 ¥ 15 x 16. E. alsa x E. caudata Hiiisty.. Fig. 11. Uppermost leaves, actual size dried, of main inflorescence-bearing axes in taxa of Euphrasia in Australia (7). 23 W.R. Barker J. Adelaide Bot. Gard. 5 (1982) | ___ Sect. Scabrae | 16. E. caudata LAE. a SHAH. 17. E. scabra Mueller Ashby 4354 MEL41609 18a. E. orthocheila ssp. orthocheila i Beckler MEL41595 Crawford 19 Rina Collie MEL41643 18b. E. orthocheila ssp. peraspera { ay | Heron F Wilcox NSW10880 WALLS SUISLEAAVARS . MEL41589 | 19. E. arguta Woolls Crawford Musson MEL41394 MEL41756 MEL41397 20. E. ciliolata bé Bag b S82 ‘4 } } } ae Fig. 12. Uppermost leaves, actual size dried, of main inflorescence-bearing axes in taxa of Euphrasia in Australia (8). 24 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia E. bella of Sect. Australes and commonly Sect. Striatae, there are only three main vascular strands arising from the base. In leaves with very shortly attenuate leaf bases abruptly expanded into the blade, the lateral pair of main strands may be branched a short distance from the base. In the long attenuate types the three strands may remain simple for their whole length or branch towards the base. The adaptive significance to parasitism of the patches of sessile glands on the lower surface of the leaves and bracts has been alluded to on p. 5. The distribution and pattern of the patches are novel characters which have been found to be of diagnostic importance in some instances in Australia. An extended, more critical study elsewhere in the genus may show further examples of differences between taxa. The sessile glands (not be be confused with the very shortly stalked subsessile glandular hairs: p. 13), occur on the lower surface in dense patches always in rows parallel to and just inside the margins; these rows are branched sporadically on their inner margin, the lateral extensions being short or very long, and are generally confined to the areas between the main veins of the leaf. In E. hookeri (Sect. Striatae) the sessile gland patches on the lower leaf surface are hidden from view as a result of the extreme recurvature of the teeth margins (fig. 31K). Whether the sessile glands occur towards the base of the leaf is related to the distribution of the teeth along the leaf. Except when the leaf teeth are very reduced (as they often are in the New Guinea species), sessile glands are always distributed along the margins from the apical tooth at least as far as the extent of the laterally toothed parts of the leaf. Thus if the leaf teeth reach the base of the leaf, as in the Australian cuneate to truncate-based leaves, so do the sessile glands. However, if the teeth are confined to the distal half of the leaf (e.g. in E. collina spp. collina), the sessile glands may extend along the margins well below the teeth and onto the lower half, but never to the base. In species with attenuate leaf bases the sessile gland patches hardly (if at all) extend onto the attenuated region. The pattern of the sessile glands on the lower surface has been used in E. collina to distinguish ssp. diemenica, the sessile glands of which are almost entirely confined to the marginal rows except for very short lateral branches (fig. 48E), from other subspecies which have long lateral extensions of the marginal rows of glands (fig. 48D). Ssp. collina shows a transition in these same differences in pattern of the sessile glands. This appears correlated with a geographical cline in leaf shape, discussed on p. 179. Its south-eastern Tasmanian populations have a pattern of sessile gland patches and short leaf teeth similar to ssp. diemenica; the populations in the Grampians, however, approach ssp. paludosa in these characters. In Australia the sessile glands in the annuals and some densely glandular hairy perennials are apparently distributed over wider areas of the lower leaf surface than in the non-glandular hairy perennials. Outside Australia the pattern of sessile glands on the leaves of Euphrasia is a character of potential taxonomic value. E. cuneata (Sect. Cuneatae), Sect. Phrag- mostomae and Sect. Paradoxae have very distinctive, reticulately patterned sessile gland patches. Yeo (1973) has indicated that Sect. Atlanticae is unique in the genus by the confinement of the sessile glands to the veins of the lower leaf surface as in related genera in Trib. Rhinantheae (p. 79). 6. Inflorescence The arrangement and number of flowers is of importance in the infrageneric classification of Euphrasia. Throughout the genus the flowers develop acropetally along the axes. In most of the genus the flowers are arranged decussately in racemes. Both a BD W. R. Barker J. Adelaide Bot. Gard. 5 (1982) flowers at one node usually develop simultaneously, but in certain conditions, such as where plants grow from the side of a dense shrub, development may be much earlier or possibly confined to the outward-facing side. Whenever plants of Euphrasia are completely prostrate (Sect. Pauciflorae Subsect. Humifusae, partly Sect. Anagospermae), flowers are distributed sporadically along the axes with usually one flower only at a node. However, the erect to ascending axes of Sect. Malesianae are terminated by racemes, which are sometimes interrupted by the absence of flowers from some nodes as well as sometimes having only one flower at a node. In perennials transformation on the one axis from inflorescence to leaf-bearing and again to inflorescence occurs rarely in E. gibbsiae ssp. kingii (Sect. Striatae) and E. phragmostoma (Sect. Phragmostomae). Yeo (1964) has also seen this phenomenon in the annual EF. pseudokerneri of Sect. Euphrasia, and has linked it with abnormal temperature fluctuations. In E. formosissima (Sect. Paradoxae), however, this apparently occurs regularly (Skottsberg 1921) as the means of perennation (p. 10). The number of flowers has been used to separate Sect. Pauciflorae from Sect. Striatae and in Australia to distinguish species and subspecies. In the Australian annuals there is a correlation between the number of flowers in the main inflorescence and the climatic range of the species in parallel with other character trends (table 1). The number of flowers produced may be linked to some extent with the relative robustness of the species and hence the capacity to maintain the production of flowers. This is possibly applicable throughout the genus. Pedicel length shortens acropetally in the inflorescence. Although not of major diagnostic importance in the genus in Australia, pedicel length is useful in separating E. bella and E. bowdeniae of Sect. Australes, and some New Zealand species. Except in plants with very few flowers, the buds and bracts of the young inflorescence are generally crowded together at the tip of the axis in what has been termed in this work the apical bud cluster. In most of the genus this is broadly ellipsoid or ellipsoid- ovoid to spherical in shape. However, in the Australian species there is a wide variation in the number of flowers produced and there is some variation in the shape and degree of exsertion of the apical bud clusters from the uppermost flowers. Two types of bud cluster are discernible which have been given diagnostic importance in the past by Wettstein (1896), Curtis (1967) and Burbidge (Burbidge & Gray 1970); whether the differences have any genetic basis, however, is open to question. In the alpine species the apical bud clusters are short and broad and soon after flowering commences become hidden in the uppermost flowers of the inflorescence (e.g. figs 44, 66, 80) This inflorescence type has been termed “capituliformis” (Wettstein 1896; Du Rietz 1948a) or ‘capitata” (Wettstein 1896; Curtis 1967). In the lowland and subalpine members of E. collina the apical bud cluster in narrowly cylindrical or narrowly conical and remains extended above the upper corollas after the flowers at many nodes (sometimes over 20) have matured (e.g. figs 71, 73, 74). This inflorescence type has been termed “elongata” (Wettstein 1896), “spiciformis” (Du Rietz 1948b), and “conica[l]” (Wettstein 1896; Curtis 1967). In the past the character has been used particularly in separating the taxa now grouped under E£. collina. It is considered that the variation may be an environmental response, particularly as the two types have never been observed to grow together. In addition, two subspecies have both character states. Ssp. paludosa is normally characterized by a narrowly subconical apical bud cluster, but in the highest parts of its range (e.g. on Mt Speculation, Victoria) it may have apical bud clusters of the other type. Ssp. diemenica typically has the short, broad type of cluster, soon hidden by the upper flowers. However, montane and even subalpine occurrences may have conical apical bud clusters. Because of this uncertainty about the genetic basis the variation, the initial shape and subsequent development of the apical bud cluster have been described in detail, but have not been used in the keys. 26 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia 7. Calyx The structure of the calyx is essentially the same throughout the genus. There is variation in size and the depth of the lateral and median clefts (the divisions between the teeth); these are of some diagnostic importance at the specific and infraspecific level. The shape of the apex of the teeth and the degree of recurvature of the margins tends to be associated with analogous characters in the uppermost leaves and bracts, although the recurvature in the calyx is not as marked as in the leaves. A special terminology, devised to described the shape of the teeth, is discussed on p. 14. The indumentum on the outer surface of the calyx is of major diagnostic importance. Its nature is closely related to that of the bracts, rachis, leaves and axis and is discussed under the general treatment of indumentum. 8. Corolla The shape, size, colour and its distribution and indumentum are of diagnostic importance at various levels of classification. a. Shape and size The shape of the corolla is useful in separating only more distantly related sections and subsections for generally closely related infrageneric taxa are connected by a series of gradually changing forms of corolla. The type of corolla characteristic of Sect. Euphrasia, as illustrated in Wettstein (1896, pl. 2), is the basic type which occurs throughout much of the range of distribution of Euphrasia (figs 30, 31, 37, 42-44, 47). Corollas of this type are two-lipped, with the tube directed obliquely away from the axis; the tube is cylindrical in the basal parts and distally expanded laterally and ventrally. The upper lip is hooded and more or less porrect with the lobes recurved sharply so that they are directed upwards and approximately lie in the same plane (i.e. they face forward). The lower lip spreads from its base away from the base of the upper lip; it is concave from above at its base. In the southern hemisphere there are a number of corolla types divergent from this basic form. In Australia the corollas of Sect. Scabrae (figs 91, 95) are the most divergent. They are characterized by a porrect flat lower lip which lies against or hardly spreads from the upper lip (except in E. caudata in distal regions). The lower side is broadly grooved and flat or slightly convex from above. Except in E. caudata, the upper lobes are sharply reflexed, but are at a sharp angle to each other because they are more or less appressed against the side of the hood. The corollas of Sect. Lasiantherae (figs 17, 85, 87, 90), E. collina (figs 63, 66, 70, 71, 74, 77-80) and possibly E. bowdeniae and E. bella of Sect. Australes and E. caudata are intermediate between those found inthe rest of Sect. Scabrae and the basic type. In these the upper corolla lobes are directed forward and lie inthe same plane and the lower lip, although initially porrect, is spreading in the distal regions. The lower side is flat and sometimes broadly grooved. Sometimes, however, the lower lip and tube may be concave from above. Study of living material is required to verify these differences. In the New Guinea perennials (Sect. Pauciflorae) and the New Zealand annuals of Sect. Anagospermae (fig. 18) the corolla tube is sometimes very long relative to the two lips. However, except in E. disperma of the latter section and possibly E. scutellarioides of Sect. Pauciflorae, plants with a relative tube length more typical of the genus occur in each of the species. The corolla ofa number of New Zealand species shows some tendency toa more regular arrangement and equality in size of the lobes with the corolla rim facing upwards; this may relate to the low stature of plants and a likelihood of pollinators detecting the flowers from above, as is proposed for the Australian Mimulinae (Barker, in press). 27 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) Finally, Sect. Trifidae of South America diverges from the rest of the genus by the production of some remarkable corolla types. Some species, e.g. E. antarctica (Vallentin & Cotton 1921, pl. 45), differ very little from the basic corolla shape, although, by the arrangement and equal size of the upper and lower lobes, the corolla appears almost regular when viewed from the front (Wettstein 1896, pl. 6 f. 472,473). However, in E. chrysantha and E. flavicans the lower corolla lip is abruptly reflexed so that the corolla lobes are in the same plane, and E. meiantha (Wettstein 1896, pl. 6 f. 482) has a very long porrect hood and a minute lower lip reflexed from its base. Corollas in this section show the greatest divergence from the basic type and the greatest diversity. A character sometimes important at all levels of classification is the shape of the corolla lobes. Sect. Euphrasia, Sect. Atlanticae (ex Wettstein 1896), Sect. Malesianae and Sect. Pauciflorae Subsect. Humifusae of the northern hemisphere and tropics always have emarginate lobes. Throughout the remainder of the genus in the southern hemisphere, including Sect. Pauciflorae Subsect. Pauciflorae in New Guinea, the lobes vary from obtuse to emarginate, sometimes even within populations; occasionally, however, a species or subspecies may be characterized by consistently emarginate lobes. In the Australian revision differences in shape and size of the corolla and its parts have been measured using a range of diagnostic parameters (fig. 13): the length along the upper side (i.e. excluding the lobes), the length of the tube from the base of the corolla tothe point of insertion of the anterior filaments, the length of the hood (upper lip), the breadth of the hood (including and excluding the lobes), the length and breadth of the . Length of upper side . Width of hood (excluding lobes) ; . Length of tube . Depth of cleft between . Length from base of corolla upper lobes to point of insertion of anterior filaments . Length of lower lip . Length of hood i. Width of lower lip . Width of hood (including j. Depth of cleft between lobes) lower lobes Fig. 13. Definition of the corolla parts of Euphrasia measured. 28 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia lower lip and the depths of the clefts between the upper and lower lobes. All are of diagnostic value. Measurements were taken from the lowermost flowers of the main inflorescence, which had been fixed in the field in FAA and transferred to 70% ethanol, or taken from herbarium specimens and boiled in a weak detergent solution, or well- pressed and untreated. There seems to be little difference in corolla size in pressed as compared with fixed or boiled material. b. Ground colour The “ground colour” of the corolla (i.e. excluding striations and yellow nectar guides) is of limited diagnostic value, at least in Australia. It is useful, however, for separating the yellow-flowered species of Sect. Scabrae, Sect. Novaezeelandiae and apparently Sect. Trifidae from related species with corollas with a ground colour of white to purple or violet. In Sect. Euphrasia the presence of a yellow corolla is apparently not diagnostic at the species level (¢.g. E. minima Jacq. ex DC.: Pugsley 1930, Yeo 1972). Ground colour in the Australian species, when not yellow, may vary from white to maroon, pink, violet or blue. A particular taxon may vary through all or part of this range, and may show different ranges of variation in different geographical regions. Elsewhere in the genus the ground colour is usually white, although sometimes lilac or purple corollas may occur. This is sometimes diagnostic at the species level in Sect. Euphrasia (Yeo 1972). c. Coloration Corolla coloration in Euphrasia has received little attention in past classifications in the genus. General reference to a basic type of coloration involving purple to violet- striated corollas with yellow blotches in the tube and at the base of the lower lip (e.g. fig. 31) have been made by Ashwin (1961), Yeo (1968) and van Royen (1972) for New Zealand species, the northern hemisphere annuals and New Guinea species, respectively. From my field observations of taxa in Australia, New Zealand and, to a limited extent, New Guinea, and from colour photographs, specimen annotations and well-preserved herbarium material of taxa elsewhere, it is clear that there is wide variation in the pattern and extent of both blotches and striations on the corolla, which is of potential diagnostic importance at a specific and infraspecific level. In the following, reference to published illustrations is given, apart from Sect. Euphrasia, of which there are many sources, and usually Salmon (1967) and Mark & Adams (1973), which give a large cover of the New Zealand species. A photograph or painting is rarely sufficient to give all details of corolla coloration. For example, that of the rear of the lobes is rarely shown. It is vital that collectors record such characters in as much detail as practicable. Geographically, the most widespread expression of the corolla striations is that usualin the northern hemisphere annuals, Sect. Lasiantherae and most species of Sect. Striatae of Australia (figs 17, 31, 37, 42, 44, 47, 85, 89, 90), Sect. Cuneatae of Australasia, in Sect. Paradoxae (Skottsberg 1921, pl.20), E. cheesemanii of Sect. Novaezeelandiae, and E. borneensis, the sole known record for Sect. Malesianae (Stapf 1894). This consists of 3(5) purple or violet striations on each lobe, those of the upper lip extending from the hood; they are most prominent on the outer face of the upper lobes and the upper face of the lower lip. In Sect. Striatae of Australia the limited extension of the striations onto the lower lip in E. gibbsiae ssp. pulvinestris (fig. 43) and E. semipicta is diagnostic. Some specimens of E. semipicta may completely lack striations. Complete absence of striations is sometimes a sectional characteristic, as in Sect. Phragmostomae, Sect. Australes and Sect. Scabrae of Australia (figs 63, 66, 70, 71, 74, 77-80, 95). In the New Guinea species (Sect. Pauciflorae) the fine purple striations on 29 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) the hood are replaced by a suffusion of purple throughout (personal observations in ¢ mirabilis, E. humifusa and E. callosa). The corolla of E. papuana also lacks prominent striations (Mr P. Kores, pers. comm. 1976). In New Zealand, the three sections other than Sect. Cuneatae show divergence from the basic form of striations. E. disperma (Sect. Anagospermae) and E. cheesemanjj (Sect. Novaezeelandiae) show the basic type. However, in many species the striations, jf present at all, are prominent on the hood, rear surface of the upper lobes and the rear oy lower surface of the lower lip, as strikingly portrayed by the 3 prominent striations on each corolla lobe in E. townsonii (pers. observ.). A number of New Zealand annuals, e. 9. E. zelandica, E. dyeri, and possibly E. repens, often have a broad purple streak along the midline of the outside of the hood. E. integrifolia rarely has a finer streak along the edge of the hood on either side of the central streak (pers. observ.). In this species the lower lobes also have a single purple to indigo streak, on the rear or lower side only, Sect. Trifidae of South America may also diverge from the basic form of corolla striation. E. antarctica has a single purple striation down the midline of the upper side of each of the lower lobes (Vallentin & Cotton 1921, pl. 45; Skottsberg 1913). The other feature of the basic coloration type is the yellow blotches, which occur at the base of the lowest lip and in the tube at the base of each anterior filament. Such coloration occurs in all cases known to the author in the northern hemisphere annuals (Yeo 1968, 1972; Sell & Yeo 1970), in the New Zealand and New Guinea species, in Sect. Paradoxae of Juan Fernandez Islands (Skottsberg 1921, pl. 20), at least E. antarctica of Sect. Trifidae of South America (Vallentin & Cotton 1921, pl. 45), and Sect. Phrag- mostomae, Sect. Cuneatae and Sect. Striatae (e.g. figs 31, 37) of Australia. There is variation in the size and shape of the blotch which may prove to be of some diagnostic importance. In particular, in some New Zealand perennials of Sect. Pauciflorae, e.g. E. revoluta and E. townsonii, the yellow blotches are continuous and extend well back towards the base of the tube. In E. dyeri, E. integrifolia and E. disperma of Sect. Anagospermae the tube is yellow throughout. In E. disperma occurs an apparently unique character of a yellow patch on the upper lobes bordering the dividing sinus. Ag remarkable is the ring of deep brown-purple at the base of the corolla tube of an undescribed taxon allied to E. townsoniiin the Tasman Mts, New Zealand (pers. observ.), The species of Sect. Euphrasia Subsect. Alpicolae of Japan have the normal distribution of yellow blotches, but have a distinctive and possibly unique deep purple blotch on either side of the hood just above its base and also at the base of each anterior filament (Pugsley 1936; Yamakei Color Guide 1967, p. 40; Takeda 1959, pl. 65, n. 35, 36). Subsect. Japonicae has the normal yellow blotch on the lower lip (Takeda 1959, pl. 65, n. 34). In Sect. Lasiantherae, Sect. Australes and Sect. Scabrae of Australia, most taxa have the basic yellow markings. In the first section, £. alsa conforms in this way but E. lasianthera (fig. 85) lacks the yellow blotch behind the lower lip, but the small yellow blotches, one at the base of each anterior filament and distinctly demarcated and highlighted by the radiating striations, occupy a central position in the open-mouthed flower. The geographically and morphologically intermediate E. eichleri may be polymorphic in these features (p. 256). In Sect. Australes, E. collina is variable in the presence or absence of yellow blotches. The alpine subspecies, ssp. diversicolor (figs 77, 78), ssp. lapidosa, and ssp. glacialis (fig. 80), and the lowland ssp. tetragona (fig. 70) consistently have a yellow blotch on the lower lip and two others deep in the throat (sometimes the blotches are continuous). The yellow blotches are apparently completely lacking from ssp. deflexifolia and possibly ssp. speciosa. In a number of other sub- species the incidence of a yellow blotch behind the lower lip is varied. It is usually present in ssp. collina. Presence and absence vary on a geographical basis in ssp. 30 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia osbornii (p. 221) and on an ecotypic basis in ssp. diemenica (p. 185). In Sect. Scabrae the nectar guide on the lower lip of at least E. caudata (figs 91, 95) and possibly E. ciliolata consists of a yellow to red streak down the middle of the lower lip, a character possibly unique in the genus. d. Corolla indumentum The distribution, length and density of the indumentum on the inner and outer surfaces of the corolla, and the composition of the indumentum on different parts of the corolla, in relation to whether or not both glandular and eglandular hairs are present, are variable and furnish characters of diagnostic importance at the level of species and below. The density and length of the glandular and eglandular hairs are described in the terms used for indumentum on the calyx, bracts, rachises, leaves and axes (see p. 13). Consistencies seen in the corolla indumentum of the Australian species are: the dense eglandular indumentum on the outer surface of the hood extending to a greater or lesser extent onto the tube; the small or large patch of glandular hairs on the outer surface extending from the region about the lateral clefts towards the point of insertion of the anterior filaments; the dense patch of long eglandular hairs on the inner surface of the hood at the base of the sinus (or cleft) between the two upper lobes; and the glabrous nature of both the inner and outer surfaces of the very base of the tube. The above regions of the corolla may vary in the incidence of other hair types not cited. In all other regions the indumentum is variable in its incidence and composition. Of particular diagnostic importance are: the presence or absence of an indumentum on the external surface of the lobes (for the lower lobes best seen in mature buds); the presence or absence of an indumentum on the inner surface of the tube; the presence or absence of hairs lining the margins of the lobes; and the presence or absence of hairs at the base of the lower lip. The indumentum of the corollas of the extra-Australian members of the genus has not been investigated. Sell & Yeo (1970) mention that the inner surface of the throat is “papillose-hairy”, presumably on the basis of their combined experience of Sect. Euphrasia of North America and Europe. 9. Androecium In most species of Euphrasia the arrangement of the stamens is very constant. The stamens are didynamous. The anthers are fused along their narrowest sides in a U- shaped configuration, with the two posterior anthers quite free from each other, but connected with the two anterior anthers, which are themselves fused. The cluster of anthers is enclosed in the hooded upper lip and the anther awns project from the base of each cell into the mouth of the corolla. The rearmost pair of awns projects further into the mouth than the other awns as it is situated closer to the lower lip (being nearest the join of the two lips) and is longer than the other awns. Sect. Trifidae and E. disperma (Sect. Anagospermae) are the only exceptions to this. In both, the anthers are free. In E. disperma (fig. 18) and some species of Sect. Trifidae the anthers occur at different - levels and still lie against or under the hood. However, in other members of Sect. Trifidae the anthers are at the same level and are exserted from the corolla. The length and indumentum of both the anterior and the posterior filaments, the length, breadth and indumentum of the anthers (excluding the awns), and the length and shape of the rearmost pair of anther awns have been described for all species (fig. 14). All except the indumentum of the filaments have been found to be of diagnostic importance, some more so than others. Measurements of the stamens have been taken from the lowermost flowers of the main inflorescence. There is some decrease in size of the filaments and anthers in 31 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) successively formed flowers, but even when the anthers are much smaller than those in the lowermost flowers, the rearmost pair of anther awns apparently remains the same length (see p. 170). Fig. 14. Definition of the anther parts of Euphrasia measured. (a = anther length; b = anther breadth; c = awn length.) In the species with fused included anthers, the length of the two sets of filaments seems directly related to the length of the hood. These characters usefully distinguish related species in several cases. The size of the anthers is also diagnostic at the specific and infraspecific level. The length of the anther is measured from the apex to the point where the awns arise (fig. 14), which is taken to be the point on the outward-facing surface of the anther where the colour abruptly changes from the light brown, orange or deep purple of the main body of the anther to the white or yellow of the awn (this is slightly higher than the actual point where the attenuation begins). The anther awns of Euphrasia are with one exception alike in morphology. They are usually very narrow and sharp (figs 31, 48, 85, 91). However, the posterior awns of E. phragmostoma are remarkable, not only in their extreme length but also in the broadened and sometimes twisted or erosulate tip (fig. 30). The length of the rearmost pair of anther awns has been found to be of diagnostic importance at the specific and infraspecific level throughout the genus in Australia. The length appears in some cases to be independent of the size of the corolla, measured, for example, by the length of the upper and lower lips. Thus, in similarly sized corollas in ssp. paludosa, ssp. collina and ssp. diemenica of E. collina, the length of the rearmost awns in the first is significantly shorter than in the other two subspecies. In other cases, however, such as in Sect. Lasiantherae, corolla size is correlated with awn length. As pointed out by Du Rietz (1932b, 1948b), Bentham (1846) was incorrect in describing the anther awns of all southern members of the genus, except E. cuneata, as being equal. Du Rietz (1932b, p. 530), however, considered that “subequally mucronate” anthers were found in apparently all the South American species (Sect. Trifidae), all species now belonging to Sect. Anagospermae, E. monroi and E. papuana now in Sect. Pauciflorae, E. striata now in Sect. Striatae, and E. merrillii now in Sect. Malesianae. In no case, however, have I seen any taxon which consistently has equal anther awns. In some taxa the difference in length may be only slight, but it nevertheless exists. The possible loss of function of the longer rearmost anther awns in the taxa with free anthers, E. disperma and Sect. Trifidae is discussed on p. 46. 32 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia The indumentum of the anthers is of major diagnostic importance. Throughout most of the genus the anthers are completely glabrous but for dense hairs which line the slits of the inner surface (e.g. fig. 31). However, Sect. Australes, Sect. Lasiantherae (fig. 85) and Sect. Scabrae (fig. 91), which are confined to Australia, have a distinctive anther indumentum. Not only are the dehiscence slits lined by hairs which are probably longer and denser than anywhere else in the genus, but the anthers are almost always hairy on the outer surface. This indumentum on the outer surface is composed of straight or flexuose eglandular hairs identical to those occurring along the slits. It is usually very dense and long, but sometimes sparse and short, as occasionally in E. collina ssp. glacialis or on the anterior pair of anthers of the annuals. On rare occasions in some taxa, for example E. scabra and E. collina ssp. lapidosa and ssp. diemenica, the anthers may be completely glabrous on their outer surface. In some New Zealand sections and those of the Juan Fernandez Islands and South America, there is a trend towards the complete loss of hairs from the anthers. Sect. Trifidae and E. disperma (Sect. Anagospermae) have completely glabrous anthers. Although no anatomical studies have been made, it seems certain that the loss of fusion of the anthers is linked with the loss of the hairs lining the uppermost parts of the dehiscence slits (see Hartl. 1972). Sect. Paradoxae, Sect. Novaezeelandiae and the other species of Sect. Anagospermae link that part of the genus characterized by hairy anther slits with Sect. Trifidae with its completely glabrous anthers. E. formosissima (Sect. Paradoxae) has fused anthers which bear a few hairs within the slits. The anthers of Sect. Novae- zeelandiae are also fused, but show a transition from the type with densely hairy slits (E. cockayniana, E. cheesemanii) to very sparsely hairy slits (E. australis, E. zelandica) as seen in Sect. Anagospermae apart from E. disperma. This may relate to their reliance on a wide array of pollinators (p. 44). 10. Gynoecium The few characters of the gynoecium which show variation in Euphrasia are the indumentum of the ovary, the ovule number, and the shape and size of the stigma. Although the shape of the capsule varies considerably, the shape of the ovary seems to show little variation. The indumentum of the ovary is identical to that on the capsule and is discussed in the treatment of the latter. The size of the stigma is of great importance at the sectional level as it is the one character which divides the northern members of the genus as a whole from all southern members. In the tropics it separates Sect. Malesianae from Sect. Pauciflorae. Care should be taken in measurements from flowers in which fertilization has taken place as the stigma soon shrivels. Although sometimes appearing capitate, the stigma is probably always bilobed with the lobing often obscure through its shortness or reduction of the upper lobe. The number of ovules has been found to be of diagnostic importance at the level of species and below. In the Australian annuals there is an increase in ovule number as the climatic preferences of taxa change from alpine through subalpine to montane and lowland (table 1). The significance of this is discussed under the treatment of seeds. The number of seeds rather than ovule number appears to have been used as a diagnostic character by previous workers in the genus (Wettstein 1896; Pugsley 1930, 1936). Du Rietz (1932b) found that E. borneensis differs from E. philippinensis and E. merrillii by its fewer ovules and seeds, but in discussing the relationships of the Philippines species with those of Taiwan, Japan and Norway, he compared seed numbers. This is probably because Takeda (1910), Nakai (1913) or Joergensen (1919) did not consider ovule number. The use of seed number as a diagnostic character is 33 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) questionable as the variation in it is not only determined by the number of ovules. Within the Australian species there is great variation in the proportion of ovules which develop into mature seed. The number of ovules seems to be of particular diagnostic importance in the New Zealand annuals (Ashwin 1961: Sect. Novaezeelandiae and Sect. Anagospermae). There is apparently a progressive reduction in ovule number in these species as the habit becomes more procumbent and the corolla lengthens, with E. disperma with its single ovule per cell (see Simpson 1977), prostrate habit and extremely long corolla tube forming a possible endpoint of an evolutionary sequence (see p. 55). Counts of ovule number are too few, however, to be certain of this trend (Ashwin 1961). Froma few samples of ovule numberin each section it seems that the Australian species may have the highest numbers inthe genus. 11. Capsules The indumentum, size and shape of the capsule in lateral view are of diagnostic importance, usually at the level of species or below, but sometimes at the level of section or subsection. Measurements have been made on the capsules at the lowest nodes of the main infruct- escence. Capsule size is often greatly reduced higher up the rachis. The last capsules formed are usually well below the last-formed flowers. Apart from the remarkable bicornute capsules which occur in single species of Sect. Atlanticae (Pugsley 1936; Yeo 1973) and Sect. Anagospermae*, the capsule shape varies within rather narrow limits. The shape of the capsule is a product of the shape in lateral view, the median view apparently being consistently ovate toellipticand usually caudate or acuminate. Capsule shape has been described excluding the base of the style which persists on the capsule summit. Although there is not a great diversity of capsule shapes in the genus, apparently no section or subsection encompasses the whole range of variation. Broadly obovate or obcordate capsules occur throughout Sect. Pauciflorae Subsect. Humifusae, Sect. Novae- zeelandiae and apparently Sect. Trifidae. The species of Sect. Anagospermae without the bicornute capsule are probably also of this type, although little fruiting material is available. Sect. Cuneatae, Sect. Malesianae and Sect. Pauciflorae (especially the New Guinea species) have some species with capsules of this type, and others (e.g. E. merrillii and EF. lamii) which are oblong with obtuse to emarginate apices. Australian members of the genus and Sect. Paradoxae of the Juan Fernandez Islands have oblong to ovate- elliptic capsules. In the latter the apex is obtuse to shallowly emarginate. However, the former shows a wide variation, some Australian taxa having oblong capsules with shallowly emarginate apices (fig. 31G), others narrowly ovate to ovate-caudate capsules with obtuse to acute apices (figs 48, 85). Most, however, fall between these extremes. Although capsules of only a few species in Sect. Euphrasia have been seen, it appears that the broadly obovate type of capsule is absent. Du Rietz’s (1932b, 1948b) notion of an “acuminate” type of capsule, which he considered to be restricted to and consistent throughout the Australian and South American species, is a misconception. With the constancy of the shape in median view, he could only have been referring to the lateral view. He apparently saw little or no fruiting material of the several Australian species with capsules broadly obtuse or emarginate in lateral view. The distribution and density of the capsule indumentum is a useful character in Australia for the separation of species and infraspecific taxa. In Sect. Euphrasia it is one of the characters used to separate Subsect. Angustifoliae from Subsect. Ciliatae (Pugsley 1930, 1936; Sell & Yeo 1970; Yeo 1972). * E. disperma (Sect. Anagospermae) is usually single-lobed through the development of only one ovule (Ashwin 1961; Simpson 1977). 34 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia 12. Seeds The nature of seed in relation to ability of Euphrasia species to disperse over long distance is discussed on p. 59. Seed number and size are very useful diagnostically at the level of species and below in the Australian taxa. Seed number is less reliable diagnostically than the related character of ovule number (p. 33.) Sell & Yeo (1970) state that (in Sect. Euphrasia) “seed size tends to be proportional to capsule size and inversely proportional to the number of seeds in the capsule”. This does not hold in the Australian species. In the Australian annuals there is a clear correlation between the number of ovules, seed size and climatic range of each taxon (table 1). The fewest ovules and largest seed are found in alpine areas, while the highest ovule numbers and smallest seeds occur in the three lowland and montane species. A similar relationship is found in the Australian perennials. In Sect. Striatae, E. striata of alpine and subalpine areas has larger seeds and fewer ovules and seeds than its close relative E. semipicta of lowland heath. In Sect. Australes seed in the alpine subspecies ssp. diversicolor and ssp. glacialis of E. collina similarly tends to be larger than that of the subspecies of montane and lowland regions. Ssp. lapidosa which inhabits fjaeldmark of the highest alpine areas is one exception; its seed size is somewhat intermediate tending more to that of the lowland subspecies than its alpine herbfield allies. Within ssp. paludosa (q.v.: note 2) there is an increase in seed size with increasing altitude. Although sampling of ovule and seed number is inadequate, the ovule and seed number in the three above subspecies of E. collina in the alpine region of the Snowy Mountains of New South Wales appears to be lower than that for lowland subspecies, such as ssp. tetragona, ssp. osbornii, ssp. collina and ssp. paludosa. The several independent parallel trends of decreased ovule and seed number and increased seed size associated with the transition from lowland/ montane to subalpine/ alpine conditions in the Australian species may relate to several factors: the energy available to the plant for seed production (there is usually a parallel decrease in plant size and presumably photosynthetic area); the seed number required to maintain a viable population; and the nutrient reserves required in seed for germination and seedling development prior to attachment to a host in the different climatic conditions. The reversal in the trend, at least in seed size, in the fjaeldmark perennial may be associated with lower water-holding capacity and nutrient availability of this community compared with the adjacent herbfield (Costin et al. 1969) and their effect on optimum habit and organ size. E. disperma of Sect. Anagospermae, with its single ovule per locule and its usual development of only one locule with a large seed, shows some tendency to vivipary in the laboratory (Simpson 1977), for seed remains green up to germination and sometimes enlarges and germinates within the split capsule while it is still attached; other seeds germinate after release. Owing to an abundance of flowers, each plant produces many seeds. Simpson considers that the persistent moisture of the wet, poorly drained habitat in a mild humid climate provides ideal conditions for successful germination. Vivipary in this apparent annual would require the absence of a seasonal break between generations. 13. Chromosome number With the exception of two counts, one diploid (n = 11) for E. mirabilis of New Guinea (Borgmann 1964: “E. rectiflora”) and the other octoploid (n = 44) for E. antarctica from the Falkland Islands (D.M. Moore, fide Yeo 1968), studies of chromosome numbers in Euphrasia have been confined to Subsect. Ciliatae and 35 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) Subsect. Angustifoliae of Sect. Euphrasia in the northern hemisphere (e.g. Yeo 1954, 1970), both of which have diploid and tetraploid species. The current knowledge of chomosome numbers in Sect. Euphrasia is summarized in Yeo (1978b) and placed in abbreviated form in table 2. Since Euphrasia shows greater morphological diversity in the southern hemisphere, material from Australian members of the genus has been investigated for chromosome numbers. Table 2, Chromosome numbers in Euphrasia, a survey of the present knowledge, with the source of information in extra-Australian taxa and the voucher specimen (in AD) for Australian taxa. Taxon Chromosome number Source Sect. Euphrasia Subsect. Ciliatae Ser. Grandiflorae E. officinalis L. n=11 (Yeo 1972, 1978b) E. rivularis Pugsley n=1] (Yeo 1954) E. anglica Pugsley n=11 (Yeo 1954) E. vigursii Davey n=11 (Yeo 1972) E. hirtella Reuter n=11 (Yeo 1954) n=c.11 (Yeo 1970) E. picta Wimmer n=11 (Yeo 1978b) E. marchesettii Marchesetti n=11 (Feoli & Cusma, fide Yeo 1978b) Ser. Alpinae E. alpina Lamk. n=!1 (Yeo 1970) (Sell & Yeo 1970; Yeo 1954, 1970, Five other series: 16 species n=22 1972, 1978b) Subsect. Angustifoliae E. cuspidata Host n=11 (Feoli & Cusma, fide Yeo 1978b) E. salisburgensis Funk n=22 (Yeo 1972) Sect. Pauciflorae Subsect. Pauciflorae E. mirabilis n=11 (Borgmann 1964) Sect. Striatae E. striata n=c.20-30 Barker 1060 E. hookeri n=c.26-30 Barker 1212 Sect. Australes E. crassiuscula ssp. eglandulosa n=c.27-30 Barker 1590 n=c.27-29 Barker 1593 E. collina ssp. paludosa n=c.27-29 Barker 1489 n=c.50-60 Barker 1504 ssp. collina n=c.28-30 Barker 1439, 1440 ssp. tetragona n=c.30 Barker 1374 ssp. trichocalycina n=c.28-32 Barker 1438 [2n]=c.56 Barker 1438 (tapetal cell mitosis) ssp. diversicolor n=c.50-60 Barker 1684 ssp. glacialis n=c.50-60 Barker 1685 Sect. Lasiantherae E. alsa n=27 Barker 1696 E. lasianthera n=c.38-40 Barker 1535 n=c.42-47 Barker 1536 Sect. Scabrae E. caudata n=27, &/or 28, &/or 2711+21, &/ or 251142111 Barker 1649 Sect. Trifidae E. antarctica n=44 (D.M. Moore, fide Yeo 1968) 36 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia a. Method Bud material from plants in the field was used in the study. Young inflorescences were fixed immediately after collection in a mixture of 3 parts ethanol : 1 part acetic acid, transferred to 70% ethanol after approximately 24 hours, and stored, wherever possible, in ice until transfer to temperatures of below 0°C. Storage for periods of a week up to 18 months after collection showed no noticeable difference in chromosome definition. In the fixed material it was found that pollen mother cell meiosis was confined to one or two consecutive bud pairs in each inflorescence. These buds were about 2mm long and usually 5-7 nodes above the youngest open flowers. Difficulties, such as described by Yeo (1954) in Euphrasia and Heckard (1968) in the North American genus Castilleja (also of Trib. Rhinantheae), were experienced in obtaining preparations suitable for accurate determination of chromosome numbers. The chromomsome exhibited poor differential staining against the cytoplasm, and overlapped in a high percentage of well-spread pollen mother cells at late prophase to anaphase of the first meiotic division, the best stages for study. Neither lactic-aceto- carmine nor lactic-propiono-carmine (see Dyer 1963; Cooperrider & Morrison 1967; Cooperrider & McCready 1970) were satisfactory in the present study. The technique adopted was recommended by Dr B.A. Barlow (pers. comm. 1972). Standard anther squash techniques were employed using a light propiono- carmine stain of concentration of 1% carmine (or slightly less) in 45% propionic acid; no iron mordant or acid hydrolysis was found necessary. Meiotic figures were studied with the aid of phase-contrast illumination. This technique was not entirely satisfactory but was sufficient to give an indication of chromosome number. b. Results Despite the inaccuracy of the chromosome counts obtained for the Australian species of Euphrasia, the information is published to help resolve questions concerning the evolution of the genus and better define the more useful areas for future karyological study. The results obtained (table 2) cover representatives of all sections in Australia except Sect. Cuneatae and Sect. Phragmostomae, each with only a single Australian species. Chromosome counts of the Australian species are sufficient to show that they differ in chromosome number from those known for the rest of the genus. Their numbers are rarely, if at all, a multiple of the generic base number, x = lI. They indicate that the species studied have had a complex history of karyotype evolution, and are unlikely to represent a primitive stock from which the rest of the genus evolved, as proposed for example by Yeo (1968) for the perennials. However, it should be noted that a number of key taxa, e.g. E. bella (Sect. Australes), Sect. Phragmostomae and Sect. Cuneatae, were not examined. It is fairly certain that the haploid number of E£. alsa is n = 27 (fig. 15). Its close relative E. caudata has n = 27 or 28 or both. In this species there is some evidence of abnormalities at least in the population Barker 1649, as within the one flower meiotic figures apparently exhibited 28 bivalents, others 27 bivalents and 2 univalents, another perhaps 25 bivalents and 2 trivalents (figs 15, 16). The counts of the subspecies of E. collina (Sect. Australes) clearly show evidence of two levels of ploidy, with E. collina ssp. paludosa showing both. The lower ploidy level in E. collina approximates to the chromosome number estimates for E. alsa (Sect. Lasiantherae), E. caudata (Sect. Scabrae), E. crassiuscula ssp. eglandulosa (possibly a hybrid species from parent species in Sect. Australes and Sect. Striatae: 37 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) see below), and two species of Sect. Striatae. Sect. Australes, Sect. Scabrae and Sect. Lasiantherae apparently form a monophyletic assemblage (p. 55), and it is therefore likely that any differences in chromosome number masked by the variable estimates obtained in the present study are of an aneuploid nature, with secondary development of chromosome number differences within these sections being either polyploid (exemplified by the subspecies of £. collina) or even aneuploid (possibly indicated by the counts for Sect. Lasiantherae). The status of E. crassiuscula, with its position intermediate between Sect. Striatae and Sect. Australes (p. 64), in the evolutionary history of Euphrasia in Australia may well be resolved in a chromosome number survey of the Australian species. Fig. 15. Sketches of meiotic chromosome configurations in pollen mother cells of E. alsaand E. caudata (not to scale). A, E. alsa (Barker 1696), diakinesis, n = 27; B, C, E. caudata (Barker 1649), diakinesis, n = 27/28; D, E. caudata (Barker 1649), part of cell at diakinesis. 38 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia Fig. 16. Sketches of meiotic chromosome configurations in pollen mother cells of E. caudata (Barker 1649) (not to scale). A, metaphase I - early anaphase I, n = 27/28; B, early anaphase I, n = 27II and 21 (arrowed); C, diakinesis, n = 2511 + ?2III (arrowed); D, early anaphase I, n = 28. 39 W.R. Barker J. Adelaide Bot. Gard. 5 (1982) In conclusion, it has become obvious that the techniques used in the study of the chromosomes need to be improved and that future investigations in Euphrasia be extended to include species elsewhere in the southern hemisphere and tropics. Such a study would assist greatly in the clarification of evolution in the genus. Chromosome numbers of the New Zealand species must be especially important because of their possible intermediacy between the tropical members (with a single record of n = 11) and on the one hand the Australian species (with chromosome numbers apparently derived by a series of steps) and on the other the South American species (with their single record of an octoploid, n = 44). A survey of the chromosome numbers particularly of the perennial sections could also assist in assessing which of the present-day species are closest to the primitive stock from which Euphrasia and allies such as Odontites (n = 10, 20: Yeo 1972) and Bartsia (n = 12, 24, 36: Yeo 1972) are derived (p. 51). 14. Pollen sterility tests The confusing taxonomy of Euphrasia in Australia has often been attributed to a possible high incidence of hybridization. A hybrid may often be detected simply by the incidence of a higher than normal proportion of sterile pollen. The technique used for tests for pollen sterility is identical to that proposed by Owczarzak (1952). Only dried herbarium material was used. Pollen was taken from mature buds or young flowers prior to anthesis. The grains were mounted on a . microscope slide in a medium of glycerol jelly containing phloxine, which stains the contents of the pollen, and methyl green, which stains the pollen wall. The slides upon which the counts are based are housed in AD, marked by the PS code given in the text. The estimates of the percentage of functionally appearing pollen have been listed in a duplicated table available on request from the author. It is stressed that the technique does not measure pollen fertility. The estimate of the proportion of grains of “functional” appearance is only an approximation of this, as some of the grains may be incapable of fertilization. Furthermore, high pollen sterility need not be caused by hybridization. Abnormal environmental conditions and genetic aberrations in pure populations may also affect pollen development. However, if high pollen sterility occurs. in plants which are intermediate between taxa which themselves produce mainly functional pollen, then this can be taken as good evidence that the intermediates are of hybrid origin. Finally, it is emphasized that-hybrids may show no loss in pollen fertility. C. FLORAL BIOLOGY The flowers of Euphrasia are clearly adapted to cross-pollination by insects, as is shown by field observation of pollination in Europe (e.g. Yeo 1966) and Australia (see below). There are two general flower types. One (type 1) is distinctly bilabiate, with a large lower lip, with anthers which are fused together and rigidly held to the upper side of the mouth under the hood and with, in most species, the stigma initially placed well in front of, and often below, the level of the anthers. The other (type 2) is sometimes erect and often with a long tube, and bilabiate but tending to be falsely actinomorphic by the lower lip being much shorter relative to the upper lip than in the first type and the upper and lower lobes being sharply reflexed and more or less equal. The stamens are free and appear to take up more room in the mouth and tube than in the first type. The stigma in most species is initially directed well above or in front of the anthers. 40 iii abi isp eee Lb, ce. J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia With few exceptions the process of flower development important to pollination is probably identical to that reviewed by Yeo (1966) for Sect. Euphrasia. Most species are apparently protogynous, with the stigma and anthers well separated at anthesis. Elongation of the corolla tube, which supports the stamens, finally brings the anthers | into contact with the stigma. Thus the flowers seem primarily adapted to cross- q pollination and, failing this, possibly self-fertilization, providing there are no self- compatibility barriers. This process apparently occurs in species with relatively tiny <— flowers such as E. alsa and E. dyeri. However, in the European species (Yeo 1966), whereas the larger-flowered species are essentially adapted to outcrossing, the smaller- flowered species are strongly adapted to self-pollination as the dehiscing anthers contact the stigma by the time the flower opens. Similarly, as the small flowers of E. antarctica (Sect. Trifidae) open, “the anthers stand exactly above the stigma, which is fully developed before the flower is opened. It is inevitable that pollen falls upon the stigma” (Skottsberg 1913, p. 51). | Flower type 1 (e.g. fig. 17) » This flower is characteristic of all species except those of South America and E. disperma of New Zealand. It is clearly adapted to pollination by bees for its morphology and coloration, usually of yellow blotches and often with purple L, striations, matches that characteristic of bee-pollinated flowers as described by Faegri & van der Pijl (1971), although other insects may occasionally act as pollinators ) (p. 44). In this flower type the structure of the anthers, which have rigid awns pro- truding into the corolla mouth and anther slits opening obliquely downwards, ensures that an insect entering a tube to gather nectar knocks pollen on to its upper parts. Fig. 17. Native bee Exoneura (Exoneura) sp. visiting the prominently striated flowers of E. lasianthera (Sect. r Lasiantherae), Macalister River headwaters, Eastern Highlands, Victoria ( Barker 1498). 41 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) The awns of the posterior pair of anthers are presumably most involved in this process as they are longer than the other awns and nearer the tube. It is not known whether difference in length of the rearmost awns between taxa has any significance in pollination. The character is not always related to variation in size of the corolla mouth (p. 32). However, the remarkably long and often oddly shaped rearmost awns of E. phragmostoma (Sect. Phragmostomae: fig. 30) may relate either to some unusual pollinator or to the particularly large corolla mouth (fig. 30) or to both. The Australian species of Euphrasia have this type of flower and all those seen in the field were found to be pollinated mainly by native bees. On all but two of the occasions on which they were seen visiting flowers of Euphrasia, one or two native bees were collected for identification. Bees were observed to enter the corolla tube, apparently to feed on nectar, and to crawl over the anthers to gather pollen. Their attraction for the flowers of Euphrasia was well-illustrated when a plastic bag containing three copiously flowering plants collected from a population of E. collina ssp. osbornii (Whibley 4155) was found to contain about twenty native bees hidden within the flowers. Bees have also been seen visiting flowers of Euphrasia in the northern hemisphere, but according to Yeo’s (1966) observations their interest was mainly confined to gathering nectar rather than pollen. The bees seen visited flower after flower in the populations of Euphrasia. In at least one location (Barker 1492: E. collina ssp. paludosa), if not all, the bees visited only Euphrasia; they ignored a Ranunculus species and two yellow composites, although they flew to these plants and hovered over them before returning to Euphrasia. At one site the same bee species may visit two species of Euphrasia, effecting hybridization. On the Central Plateau of Tasmania (Barker 994) where bees visited both E, collina ssp. diemenica and E. striata, which were flowering simultane- ously and growing together, the two plant species hybridize (see p. 287). It is possible that bees (fig. 17) in the Mt Howitt region are responsible for hybridization between E. lasianthera and E. collina ssp. paludosa, which are sympatric and have overlapping flowering times (p. 288), for bees similar to those collected from the flowers of E. lasianthera (Barker 1498) were seen (but not collected) among the flowers of E. collina ssp. paludosa at localities several kilometres away (Barker 1492, 1495). Dr T. Houston of the Western Australian Museum has kindly identified the bees collected (table 3). Seven different species of native bee from the three genera Lasioglossum (5 species), Leioproctus (1) and Exoneura (1) were collected at eight different sites, which encompassed lowland to alpine regions from Tasmania, the Australian Alps and South Australia. There were four cases of the same bee species being found in the flowers of Euphrasia at two separate sites in the same geographical region but often widely apart (table 3). Accordingly, despite the small number of collections made (a total of 35 bees), it would appear that out of 3,000 species of bee estimated to occur in Australia (Michener 1970), a significant proportion of these being polylectic (although there is little data on this: Dr T. Houston, pers. comm. 1977), only a limited number of taxa frequent the flowers of Euphrasia. Three of these twice-collected species were Lasioglossum, the other was the only species of Leioproctus collected. The purple striations and yellow blotches on the corollas of many species with this flower type are considered to be nectar guides for the bee pollinators. Yeo (1968) considers that in Euphrasia the bee is guided to the nectar by the striations and the pollen by the yellow blotch on the lower lip onto which most pollen falls. However, it also seems likely that the yellow areas in the tube, which are often present when the yellow blotch on the lower lip is lacking, serve as a final guide to the nectary, for these blotches are well behind the anthers. 42 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia Table 3. The native bees collected from populations of Euphrasia in Australia, with corolla coloration of flowers visited. (Lasioglossum = Family Halictidae Subfam. Halictinae; Leioproctus = Family Colletidae Subfam. Paracolletini; Exoneura = Family Xylocopidae Subfam. Ceratininae.) Euphrasia species Euphrasia Corolla Bee species Sex Number collection coloration of of bees number "Purple Yellow. bees collected stria- blotch tions E. striata Barker 994 ot + Lasioglossum (Austrevylaeus) sp. female 2 Tas. highlands ’ E. lasianthera Vict. alps location A Barker 1498 + + Lasioglossum (Parasphecodes) sp. female 1 Lasioglossum (Chilalictus) sp. female 1 Exoneura (Exoneura) sp. female 6 location B Barker 1535 + + Lasioglossum (Parasphecodes) sp. (= 1498) ‘female 2 Leioproctus (Leioproctus) sp. female 2 E. collina . —ssp. diversicolor. Barker 1665 - + Leioproctus (Leioproctus) sp. NSW alps (= 1535) female 1 —ssp. diemenica Sympatric with - + Same bees as 994 Tas. highlands Barker 994 —ssp. deflexifolia Barker 942 - - Lasioglossum (Austrevylaeus) Tas. east coast sp. (= 994) female 2 Lasioglossum (Lasioglossum) sp. _ female 2 —ssp. osbornii SA location A Barker 858-869 ~ - Lasioglossum (Chilalictus) male 9 lanarium location B Whibley 4155 - - Lasioglossum (Chilalictus) male 6 lanarium (= 858) location C Barker 1346 - - Lasioglossum (Chilalictus) male ] lanarium (= 858) Some sections of Euphrasia in the southern hemisphere diverge by the complete lack of purple striations and common lack of yellow blotches on the lower lip. In Australia this is especially characteristic of Sect. Australes, Sect. Scabrae and Sect. Phragmostomae. From table 3, both of the genera of bee collected more than once, Lasioglossum and Leioproctus, apparently show no preferences for any one type of flower coloration. Thus Lasioglossum was found in striated and non-striated flowers and flowers with and without yellow blotches on the lower lip. Such non-specificity for coloration is shown within single species of both bee genera, and is supported by each of the few examples of interspecific hybridization in Euphrasia in Australia (table 4), in which corolla coloration and sometimes shape or size differ markedly. The absence of purple striations and yellow blotches in Australian species of Euphrasia occurs in several unrelated groups. This is concordant with the fact (Dr T. Houston, pers. comm. 1973) that Australian native bees can recognize suitable flowers without such guides. For example the Myrtaceae, which provide the major source of nectar and pollen for these bees (Michener 1965, 1970), lack such stripes and blotches. The development in Euphrasia in Australia of special features of coloration, such as the wide variation in ground colour of the corollas of many species and the prominent pair of yellow blotches in the centre of the flower of E. Jasianthera (Sect. Lasiantherae), and the diversity of floral shape can probably be linked to competition in the presence of a diverse Australian bee fauna and the wide diversity of floral shape and coloration in Australia’s bee-pollinated plants. 43 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) Table 4. Interspecific hybrids in Euphrasia in Australia with the floral characters of the parent species. E. alsa (open corolla, purple-striated, yellow-blotched) x E. caudata (closed corolla, non-striated, reddish-blotched) E. lasianthera (purple-striated, yellow-blotched) x E. collina ssp. paludosa (non-striated, non-blotched) E. striata (purple-striated, yellow-blotched) x E. collina ssp. diemenica (non-striated, yellow-blotched) E. orthocheila ssp. orthocheila (corolla closed, yellow) x? E. collina ssp. paludosa (corolla + open, white to lilac) There is a similar lack of prominence in striations and yellow patches in the corollas of the New Guinea and New Zealand species of Euphrasia (p. 29). I was able only to see E. humifusa and E. mirabilis in the field in New Guinea and found no pollinators visiting these species during my relatively short time studying them. The corolla lips of such species as E. mirabilis and E. papuana, however, are large relative to their small calyces, and the stigma initially projects in front of the anthers. Both attributes point to the species being cross-pollinated. It is noteworthy that the New Guinea bee fauna, which is very depauperate compared to the rest of the world, contains representatives of Lasioglossum and Leioproctus (Michener 1965). In New Zealand the perennial Sect. Pauciflorae and the two annual species E. zelandica and E. australis (Sect. Novaezeelandiae) were seen to lack the prominent purple striations which characterize other New Zealand species, but to possess striking yellow coloration on the mouth and tube. Remarkably, prominent striations on the outside (rear) of the corolla lobes, out of view of the visiting insect, occur frequently in these species (p. 30). This may reflect the depauperate nature of the New Zealand bee fauna (Michener 1965, 1979), and the reliance of New Zealand plant groups with characteristic bee-pollinated flowers on a more generalized set of insect pollinators (Godley 1979; Wardle 1978). Furthermore, it seems clear that the absence of corolla striations on the front of the lobes is a derived condition for sometimes a vestige of pigmentation is apparent. There are no patterns in ultra-violet in the places where striations are lacking. The gradual loss of hairs on the anthers of a series of species in the New Zealand annuals culminating in the free completely glabrous anthers of E. disperma (p. 33), may also reflect a reduced dependence on bee pollination, to which the position, arrangement and structure of the anthers typical of Flower Type | are clearly adapted. Finally, it is noted that the New Zealand bee fauna includes Lasioglossum and Leioproctus (Michener 1965), species of which visit Euphrasia in Australia. My collections of bees in Australia are confined to those visiting E. collina (Sect. Australes), E. striata (Sect. Striatae) and E. lasianthera (Sect. Lasiantherae), all perennials with similar corolla shape. A study of pollinators of Sect. Scabrae, which consists of annuals with a more closed and elongated corolla mouth (p. 27), is desirable. Of further interest would be a comparison between E. caudata (Sect. Scabrae), with a non-striated corolla of sucha shape and with a long narrow blotch on the lower lip, and E. alsa (Sect. Lasiantherae), a closely related annual from the same geographical region, with a tiny striated corolla of the same shape as the perennials studied above. That such different corolla shapes at least sometimes attract the same pollinators is indicated by the hybridization between the two species (p. 290). Whereas the native bees seem to be the most prevalent visitor to the flowers of Euphrasia in Australia, several other flying insects were seen visiting them. Among these were several species of butterfly. Graphium macleayanus was recognized visiting E. crassiuscula ssp. crassiuscula. It alternated between the flowers of Euphrasia and 44 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia those of other plants. The variant of E. collina related to ssp. diversicolor on The Cobberas was also visited by a butterfly. A hover-fly (Syrphidae) was seen visiting flowers of E. collina ssp. collina (Barker 982). Butterflies and hover-flies have also been observed visiting the flowers of the same type in the northern hemisphere (Yeo 1966). Flower type 2 (e.g. fig. 18) It is uncertain whether the obvious differences between the two floral types reflect a general change in pollinator. This second type is found only in Sect. Trifidae of South America and the closely related E. disperma (Sect. Anagospermae) of New Zealand (fig. 18). E. scutellarioides (Sect. Pauciflorae) of New Guinea, a species bearing no close relationship with the above sections, has flowers of an intermediate type with a long-tubed corolla with much-reduced lips but retaining the fused anthers. It seems possible that these flowers are adapted to pollination by butterflies or moths (Yeo 1968). On morphology this seems very likely for the extremely long and narrowly tubed flowers of E. disperma. Yet in this species the indigo striations and yellow markings are very marked (p. 30) and seem to indicate bee pollination, although it would have to be a small and adventurous or very long-tongued bee if it gathered nectar. In some of the species of Sect. Trifidae the mouth of the flower opens down- wards because of the long upper lip and the very short lower lip. These seem indicative of a type of flower pollinated by moths as described by Faegri & van der Pijl (1971). Only field studies of flowers of this type can show to what extent the different morphology has been paralleled by change in the nature of the pollinator, and whether such characters as the glabrous anthers have any significance in the pollination process. The lack of hairs along the anther slits in the former two sections may enable the pollen to be dispersed in larger masses. It seems unlikely that the anther awns would have much significance in these flowers, although they are long in some species. Fig. 18. The flowers of E. disperma (Sect. Race er near Denniston, New Zealand (Barker 3716), showing the slender long erect corolla tube, striated lobes and free subexserted anthers. 45 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) Two of the anthers always possess one awn longer than the other six awns in the flower. These longer awns are the rearmost pair of awns in the bilabiate bee-pollinated flower from which this flower type has almost certainly evolved, but they have possibly lost any special functional significance. III. THE ORIGIN AND DIVERSIFICATION OF EUPHRASIA The means by which Euphrasia attained its distribution (fig. 19) in the temperate regions of both hemispheres with its greatest diversity in the southern hemisphere and a series of “connecting species” on the isolated mountain peaks of Malesia has been the subject of controversy. The genus has been considered either to be old and to have migrated by land from various places of origin to its widely scattered present day location, or to be young and to have spread by long-distance dispersal from its origins in the northern hemisphere through the young Malesian mountains into Australasia and South America. After a brief survey of past proposals, this author presents evidence for his view of the age and means of dispersal of the genus, and then how the genus attained its present range and diversity both within Australia and on a world scale. A. PAST HYPOTHESES Perhaps the first attempt at explaining the remarkable bihemispheric distribution of Euphrasia was by J.D. Hooker (1859b), who was struck by the many European features of the Australian flora (p.xciv). He (p.xvili) proposed that partially submerged former mountain chains could have provided a means of dispersal between Japan and Australia, the existence of Euphrasia on “the lofty mountain Kini Balou in Borneo” (material probably cited in Stapf 1894) providing evidence of this. Until the relatively recent wide acceptance of the reality and applicability of plate tectonics to biogeography, it has been believed that Euphrasia was an old Tertiary genus which had attained its world distribution by migration over land. Wettstein (1896) pointed to the lack of affinity between the North and South American populations, and proposed that the genus arose on a now submerged South Pacific land mass connecting Australasia, South America and Asia. Like Hooker, Du Rietz (1932b, 1948a) proposed that the genus was linked by more continuous bridges between Asia and Australia, and between Australia, New Zealand and South America, the Australasian representatives being morphologically intermediate between those of the northern hemisphere and South America. More recently it has been suggested that Euphrasia, along with other advanced groups, attained its distribution by long-distance dispersal since the Neogene formation of subalpine and alpine outposts through the uplift of mountains surrounding the South Pacific and in Malesia and the Plio-Pleistocene glaciations (Raven & Axelrod 1972; Raven 1973). The genus belonged to the advanced sympetalous order Tubiflorae which is known in the fossil record largely from Neogene times when direct migration via Antarctica was impossible (Raven 1973). Wardle (1978) apparently also believed that Euphrasia could not have existed when Gondwanaland was whole, but disputed the need for genera such as Euphrasia being of Neogene age, commenting that a fragmented trans-Antarctic bridge would have been available for migration involving dispersal over ever-widening seas until the late Tertiary spread of the Antarctic ice sheet. Van Steenis (1971), however, has continued to believe that the number of bihemispheric distribution patterns similar (homologous) to Euphrasia, including the Nothofagus-Fagus and Winteraceae-Magnoliaceae pairs, both of which are widely recognized to date back to the late Cretaceous, reflect a synchronous development of 46 Studies in Euphrasia J. Adelaide Bot. Gard. 5 (1982) §. Atlanticae Fig. 19. The geographical distribution of the sections of Euphrasia, with the endemic Australian and New Zealand sections occurring within the blackened areas. The number of species in each section is shown, those largely comprising “microspecies”, equivalent to subspecies in the Australian revision, are bracketed. Euphrasia (c. 90) GP Paradoxae 1 AUSTRALIA E Phragmostomae | Cuneatae 2 Striatae 4 Australes 5 NEW ZEALAND Lasiantherae 3 plovaczeclandiae 4 Scabrae 5 nagospermae 4 (Distribution of Sect. Euphrasia after Hulten 1976). 47 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) all such groups from the Cretaceous. In reaffirming these assertions (van Steenis 1979), he argued that the repeated use of long-distance dispersal for explaining discrepancies between widely disjunct ranges and current geophysical theory was fallacious, and suggested a late Cretaceous-Paleocene or older tectonic connection between the Asia and Australia plates to accommodate bihemispheric distributions with present-day links in Malesia. There has been controversy over the site of origin of the genus. Wettstein (1896) and Hartl (1972) have both suggested that the genus has a common origin with its close relatives in Trib. Rhinantheae, in particular Bartsia and Odontites. Wettstein believed it to have an origin on his South Pacific land mass, while van Steenis (1962) proposed that it originated in the south-west Pacific region for in New Guinea and New Zealand the genus showed its greatest morphological diversity, in marked contrast to the poorly differentiated species of the northern hemisphere. The long-distance dispersalists (Raven & Axelrod 1972; Raven 1973) have proposed that a young Euphrasia had a northern origin, presumably on the grounds that this is where its present generic relatives occur. Du Rietz (1932b, 1948a), Hartl (1972) and Moore (1972) have not alluded to a centre of origin, but all believed that the southern hemisphere contained a number of old primitive or relictual types. Yeo (1968) also alluded to the primitiveness of the large-flowered Australian perennials. Compared with the diversity in Euphrasia in the southern hemisphere it has been widely recognized that the northern hemisphere annuals are a group showing great uniformity (Hartl 1972; van Steenis 1962) and of recent origin since the onset of the Plio-Pleistocene glacial periods (Yeo 1968, 1978a; Moore 1972; Hartl 1972; Karlsson 1974, 1976; Hulten 1976). Karlsson (1974) suggested that the complex ecotypic variation in these annuals in anthropogenic non-forest vegetation of temperate Europe may have developed in step with man’s influence. The arctic and warmer Mediterranean regions showed no comparable ecotypic variation. Before man modified the vegetation, Euphrasia may have existed there in isolated open habitats such as on cliffs and fens and in alpine regions. Sell & Yeo (1970), Karlsson (1976) and Hulten (1976) have pointed to the recent dispersal of some of these northern annuals, Karlsson suggesting that this capacity was confined to taxa which are adapted to man’s influence on the landscape. B. PROBLEMS IN ESTABLISHING CLADISTIC RELATIONSHIPS OF THE SECTIONS AND SUBSECTIONS OF EUPHRASIA The revision of the infrageneric classification of Euphrasia has produced a series of sections and subsections separable by about 25 characters of diagnostic importance (table 5), a few, e.g. chromosome number and corolla coloration, only potentially so because of poor data. The infrageneric taxa are separated entirely on phenetic grounds. Delimitation using only unique derived states, as required by the cladistic techniques of Hennig (1966), is not only impossible at this stage of our knowledge, but may also in theory remove from recognition distinctive species groupings such as Sect. Lasiantherae and Sect. Scabrae or, alternatively, dictate the division of Sect. Australes into two sections, as well as prevent the separation of E. hookeri from E. gibbsiae, since paraphyletic groups may be involved. However, the use of cladistics as the sole basis for production of classifications is questionable (e.g. McNeil 1978). The phenogram (fig. 20) derived from the number of absolute character differences between each pair of sections and subsections shows a central group of poorly differentiated sections. In view of the possible occurrence of convergence and parallelism in several characters used to delimit the sections, for example in a number of leaf and habit characters and in the origin of annual duration, a number of the 48 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia Table 5. The character states for each section and subsection of Euphrasia. “+” indicates the possession of a character state, “-” its absence, and “2” its possible presence. States in italics are considered as primitive. B. Subsect. Angustifoliae C. Subsect. Japonicae D. Subsect. Alpicolae Il. Sect. Atlanticae III. Sect. Malesianae IV. Sect. Cuneatae A. Subsect. Pauciflorae B. Subsect. Humifusae A. Subsect. Ciliatae VII. Sect. Striatae VIII. Sect. Australes (I. Sect. Euphrasia) V. Sect. Phragmostomae (VI. Sect. Pauciflorae) IX. Sect. Lasiantherae X. Sect. Scabrae XI. Sect. Novaezeelandiae XII. Sect. Paradoxae XIII. Sect. Anagospermae XIV. Sect. Trifidae LIFE-SPAN 1, Annual 2. Perennial 1+ + 1+ 1+ + + + +o + re bd 1+ 1+ ++ MAIN INFLORESCENCE-BEA RING AXIS Direction : 1. + entirely prostrate ee eo Srp > Fe So og o oe s 2. Initially prostrate, distally erect, simple above ground level we mm ee 3. + erect, branched well above ground level Incidence of branching 1. Consecutive nodes 2. Sporadic nodes 3. Separated groups of nodes, consecutive in groups - ee ee ee Order of branch development 1. + basipetal 2. No fixed order ee ie 3. + acropetal S54 =» = Bo as a rt + rt + rt + 1+ + th o+ ++ + 1+ + +i + + ++ +o o+ + ay r+ + ++ + ++ + + + + + + + + (+o + 1+ UPPERMOST LEAVES Depth of toothing 1. Crenate 2. Serrate 3. (Pinnati)fid Number of pairs of teeth 1. (0)1(2) ee eee 2. (1)2-6(more) ++4+4+-4++ + + 3. c. 6-many a 6 so deen ke z = Base 1. Attenuate, abruptly expanded into blade woes Woaear es aoa oe ee we So aes 2. Attenuate, gradually expanded into blade Sap gr oe = (Aap oe SF ar ae sr 3. Narrow-cuneate to truncate Sse Ss + & Ss = a Se + 4 + 44 ~2 +4 ot +. ot ' ydeak bapa 1+ +44 r++ +45 Blade (defined p. 14) 1. Large cf. base and teeth ap Se ae 2. Small cf. base and teeth - = Distribution of teeth 1. Distal !/, or less 7 oe 2. Distal !/, or more Trap Fe Number of main veins produced from base 1. (1)3 or 3 arr ar oe & ae ae 2. 3(5) CP SS ee = 3. 3-5 or more - - = = te. - - = = 1+ + + 1+ + eet +4 41 +4 1+ 1+ fae 33 + + 1+ Continued on page 50 49 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) Table 5 (continued) (I. Sect. Euphrasia) A. Subsect. Ciliatae B. Subsect. Angustifoliae C. Subsect. Japonicae D. Subsect. Alpicolae II. Sect. Atlanticae III. Sect. Malesianae IV. Sect. Cuneatae V. Sect. Phragmostomae (VI. Sect. Pauciflorae) A. Subsect. Pauciflorae B. Subsect. Humifusae VII. Sect. Striatae VIII. Sect. Australes IX. Sect. Lasiantherae X. Sect. Scabrae XI. Sect. Novaezeelandiae XII. Sect. Paradoxae XIII. Sect. Anagospermae XIV. Sect. Trifidae FLOWERS Occurrence 1. Sporadic along axes my fal eee 2. Racemes usually with less than 10 flowers 3. Racemes usually with 10-20 flowers 4. Racemes usually with more than 20 flowers COROLLA Striations 1. Present, but absent along midline of hood ; te inet ar ff an re geogro ae sec) oY ar Y 2. Present, at least aiong midline : of hood Se SS oo hE 3. Absent ee ee Blotch on lower lip l. Present +4+4+ H+ 2. Absent ee Lower side 1. Concave from above 2. Flat or grooved Lobes (lower) 1. Emarginate 2. Acute or obtuse to truncate ANTHERS Arrangement 1. Fused 2. Free Area about connectives 1. Glabrous ae ap ap awe ae fae op a Ca et 2. Hairy, at least on posterior anther pair me ee Slits 1. Densely hairy ap Sr Geese ar ap car se ar ar) ae ae 2. One or two hairs ee Be Ee ws er er ts Si ts 3. Glabrous iS pe es es Blends ee 4 Awns 1. All needle-like Brae Te Ge aR ae ar 23 ++++4¢+44+4 2. Rearmost pair flattened or with margins erosulate or twisted ee ee ine Se Bes te ee Pe Shs STIGMA 1. 0.1-0.3mm long os os oe Ss So SO a ee ee oe eee 2. (0.2)0.3-0.5mm or more long en Ss ae dk et a, a Te de ae CAPSULE Lateral view (overall) 1, Oblong order ase apy Y 2. Broad-ovate order Sc pe, ar Rar ay Aen ~ oP + + ' ' a ap tear Shap ou os 1 1 at ' + ipteet at + ++ +: ++ a+ +: 1+ + ~ 1+ +4 1+ 1+ 1+ 4. S a 1+ v+ 1+ 1+ 1+ 1+ 1+ 1+ + + 1+ 1+ ++ t+ ++ t+ +4 ~~ + + 4 + + + + + + a + + + + -$ + a + ' +. t+ ote + ++ ++ t+-++4+ 4-4-2 tHt-- +--+ - 4H ++ Continued on page 51 50 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia Table 5 (continued) 3 $e a) ~ hk) 2; v iS} aSEcES = $8 SS SS § 5 a SS gy = = SEES iS b= 2 Sy Ss 8 ei AP asy IS Se NHS ES 8 a SE AE m & vo 8s 8 ao 5 8 Se & Se Sas Sk 8 PS SPSS Be EUs DS aS MS soe og ES OPS Ae FETE FS FHS SSRESSSSSESPRESESSERE ere ~ HBeESBISORA ESR XIGC SRE EERE) el cea aiiabes Pde hea cp ar hbo pete oe Be ae Co Bod 2 ALMUANRANRHAUH IF ANANRUAANRHAAR 3 Hees Hoxxdede see S SERS KEES CAPSULE continued Lateral view of apex ; 1. Acute (acuminate) to obtuse qe oe 2 OR spar SP Gr Tee ae ap ae Se ap) SF 2. Shallowly emarginate toemarginate + + + + - + + - ah oar oar ar ae ape tae, 3. Deeply cleft ee ee tl Od eee, SS ES CHROMOSOME NUMBER 1. n=11 aro Pe EY ae ee ae a 2. n=22 gear 4 WY 1 ge eee Rony yy Pe 4 3. n=44 aD SP Sage) TR) al? Be? 2. gate =) ESE A Et 4. n=27-33 Se EEG) EN Me yan teers ay PN EOE 5. n=45-60 ih he UE 8 YP oa se Wy) a sections are likely to be phyletically more distinct than shown. It is possible to distinguish several lineages from the central group, but unless the direction of evolutionary trends in characters is defined, i.e. primitive and derived states are deter- mined, the direction of the lineages is questionable. In considering the biogeography of any group, the formulation of evolutionary trees is of great use, and undoubtedly cladistic theory has conceptually many advantages here. However, to use cladistic techniques properly unique derived character states must be discerned (Hennig 1966). These define monophyletic groupings. In a study of the Australian Mimulinae (Scrophulariaceae: Barker, in press) sufficient unique states were available to offset the phenomena of convergence and parallelism which are widespread in the family. However, it is difficult to define such states in Euphrasia. Characters such as the hairy anther backs and sessile leaves of three Australian sections, and the glabrous free anthers and trifid leaves of the South American species and FE. disperma of New Zealand are peculiar to these taxa within Euphrasia, but like their alternative states, are found elsewhere in the tribe and, indeed, other tribes of Scrophulariaceae. Such characters, nevertheless, usefully define closely allied groups of species, and provide a firm basis for consideration of infrageneric relationships. Consideration of relationships with allied genera could help to define an archetype of the genus from which to erect a detailed scheme of diversification in the genus. This is not possible at this stage owing to problems of heterobathmy; apparently primitive attributes are spread over a number of genera in Trib. Rhinantheae, such that no one genus or group of genera can be designated as primitive within the tribe. Thus Euphrasia is closely allied to Bartsia and Odontites, and more distantly to Parentucellia and Bellardia in Trib. Rhinantheae (Hartl 1972). Whereas Hartl considers Euphrasia to be the most derived of these genera, Yeo (1968) suggests the corolla morphology of the genus is primitive in the tribe. Weber (1980) considers that this group of genera has a primitive haustorial anatomy; they lack the “wart haustoria” of the more advanced semiparasitic genera in the tribe. On the other hand, in considering the primitive traits of the tribe attention should be given to the genera 51 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) Sca Character differences All Fig. 20. A phenogram showing the degree of phenetic similarity between the sections and subsections of Euphrasia based on the total number of absolute character differences between each pair of infrageneric taxa derived from table 5. Sections and subsections are abbreviated to their initial three letters. The subsections of Sect. Euphrasia are not shown. Schwalbea of North America and Cymbaria (Pennell 1935) as well as Bungea, Monochasma and Phtheirospermum of central and east Asia. These genera have one or several traits considered by Pennell (1935) to be primitive in the Scrophulariaceae, namely a pair of bracteoles, a 5-partite calyx, and a septicidal capsule. It is noteworthy when considering possible derived character states in Euphrasia that the first three genera are perennials with basal branching and entire leaves, that at least Cymbaria and Bungea have glabrous, apparently free, but didynamous anthers, and that all apparently possess the porrect upper corolla lip which, apart from in Euphrasia, prevails in the tribe. It is possible that critical morphological and anatomical studies may resolve a number of the problems of possible convergence within the genus and the tribe, and provide other characters, e.g. in the pattern and distribution of the sessile gland patches on the leaves, but only in wide family based studies are sufficient unique derived states likely to be defined. Such studies seem far off. Primitive character states could arguably be defined as those which are the most frequent within the genus. Such a practice is dangerous as one (derived) part of a genus may diversify while the rest may remain stable or become extinct. Indeed, it is 52 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia proposed below that most of the extant taxa of Euphrasia have derived from a phase of diversification separate from the early radiation of the genus from which only a few, often very localized taxa survive. However, it has been possible to propose a series of relict species in Euphrasia, and on this basis a hypothetical set of primitive character states and in turn evolutionary lineages within the genus have been established. C. RELICTS OF AN EARLY PHASE OF WORLD EVOLUTION Some attributes, which occur in several distantly related sections, seem hardly able to have been derived by convergence and therefore are likely to be primitive states. It follows that some of the characteristics of a possible archetype of Euphrasia are as follows. Undoubtedly it was a perennial. Its pattern of branching was basipetal in consecutive nodes of erect to ascending main branches. Its leaves had a large, shallowly toothed blade gradually attenuated (subpetiolate) at the base. Very few of the species of Euphrasia existing today possess all or most of these proposed archetypal attributes. Those species that do tend to be morphologically isolated and geographically restricted, and are scattered through much of the range of the genus. These factors indicate that such species are likely to be relicts, as suggested for some of the species in the past (e.g. Skottsberg 1921; Du Rietz 1932b, 1948a). Sect. Cuneatae comes closest to the proposed archetype, bearing all the character states, while the endemic Japanese subsections of Sect. Euphrasia, Sect. Atlanticae, afew species of Sect. Malesianae, E. papuana of Sect. Pauciflorae, Sect. Paradoxae, E. bella of Sect. Australes, and Sect. Phragmostomae have all but one of the proposed primitive states. Among these Sect. Cuneatae (two very distinct species), Sect. Phragmostomae (monotypic), Sect. Atlanticae (two very distinct species), and Sect. Paradoxae (mono- typic) comprise morphologically very isolated perennials. E. bella of Sect. Australes is phyletically connected by one or probably two other geographically isolated perennial species to the rest of the section, and it must be considered a relict, possibly along with the other two species (p. 62). The two Japanese subsections Alpicolae and Japonicae differ from the hypothetical archetype only in their annual duration, and it is likely that they are derived in more recent times directly from archetypal progenitors. The world distribution of these proposed relicts is remarkable in the widely disjunct out- posts of the Azores, Taiwan and Borneo in the northern hemisphere and northernand central montane New South Wales, Tasmania, New Zealand, southeast New Guinea (Mt Victoria), Ceram and the Juan Fernandez Islands in the southern hemisphere (fig. 21). It seems, therefore, that the diversification in the genus was in at least two phases. The first produced an array of forms, some of the characters of which are likely to be retained in the proposed relicts. It is noteworthy that five of the evolutionary lineages proposed below (fig. 22) contain at least one relict species. The second phase involved the development of the present-day diversity of species with modification particularly to habit, leaf and floral characters. D. MAJOR EVOLUTIONARY LINEAGES The proposed relict species were surveyed to determine the state of each character diagnostic at the sectional level which is the most common amongst them. For the purposes of producing a phyletic scheme, this state has been assumed to represent the primitive state for each character (table 5). Remarkably, Sect. Cuneatae possesses all such states and is accordingly close to the consequential archetype of the genus. For the very few characters (e.g. presence of corolla striations) which were shared equally amongst the relict groups, the state possessed by Sect. Cuneatae (i.e. striated corollas) was accordingly taken as primitive. In a few cases there are inadequate data (e.g. chromosome number) or Sect. Cuneatae had each state (e.g. of corolla lobe apex). 53 J. Adelaide Bot. Gard. 5 (1982) W.R. Barker ‘(L681 “S6-168] W191SII9A\ WOIJ Peljpour Bjep) oUT] 24) MOTaq PUL 2A0Qe ATaANoadsar UMOYS UOTSa1 Yous Joy BOUT O1WapUD PUk [eI0} YIM ‘Ppom gy} Jo suOIBaI sNOLeA ul (avaoeLE(NYydolog) seayjUEUIyY “qu UI AyISIOAIP o119UE8 Jo sa1Sap ou]. ‘(ZL-6961 [48H 401Je) vIs14pg JO UONNGISIP plIOM oyL “pispdydng jo saioads jo1a1 pasodoid oy} Jo uoneo07T “17 ‘314 avoyueulyY “quy (_) DIS1Dg eS pispiydny jo satoads yoay > : “ J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia Using shared derived states six lineages have been produced (fig. 22), which on the above assumptions appear monophyletically defined. They conform with those generally obtained from a series of “Wagner Trees” using the methodology of Whiffin & Bierner (1972). Six sections (Striatae, Australes, Pauciflorae, Malesianae, Novaezeelandiae, Cuneatae) forming a central group in the genus on phenetic grounds (fig. 20) were used in turn as the taxon closest to an archetype of the genus in formulating the Wagner Trees. Such trees are not wholly cladistic, for they are derived through an assessment of relationships between the sections involving primitive as well as derived states. Most of the lineages have throughout or in a substantial part one or more characters unique in the genus. Although Sect. Cuneatae and Sect. Phragmostomae represent an entirely relictual Australasian lineage, the latter section is unique in its groups of branches and remarkable pair of rearmost anther awns. The character linking the northern hemisphere sections, Malesianae, Atlanticae and Euphrasia, is the small stigma. The Australes-Lasiantherae-Scabrae lineage of Australia is characterized by hairy anther backs and the usually sessile leaves with 3-5 or more main veins. In a south-west Pacific lineage Sect. Paradoxae of the Juan Fernandez Islands shares with Sect. Novaezeelandiae and Sect. Anagospermae of New Zealand acropetal branch development and sparsely hairy anther slits, while E. disperma of Sect. Anagospermae shares with Sect. Trifidae of South America trifid leaves and free glabrous anthers (see below). In contrast, the two lineages comprising Sect. Striatae of Australia and Sect. Pauciflorae of New Zealand and New Guinea are defined by derived character states of habit, leaves and inflorescence which are shared with more than one other lineage. The geographical restriction of the six lineages to different parts of the world supports their naturalness. Phyletic arrangements of these lineages, such as the one proposed in fig. 22, are debatable in some areas at this stage, but should be able to be tested by additional data, particularly chromosome numbers. In particular, morphological similarity between the sections of west Malesia (Sect. Malesianae) and east Malesia-New Zealand (Sect. Pauciflorae) presented under the proposed scheme (fig. 22) as reflecting close phyletic affinity, may be a product of convergence; the similar reduced habit with sporadic irregular branching could well have arisen in this way under similar ecological conditions in the high tropical mountains. The large leaf blade of Sect. Malesianae is evident in Sect. Pauciflorae only in E. papuana in New Guinea and in some New Zealand species. The other tropical species of Sect. Pauciflorae have a peculiar hooded leaf. Stigma size remains an important character, not only in dividing the genus into northern and southern hemisphere groups, but also possibly at a wider level in the tribe. A second problem is the phyletic relationships of E. disperma (Sect. Anagospermae). A placement as a sister group of Sect. Trifidae by the shared attributes of trifid leaves and free glabrous anthers would not account for their apparently large difference in ovule number, with its possibly associated difference in capsule shape, and the absence of the prostrate habit and extremely long corolla tube of £. disperma within Sect. Trifidae. The adopted arrangement linking E. disperma phyletically with other New Zealand annuals assumes that trifid leaves and free glabrous anthers are convergent traits. As Du Rietz (1932a) and Ashwin (1961) have indicated, the species apparently represents an end-point in transitions involving these and other characters, i.e. involving leaf incision, anther slit indumentum, ovule number, corolla tube length and development of prostrate habit, this last character being shared with two other species. 55 J. Adelaide Bot. Gard. 5 (1982) W. R. Barker Aust., N.Zeal. Celebes, N.Guin., N.Zeal. W.Males., Eurasia, N.Am., Azores oT! ——o —_—_—_—_—A_ Cuneatae Phragmostomae Pauciflorae Humifusae Malesianae Euphrasia_ Atlanticae Branching Prostrate. sporadic. Flowers Flowers few sporadic Branches in groups. Rearmost anther awns very long, flattened at tip Leaves: base abruptly attenuate, Branching teeth many sporadic, no fixed order. Flowers few. 1 Corolla non-striated 3 Stigma small Aust. Aust. Juan Fernandez Is., N.Zeal., S.Am. Trifidae Striatae Australes Scabrae Lasiantherae Paradoxae Novae- Anagospermae zeelandiae \ Leaves \ deeply incised \ Corolla non- striated Leaves sessile, 3-5 or more veined. Inflor- escences dense. Ovules many Branching sporadic, no fixed order. Leaves 3-5-veined Annual. Leaves abruptly attenuate Branching ete Leaves trifid, sporadic. Anther (to no fixed SEAN Ta: connectives hairy anther slits sequence) glabrous Anther slits (densely, Novae- zeelandiae p.p.) sparsely hairy Fig. 22. Six evolutionary lineages evident among the sections and subsections of Euphrasia, based on the assumption that the character states predominating amongst the proposed relict species are primitive, Sect. Cuneatae possessing all such primitive states. Derived attributes which are shared by the included taxa at each branch point, and distribution of each lineage are shown. Those sections which include proposed relicts are in italics. One possible phyletic arrangement of the lineages is shown in the inset. 56 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia Sect. Australes presents a third problem. At this stage evidence points to this section comprising two paraphyletic groups within the lineage ‘defined by the presence of hairy anther backs. Further work is needed to determine if the non-striated state of the corolla is a convergent trait, for the section as presently constituted seems natural phenetically. The phyletic placement of Sect. Striatae provides a further problem, for it differs only in the number of flowers in the inflorescence and the number of veins at the leaf base from Sect. Pauciflorae, and yet its two character differences from Sect. Australes are transcended by a species in the Australian Alps (p. 64). Clearly there is still much comparative morphological and cytological work to be done within as well as outside the genus before a stable evolutionary arrangement of the lineages can be defined. E. TIMING OF DISPERSAL EVENTS ACROSS BARRIERS TO MIGRATION This survey of the distribution patterns of the subspecies and species or natural groups of these taxa shows that two phases of dispersal of different chronology are involved. Sect. Euphrasia is excluded from this discussion because of its recently derived ready ability to disperse widely (p. 59). If Euphrasia attained its present world distribution by long-distance dispersal during or since the late Tertiary, as proposed by Raven & Axelrod (1972), it must have crossed at least 13 ancient oceanic barriers (table 6). No taxon of species or subspecies rank crosses such a barrier today, although the sections Malesianae, Pauciflorae and Cuneatae do (fig. 19). Table 6. Regions between which long-distance dispersal is required for Euphrasia to have attained its distribution in the last few million years (excluding Sect. Euphrasia). Eurasia - Azores North Borneo - Cream Australia - New Zealand (probably Asia - Taiwan Ceram - Celebes 3 times) Celebes - New Guinea New Zealand - Juan Fernandez Is. Taiwan - North Philippines North Philippines - North Borneo New Guinea - Australia or _ New Zealand - South America New Zealand Within the continental boundaries, the distribution patterns in Euphrasia provide a substantially different picture. Such distributions encompass generally narrower barriers to migration, bridged by suitable vegetation in Plio-Pleistocene times (e.g. southern Australia: Costin 1959, Hope 1978; New Zealand: Grant-Taylor 1966, Wood 1966, Shaw & Steven 1966; bridging between Fuegia and the Falkland Islands from bathymetric contours). Table 7 sets out taxa which cross such barriers of substantial width. Within Australia a number of disjunctions, which are narrower or based on insufficient distribution data, have been omitted. Data for taxa outside Australia are limited by lack of revisional studies and lack of sufficient information on distribution. Table 8 shows vicarious groups in which the taxa are separated by barriers bridged periodically in Plio-Pleistocene times. It is limited to Australian examples for the above reasons, although it is likely that New Zealand has no more than one such vicarious pattern. This involves E. drucei, which being on the North Island is separated from most other species of Sect. Pauciflorae by Cook Strait; all other species on North or Stewart Islands also occur on South Island. For the now well-documented Australian distribution patterns, out of 14 separate crossings of Plio-Pleistocene barriers, 9 are encompassed by a species or subspecies and 5 involve vicariance. Making a statistical comparison using the exact test for 2 x 2 contingency tables for small samples (Bailey 1959), the probability that patterns across 57 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) Table 7. Species and subspecies of Euphrasia which occur on either side of sea or other at present inhospitable regions in the South Pacific region. Taxon Geographical barriers to present-day migration Australia E. collina ssp. tetragona Bass Strait Nullarbor Plain Investigator Strait (Kangaroo Isl.) E. collina ssp. osbornii Investigator Strait (Kangaroo Isl.) E. collina ssp. paludosa Semi-arid Upper South-East, South Australia E. collina ssp. collina Bass Strait E. scabra Bass Strait Nullarbor Plain Semi-arid Upper South-East, South Australia New Zealand E. cuneata Cook Strait Region between Marlborough Sounds and Lake Ellesmere E. laingii Region between Marlborough Sounds and more southerly locations E. revoluta Cook Strait E. zelandica Cook Strait E. dyeri Foveaux Strait E. repens Foveaux Strait South America E. antarctica Strait west of Falkland Islands. a, Table 8. Species and section of Euphrasia with disjunct vicarious ranges on either side of at present inhospitable regions in Australia bridged by suitable vegetation in the Plio-Pleistocene. Taxon Nature of disjunction Sect. Lasiantherae Three high montane-alpine species separated by low montane regions. E. gibbsiae ssp. subglabrifolia Subalpine taxon separated from rest of E. gibbsiae by Bass Strait and lowland-montane vegetation. E. bella - E. sp. ‘Tamworth’ - Three closely related isolated montane species of nore rn and central E. bowdeniae New South Wales. the ancient oceanic barriers reflect an identical set of conditions is extremely low (P = 0.0007). The New Zealand situation (not differing significantly from the Australian) would lower the P value even further. Since no taxon of species rank or below crosses the oceanic barriers, the lowness of the P value seems limited only by the number of oceanic barriers associated with the range of the genus. The relatively large number of instances of specific or infraspecific taxa occurring on either side of the Plio-Pleistocene bridges can only be explained by overland migration, in view of the obvious rarity of successful long-distance dispersal across barriers, if it has occurred at all. It is clear that the spread of the genus across most, if not all, oceanic barriers has occurred at a much earlier time than the most recent intra- continental migrations. F. MEANS OF DISPERSAL It is possible to show that a plant group is capable of long-distance dispersal by the detection of propagules in transit and observation of establishment. However, it is impossible to prove a lack of such a capacity. One can only deal with probabilities based on the available data. It is important to remember that at least 13 independent instances of long-distance dispersal and successful establishment are required for the 58 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia genus to have attained its present world distribution in the last few million years. The following summary of various biological, morphological and chorological characteristics of Euphrasia shows no evidence for its present-day world distribution being attained by long-distance dispersal. Notwithstanding this, one portion of Euphrasia, comprising the two subsections, Angustifoliae and Ciliatae of Sect. Euphrasia, widely distributed in temperate Eurasia and North America, is likely to have a capacity for long-distance dispersal. There are strong grounds for considering these groups to be exceptional in the genus. It is widely believed (e.g. Yeo 1968; Hartl 1972; Karlsson 1974, 1976; Hulten 1976) that these are a derived group of which many taxa have become adapted to grazing and mowing and other anthropogenic changes to the vegetation. Hulten (1976) has indicated that these taxa are easily spread, for example in hay, and has suggested that a number of the most northerly arctic occurrences may have been attained through the agency of man. Seed of some of these taxa has been found in excreta of horses, cattle and reindeer (Ridley 1930), which graze their pasture and meadow habitats. A number of taxa are adventive in parts of Europe (Karlsson 1974) and North America (Sell & Yeo 1970). It is, therefore, remarkable that Sect. Euphrasia has only rarely become naturalized outside its natural range. I know only of a localized New Zealand record of E. nemorosa (Sykes 1981). No taxon of any other section is known to have been recorded as adventive anywhere in the world. The specialized ecological characteristics of these subsections are derived features which cannot be taken as indicative of similar capabilities in the remaining members of the genus and their progenitors. Seed morphology in Euphrasia shows no special adaptations for long-distance dispersal. Seeds are unlikely to be light enough to be dispersed by any but the most extreme winds. They are small but not minute and are, therefore, unable to be dispersed in a dust-like mass. Smith (1977) has reported a “splash-cup” method of short-distance dispersal in E. mirabilis of New Guinea, in which the hygroscopic capsule opens when wet and it is proposed that seed is splashed out by rain drops. It is difficult to see how this method of dispersal can significantly increase the chances of long-distance dispersal of this low-growing species, as proposed by Smith. The seed surface is characteristically multi-sulcate and scalariform. There are no appendages or sticky substances which would facilitate dispersal externally by animals. Apart from Ridley’s (1930) references to seed of Sect. Euphrasia, there is no evidence of animals, in particular birds, eating seed of Euphrasia, nor is there any apparent attraction for them to do so. Raven (1973) has suggested that seeds may have been dispersed between the lands of the southern hemisphere by mud caked on the feet of birds. The species of Euphrasia commonly occupy areas of grassland and low herbage, and some may associate with swamps or boggy sites, but unlike some widely distributed members of Scrophulariaceae, e.g. in Mimulus, Peplidium and Glossostigma (Barker, in press), few species, if any, grow in mud. The chances of such dispersal are therefore reduced considerably. The species of Euphrasia are apparently all bisexual and cross-pollinated by insects, but probably able to be self-fertilized in the absence or cross-pollination. Breeding experiments showing this have been confined, however, to Sect. Euphrasia (p. 41). The plants are all able to parasitize roots of other plants, but this is not obligatory nor is there any specificity known in the choice of host. There is no known symbiotic relationship important in the establishment and maintenance of populations of the genus. The high degree of endemism which characterizes the genus throughout its range points to an inability to disperse across even small disjunctions. The Australian taxa provide many examples, such as the local endemics E. phragmostoma, E. semipicta, 59 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) E. bowdeniae and E. bella, and the three species of Sect. Lasiantherae which are separated from suitable habitats by distances of only 40-80km. If any seed were transported by wind or birds over the long distances required, it may have little chance of forming an established population. Taxa of Euphrasia are strongly linked to their particular habitat, as exemplified by the strong ecotypic differentiation of the alpine and subalpine taxa of E. collina and E. gibbsiae in Australia. While Yeo (1964, 1966) has had considerable success cultivating European species of Sect. Euphrasia, reports of cultivation of Australian taxa are exceedingly rare despite their showiness, and the only record known to me (Malahide 1973) relates to transplantation. That no adventives are known from the genus apart from Sect. Euphrasia also attests to their strong ecotypic fixation. Moreover, from the general preponderance of old (pre-1900) herbarium specimens of montane and lowland taxa, no Australian taxon has flourished with the advent of European man, most if not all have diminished in range, and a number, both annual and perennial, may be extinct or threatened with extinction. It seems likely that successful germination and establishment after long-distance dispersal would require that the new habitat conforms with a narrow set of ecological conditions similar to the seed source. It is concluded that apart from the recently derived, highly specialized species of two subsections of Sect. Euphrasia, the present-day members of the genus exhibit no evidence of a capacity for dispersal over the long distances required for Euphrasia to have attained its world distribution. In the absence of even one example, outside Sect. Euphrasia, of a species or subspecies presently encompassing one of the ancient wide oceanic barriers which species in the genus would need to have crossed in the past, it is concluded that the genus is far more likely to have migrated over land with plant associations to which it was closely adapted. G. DIVERSIFICATION OF EUPHRASIA IN AUSTRALIA The present-day Australian taxa have apparently been derived from three separate evolutionary lineages. The first lineage, involving two relictual sections, has shown no radiation in Australia, but the other two have diversified into a wide range of habitats, with the high polymorphy of two species in particular indicating that speciation is still going on today. In these last two lineages there is evidence for independent parallel evolution in a number of characters involving habit, ovule number and seed size commonly in association with a transition between montane/ lowland and subalpine/alpine conditions. Speciation has apparently resulted primarily from geographical or ecological isolation. There is evidence for some speciation through hybridization between unrelated species, but this seems of secondary importance. The situation thus conforms to some extent with the picture presented by Karlsson (1976) for the European species. He refers to strong ecotypic differentiation in the indigenous Swedish species, parallelism in a range of characters, including some involved in the Australian situation, related to climatic and habitat factors, and a role for hybridization much more limited than proposed by Yeo (1978a, b). Even more so than at the infrageneric level, the establishment of uncontroversial evolutionary trees on strict cladistic grounds at a level of species and subspecies seems impossible, owing not only to the almost complete absence of unique derived character states, but also to difficulties in defining primitive and derived states for almost all characters through their often quantitative nature and the clearly frequent possibilities of convergence or parallelism. The schemes are therefore by necessity a subjective assessment based on phenetic relationships. 60 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia Evolutionary line 1: Sect. Cuneatae - Sect. Phragmostomae (fig. 23). This group comprises three relict species which are morphologically very distinct and widely disjunct (fig. 27). Of the two species of Sect. Cuneatae, E. cuneata is the more closely allied to E. phragmostoma (Sect. Phragmostomae). E. cuneata of New Zealand is polymorphic and has a wide ecological range, including Nothofagus forest and ranging from coastal to subalpine situations. It encompasses the different climatic ranges of the other two species, E. ramulosa extending from wet sclerophyll forest into subalpine grassland in northern New South Wales, and £. phragmostoma occupying coastal scrub in southern Tasmania. Sect. Phragmostomae Sect. Cuneatae Fig. 23. Evolutionary diversification of Euphrasia E. phragmostoma _ E. cuneata Eararilosa in Australia (1). Postulated evolution of the species of Sect. Cuneatae and Sect. Phrag- mostomae. Branches in spaced groups. Rearmost anther awns very long, flattened at tip Evolutionary line 2: Sect. Striatae (fig. 24). Sect. Striatae is represented today by five species, mostly occupying alpine and subalpine habitats on the summits of the many mountains of southern, central and western Tasmania. Two distinct lineages are evident while there may have been a third, now extinct. The presence of E. gibbsiae ssp. subglabrifolia on Mt Baw Baw of eastern Victoria attests to migration between the present mountain outposts via land across Bass Strait during the Plio-Pleistocene glacial periods. The “£. gibbsiae line” appears to be undergoing active speciation in response to the shrinkage and expansion of the subalpine/alpine zones during the Plio-Pleistocene period. Subspecies such as ssp. wellingtonensis and ssp. pulvinestris, which are restricted to single massifs, may have evolved during the present interglacial period, while wider ranging subspecies such as ssp. comberi, ssp. gibbsiae and ssp. discolor may have evolved in earlier interglacial periods. The narrow ecological preferences and evidence for some development of barriers to interbreeding indicate that some subspecies have potential for wider morphological divergence. E. hookeri, with its remarkable hand-like leaves, may have evolved in this way from the progenitors of one group of present-day sub- species of E. gibbsiae. Its past placement (Wettstein 1896; Du Rietz 1932b, 1948a), with other “subdigitate”-leaved species of the section, close to Sect. Trifidae and Sect. Anagospermae is misconceived. The “E. striata line” shows little evidence of active speciation. Of the three distinctive vicarious species, E. striata occurs on mountain tops in the subalpine zone, while the other two occur near sea level, with E. semipicta in coastal heath. Data are required on the morphology and ecology of the unnamed Southport species, but it is possible that E. semipicta is derived from hybridization with E. collina (p. 105). 61 J. Adelaide Bot. Gard. 5 (1982) W. R. Barker E. gibbsiae “2 zg 8 B 8 3 iS} vy Be} S S £ = gE S$ = Soy aes SS > eee eS eS lel So Hee EHS 3 = . o as So S < = ed = a) Q aS) FE 5 i= 3 S = tS 3 & s 8 8 i 25 mS 2 = S = Ss 2 = $ nee} = g& Saw pln oa a GS guy Cee ett anh ee OY » Chea atts \ ; 1 i “ ' Branching H i basal Branching y I 1 ! Plant i 4 base ' Branching glandular H basal. hairy ! Leaf teeth } Chon Corolla Glandular sub- | hairs long striated! ! Branching ‘ 1 B’ H ' 'Bract and 1 calyx Leaf and ! margins bract margins 1 woolly greatly reflexed, ‘eglandular teeth many hairy Plant glandular hairy B” Fig. 24. Evolutionary diversification of Euphrasia in Australia (2). Postulated evolution of the species and subspecies of Sect. Striatae, showing some derived states at the branch points. (* indicates a taxon possibly derived by hybridization with Sect. Australes; A is a possible alternative position for the origin of E. gibbsiae ssp. wellingtonensis; B’ and B” are alternatives for the origin of E. hookeri.) The former existence of a lineage of Sect. Striatae in the eastern highlands of Victoria is discussed on p. 64. Evolutionary line 3: Sect. Australes - Sect. Lasiantherae - Sect. Scabrae (figs 25, 26). The taxonomic, morphological and ecological diversity of this natural group of sections is probably greater than any comparable group in the genus. It extends across temperate southern Australia, ranging from alpine fjaeldmark to semi-arid mallee communities, and from montane Nothofagus rainforest to coastal cliffs. A relict of the early phase of diversification of the genus, EF. bella of the perennial section Australes, is considered closest to the progenitors of the group. With two other morphologically distinct and geographically isolated species, E. sp. ‘Tamworth‘ and E. bowdeniae, it forms a vicariant progression in montane eastern Australia (fig. 49). This lineage may itself be old. From their poor collection, the species are probably very rare, and with E. bella and E. bowdeniae being apparently confined to cliffs, they give the impression of struggling to survive under present conditions. The apparent paraphyletic relationship of this group of species with the rest of the section is indicated on p. 57. 62 3 E. crassiuscula E. collina 8 [eee ee i « i] x x = er Sj 3 NI Q ~ 3 Sy 3 ost ~ ~ s 3 S a = ma 3 2 2 aS) xs a S oN sh Sas SF oS Gs Roo fy Ga? ga en eg Se S FS = = ep ee SS 4 s 8 = S > So &. = 2 S mB OS 5 & co 5 8 Bs = R & 3 8 S 8 8 2 3 & x & S 2 3 os) 5 ei Sf ESS Beas PS 8 Se a Re a 2 = OS = iS} g SS § = S$ 2 3 ies Fc tee Si Sige re genes So ED Qa eS! ae \ \ Inter- Plant Branching ae Be nodes glandular basal eglandular- aerial) short. ' scabrous Seeds Corolla hairy Plant base small sub-striated Capsules Rear of glandular Leaves Anthers SHAMS usually corolla hairy. Leaves few-toothed sub-glabrous hairs densely lobes : 1 glabrous many-toothed ong setose Rear Leaves Plant \ anther few-toothed glan Ry é awns Upper hairy 8 parts Plant glandular @@ Rear anther glandular hairy. f awns very short ‘ hairy Pedicels pers ' Leaves wy long i ae . few-toothed Branching argins o: 1 bracts and ES usually basal calyces woolly eglandular hairy ij Annual Corolla non-striated Leaves: base sessile, 3-5 or more veined. Inflorescences dense. Ovules many Fig. 25. Evolutionary diversification of Euphrasia in Australia (3). Postulated evolution of the species and subspecies of Sect. Australes, showing some derived States at the branch points. (* indicates a taxon possibly derived by hybridization with Sect. Striatae.) J. Adelaide Bot. Gard. 5 (1982) 63 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) The bulk of the remaining taxa of Sect. Australes belong to the highly polymorphic E. collina which encompasses almost the whole ecological and geographical range of the genus in Australia. Morphological differences from the E. bella - E. bowdeniae lineage, may reflect its adaptation to seasonal conditions in sclerophyllous and alpine habitats and to pollinators possibly not available to its progenitors. A capacity for more vigorous growth in deeper soil and more open vegetation may have contributed to development of a more robust habit, and an increased seed production by each plant in response to seasonal drought or snow through a greater ovule number and the production of an abundance of flowers. : A number of lineages are evident in E. collina, which all centre on a group of subspecies occupying the sclerophyll forests of south-eastern Australia. It is proposed that the species arose in this habitat, possibly from a stock resembling ssp. paludosa. The present lineages appear to be derived from a series of episodes of migration in the Plio-Pleistocene glacial periods (using in some cases exposed land bridges into Tasmania and Kangaroo Island) and contraction in the interglacial periods as the forest and subsequently subalpine and alpine habitats spread and contracted. Disjunct and vicarious distributions in the present-day ranges of the subspecies (tables 7, 8) are evidence of this. At all stages new ecotypes would have been able to develop sympatrically, as seen today in the subspecies on Mt Kosciusko and in the local races of ssp. diemenica. In addition, during the migrational episodes, morphological divergence may have occurred on a geographical basis, as seen today in the geographical clines of ssp. tetragona and ssp. collina, which through extinction of intermediate forms in the contraction episodes, would enhance the creation of isolated variants. The isolated populations on mountains of eastern Victoria which, in a stepped morphological transition, connect the widespread ssp. paludosa and ssp. speciosa with ssp. diversicolor of the Mt Kosciusko region of southern New South Wales (p. 168), have arisen apparently in this way, as may have the vicarious species of Sect. Lasiantherae in the same region (see below). The origins of the one remaining species of Sect. Australes, E. crassiuscula are problematical, for it possesses the characteristics of both Sect. Striatae and Sect. Australes. Two subspecies are variable in both characters separating the two sections. Ssp. eglandulosa more commonly possesses those of Sect. Striatae and in keeping with the climatic range of this section, occupies a higher altitudinal range, while ssp. crassiuscula more frequently has the character states of Sect. Australes, and, consistent with this, occupies a lower range.* The third subspecies, ssp. glandulifera, invariably has the character states of Sect. Australes. E. crassiuscula is restricted in eastern Victoria to the Bogong-Hotham mountain complex with an outlier on Mt Buffalo. It is proposed that the species evolved from the original sclerophyll forest stock of E. collina or from subalpine populations of ssp. paludosa with which it is parapatric, and that its characteristics of Sect. Striatae have been acquired through introgression, possibly from an extinct lineage, of which E. gibbsiae ssp. subglabrifolia of Mt Baw Baw may be the sole mainland remnant, and which extended along the higher parts of the eastern Victorian highlands in a former glacial period. The three species of Sect. Lasiantherae, linked by their common attribute of striated corollas, form a vicarious evolutionary series in the Australian Alps (figs 26, 86) possibly derived from the progenitors of the ssp. paludosa-ssp. speciosa-ssp. diversicolor lineage of E. collina, to which the perennial E. Jasianthera is the most closely related. The other two species, being annuals, are clearly derived. Because the * Sect. Striatae occupies mainly alpine and subalpine environments, extending into lowland areas only in the extreme south of Tasmania where similar conditions occur. Sect. Australes also extends into subalpine and alpine habitats, but generally occupies lower (milder) altitudes when the two sections are sympatric. 64 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia differences between the species are very large in comparison to other allied species in Euphrasia, divergence may have occurred over a relatively long period. The origins of Sect. Scabrae, a group of five robust annual species, are a little uncertain but it seems most likely that it was derived from the same stock from which particularly E. eichleri and E. alsa of Sect. Lasiantherae were derived (fig. 26). E. caudata (Sect. Scabrae) and E. alsa show strong similarities and occur sympatrically in the Australian Alps, with the former occupying montane and subalpine habitats, the latter alpine, rarely subalpine locations. E. scabra presumably spread during a glacial period across southern Australia, including south-west Western Australia and Tasmania. Relictual populations of its progenital links with E. caudata may still be evident in the Australian Alps (p. 276). The section has radiated in the northern and central New South Wales tablelands. A linking species may have occurred in the Blue Mountains (p. 284). Two lineages are apparent in this radiation, and it is likely that E. orthocheila ssp. peraspera, which shares derived characters of either line, has originated from hybridization between E. orthocheila ssp. orthocheila and E. ciliolata. Sect. Scabrae Sect. Lasiantherae ey pe E. orthocheila = 2 S 8 = = ~~ cS o. ”n an 2 Sect. Australes, p.p. E. scabra E. eichleri E. lasianthera M=-- SSp. peraspera E. ciliolata E. arguta E. caudata E. alsa Plant small. Leaves broader, Upper leaves Upper leaves fewer toothed. few-toothed few-toothed Floral parts smaller Branching basal. Anther awns long Annual, Ovules few hairs tiny or absent Upper corolla lobes facing sideways. Seeds small emarginate Corolla non-striated Fig. 26. Evolutionary diversification of Euphrasia in Australia (4). Postulated evolution of the species and subspecies of Sect. Lasiantherae and Sect. Scabrae, showing some derived states at the branch points. 65 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) H. EUPHRASIA—AN OLD TERTIARY GENUS? The following facts are considered to lend support to Euphrasia existing in the early Tertiary or before. 1. The bihemispheric distribution of the genus is matched by a number of homologous distributions, including both primitive groups, e.g. Magnoliaceae-Winteraceae, Fagus- Nothofagus, Cunoniaceae-Dilleniaceae, and advanced groups, e.g. Veronica-Detzneria- Hebe-Parahebe-Chionohebe alliance of the Veroniceae (van Steenis 1971, 1979). The Nothofagus-Fagus generic pair is particularly remarkable for its close-matching range. Van Steenis (1971, 1979) concludes that these common patterns have arisen through an ancient synchronous development and migration across land. Like many of these groups, there is a division in Euphrasia about “Wallacea”, for the northern and southern representatives of the genus are divided on a single character of stigma size, and it is possible that Sect. Malesianae and Sect. Pauciflorae are convergent reduced woody forms. Certainly the oddly hooded leaf of the New Guinea species is not evident in the west Malesian members, and Sect. Euphrasia and Sect. Atlanticae are morphologically more isolated from the southern sections than the latter are from each other (fig. 20). 2. As has been demonstrated, there is no evidence that the genus is capable of long- distance dispersal. Much evidence points to the need for migration across land. 3. Bartsia, the close generic relative of Euphrasia (Hartl 1972) has a somewhat complementary range; it occupies the mountains of tropical Africa (c. 8 species) and the northern and central Andes (c. 30 species) (fig. 21; Hedberg 1957; Hartl 1969-72). Diploid populations of B. alpina occupy the European Alps, with derived tetraploids and hexaploids occurring across formerly glaciated areas of the arctic regions. No species cross between the various continents. The two genera are the only members of Trib. Rhinantheae which extend into the tropics and the southern hemisphere (fig. 21). Just as with Euphrasia, the disjunct range of Bartsia appears difficult to explain by long-distance dispersal since the Pliocene. 4. The proposed relicts of the earliest phase of evolution of Euphrasia are widely scattered and often very restricted in their range (fig. 21). 5. At least two relicts show-association with Norhofagus forest. E. cuneata of New Zealand is polymorphic and extends into other communities from sea level to subalpine tracts, while £. bella is known only in association with Nothofagus, but is poorly known. This link with Nothofagus is important in view of the late Cretaceous age of the genus (e.g. Muller 1970, 1981; van Steenis 1971; Raven 1979) and the similarity of the range of the Fagoideae and Euphrasia (see point 1). 6. At least some of the relicts occupy habitats which conform with conditions expected between the late Cretaceous and mid Tertiary. The early angiosperms are believed to have radiated into niches unoccupied by the dominant gymnosperms (Doyle & Hickey 1976; Doyle 1977). Doyle specifically mentions the forest floor, a habitat in which I found E£. cuneata growing, with little other ground cover under Nothofagus mixed with other species near Wellington, New Zealand. E. bella, E. phragmostoma, E. cuneata, E. grandiflora (Wettstein 1896) and E. formosissima (Skottsberg 1921) are all known from cliff-faces, with the first two possible confined there. Such a niche might well have been occupied by angiosperm shrubs and woody herbs in the late Cretaceous. 7. The relicts in Euphrasia possess leaves with a broad blade and poorly differentiated petiole, again characteristics of early angiosperms (Doyle 1977; Doyle & Hickey 1976). Euphrasia and other Scrophulariaceae have tricolpate and tricolporate types of pollen; 66 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia such generalized types were already known from the late Cretaceous (Muller 1970, 1981; Doyle 1977). 8. Other groups of Australasian Scrophulariaceae show widely disjunct ranges difficult to explain by recent dispersion either by long-distance dispersal or across-land migration. The Veroniceae are similar to Ewphrasia in their bihemispheric distribution (see point 1), and their southern representatives showing great diversity and including both primitive and derived attributes (Ehrendorfer 1971; Wardle 1978). The two other major extra-tropical groups, Stemodia and the Mimulinae, in particular the group of Australian-centred genera Elacholoma, Peplidium, Microcarpaea and Glossostigma, have disjunct trans-Pacific connections which may be as ancient (Barker, in press). To these groups can be added Ourisia and Jovellana which are the only representatives of their South American-centred tribes to extend from South America into Australasia and New Zealand, respectively. No wide-ranging species are involved, and their distributions may date back to when links across Gondwanaland existed. The ranges of Gratiola and Limosella remain of doubtful age as these groups need revision and they may include wide-ranging taxa which are associated with mud and are of indefinite evolutionary advancement within the genera. For Euphrasia to have existed as far back as the late Cretaceous, not only must other representatives of the family have existed at that time, but so must have other advanced angiosperm families. A number of these families are cosmopolitan and, like Scrophulariaceae, should be considered in terms of infrafamilial taxa. Others, although small, nevertheless contain distribution patterns with wide disjunctions (e.g. Bontia in Myporaceae, Mr R.J. Chinnock, pers. comm. 1980; Josephinia in Pedaliaceae). The fossil record as yet contains no sufficiently authenticated Tertiary Scrophul- ariaceae, although (Muller 1981) there are unsubstantiated records from the Paleocene in England and Pliocene (Hebe) in New Zealand. The earliest identified pollen of the Scrophulariales is from the Eocene; most families date from the Miocene or more recently (Muller 1970, 1981). An advanced group such as the Compositae dates from the Oligocene. How can these groups be much older? There is a sampling bias in the fossil pollen record. The chances of including plants which produce relatively little pollen at a given site are severely limited. Pollen production may be restricted by small population size, a limited seasonal production of flowers, and by efficient pollination mechanisms. All these attributes apply to most advanced angiosperms which are herbaceous and bee-pollinated. Furthermore, the ecological range of a plant group significantly affects its chances of turning up in the fossil record (Martin 1978; Truswell & Harris, in press; Smith- White, in press). Plants such as bee-pollinated types, with a limited range of pollen dispersal, would have to have grown in or close to fossil-forming conditions, such as swamps, lakes or peat-forming sites, for pollen to be deposited in sufficient quantity there. The proposed relicts of Euphrasia rarely, if at all, grow in such sites, for below the tree-line their scattered populations may occur in Nothofagus forest which occurs on ridges or slopes, or they may grow on cliff-faces. The poor coverage of the fossil record is demonstrated in several ways. It is deficient in transitional forms of pollen linking for example the elaborate and diverse types in the Acanthaceae to simpler more primitive types (Muller 1970). Furthermore, for a continent such as Australia, the total of about 50 taxonomic identifications for the Tertiary, some only to family (Martin 1978), indicates a very sparse knowledge of the flora of that period. The discrepancy between the macrofossil and pollen records at given sites also reflects this (Smith-White, in press). Finally, Truswell & Harris (in press) refer to the substantial amounts of fossil pollen belonging to generalized types 67 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) which are able to be assigned to a number of modern families. The pollen of Euphrasia is of one such type (Mr W.K. Harris, pers. comm. 1979). Michener (1979) proposes that bees existed in the latest Cretaceous, when pre- dominantly bee-pollinated groups such as //ex and Myrtaceae are first recorded as fossils. The earliest fossil bees, including the most advanced group, the long-tongued Apidae, are known from Eocene deposits in North America and Europe, and several widely disjunct ranges of distribution of bee genera are inexplicable on long-distance dispersal and the land connections available from that time. To accommodate these facts, Michener postulates that bees arose in the mid Cretaceous and became abundant in the late Cretaceous in the xeric interior of Gondwanaland, a view supported by Raven (1979). A radiation of bees in the late Cretaceous must have been paralleled by a radiation of the plants they pollinated. In other words, advanced largely bee-pollinated groups such as the Scrophulariales and Compositae may well have attained their family and much of their generic make-up during that time. Michener (1979) suggests that the bees diversified in the xeric interior of Gondwanaland, in particular West Gondwanaland where (Raven 1979) the angiosperms are considered to have evolved. In such conditions there is little likelihood of gaining direct fossil evidence for either the bees or the flora. Accordingly, it is suggested that the bee-pollinated groups radiated in similar conditions in the late Cretaceous, but had only limited success invading valley forest or swampy sites until their explosive radiation in Neogene times, linked with newly formed alpine and subalpine sites and the large fluctuations of world climate which favoured diversification of such herbaceous groups. I. CONCLUSION It is accordingly proposed that the present day range and diversity of Euphrasia has been attained in two main phases. Firstly, the genus evolved with other bee-pollinated groups, possibly under seasonally dry or xeric conditions in the late Cretaceous in Gondwanaland. At this time suitable migration routes were available into all continental regions, including the present northern and equatorial land masses, whether via Africa alone (Raven & Axelrod 1972; Raven 1979) or also via a subsequently severed link with south-east Asia (van Steenis 1979). Relictual forms, possibly closely allied to early Euphrasia, are scattered in the northern and southern temperate zones (fig. 21). A route via Africa would accommodate the occurrence of Euphrasia in the Azores and the allied genus Bartsia in the Andean and equatorial mountains of Africa (fig. 21), together with a division of Euphrasia into northern and southern hemisphere groups about “Wallacea” (p. 55). The similarity of the sessile gland patterns of the Azorean species and Bartsia (p. 79; Yeo 1973) provides confirmation of this. Until the Miocene, Euphrasia may have existed in temperate or subtropical conditions on the forest floor or in open sites such as cliff faces, habitats occupied by the proposed relicts today. Indeed, the semi-parasitic habit of Subfam. Rhinanthoideae of the Scrophulariaceae may have developed in response to seasonally dry conditions or to poor water retention by shallow soil in cliff face habitats. Flowers apparently pollinated by a more general group of insects (p. 44) or by Lepidoptera (p. 45), represented today in New Zealand and South America, possibly developed also at this time in habitats rarely frequented by bees. Michener (1979) proposes that the temperate forest which covered much of Gondwanaland was occupied then, as today, by few bees, New Zealand’s depauper- ate bee fauna being explained by an almost complete forest cover at the time of its separation from Gondwanaland about 80 million years ago. However, Smith-White (in press) questions the reality of this broad Gondwanan forest cover as being an 68 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia artefact of a deficient fossil record. The continued existence of apparently functionless striations on the rear of the corolla limbs of some New Zealand species of Euphrasia (p. 44) may point to a relatively recent demise of bees there. The uplift of mountains and fall in world temperatures in the late Tertiary apparently heralded a second phase of evolution in Euphrasia. Euphrasia was one of the groups, already of predominantly herbaceous or shrub habit, which migrated into newly formed subalpine and alpine regions. During the climatic fluctuations of the Plio-Pleistocene period, the genus radiated into open communities through its capacity to form ecotypes. Many old forms of Euphrasia became extinct during this general phase of diversification in the angiosperms, for it has been proposed that only a few scattered relicts remain. In Australia, the newly diversifying sclerophyllous communities dominated by Eucalyptus which may have developed from late Pliocene times at the earliest (Truswell & Harris, in press) provided an opening for further radiation of Euphrasia, while in Europe the genus spread into man-made grassland and other communities which developed during the evolution of man’s agricultural practices (Karlsson 1974, 1976; Hulten 1976). The new habitats produced different demands on the morphology of the plant. In alpine conditions vegetative buds were limited to ground level, while pressure of grazing and mowing and/or shortness of growing seasons suppressed plant height and possibly the development of annuals. Reduction of the petioles may have given added strength to the leaf in exposed sites. To what extent floral form and coloration diversified from the late Tertiary in competition with the abundant herb and shrub floras for the available pollinators can only be speculated upon. The flower type restricted to Sect. Trifidae of South America and Sect. Anagospermae of New Zealand and that of Sect. Scabrae are not represented among the proposed relict species. However, if the first two sections are monophyletic (p. 55), it is likely that their flower type evolved during the early phase of generic diversification owing to their occurrence on either side of the south Pacific. Today, in a region such as Australia where the vegetation has altered radically in the past two centuries, we see a third phase. Euphrasia was able to co-evolve with the relatively slow spread of man’s influence in Europe during the Quaternary, but it has been incapable of coping with the rapid advent of European man in Australia and the associated pressures of clearing, ploughing and grazing. As a result many taxa are now rare and at risk. Some, indeed, may have become extinct. IV. A REVISED INFRAGENERIC CLASSIFICATION OF EUPHRASIA A. PREVIOUS INFRAGENERIC CLASSIFICATIONS Although a number of botanists have discussed the infrageneric classification of Euphrasia, only Bentham (1846), Wettstein (1896), and Hartl (1972) have dealt with the classification on a world-wide basis. The two earlier workers are the only botanists to revise the entire genus. Except for changes in rank, the addition of two small morphologically and geographically isolated infrageneric taxa, and the segregation of two monotypic genera in New Zealand, which were soon reduced to synonymy, a basic framework of three large infrageneric taxa has been maintained since Bentham’s time. Bentham (1846) divided the genus into three categories of equal status, but with no clear indication of rank (see Sell & Yeo 1970). His S Semicalcaratae, characterized by “Antherae pilosae, mucronatae, duarum breviorum posticarum loculus alter longius calcaratus... Folia inferiora crenata, floralia summa saepe acutius incisa vel dentata”, 69 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) was confined to the northern hemisphere except for one species in New Zealand (E. cuneata). S Australes, distinguished by its ““Antherae pilosae, omnes subaequaliter mucronatae . . . Folia apice paucicrenata rarius fere a basi crenata”, was restricted to “Australasia” (which is probably Australia, as he cited only Australian species and localities). S Trifidae of South America, with one questionable record from the Himalayas, differed by its “Antherae glabrae, aequaliter mucronatae . . . Folia sessilia cuneato-trifida vel tripartita”. Of the Himalayan plant resembling the South American species in habit and leaf shape, Bentham saw only one corolla and was doubtful of its placement even within the genus (he wrote “E. ? glandulosa”). The specimen was later placed by Bentham & J.D. Hooker (1876) in the related genus Phtheirospermum. J.D. Hooker (1879) described a new species, E. disperma, from New Zealand which, together with E. repens, he placed in a new subgenus Anagospermae. The subgenus, characterized by its 2-4-ovulate ovaries, solitary flowers, prostrate habit and remarkably long corolla tube, was possibly a distinct genus providing the capsules were found to be indehiscent. Two years later on the basis of meagre material containing a single flower, Armstrong (1881) described a new genus, Siphonidium, from New Zealand, closely related to Euphrasia but differing “in the long curved gibbous tube and bi-lobed stigma”. He did not mention Hooker’s subgenus even though the two descriptions are similar. Wettstein (1895), having seen fruiting material of what he considered to be E. disperma, elevated Hooker’s Subgen. Anagospermae to a genus: “..... differt ab Euphrasia tubo corollae elongato, labio corollae superiore non excavato, antheris patentibus et glaberrimis, imprimis vero loculis germinis uniovulatis et fructu bicorni bispermo. A Siphonidio Armst. differt tubo corollino recto, stigmate non bilobo, probabiliter etiam fructu et germine”. At length the status of these genera was questioned. Certain that Anagosperma and Siphonidium were identical, Cheeseman (1925) united them under the latter, earlier- published name. Then, in his detailed account of “The long-tubed New Zealand species of Euphrasia (= Siphonidium Armstr.)” Du Rietz (1932a) showed that: “there is such a gradual transition between Siphonidium and Euphrasia sens. strict., that it is not possible at our present stage of knowledge to retain Siphonidium as a separate genus. . . it seems even doubtful whether Siphonidium should be retained as a subgenus or section”. Wettstein (1896) saw much more material of the extra-European members of Euphrasia than Bentham, but like his fellow-European botanist, he knew nothing apart from a Bornean record in his addendum, of the Taiwanese and Malesian species. He saw material of most of the New Zealand species, whereas Bentham apparently saw only E. cuneata. His major classification of the genus was similar to Bentham’s using identical characters. He treated S Semicalcaratae (which included all the northern hemisphere occurrences in the genus) and S Australes (comprising all Australian and New Zealand species, including E. cuneata) as subsections of Sect. Eueuphrasia, Bentham’s S Trifidae excluding the Himalayan species, was treated as a section. Sect. Trifidae and Subsect. Australes were divided into annual and perennial groups which were not given formal taxonomic recognition. Wettstein also divided Subsect. Semicalcaratae into three unranked groups, § Parviflorae, § Grandiflorae and § Angustifoliae on the basis of differences in the time of elongation of the corolla tube relative to anthesis, the breadth of the leaves and the indumentum of the capsule. Subsequent European botanists (e.g. Joergensen 1919; Pugsley 1930, 1936; Sell & Yeo 1970; Yeo 1972) have considered the character of the elongation of the corolla tube to be of lesser, or even doubtful, diagnostic use. The upgrading by Joergensen (1919) of Wettstein’s sections and subsections to subgenera and sections respectively has been followed in all subsequent works on the genus up to Hartl’s (1972) publication (see below). Joergensen’s recognition of two 70 J. Adelaide Bot. Gard. 5 (1982) ; Studies in Euphrasia subsections in Sect. Eueuphrasia, namely Subsect. Angustifoliae (equivalent to Wettstein’s § Angustifoliae) and Subsect. Ciliatae (covering Wettstein’s § Parviflorae and § Grandiflorae) on the same characters of leaf shape and capsule indumentum used by Wettstein, has been universally accepted. Joergensen refined the character of leaf shape by recognizing the apparently reliable differences between the two subsections in the proximity of the teeth along the margins. Since Joergensen’s time Subsect. Ciliatae has been progressively divided into a number of series, especially by Pugsley (1930, 1936) and Sell & Yeo (1970). These series are distinguished by the combination of a number of often overlapping characters including habit, indumentum, leaf and bract shape, corolla size, and capsule size and shape. Pugsley (1936) erected two new subsections of Sect. Semicalcaratae, Subsect. Japonicae and Subsect. A/picolae, both confined to Japan, on the basis of habit, leaf shape, calyx shape, corolla size and coloration, and the indumentum on the corolla, anthers and capsule. In the same paper, Pugsley proposed two newsections. One Sect. Atlanticae, endemic to the Azores, was distinguished on the basis of its perennial life-span, rounded leaves and deeply emarginate capsules. Yeo (1972, 1973) has recently modified the diagnostic characters of this section, using its perennial life-span, more numerous leaf teeth, distinctive leaf indumentum and large flowers to distinguish it from the rest of the European and North American species. The distinctive capsule shape described by Pugsley for one species of the section was found not to occur in the other Azorean species. Pugsley’s other new section, Sect. Paradoxae, was based on a single species E. formosissima of the Juan Fernandez Islands west of South America. The section was characterized by a unique method of perennation and also by “entire (not emarginate) lobes of the lower lip of the corolla, . .. glabrate, unequally spurred anther-cells and... scarcely retuse, setulose-edged capsules”. Skottsberg (1921), who described the species, and Wettstein (1921) had previously remarked on its isolated morphological characteristics and its apparent affinities to the Japanese and Australasian members of the genus. There have been no major changes in the infrageneric classification of Euphrasia in the northern hemisphere since Pugsley’s works. Sell & Yeo (1970) have correctly called the section containing the type species of the genus Sect. Euphrasia in place of Sect. Semicalcaratae (Benth.) Joerg. In addition they have revised Pugsley’s classification of Subsect. Ciliatae at the level of series. Although there has been little change in the formal classification of the genus in the southern hemisphere and Malesia since Wettstein’s (1896) monograph, knowledge of the characters potentially useful in an infrageneric classification has been greatly increased by the work of Du Rietz (1932a,b; 1948a,b). Du Rietz never published his proposed paper (see Du Rietz 1932a, p.121) on the general subdivision of the genus. Considering the depth and breadth of his work he did little formal taxonomic work. He proposed on the basis of capsule shape two subsections, Subsect. Australienses and Subsect. Novaezeelandiae (Du Rietz 1948b), of (presumably) Sect. Australes, covering the Australian and New Zealand species respectively. He also divided the Australian species into four series, Striatae, Collinae, Hookeriae and Scabrae. For this he used characters of lifespan, habit and leaf shape (Du Rietz 1948a,b). In his critical discussions of the relationships between the species of Euphrasia in South America, the Juan Fernandez Islands, Australia, Tasmania, New Zealand, New Guinea, Borneo, the Philippines, Taiwan and Japan, Du Rietz discerned several characters to be useful in distinguishing groups of species. Among these were the pilosity and colour of the anthers, shape of the corolla lobes, leaf shape and habit. It is not clear 71 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) whether he recognized the importance of the free anthers characteristic of the South American species and E. disperma of New Zealand. His term “patent anthers” seems to refer more to the orientation of the cells of each anther in these species (see Du Rietz 1932a, p.118, especially in his comparison with f.1). In descriptions of the long-tubed species of Euphrasia in New Zealand (E£. disperma sensu Ashwin 1961) he referred to the anthers as “non cohaerentes”. Du Rietz also questioned usage since Bentham’s time of the length of the rearmost pair of anther awns relative to the other awns to distinguish between the species of the southern and northern hemisphere, but he still considered that “subequally mucronate anthers” were characteristic of all the South American species, some New Zealand species and E. striata of Tasmania (Du Rietz 1932b, p.532). Ashwin (1961) gave a synopsis of the main character differences in Euphrasia in New Zealand without proposing any formal infrageneric classification. She divided the perennial species into two main groups, one containing only E. cuneata and characterized by much-branched inflorescences and small bracts, the other with inflorescences simply racemose and with leaf-like bracts. This latter group, which contained the six remaining perennial species, was further divided on the indumentum of the calyx and the relative lengths of the calyx clefts. The annuals were divided into two main groups on leaf and habit characters, the species of the first group, E. cockayniana, E. zelandica, E. australis, and E. cheesemanii, having leaves of the “ovate order, crenate or toothed” and erect branches which never root, and those of the second group, E. dyeri, E. repens, E. integrifolia and E. disperma having mostly decumbent or prostrate branches, which sometimes root, and leaves either deeply divided or entire and of the lanceolate type. This second group was further split into three on the basis of the same characters of leaf and habit type. In 1972 Hartl revised the higher classification of Euphrasia on a world-wide basis. Of significance is his treatment of all Joergensen’s (1919) and Pugsley’s (1936) taxa of the rank of subgenus or section at the one level of section. Thus his major classification of the genus was composed of five sections. He followed Pugsley’s division of Sect. Semi- calcaratae (Hartl’s Sect. Euphrasia) into four subsections without alteration of the somewhat cbscure diagnostic characters of the Japanese subsections. Sect. Australes was expanded to include the Malesian and Taiwanese species, which had been previously unplaced in the infrageneric classification. He believed that both Sect. Australes and Sect. Trifidae could be subdivided into a number of subsections. B. THE REVISION 1. Introduction The proposed classification of Euphrasia represents a radical departure from the concepts in previous works, in which the genus has been divided into three major groups of varying rank, together with, from 1936, two peripheral sections. In the new classification, with the exception of Sect. Pauciflorae (2 subsections) and Sect. Euphrasia (the 4 subsections recognized by Pugsley being retained), the infrageneric groups recognized have been given equal status as sections (the many series in Subsect. Ciliatae have not been considered). Each section is separated by only one or two distinct characters from at least one other section. However, sometimes there is quite remarkable divergence of a transitional nature, away from the most closely linked species of related sections. There are a number of differences in usage of characters from previous classifications, and several new diagnostic characters have been discerned. The characters are discussed in detail in the chapter dealing with morphology. The very obvious free anthers of the South American species and E. disperma of New Zealand have surprisingly never been used diagnostically before, except in the distinction of E. disperma from its related 72 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia species (Ashwin 1961). The presence or absence’ of corolla striations, the presence or absence of hairs on the back of the anthers, the number of main veins arising from the very base of the leaf, the distinctive corolla shape of Sect. Australes and especially Sect. Scabrae, and the size of the stigma are all characters which have previously not been used in the infrageneric classification of Euphrasia. Some of the characters, however, have been used to a small extent at the species level. Other characters have been refined in their definition. The distinction between “petiolate” and subsessile leaves used by Pugsley (1936) for E. formosissima and Ashwin (1961) for E. cuneata has not been considered an accurate statement of the differences. In this work the shape of the leaf base has been described adhering to a rigid terminology, and its diagnostic value is not in the areas used previously. The depth of the leaf toothing has been treated differently from the rather vague terminology used by Du Rietz (1948a,b) in which he separated “digitate” and “subdigitate” types from the rest of the genus. The “blade” (defined on p. 14) is either small (little of the area within the leaf outline) or large (much of the area within leaf outline). Two characters used in the past to distinguish infrageneric taxa have been found to be of little use at that level. Contrary to observations of all previous workers on the infra- generic classification, the rearmost pair of anther awns has been found to be consistently longer than the other six awns in all the material seen (p. 32). Similarly, Du Rietz’s opinion that the Australian and South American species have a unique “acuminate” capsule shape is a misconception (p. 34). 2. Revision The extra-Australian species cited in the text are. those recognized in the most recent regional treatments of the genus. These are Wettstein (1896) and Reiche (1911) for South America, Ashwin (1961) for New Zealand, van Royen (1972) for New Guinea, Du Rietz (1932b) and van Royen (1971) for Malesia, Ohwi (1933) and Li (1950, 1978) for Taiwan, Skottsberg (1921) for the Juan Fernandez Islands, and Yeo (1972, 1973) for the Azores. The many species in the remaining regions of the northern hemisphere have not been listed in this conspectus. Where herbarium collections which are not types of a species have been seen, the species is designated by “!”. If type material has been seen this is designated by “T”. Sections and subsections are ordered in a sequence which as far as possible places closest allies adjacent, commencing with the autonymous taxon. EUPHRASIA L. Sp. PI. (ed.1) (1753) 604, p.p. (see Typification); L., Gen. Pl. (ed.5) (1754) 263, p.p.; Benth. in DC., Prodr. 10 (1846) 552, p.p. (excl. “E. ? glandulosa”); Wettst. in Engler & Prantl, Nat. Pflfam. IV3b (1893) 100; Wettst., Monogr. Gatt. Euphrasia (1896) 9; Hartl in Hegi, Ill. Fl. Mitteleur. (ed.2) 6 (1) (1972) 335. Lectotype (Britton & Brown 1913): E. officinalis L., s.str. (syn. E. rostkoviana Hayne). See Typification. Siphonidium Armstr., Trans. Proc. N.Z. Inst. 13 (1881) 341 (see p. 88). Anagosperma (Hook.f.) Wettst., Ber. dtsch. bot. Ges. 13 (1895) 242 (see p. 88). Annual or perennial terrestrial herbs or undershrubs; single main root branched laterally, the lateral rootlets often connected to the roots of other plants by haustorial swellings. Axes with indumentum consisting entirely or partly of eglandular hairs which are often in two bands or four lines decurrent from between leaf bases, sometimes all around, sometimes lacking in lower parts. Cotyledons entire, fleshy, glabrous. Leaves decussate, appressed to axis at base, usually then recurved, fleshy but usually brittle, rarely (E. bowdeniae) pliant when dry; abaxial surface usually with patches of sessile glands symmetrical about midrib, confined to marginal rows or extending over most of blade between veins, rarely (Sect. At/anticae) with sessile glands confined to veins; 73 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) margins somewhat revolute, usually shallowly to deeply incised, rarely entire; main veins prominent on abaxial surface, submerged below adaxial surface and topped by narrow grooves. Inflorescences simple, terminal, sometimes spike-like racemes, or consisting of solitary flowers. Bracts similar in morphology to leaves immediately below inflorescence, but often differing slightly in shape and indumentum. Calyx zygomorphic, campanulate or + cylindrical, slightly recurved, 4-lobed, with median clefts equal to or longer than lateral clefts. Corolla bilabiate, with tube proximally cylindrical, distally expanded and then divided into hooded upper lip, which usually encloses the anthers and is terminated by two abruptly reflexed lobes, and a three- lobed lower lip; with at least outer surface and front of inner surface of hood pilose. Stamens 4, didynamous, the posterior pair inserted higher on the corolla tube than anterior pair; filaments straight or curved; anthers free, or fused to each other along margins into a U shape with the posterior pair free from each other, with each cell clavate, dehiscing introrsely by longitudinal slit which is widest towards anther base and terminated in sharp awn, the rearmost pair of awns longer than the other six awns. Gynoecium: ovary 2-celled, slightly compressed laterally, with a nectary at base on abaxial side; sry/e filiform, passing above or (sometimes when anthers free) between anthers, setose in upper half; stigma capitate, oblong or unequally bilobed; ovules 2-200. Capsule dehiscent loculicidally, with base of style persistent for short length; seeds 0-150, obliquely + ellipsoid, longitudinally ribbed or (E. azorica) winged, scalariform between, whitish. Chromosome number: variable (see infrageneric taxa). Typification Euphrasia L. Linnaeus (1753) included under the protologue of Euphrasia the currently recognized genera Odontites (his E. odontites, E. linifolia and E. lutea) and Parentucellia (his E. latifolia). The remaining two species in the protologue, E. officinalis and E. tricuspidata, have been shown to fall closest to Linnaeus’s (1754) generic description of Euphrasia (Pennell 1930), which must be taken into account for typification (ICBN: Art. 13.4, Art. 41), and fall into the limits of the genus in the strict sense in which it has been generally treated since 1830 (Pugsley 1930). The lectotypification of the genus began with Britton & Brown’s (1913) selection of E. officinalis. Despite the arbitrary nature of their choice, Pennell (1930) argued that this was the logical decision taking Linnaeus’s (1754) generic description of Euphrasia into account. The selection aiso accords with Linnaeus’s apparent generic concepts and methodology, for Stearn (1957) indicates that Linnaeus based his generic descriptions on the “best known and officinal plant”. According to Pugsley (1930) material of the two Linnean species of Euphrasia s.str. is contained in the Linnean herbarium (LINN), “the sheet of the former [E. officinalis] showing three specimens, of which two are the glandular E. rostkoviana Hayne, and the third* an eglandular form that has been referred to EF. nemorosa Pers. but is more probably the Scandinavian £. curta Fr. E. tricuspidata is represented by a single unmistakable example”. Sell & Yeo (1970) selected as lectotype of E. officinalis the element of the type which corresponds to E. rostkoviana. Because of the difficult taxonomy of the northern hemisphere annuals (Sect. Euphrasia), E. officinalis has often been used in a very broad sense encompassing many described species accepted as distinct by others, and it is for this reason that the name has been rejected as a nomen ambiguum in recent times (Smejkal 1963; Sell & Yeo 1970; Yeo 1972, 1978b). On this basis also Smejkal (1963) rejected the species E. officinalis as the lectotype of Euphrasia in favour of E. tricuspidata, but the decision was reversed (without discussion) in favour of the initial choice E. officinalis by Sell * Now considered to be E. stricta Wolff ex Lehm. (Yeo 1972). 74 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia & Yeo (1970), as lectotypified by them but with the name rejected in favour of E. rostkoviana. Article 69 of ICBN, however, places in serious question the rejection of the name E. officinalis as a nomen ambiguum, for it does not appear that the name has been used widely and persistently in a sense excluding its type, as is now required for rejection of a name under this article. Distribution (fig. 19): Euphrasia is one of the few groups of rank higher than species with a centre of diversity in the temperate zone of both hemispheres and a connection through Malesia. Euphrasia is usually a microtherm (cold-inhabiting) genus spread widely in the northern temperate zone, with outposts in the Azores and North Africa. Its southern temperate distribution is in southern Australia, New Zealand and southern South America, including the Juan Fernandez and Falkland Islands. These zones of distribution in either hemisphere are connected by a relatively continuous series of localities on the highest mountains of Taiwan, the Philippines, Borneo, Celebes, Ceram and New Guinea. The genus is divided in this revision into 14 sections. Twelve of these are well- defined homogeneous groups (Sect. Pauciflorae being subdivided into two subsections), and constitute all representatives of the genus between Taiwan and the Juan Fernandez Islands, as well as the distinctive species of the Azores. Together they comprise 59 species, although future revision adopting the species concepts of the present Australian revision, particularly in Sect. Pauciflorae and the Taiwanese species of Sect. Malesianae, may reduce this number to about 50. Comparable numbers of subsections and species in the remaining two sections, Sect. Euphrasia and Sect. Trifidae, are difficult to estimate. While constituting natural groupings, they require revision at both the infrageneric and species level. There have been recent revisions at the species and series level in Sect. Euphrasia in parts of its wide range, for example for Europe (Yeo 1972, 1978b), North America (Sell & Yeo 1970), Russia (Juzepcuk 1955), China (Li 1953), Western Himalayas (Pennell 1943), and Japan (Kimura 1941, 1948; Ohwi 1965), but as the species concept has been very narrow and there are a number of widely distributed species (Sell & Yeo 1970; Hulten 1976), a revision encompassing the whole section seems desirable. This would also provide the basis for a review of the infrasectional taxonomy, in particular in relation to the segregation of the two Japanese endemic subsections (see Sect. Euphrasia: note). Sect. Trifidae has not been revised since Wettstein’s (1896) monograph of the genus and Reiche’s (1911) reappraisal of the Chilean representatives. The section shows diversity in duration (both annuals and perennials) and floral morphology (pp. 28, 45), characters of potential importance at the sectional or subsectional level. KEY TO THE SECTIONS AND SUBSECTIONS OF EUPHRASIA la. Anthers, at least posterior pair, hairy about connectives. Leaves usually truncate to cuneate at base, rarely (£. bella) attenuate, with (1)3-7 main veins arising from base. 2: TOCTCTE GTEC oi sco oopobyoanudnobondntosacdocasunvegenmuom IX. Sect. Lasiantherae (p. 85) 2b. Corolla lacking striations. 3a. Perennial. Branches on main inflorescence-bearing axis developing in no fixed sequence, not consistently in consecutive nodes if high above ground level. Upper corolla lobes + coplanar, facing forward ..............+..4- VIII. Sect. Australes (p. 84) 3b. Annual. Branches on main inflorescence-bearing axis developing basipetally in consecutive nodes high above ground level from I-few nodes below inflorescence. Upper corolla lobes usually angled sharply to each other, facing to side. MCUs O POs Ba bedArbo onc soh dpbocasupuDh contopgdune sano X. Sect. Scabrae (p. 86) W. R. Barker J. Adelaide Bot. Gard. 5 (1982) 1b. Anthers glabrous about connectives. Leaves attenuate at base, with (1)3(5) main veins arising from base. 4a. Anther slits glabrous or with | or 2 tiny hairs along margins. 5a. Perennial. 6a. Leaves shallowly lobed, with (3)4(5) pairs of teeth. Anthers fused, with slits very sparsely hairy. Corolla lobes obtuse. ............. XII. Sect. Paradoxae (p. 88) 6b. Leaves deeply divided with 1(2) pairs of teeth, Anthers free, with slits glabrous. Corolla lobes emarginate or shallowly so. ..... XIV. Sect. Trifidae (p. 89) 5b. Annual. 7a. Leaves crenate to serrate. [Anthers fused.]........ XI. Sect. Novaezeelandiae (p. 87) 7b. Leaves pinnatifid, trifid or entire. 8a. Main inflorescences with more than 10 flowers. Anthers free. Plant erect, with uppermost leaves of main inflorescence-bearing axis with 1(2) pairs ofiteeth. ......0 0. esse ces eeeesesere ress XIV. Sect. Trifidae (p. 89) 8b. Main inflorescences with less than 10 flowers, or flowers sporadic along axes. Anthers fused or free. Plants either erect with uppermost leaves of main inflorescence-bearing axis with 1-3 pairs of teeth or prostrate with leaves entire or with | pair of teeth........ XIII. Sect. Anagospermae (p. 88) 4b. Anther slits distinctly hairy. 9a. Stigma (0.2)0.3-0.5mm long or longer. Corolla lobes emarginate to obtuse or acute. t LOateA nmitial frees tem eretsrses trelses ot stakstotetst stots ylot=svee] fete otette g XI. Sect. Novaezeelandiae (p. 87) 10b. Perennial. lla. Inflorescence-bearing axes prostrate for entire length. Flowers sporadic along axes. ....... VI.B. Sect. Pauciflorae Subsect. Humifusae (p. 83) 11b. Inflorescence-bearing axes, at least in distal parts, erect. Flowers in terminal inflorescences. 12a. Rearmost pair of anther awns (1.5)2.5-3.2mm long, as long as or longer than anthers, sometimes needle-shaped and entire, some- times distally flat and twisted or erosulate. [ Branches or shoots on main inflorescence-bearing axes at widely spaced groups of nodes, developing within groups in basipetal sequence in con- secutive nodes. Corolla lacking striations.| V. Sect. Phragmostomae (p. 81) 12b. Rearmost pair of anther awns to 0.8mm long, shorter than anthers, needle-shaped, entire. ' 13a, Plant tall. Branches or shoots on main inflorescence- bearing axis(es) developing in consecutive axils from I-few nodes below inflorescence in basipetal sequence. [Corolla HLA | bistbodd boasieopsonuormsuuotd .. IV. Sect. Cuneatae (p. 80) 13b. Plant short or tall. Branches or shoots on main inflorescence- bearing axis(es) developing sporadically in axils, often only near ground level, and in no fixed sequence. 14a. Flowers in main inflorescence 2-8(12). Corolla striated or lacking striations. .. VI.A. Sect. Pauciflorae Subsect. Pauciflorae (p. 82) 14b. Flowers in main inflorescence (7)10-24(36). Corolla striated, at least partially. ............ VII. Sect. Striatae (p. 83) 9b. Stigma 0.1-0.3mm long. Corolla lobes emarginate. 15a. Perennial. 16a. Uppermost leaves on main inflorescence-bearing axis(es) with (5)7-12 pairs of teeth. Corolla 13-17mm long along upper side.II. Sect. Atlanticae (p. 78) 16b. Uppermost leaves on main inflorescence-bearing axis(es) with 2-4 pairs of teeth. Corolla c. 6-10mm long along upper side. . III. Sect. Malesianae (p. 79) 15b. Annual. [Uppermost leaves on main inflorescence-bearing axis with (1)2-6(7) pairs of teeth. Corolla c. 3-13.5mm long along upper side.) .1. Sect. Euphrasia (p. 77) 76 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia I. Sect. Euphrasia Wettst. in Engler & Prantl, Nat. Pflfam. IV 3b (1893) 100, p.p. (excl. E. grandiflora and species from southern hemisphere) “Eueuphrasia”; Wettst., Monogr. Gatt. Euphrasia (1896) 68, p.p. (as to Subsect. Semicalcaratae) “Eueuphrasia”; Sell & Yeo, Bot. J. Linn. Soc. 63 (1970) 203; Hartl in Hegi, Ill. Fl. Mitteleur. (ed.2) 6 (1) (1972) 344; Yeo, Fl. Europaea 3 (1972) 260; Yeo, Bot. J. Linn. Soc. 77 (1978) 227. S Semicalcaratae Benth. in DC., Prodr. 10 (1846) 552, p.p. (as to E. officinalis and E. tricuspidata).—Subsect. Semicalcaratae (Benth.) Wettst., Monogr. Gatt. Euphrasia (1896) 68; Du Rietz, Sv. Bot. Tidskr. 42 (1948) 361.—Sect. Semicalcaratae (Benth.) Joerg., Berg. Mus. Aarb. 1916-17 Naturvid. raekke 2 (1919) 5, p.p. (excl. the Azorean representatives; as to lectotype) “Subgen. Eueuphrasia Sect. Semicalcaratae”; Pugsley, J. Linn. Soc. Bot. 48 (1930) 484; Pugsley, J. Bot. (Lond.) 74 (1936) 284. Lectotypus hic designatus: E. officinalis L. (as lectotypified by Sell & Yeo 1970: syn. E. rostkoviana Hayne, see p. 74). See Typification. Annual. Main inflorescence-bearing axis single erect stem, developing branches + baispetally in consecutive nodes from the node below the inflorescence. Uppermost leaves of main axis pinnatifid-serrate to crenate, with base attenuate, abruptly (Subsect. Ciliatae, 2? Subsect. Japonicae p.p.) or gradually (Subsect. Angustifoliae, Subsect. Alpicolae) expanded into large blade, with (1) 2-6 (7) teeth along distal c. 4; - 7, of each margin; with 3 main veins arising from base of leaf, branched distally. Flowers 4-50 in (main) racemes. Corolla striated, with yellow blotch on lower lip, or (in Subsect. A/picolae) with purple blotches behind upper lip and in tube, with lower side apparently concave from above, spreading from base of lower lip; Jobes emarginate. Anthers fused, glabrous around connectives, hairy along slits; awns entire. Ovary with stigma capitate, 0.1-0.3mm long. Capsules in lateral view usually emarginate, sometimes obtuse or obliquely so. Chromosome number: n=11, 22. Typification S Semicalcaratae Benth. E. cuneata of New Zealand is excluded from consideration as a possible lectotype since it was omitted by Wettstein (1896) and all subsequent authors from infrageneric taxa based on S Semicalcaratae. Similarly, E. grandiflora has not been considered in view of its segregation as a separate section since Pugsley’s (1936) work. E£. officinalis is chosen as lectotype as it must almost certainly have been the most abundant of the remaining two species seen by Bentham. Under Bentham’s concept it covered a multitude of variants which are now considered to be distinct species. Distribution: Sect. Euphrasia is widespread in the northern temperate and holarctic zones in North America, Eurasia as far south as the Himalayas, north-west Africa (Morocco) and Japan. Hulten (1976) gives a detailed account of the distribution of the section, including maps. Of particular biogeographical importance is the occurrence of the genus in North America and northern Eurasia in areas glaciated in the Pleistocene, with pre-glacial localities in the southern areas of Eurasia containing close relatives of the northern taxa. The taxonomy of this section is notoriously difficult. Its species are perhaps equivalent to geographical and ecological races and are often termed “microspecies” (Yeo 1967, 1978a,b; Sell & Yeo 1970; Hulten 1976; Karlsson 1974, 1976). The problem of classification is compounded by the phenomenon of “seasonal dimorphism” within species (p. 6). Note: Pugsley (1936) divided Sect. Euphrasia into four subsections. From breeding and chromosomal studies (Yeo 1966, 1967, 1978b; Sell & Yeo 1970), the two subsections occurring in Europe, Subsect. Ciliatae and Subsect. Angustifoliae, represent natural groupings. Pugsley (1936) divided Subsect. Ciliatae, the largest of the two subsections, into a number of series. Sell & Yeo (1970) have found this useful, and have made a number 77 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) of modifications to Pugsley’s classification including the description of several new series. Hartl (1972) retained Pugsley’s four subsections of Sect. Euphrasia but has not clarified the status of the Japanese subsections. I have seen very few specimens of the Japanese species of these subsections, but a check of their descriptions by Ohwi (1965) shows some of Pugsley’s diagnostic characters for the two subsections to break down, particularly the pilosity of the capsules and the calyx lobing. I am also doubtful of the diagnostic reality of the relative shape and depth of toothing of bracts and upper leaves. Nevertheless, there is clearly diagnostic potential at the infrageneric level in the peculiar purple blotching of the base of the corolla lip used by Pugsley (1936) in his diagnosis of Subsect. Alpicolae. The distinction of abruptly and gradually attenuated types of leaf base and the “glandular-fimbriate” corolla hood attributed by Pugsley to Subsect. Alpicolae may also prove useful diagnostically. A. Subsect. Ciliatae Joerg. : ! In Europe 42 species (Yeo 1972, 1978), 15 (including possible introductions from Europe) in North America (Sell & Yeo 1970), and an undetermined number in Asia. B. Subsect. Angustifoliae (Wettst.) Joerg. : ! In Europe 6 species (Yeo 1972, 1978) and one possibly introduced species in North America (Sell & Yeo 1970). C. Subsect. Japonicae Pugsley : ! Undetermined number of species, endemic to Japan. D. Subsect. Alpicolae Pugsley Undetermined number of species, endemic to high mountains of Japan. II. Sect. Atlanticae Pugsley, J. Bot. (Lond.) 74 (1936) 284 Du Rietz, Sv. Bot. Tidskr. 42 (1948) 360; Hartl in Hegi, Ill. Fl, Mitteleur. (ed.2) 6 (1) (1972) 344; Yeo, Fl. Europaea 3 (1972) 259; Yeo, Bol. Mus. Funchal 17, Art. 121 (1973) 76. Lectotype (Yeo 1973): E. grandiflora Hochst. Sect. Euphrasia: Wettst. in Engler & Prantl, Nat. Pflfam. 1V3b (1893) 100, p.p. (as to E. grandiflora) “Eueuphrasia”; Wettst., Monogr. Gatt. Euphrasia (1896) 68, p.p. (as to Azorean species of Subsect. Semicalcaratae) “Eueuphrasia”. S Semicalcaratae auct. non Benth.: Benth. in DC, Prodr. 10 (1846) 552, p.p. (as to E. grandiflora).—Subsect. Semicalcaratae auct. non (Benth.) Wettst.: Wettst., Monogr. Gatt. Euphrasia (1896) 68, p.p. (as to E. grandiflora).—Sect. Semicalcaratae auct. non (Benth.) Joerg.: Joerg., Berg. Mus. Aarb. 1916-17 Naturvid. raekke 2 (1919) 5, p.p. (as to Azorean representatives; lectotype excl.) “Subgen. Eueuphrasia Sect. Semicalcaratae”. Perennial, of “Cuneata” habit type; main inflorescence-bearing axes 2one to several, erect, developing branches basipetally in consecutive nodes from c. 1-2 nodes below the inflorescence. Uppermost Jeaves of main axes crenate to serrate, with base shortly attenuate, abruptly expanded into large blade, with (5)7-12 teeth along greater part of each margin; with 3-5 main veins arising from base of leaf, branched distally. Flowers c. 15-20 or more in (?main) racemes. Corolla with presence of striations unknown, with yellow spots on the lower lip; Jobes emarginate. Anthers fused, glabrous around the connectives, hairy along slits; awns entire. Ovary with stigma capitate, 0.2-0.25 mm long, Capsule in lateral view deeply emarginate (E. grandiflora) or acuminate to cuspidate (E. azorica). Chromosome number unknown. Distribution: The section consists of two species which are endemic to the Azores of the Atlantic Ocean. They are apparently confined to the mountain regions (Yeo 1972) and geographically separated on two distinct groups of islands about 200km apart (Yeo 1973). E. grandiflora occurs on cliffs (Wettstein 1896) and on “shallow soils near lava flows” (Tutin 1953). 78 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia Notes 1. No material of this section has been seen. The description is compiled largely from information kindly provided by Dr P.F. Yeo (pers. comm. 1976) as well as from Wettstein (1896), Pugsley (1936) and Yeo (1972, 1973). 2. Yeo (1973) refers to the distinctive characters in Sect. Atlanticae of the type of leaf venation (a prominent reticulum of veins visible on the lower side of the leaf between the main veins) and leaf indumentum (sessile glands confined to veins on lower side, and eglandular hairs when present confined to the main veins and the spaces between the veins). He states that these characters are present nowhere else in Euphrasia, but occur in related genera of Trib. Rhinantheae. Unfortunately, little extra-Australian material was available to me when I learnt of these characters. Certainly such attributes are not present in any of the Australian species, E. cuneata of New Zealand, Sect. Novaezeelandiae, Sect. Pauciflorae and Taiwanese material of Sect. Malesianae. On the lower side of the leaves in all of the species studied, sessile glands occur between the veins, eglandular hairs may be spread generally over the leaves when they are present in profusion, and veins other than the main veins are not particularly, if at all, prominent. 3. Yeo (1973) draws attention to the unequally 5-6-winged seed, a character known only in £. azorica. As Yeo points out, with seed morphology unknown in E. grandiflora, it is not known whether this character may be diagnostic for Sect. Atlanticae. If, however, E. grandiflora has the typically many-ribbed seeds of the genus, then there would be strong grounds for separating E. azorica, also peculiar in its bicornute capsules, as a separate section or, at the least, a subsection of Sect. Atlanticae. E. grandiflora Hochst. E. azorica Watson III. Sect. Malesianae Barker, sectio nova Sect. Australes auct. non (Benth.) Joerg.: Hart! in Hegi, II]. Fl. Mitteleur. (ed.2) 6 (1) (1972) 343, p.p. (as to species of Formosa and the Philippines). Herbae perennes. Axes principales inflorescentigeri singularis usque multi, erecti ascendentesve, ramos inordinatos in nodis sporadicis continuisve usque |-aliquot nodos infra inflorescentiam crescentes. Folia summa axis principalis crenata usque serrato-crenata, base breve attenuata, in laminam amplam abrupte expansa, (1)2-4 dentibus secus c. ('/s)?/,-4/; partes distales cuiusque marginis; 3 venis principalibus ad basem folii. Flores 2-20 in racemis principalibus interdum interruptis. Corolla c. 6-10mm longa, striata in labio supero, labio infero e base de cucullo patenti; Jobis emarginatis vel non profunde. Antherae coniunctae, circa connectivas glabrae, secus rimas pilosae; aristis integris. Ovarium stigma oblique capitata vel fere inaequaliter bilobata, 0.15-0.3mm longa. Capsulae in aspectu laterali obovatae vel late obovatae, apice obtuso vel non profunde ita, saepe oblique latere ita. Chromosomatum numerus ignotus. Holotypus: E. philippinensis Du Rietz. Perennial, of “Malesianae” habit type; main inflorescence-bearing axes one to many, ascending or erect, developing branches in no fixed sequence in occasional or consecutive nodes high above ground level, up to 1-few nodes below inflorescence. Uppermost Jeaves of main axes crenate to serrate-crenate, with base shortly attenuate, usually abruptly, rarely gradually expanded into large blade, with (1) 2-4 teeth along distal c. (1/5)%,- ‘/; of each margin; with 3 main veins arising from base of leaf. Flowers 2-20 in sometimes interrupted (main) racemes. Corolla striated (data on type of E. borneensis and from dried material of E. merrillii and E. philippinensis) on upper lip, with lower lip spreading from base away from hood; /obes emarginate but sometimes 79 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) (in E. bilineata, E. nankotaizanensis) very shallowly so. Anthers fused, glabrous about connectives, hairy along slits; awns entire. Ovary with stigma obliquely capitate or almost (unequally) bilobed, 0.15-0.3mm long. Capsules in lateral view obovate or broadly so, laterally compressed; apex in lateral view obtuse or shallowly emarginate, often obliquely or broadly so. Chromosome number unknown. Distribution: Sect. Malesianae occurs in the highest parts of the mountains of Taiwan, the Philippines (Luzon), Borneo (Mt Kinabalu) and Ceram. E. nankotaizanensis E. pumilio Ohwi: ! E. merrillii Du Rietz : T Yamamoto: !, T E. exilis Ohwi E. philippinensis Du Rietz : !, T E. durietziana Ohwi : ! E. bilineata Ohwi : ! E. borneensis Stapf : !, T E. tarokoana Ohwi : ! E. matsudae Yamamoto : !, T E. ceramensis van Royen : !, T E. transmorrisonensis E. masamuneana Ohwi Hayata : ! IV. Sect. Cuneatae Barker, sectio nova Sect. Euphrasia: Wettst. in Engler & Prantl, Nat. Pflfam. IV3b (1893) 100, p.p. (as to E. cuneata) “Eueuphrasia”; Wettst., Monogr. Gatt. Euphrasia (1896) 68, p.p. (as to Subsect. Australes p.p.) “Eueuphrasia”. S Semicalcaratae auct. non Benth.: Benth. in DC., Prodr. 10 (1846) 552, p.p. (as to E. cuneata). Subsect. Australes auct. non (Benth.) Wettst.: Wettst., Monogr. Gatt. Euphrasia (1896) 70, p.p. (as to E. cuneata).—Sect. Australes auct. non (Benth.) Joerg.: Joerg., Berg. Mus. Aarb. 1916-17 Naturvid. raekke 2 (1919) 5, p.p. (as to some New Zealand occurrences) “Subgen. Eueuphrasia Sect. Australes”; Du Rietz, Sv. Bot. Tidskr. 42 (1948) 361, p.p. (as to some New Zealand occurrences) “Subgen. Eueuphrasia Sect. Australes”; Hartl in Hegi. Ill. Fl. Mitteleur. (ed.2) 6 (1) (1972) 343, p.p. (as to some New Zealand occurrences). Herbae perennes. Axis principalis inflorescentiger caulis erectus singularis vel aliquot rami erecti ascendentesve cauli similares, ramis in nodis continuis de l-aliquot infra inflorescentiam basipete crescentibus. Folia summa axis principalis crenata, base attenuata, in laminam gradatim expansa, (1)2(3) dentibus secus !/,-!/, partes distales cuiusque marginis; 3 venis principalibus ad basem folii. Flores c. 15-50 in racemis principalibus. Corolla striata, macula flava in tubo et (in E. cuneata solum) in labio infero, lato infero e base labii inferi de cucullo patenti; /obis vix usque profunde emarginatis. Antherae coniunctae, circa connectivas glabrae, secus rimas pilosae; aristis integris. Ovarium stigma oblique oblonga usque inaequaliter bilobata, (0.2)0.3-0.6mm longa. Capsulae in aspectu laterali ovatae usque obovato-ellipticae, apice emarginato usque obtuso. Chromosomatum numerus ignotus. Holotypus: E. cuneata Forst. f. Perennial, of “Cuneata” habit type; main inflorescence-bearing axis single erect stem or several erect or ascending stem-like branches, with branches developing basipetally in consecutive nodes, from 1-few nodes below inflorescence. Uppermost leaves of main axis crenate, with base attenuate, gradually expanded into large blade, with (1)2(3) teeth along distal 1/,-, of each margin; with 3 main veins arising from base of leaf. Flowers c. 15-50 in (main) racemes. Corolla striated, with yellow blotch and (in E. cuneata only) in tube at base of lower lip, lower side spreading from base of lower lip away from hood; lobes shallowly to deeply emarginate. Anthers fused, glabrous around connectives, hairy along slits; awns entire. Ovary with stigma obliquely oblong to unequally bilobed, (0.2) 0.3-0.6mm long. Capsules slightly compressed laterally, in lateral view ovate to obovate-elliptic; apex emarginate to obtuse. Chromosome number unknown. Distribution: Sect. Cuneatae comprises two widely disjunct species, E. cuneata of lowland to subalpine regions of New Zealand (Ashwin 1961), and E. ramulosa of montane to subalpine regions of north-eastern New South Wales. E. cuneata Forst.: !, ? T E. ramulosa Barker !, T 80 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia V. Sect. Phragmostomae Barker, sectio nova Herbae perennes. Axes principales inflorescentigeri haud satis noti, probabiliter aliquot usque multi, primum prostrati, postremo erecti, ramis ad catervas dissitas nodorum, intra catervas in nodis continuis basipete crescentibus, caterva summa 0-10(18) nodis infra inflorescentiam posita. Folia summa axium principalium serrato-crenata usque serrata, base attenuata, in laminam gradatim expansa, 1-2 dentibus secus 0.1-0.4 partes distales cuiusque marginis; 3 venis principalibus ad basem folii, distale ramosis. Flores usque ad c. 15 in racemis principalibus. Corolla striis carens, macula flava in labio infero et in tubo, lato infero e base labii inferi de cucullo patenti; /obis obtusis usque profunde emarginatis. Antherae coniunctae, circa connectivas glabrae, secus rimas dense pilosae; paribus posterioribus aristarum versus apicem complanatis vel marginibus erosulatis tortis. Ovarium stigma oblique capitata, (0.3)0.4-0.5mm _ longa. Capsulae in aspectu laterali obovato-ellipticae, apice obtuso. Chromosomatum numerus ignotus. Holotypus: E. phragmostoma Barker Perennial, of “Phragmostoma” habit type; main inflorescence-bearing axes inadequately known, probably several to many, initially prostrate, finally erect, with branches at widely-spaced groups of nodes developing within groups in basipetal sequence in consecutive nodes, the uppermost group 0-10(18) nodes below the inflorescence. Uppermost Jeaves of main axes serrate-crenate to serrate, with base attenuate, gradually expanded into large blade, with 1-2 teeth over distal 0.1-0.4 of either margin; with 3 main veins arising from base of leaf, branched distally. Flowers up to c. 15 in (main) racemes. Corolla lacking striations, with yellow blotch on lower lip and in tube, with lower side spreading from base of lower lip away from hood; lobes obtuse to deeply emarginate. Anthers fused, glabrous around connectives, densely hairy along slits; rearmost pair of awns towards apex flattened or twisted with erosulate margins. Ovary with stigma obliquely subcapitate, (0.3)0.4-0.5mm long. Capsule in lateral view obovate-elliptic, with apex obtuse. Chromosome number unknown. Distribution: Sect. Phragmostomae is monotypic and confined to coastal cliffs in the south-east corner of Tasmania. Note: The section is unique in the genus in both its habit and the size and shape of the rearmost anther awns. It is closest to Sect. Cuneatae of Australia and New Zealand by its basipetal development of minor branches on the main axes, large stigma, anthers with glabrous connectives but densely hairy slits, and leaves with a large blade gradually attenuated at the base. Sect. Cuneatae differs, however, in addition to the unique characters described above, by its striated corollas and many-flowered inflorescences. E. phragmostoma Barker: !, T VI. Sect. Pauciflorae Barker, sectio nova Sect. Euphrasia: Wettst. in Engler & Prantl, Nat. Pflfam. 1V3b (1893) 100, p.p. (probably as to one or two N. Zeal. species) “Eueuphrasia”. Subsect. Australes auct. non (Benth.) Wettst.: Wettst., Monogr. Gatt. Euphrasia (1896) 68; p.p. (as to E. monroi and E. revoluta).—Sect. Australes auct. non (Benth.) Joerg.: Joerg., Berg. Mus. Aarb. 1916-17 Naturvid. raekke 2 (1919) 5, p.p. (as to some N. Zeal. species) “Subgen. Eueuphrasia Sect. Australes”; Schlechter, Bot. Jb. 59 (1924) 117; Hartl in Hegi, Ill. Fl. Mitteleur. (ed.2) 6 (1) (1972) 343, p.p. (as to N. Guinea, Celebes and some N. Zeal. occurrences). Subsect. Novaezeelandiae auct. non Du Rietz: Du Rietz, Sv. Bot. Tidskr. 42 (1948) 361, p.p. Herbae perennes. Axes principales inflorescentigeri singulares usque multi, prostrati usque erecti, ramos inordinatos in nodis sporadicis crescentes. Folia summa axis principalis crenata usque crenato-serrata, base attenuata, in laminam parvam usque amplam abrupte gradatimve expansa, 1-2(3) dentibus secus c. !/,-7/, partes distales cuiusque marginis, saepe dente apicali magno cucullatoque; 1-3(5) venibus principalibus ad basem folii. Flores 2-8(12) in racemis principalibus vel, in speciebus ramis prostratis, solitarii dispersi. Corolla striata vel non-striata; /obis emarginatis obtusisve. Antherae coniunctae, circa connectivas glabrae, secus rimas pilosae; aristis integris. Ovarium stigma oblique capitata vel inaequaliter bilobata (0.2)0.3-0.55mm 81 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) longa. Capsulae in aspectu laterali obovatae vel ita late, obcordataeve oblongaeve, apice obtuso vel emarginato, saepe ita late. Chromosomatum numerus: n = 11. Holotypus: E. revoluta Hook.f. Perennial, with habit variable. Uppermost /eaves of main axes crenate to serrate- crenate, with base attenuate, abruptly or gradually expanded into blade, otherwise of variable shape; with 1-3(5) main veins arising from base. Flowers variable in number and arrangement. Corolla with variable coloration, with lower lip concave from above, spreading from base away from hood; lobes variable. Anthers fused, glabrous around connectives, hairy along slits; awns entire. Ovary with stigma obliquely capitate or unequally bilobed, (0.2)0.3-0.55mm long. Capsules variable. Chromosome number: n = 11 (known from only one species, E. mirabilis: Borgmann 1964). Variable characters: see under subsections. Distribution: Sect. Pauciflorae, which consists of 16 species in two subsections, occurs in the mountains of three widely disjunct regions, New Zealand, New Guinea and the Celebes. A. Subsect. Pauciflorae Barker, subsectio nova Subsectio nova Euphrasiae Sectionis Pauciflorae differt a Subsectione Humifusae floribus in racemis axibusque principalibus inflorescentigeris non prostratis. Holotypus: E. revoluta Hook.f. Habit of the “Malesianae” and “Striata” types; main inflorescence-bearing axes either several to many and ascending with simple erect parts, or single to few and erect with branches developing with no fixed sequence high above ground level in occasional nodes. Uppermost Jeaves of main axes with base attenuate, abruptly or gradually expanded into small to large blade, with 1-2(3) teeth on distal ¥/,-¥, of each margin, often (in most New Guinea species and ? sometimes E. revoluta) with apical tooth large and hooded and with small pair of lateral teeth; with 1 or 3 (in E. papuana rarely 5) main veins arising from base. Flowers 2-8(12) in (main) racemes. Corolla conspicuously striated, sometimes only on upper lip or on rear of lobes, or with striations absent, sometimes replaced by flush of colour, with yellow patches on lower lip and tube; /Jobes emarginate or obtuse. Ovary with stigma obliquely capitate or unequally bilobed, (0.25)0.3-0.55mm long. Capsules in lateral view obovate or broadly so, or obcordate to oblong; apex in lateral view obtuse or emarginate, often broadly so. Chromosome number: n= 11 (known from only one species, E. mirabilis: Borgmann 1964). Distribution: Subsect. Pauciflorae occurs in the high montane to alpine zones of New Guinea (7 species) and New Zealand (6 species). Note: The New Guinea and New Zealand members of this subsection form two divergent groups linked by E. papuana of New Guinea and E. revoluta of New Zealand. The New Guinea species have corollas lacking prominent striations, although data on this are incomplete (see p. 29), corolla lobes tending to be small relative to the length of the tube (measurements of the New Zealand species overlap, however), and leaves usually with only a single pair of teeth, often producing the ‘hooded’ leaf distinctive of these species. E. papuana differs by its non-hooded leaves with 1-2 pairs of teeth, a characteristic of the New Zealand representatives. The New Zealand species all have large-lobed corollas which tend to be striated and upward-directed, and, except for E. revoluta, have leaves with 1-2 pairs of teeth. E. revoluta vegetatively resembles E. mirabilis of New Guinea; its small leaves with a single pair of teeth are, however, not conspicuously hooded. Since there is no one pair of correlated characters 82 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia to divide the species, they are combined in the one subsection. New Guinea E. mirabilis Pennell: !. T E. papuana Schlechter: !, T E. lamii Diels: ! E. scutellarioides Wernham: ?!, TE. culminicola Wernham: ?!, T E. versteegii (Diels) Du Rietz: ! E. spatulifolia Pennell: !, T New Zealand E. revoluta Hook.f.: !, T E. monroi Hook.f.: ! E. townsonii Petrie: !T E. drucei Ashwin: ! E. laingii Petrie: !, T E. petriei Ashwin: !, T. B. Subsect. Humifusae Barker, subsectio nova Subsectio nova Euphrasiae Sectionis Pauciflorae differt a Subsectione Pauciflorae axibus principalibus inflorescentigeris prostratis floribusque sporadicis secus axes. Holotypus: E. humifusa Pennell Habit of the “Humifusa” type; main inflorescence-bearing axes several to many, prostrate and rooting at occasional nodes, developing branches along whole length in no fixed sequence, sporadically or in more or less consecutive nodes which are free of flowers. Uppermost /eaves of main axes with base attenuate, abruptly or gradually expanded into small blade, with 1(2) teeth, often small, on distal 1/,-4, of either side, with apical tooth often large and hood-like; with 1 or 3 main veins arising from base. Flowers sporadic in axils of leaves along axes, one at each node, developing acropetally. Corolla purple on upper lip, from dried material sometimes striated faintly on lower lip (E. callosa), possibly (in E. humifusa: van Royen 1972) lacking striations, with yellow blotch on lower lip (at least in E. callosa); lobes emarginate or shallowly so. Ovary with stigma capitate or unequally bilobed, (0.2)0.3-0.5mm long. Capsules in lateral view broadly obovate, laterally compressed; apex in lateral view broadly obtuse to shallowly emarginate. Chromosome number unknown. Distribution: The subsection comprises three species, two from the alpine and subalpine regions of New Guinea, and one from the summit of Mt Loemuet in the Celebes. E. humifusa Pennell: !, T E. callosa Pennell: !, T E. celebica van Royen: T VII. Sect. Striatae (Du Rietz) Barker, stat. nov. Ser. Striatae Du Rietz, Sv. Bot. Tidskr. 42 (1948) 113, 359 BASIONYM; Willis, Muelleria 1 (1967) 147. Holotype: E. striata R.Br. See Typification. Sect. Euphrasia: Wettst. in Engler & Prantl, Nat. Pflfam. IV3b (1893) 100, p.p. (as to E. cuspidata Hook.f.) “Eueuphrasia”. Ser. Hookerae Du Rietz, Sv. Bot. Tidskr. 42 (1948) 359 “Hookeriae”. Holotype: E. hookeri Wettst. S Australes auct. non Benth.: Benth. in DC., Prodr. 10 (1846) 553, p.p. (as to E. striata and E. alpina var. humilis); Pugsley, J. Bot. (Lond.) 74 (1936) 276; Sell & Yeo, Bot. J. Linn. Soc. 63 (1970) 203.—Subsect. Australes auct. non (Benth.) Wettst.: Wettst., Monogr. Gatt. Euphrasia (1896) 68, p.p. (as to E. striata and E. hookeri).—Sect. Australes auct. non (Benth.) Joerg.: Joerg., Berg. Mus. Aarb. 1916-17 Naturvid. raekke 2 (1919) 5, p.p. (as to some Australian species) “Subgen. Eueuphrasia Sect. Australes”; Schlechter, Bot. Jb. 59 (1924) 117; Hartl in Hegi, Ill. Fl. Mitteleur. (ed.2) 6 (1) (1972) 343, p.p. (as to some Australian and Tasm. occurrences). Subsect. Australienses auct. non Du Rietz: Du Rietz, Sv. Bot. Tidskr. 42 (1948) 361, p.p. (as to Ser. Striatae, Ser. Hookerae and “E. milliganii”’). Perennial, of the “Striata” or “Collina” habit type; main inflorescence-bearing axes either several to many and ascending with simple erect distal parts, or single to few and erect with branches developing high above ground level in occasional nodes in no fixed sequence. Uppermost /eaves of main axes crenate to serrate, with base gradually attenuate to cuneate, with blade small to large, toothed in distal '/,-',(4;), in Ser. Striatae with 1-2(5) teeth along each margin, in Ser. Hookerae with (3)4-5(7) teeth 83 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) along each margin; with 3-(sometimes in E. gibbsiae) 5 main veins arising from base. Flowers (7)10-24(36) in (main) racemes. Corolla conspicuously striated, sometimes only on lateral lobes or on tube, hood and base of lower lip but hardly extending onto lobes, with lower side concave from above, spreading from base or lower lip away from hood; /obes broadly obtuse to emarginate. Ovary with stigma obliquely capitate to oblong or unequally bilobed, 0.35-0.8mm long. Capsules in lateral view obovate to ovate-elliptic, sometimes broadly so; apex in lateral view emarginate to obtuse. Chromosome number: n = c. 20-30. Typification Ser. Striatae Du Rietz Although no type was cited by Du Rietz (1948a), the selection is automatic as his series name was based upon the epithet of one of the species described in the protologue (ICBN, Art. 22). Distribution: Sect. Striatae is restricted to Tasmania except for an outlier on the Baw Baw plateau of the eastern highlands of Victoria. In a mountainous region further north in Victoria is a polymorphic species, E. crassiuscula, which varies in the characters distinguishing Sect. Striatae from Sect. Australes (p. 152). Ser. Striatae Du Rietz E. gibbsiae Du Rietz: !, T E. striata R.Br.: !, T E. semipicta Barker: !, T Ser. Hookerae Du Rietz E. hookeri Wettst.: !, T VIII. Sect. Australes (Benth.) Joerg., Berg. Mus. Aarb. 1916-17 Naturvid. raekke 2 (1919) 5, p.p. (as to lectotype and some other Australian and some N. Zeal. species) “Subgen. Eueuphrasia Sect. Australes” Du Rietz, Sv. Bot. Tidskr. 42 (1948) 361, p.p. (as to some Australian and some N. Zeal. species); Hart] in Hegi, Ill. Fl. Mitteleur. (ed.2) 6 (1) (1972) 343, p.p. (as to some Australian and Tasm. occurrences). —S Australes Benth. in DC., Prodr. 10 (1846) 553, p.p. (excl. E. alpina var. humilis, E. striata, E. scabra and probably E. paludosa var. pedicularoides) BASIONYM; Pugsley, J. Bot. (Lond.) 74 (1936) 276; Sell & Yeo, Bot. J. Linn. Soc. 63 (1970) 203.—Subsect. Australes (Benth.) Wettst., Monogr. Gatt. Euphrasia (1896) 70, p.p. (as to perennials excl. E. cuneata, E. monroi, E. striata, and E. repens). Lectotypus hic designatus: E. alpina R.Br. non Lamk., nom. illeg. = E. diemenica Spreng. See Typification. Subsect. Australienses Du Rietz, Sv. Bot. Tidskr. 42 (1948) 361, p.p. (as to Ser. Collinae s. lat., i.e. incl. E. paludosa, E. muelleri, “E. collinoides” and “E. osbornii’, but excl. “E. milliganii’). Lectotypus hic designatus: E. collina R.Br. See Typification. Ser. Collinae Du Rietz, Sv. Bot. Tidskr. 42 (1948) 359; Willis, Muelleria 1 (1967) 147. Holotype: E. collina R.Br. See Typification. Sect. Euphrasia: Wettst. in Engler & Prantl, Nat. Pflfam. 1V3b (1893) 100, p.p. (as to E. brownii, ? p.p.) “Eueuphrasia”; Wettst., Monogr. Gatt. Euphrasia (1896) 68, p.p. (as to Subsect. Australes) “Eueuphrasia”. Perennial, of the “Striata” or “Collina” habit types; main inflorescence-bearing axes either several to many and ascending with distal erect parts simple, or single to few and erect with branches developing high above ground level in occasional nodes in no fixed sequence. Uppermost /eaves of main axes crenate to serrate-pinnatifid, with base gradually attenuate (E. bella) or cuneate to truncate (E. collina), with blade large, with (0)1-6(8) teeth along distal '/. to whole of each margin; with 3-7 main veins arising from base. Flowers (4)6-60(80) in (main) racemes. Corolla lacking striations, with or without yellow spot on lower lip, with lower side (? always) flattened, sometimes forming a broad groove, with lower lip + porrect proximally, spreading distally in region of lobes; /obes obtuse to deeply emarginate. Anthers fused, connectives almost always hairy, in a few subspecies of E. collina rarely glabrous, slits hairy; awns entire. Ovary with stigma obliquely oblong to capitate or unequally bilobed, 0.5-1.0mm long. Capsules in lateral view ovate to obovate, sometimes broadly so, or shortly caudate, 84 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia rarely linear; apex shallowly emarginate to acute or obtuse, rarely shortly broadly acuminate. Chromosome number: variable estimates (p. 36). Typification 1. S Australes Benth. Except for E. tetragona R.Br. and E. striata R.Br., the descriptions of which are rearrangements of Brown’s (1810) originals, any one of the other six species included by Bentham (1846) under S Australes would qualify equally as lectotype. In considering the characters used by Bentham to delimit S Australes, none of the species has the allegedly “subequally mucronate anthers” wrongly attributed to this group by Bentham (and other authors), while all bear hairs on the anthers to a greater or lesser extent. The final choice was based on the fact that Bentham saw much material of Brown’s “E. alpina”. 2. Subsect. Australienses Du Rietz Any of the four species, E. collina, E. gunnii, E. striata and E. gibbsiae, described in great detail by Du Rietz (1948a,b) would have been suitable as lectotype. E. collina was chosen as it was a type of one of the two series Collinae and Striatae which encompass these four species. It also appears to have capsules corresponding to the “acuminate” type which was used by Du Rietz to distinguish the subsection from his Subsect. Novaezeelandiae (but see p. 34). 3. Ser. Collinae Du Rietz. Although no type was cited, E£. collina is automatically holotype as the series name is based upon it (ICBN, Art. 22). Distribution: The section is spread throughout temperate southern Australia (including Tasmania), the entire area of which is covered by one extremely polymorphic species, E. collina. Also included in the subsection are three species which are confined to small mountain regions; one of these, FE. crassiuscula, intergrades with Sect. Striatae (see p. 152). Note: This section, which appears so natural on phenetic grounds, may consist of two paraphyletic groups (pp. 57, 62). Confirmation by further study, particularly of chromo- some numbers, is needed before the desirability of separating E. bella, E. bowdeniae and an unnamed close relative as a separate section can be considered. E. bella Blake: !, T E. collina R.Br.: !, T E. crassiuscula Gandoger: !, T E. bowdeniae Barker: !, T IX. Sect. Lasiantherae Barker, sectio nova Sect. Euphrasia: Wettst., Monogr. Gatt. Euphrasia (1896) 68, p.p. (as to Subsect. Australes p.p.) “Eueuphrasia”. Sect. Trifidae auct. non (Benth.) Wettst.: Wettst. in Engler & Prantl, Nat. Pflfam. IV3b (1893) 101, p.p. (as to Australian plants of E. antarctica). Subsect. Australes auct. non (Benth.) Wettst.: Wettst., Monogr. Gatt. Euphrasia (1896) 70, p.p. (as to E. alsa). —Sect. Australes auct. non (Benth.) Joerg.: Joerg., Berg. Mus. Aarb. 1916-17 Naturvid. raekke 2 (1919) 5, p.p. (as to some Australian occurrences) “Subgen. Eueuphrasia Sect. Australes”; Du Rietz, Sv. Bot. Tidskr. 42 (1948) 361, p.p. (as to some Australian occurrences); Hartl in Hegi, Ill. Fl. Mitteleur. (ed.2) 6 (1) (1972) 343, p.p. (as to some Australian occurrences). Subsect. Australienses auct. non Du Rietz: Du Rietz, Sv. Bot. Tidskr. 42 (1948) 361, p.p. (as to E. alsa). Ser. Scabrae auct. non Du Rietz: Du Rietz, Sv. Bot. Tidskr. 42 (1948) 360, p.p. (as to E. alsa); Willis, Muelleria | (1967) 147, p.p. (as to E. alsa). Herbae annuae perennesve. Axes principales inflorescentigeris vel (in herbis annuis) caulis singularis erectus ramis paucis in nodis continuis infra inflorescentiam basipete crescentibus, vel (in herbis perennibus) rami multi ascendentes, proxime prostrati et saepe ramosi, distale integri erectique. Folia summa axis principalis creniato-serrata usque pinnatifida, base anguste cuneata usque truncata, lamina magna, (1)2-4(7) dentibus secus ?/;-totas partes cuiusque marginis; 3-7 venis principalibus ad basem folii. Flores (14)15-26(30) in racemis. Corolla manifeste striata, sum vel sine macula flava in labio infero, labio infero ad basem plus 85 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) minusve porrecto, distale patenti; /obis plerumque emarginatis, interdum obtusis. Antherae coniunctae, circa connectivas et secus rimas pilosae; aristis integris. Ovarium stigma oblique capitata usque oblonga vel inaequaliter bilobata, 0.3-0.7mm longa. Capsulae in aspectu laterali ovatae usque obovatae, apice plerumque emarginato usque late obtuso, raro acuto. Chromosomatum numerus: n=27 et circa 38-47. Holotypus: E. lasianthera Barker Perennial of the “Striata” habit type, or annual; main inflorescence-bearing axes (in perennials) many ascending branches, initially prostrate and often branched, distally simple and erect, or (in annuals) a single erect stem with few branches developing basipetally in consecutive nodes below inflorescence. Uppermost Jeaves crenate-serrate to pinnatifid, with base narrowly cuneate to truncate, with large blade, with (1)2-4(7) teeth distributed over distal 7,-entire length of each margin; with 3-7 main veins arising from base. Flowers (14)15-26(30) in (main) racemes. Corolla prominently striated, with or without yellow blotch on lower lip, with lower side + flat crosswise, with lower lip + porrect near base, spreading distally; /obes usually emarginate, sometimes obtuse. Anthers fused, hairy about connectives and along slits; awns entire. Ovary with stigma obliquely capitate to oblong, or unequally bilobed, 0.3-0.7mm long. Capsules in lateral view ovate to obovate, slightly compressed laterally, apex in lateral view usually emarginate to broadly obtuse, rarely acute. Chromosome number: n=27 and c. 38-47. Distribution: Sect. Lasiantherae is confined within montane to alpine zones of three disjunct mountain areas of south-east mainland Australia. E. lasianthera Barker: !, T E. eichleri Barker: !, T E. alsa FvM.: !, T X. Sect. Scabrae (Du Rietz) Barker, stat. nov. Ser. Scabrae Du Rietz, Sv. Bot. Tidskr. 42 (1948) 360, p.p. (excl. E. alsa) BASIONYM; Willis, Muelleria 1 (1967) 147, p.p. (excl. E. alsa). Holotype: E. scabra R.Br. See Typification. S Australes auct. non Benth.: Benth. in DC., Prodr. 10 (1846) 553, p.p. (as to E. scabra and possibly E. paludosa var. pedicularoides).—Subsect. Australes auct. non (Benth.) Wettst.: Wettst., Monogr. Gatt. Euphrasia (1896) 70, p.p. (as to E. scabraand E. arguta).—Sect. Australes auct. non (Benth.) Joerg.: Joerg., Berg. Mus. Aarb. 1916-17 Naturvid. raekke 2 (1919) 5, p.p. (as to some Australian occurrences) “Subgen. Eueuphrasia Sect. Australes’; Du Rietz, Sv. Bot. Tidskr. 42 (1948) 361, p.p. (as to some Australian occurrences); Hartl in Hegi. Ill. Fl. Mitteleur. (ed.2) 6 (1) (1972) 343, p.p. (as to some Australian occurrences). Subsect. Australienses auct. non Du Rietz: Du Rietz, Sv. Bot. Tidskr. 42 (1948) 361, p.p. (as to Ser. Scabrae, but excl. E. alsa). Annuals. Main inflorescence-bearing axis a single erect stem, with branches developing basipetally in consecutive nodes from 1-few nodes below inflorescence. Uppermost Jeaves of main axis crenate to deeply pinnatifid, with base narrowly cuneate to rounded, with blade prominent, with 0-3(5) teeth over distal 1/,-entire length of each margin; with 3-7 main veins arising from base. Flowers (10)14-90 or more in racemes. Corolla lacking prominent striations, with (or ?without) yellow to red patch on lower lip or completely yellow, with lower side + flat, broadly grooved, with lower lip porrect at least at base, distally in region of lobes sometimes spreading; Jobes obtuse, sometimes shortly apiculate, to emarginate. Anthers fused, hairy about connectives, at least of posterior pair, and along slits; awns entire. Ovary with stigma obliquely oblong to capitate, or unequally bilobed, (0.2)0.35-1.1mm long. Capsules in lateral view oblong-ovate to obovate-elliptic, often obliquely so, sometimes broadly so, slightly compressed laterally; apex in lateral view emarginate to obtuse or acute, often obliquely so. Chromosome number: n=27 &/or 28 &/or 271I+21 &/or ? 251I+211I (E. caudata). 86 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia Typification Ser. Scabrae Du Rietz Although no type species was cited by Du Rietz (1948b), E. scabra must be considered the holotype since Du Rietz’s series name was based upon the specific epithet (ICBN, Art. 22). Distribution: Sect. Scabrae is confined to the temperate lowland to alpine regions of southern and eastern Australia, including Tasmania. E. caudata (Willis) Barker: !, T E. orthocheila Barker: !, T E. ciliolata Barker: !, T E. scabra R.Br.: !, T E. arguta R.Br.: !, T XI. Sect. Novaezeelandiae (Du Rietz) Barker, stat. nov. Subsect. Novaezeelandiae Du Rietz, Sv. Bot. Tidskr. 42 (1948) 361, p.p. BASIONYM. Lectotypus hic designatus: E. zelandica Wettst. See Typification. Sect. Euphrasia: Wettst., Monogr. Gatt. Euphrasia (1896) 68, p.p. (as to Subsect. Australes p.p.) “Eueuphrasia”. Sect. Trifidae auct. non (Benth.) Wettst.: Wettst. in Engler & Prantl, Nat. Pflam. 1V3b (1893) 101, p.p. (as to E. antarctica in N. Zeal.). Subsect. Australes auct. non (Benth.) Wettst.: Wettst., Monogr. Gatt. Euphrasia (1896) 70, p.p. (as to E. zelandica and E. berggrenii).—Sect. Australes auct. non (Benth.) Joerg.: Joerg., Berg. Mus. Aarb. 1916-17 Naturvid. raekke 2 (1919) 5, p.p. (as to some N. Zeal. occurrences) “Subgen. Eueuphrasia Sect. Australes”; Hartl in Hegi, Ill. Fl. Mitteleur. (ed.2) 6 (1) (1972) 343, p.p. (as to some N. Zeal. occurrences). Annual. Main inflorescence-bearing axis a single erect stem, with branches developing apparently acropetally or in no fixed sequence in the few consecutive nodes available from 1-few nodes below inflorescence. Uppermost Jeaves of main axis crenate to crenate-serrate, with base attenuate, gradually or + rapidly expanded into large blade, with 2-5 pairs of teeth distributed over distal '/,-entire margin; with 3 main veins arising from base. Flowers 6-14 in (main) racemes. Corolla either conspicuously striated, with yellow patches or (in E. cockayniana) completely yellow, with lower lip spreading from base away from upper lip; /obes obtuse to acute or (in E. cockayniana) shallowly emarginate. Anthers fused, glabrous about connectives, very sparsely to densely hairy along slits; awns entire. Ovary with stigma obliquely capitate to linear, or unequally bilobed, 0.3-0.6mm long. Capsules in lateral view broadly obovate to shallowly obcordate, greatly compressed laterally; apex in lateral view broadly obtuse to emarginate. Chromosome number unknown. Typification Subsect. Novaezeelandiae Du Rietz Du Rietz (1932a; 1948a,b) clearly saw material of all the annual species and, among the perennials of New Zealand, E. revoluta, E. monroi, E. laingii, E. townsonii and E. cuneata. His Subsect. Novaezeelandiae was stated to comprise all the New Zealand species (Du Rietz 1948b). Possibly the species of which he saw most material was E. zelandica Wettst. Because it has the typical capsules of the New Zealand and tropical species, referred to as “emarginate or truncate” by Du Rietz and providing the basis for his distinction of Subsect. Novae- zeelandiae from Subsect. Australienses (but see p. 34), it has been selected as lectotype. Distribution: Sect. Novaezeelandiae is confined to the montane to alpine zones of the North and South Islands of New Zealand (ex Ashwin 1961). E. cheesemanii Wettst.: !, T E. australis Petrie: !, T E. zelandica Wettst.: !, T E. cockayniana Petrie: !, T 87 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) XII. Sect. Paradoxae Pugsley, J. Bot. (Lond.) 74 (1936) 284. Du Rietz, Sv. Bot. Tidskr. 42 (1948) 360; Hartl in Hegi, Ill. Fl. Mitteleur. (ed.2) 6 (1) (1972) 344. Holotype: E. formosissima Skottsb. Perennial, of the “Paradoxae” habit type; main inflorescence-bearing axis a single erect stem, with branches developing in all nodes apparently in an acropetal sequence, with growth continuing from some apices in later years. Uppermost Jeaves of main axis crenate, with base attenuate, abruptly to + gradually expanded into large blade, with 3-5 teeth along distal ;-3/, of each margin; with 3 main veins arising from base. Flowers c. 10-15 in (main) racemes. Corolla striated on upper and lower lip, with yellow blotches on lower lip and in tube, with lower side concave from above, spreading away from hood from base of lower lip; Jobes obtuse. Anthers fused, glabrous around connectives, slightly hairy on slits; awns entire. Ovary with stigma obliquely capitate or oblong, c. 0.4mm long. Capsule in lateral view ? obovate or broadly so, laterally compressed; apex in lateral view shallowly emarginate. Chromosome number unknown. Distribution: Sect. Paradoxae in monotypic and restricted to the Juan Fernandez Islands. It occurs in the higher parts of Masafuera, commonly in the alpine zone (Skottsberg 1921). Note: Opinions as to the affinities of Sect. Paradoxae have been varied. Skottsberg (1921) pointed to the generally agreed remote relationship to the neighbouring South American species (herein Sect. Trifidae). Wettstein (1921) pointed to it being closest to both the New Zealand perennial E. cuneata (Sect. Cuneatae) and the Japanese annual E. yabeana Nakai (Sect. Euphrasia Subsect. Alpicolae). Du Rietz (1932b, 1948a), however, considered the section to be most closely allied to the Australasian species, in particular several New Zealand species. In my view (p. 55) E. formosissima is un- doubtedly closest to Sect. Novaezeelandiae, a group of small New Zealand annuals with which it shares not only a unique acropetal pattern of branch development, but also sparsely hairy anther slits and shallowly toothed leaves. E. formosissima Skottsb.: !, T. XIII. Sect. Anagospermae (Hook.f.) Barker, stat. nov. Euphrasia Subgen. Anagospermae Hook.f., Ic. Pl. 13 (1879) 65, t. 1283 BASIONYM.—Anagosperma (Hook.f.) Wettst., Ber. dtsch. bot. Ges. 13 (1895) 242; Wettst., Monogr. Gatt. Euphrasia (1896) 10; Cheeseman, Man. N.Z. FI. (ed.1) (1906) 557. Holotype: E. disperma Hook.f. © Siphonidium Armstr., Trans. Proc. N.Z. Inst. 13 (1881) 341; Wettst. in Engler & Prantl, Nat. Pflfam. 1V3b (1893) 101; Wettst., Monogr. Gatt. Euphrasia (1896) 10; Cheeseman, Man. N.Z. FI. (ed.1) (1906) 558; Cheeseman, Man, N.Z. Fl. (ed.2) (1925) 844. Holotype: S. longiflorum Armstr. (= E. disperma Hook.f.). Sect. Euphrasia: Wettst. in Engler & Prantl, Nat. Pflfam. 1V3b (1893) 100, ?p.p. (probably as to one of the four N. Zeal. species) “Eueuphrasia”; Wettst., Monogr. Gatt. Euphrasia (1896) 70, p.p. (as to Subsect. Australes p.p.) “Eueuphrasia”. Subsect. Australes auct. non (Benth.) Wettst.: Wettst., Monogr. Gatt. Euphrasia (1896) 70, p.p. (as to E. repens and E. dyeri).—Sect. Australes auct. non (Benth.) Joerg.: Joerg., Berg. Mus. Aarb. 1916-17 Naturvid. raekke 2 (1919) 5, p.p. (as to some N. Zeal. occurrences) “Subgen. Eueuphrasia Sect. Australes”; Hartl in Hegi, Ill. Fl. Mitteleur. (ed.2) 6 (1) (1972) 343, p.p. (as to some N. Zeal. occurrences). Subsect. Novaezeelandiae auct. non Du Rietz: Du Rietz, Sv. Bot. Tidskr. 42 (1948) 361, p.p. (as to some N. Zeal. occurrences). Annual. Main inflorescence-bearing axes usually several branches, either initially prostrate and gradually ascending or completely prostrate, sometimes (£. dyeri p.p.) a single + erect stem, with branches developing + acropetally or sporadically, in occasional or consecutive axils. Leaves either entire and narrow-acuminate (E. integri- folia) or pinnatifid to pinnatifid-serrate, with base narrowly cuneate or gradually 88 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia attenuate, with only a small blade, with 1-3 teeth distributed over distal '/,-}, of each margin; with 1-3 main veins arising from base. Flowers 2-6 in (main) racemes or (especially in prostrate plants) sporadic. Corolla with striation on the midline of hood and sometimes either side, sometimes with 1-5 striations on the lobes, with tube and lower side of mouth yellow, with lower lip spreading from base away from hood; lobes obtuse to truncate. Anthers fused or (E. disperma) free, glabrous around connectives, glabrous or with a few hairs along slits; awns entire. Ovary with stigma narrowly oblong, unequally bilobed, 0.35-0.8mm long. Capsules laterally compressed, broadly obovate to obcordate or bicornute in lateral view; apex in lateral view broadly obtuse to very deeply emarginate. Chromosome number unknown. Distribution: The section, which contains four species, is endemic to the montane and wet coastal regions of the South Island of New Zealand. Notes 1. The section, as here constituted, may be polyphyletic, with E. disperma possibly having a common origin with Sect. Trifidae, the rest sharing a common ancestry with Sect. Novaezeelandiae (p. 55). In addition, E. integrifolia, in my view; may be a perennial, in which case it would be misplaced with these species. It seems preferable to group these species together pending further study. 2. From J.D. Hooker’s description of Subgen. Anagospermae in 1879 to the present day there has been speculation whether the capsules of E. disperma and of its closest relatives, which are included in the section proposed herein, are indehiscent (see Du Rietz 1932a, Ashwin 1961). Simpson (1977) has shown that the capsules are truly dehiscent although there is a tendency to vivipary. 3. Wettstein (1895), in elevating J.D. Hooker’s Euphrasia Subgen. Anagospermae to generic level, incorrectly stated that Hooker had given it the status of section. E. dyeri Wettst.: !, T E. integrifolia Petrie: !, T E. disperma Hook.f.: !, T E. repens Hook.f.: !, T XIV. Sect. Trifidae (Benth.) Wettst. in Engler & Prantl. Nat. Pflfam. 1V3b (1893) 101, p.p. (excl. E. antarctica in Austral. and N. Zeal.). Wettst., Monogr. Gatt. Euphrasia (1896) 70; Du Rietz, Sv. Bot. Tidskr. 42 (1948) 114, 361; Hartl in Hegi, Ill. Fl. Mitteleur. (ed.2) 6 (1) (1972) 344.—S Trifidae Benth. in DC., Prodr. 10 (1846) 554, ?p.p. (excl. “E. ?glandulosa”, but material n.v.) BASIONYM; Sell & Yeo, Bot. J. Linn. Soc. 63 (1970) 203.— Subgen. Trifidae (Benth.) Joerg., Berg. Mus. Aarb. 1916-17 Naturvid. raekke 2 (1919) 5. Holotype: E. trifida Poepp. ex Benth. See Typification. Perennial, of the “Striata” and ? “Malesianae” habit types, or annual; main inflorescence-bearing axes either several to many and ascending, with branches developing in consecutive nodes up to several to many nodes below inflorescence, simple above, or single and erect with branches developing in no fixed sequence in consecutive nodes from 1-few nodes below inflorescence. Uppermost /eaves of main axes deeply divided, with base attenuate to cuneate-attenuate, gradually expanded into small blade, with 1(2) pairs of teeth distributed over distal '/,-7, of leaf; with 3 main veins arising from base of leaf. Flowers c. 10-30 in (main) racemes. Corolla striated with yellow area on lower lip or yellow throughout (Reiche 1911), with lower side apparently concave from above, with lower lip spreading from base away from hood; lobes emarginate, often shallowly so. Anthers free, glabrous around connective and along slits; awns entire. Ovary with stigma obliquely subcapitate, c. 0.3-0.5mm long. Capsules in lateral view broadly obovate, greatly compressed laterally; apex in lateral view broadly obtuse. Chromosome number: n=44 (known for a single species: D.M. Moore fide Yeo 1968). 89 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) Typification Under Article 22 of the ICBN, E. trifida, which was described in the protologue (Bentham 1846), is automatically the type of S Trifidae Benth. Distribution: Sect. Trifidae is confined to the Andes mountains and their surroundings in the southern part of South America, with one species E. antarctica extending to the Falkland Islands to the east of Fuegia (Skottsberg 1913; Vallentin & Cotton 1921). Eighteen species are currently recognized, on the basis of the works dating from Wettstein’s (1896) monograph, but revision is needed at both the species level (Reiche 1911) and the infrageneric level (see p. 75). There is controversy about the northern limit of the genus in South America. Wettstein (1896) shows it extending into Peru, but the northern half of this range is based upon the record of a single species E. pubescens described by Bentham (1846) and apparently only represented in herbaria (K, B, P: n.v.) by the type material which lacks a specific locality (Wettstein 1896; Edwin 1971). Thus with the Peruvian occurrence in question, Du Rietz (1940), Burbidge (1960), van Balgooy (1966), van Steenis (1971) and Hulten (1976) have shown the northern limits of the genus much further south at a latitude in the range 29-35°S. The last major work on the genus in South America was a revision of the Chilean species by Reiche (1911), who cited the northern limits at about 30- 31°S. His northernmost locations in the Province of Coquimbo were of an annual which he called E. antarctica, a species widespread in the southern regions. Johnston (1929) described E. adenotata from about 180km further north at about 29°S; he believed that Reiche’s northern occurrences of E. antarctica were actually records of his new species. Descole & Borsini (1955) described a new annual species EF. piossekii, which is undoubtedly a member of Sect. Trifidae from the description and illustration. Its location in Jujuy Province, Argentina extends the range of the genus much further north to about 24°S. Finally, to add to the controversy of a Peruvian occurrence of the genus, Ruiz & Pavon (1959) published posthumously (Stafleu 1967) seven new species of Euphrasia from Peru. Edwin (1971) considered that all but two of these species belonged not to Euphrasia, but to a related genus, probably Bartsia, which is prevalent in the northern Andes. In my view six of the species, including the two which Edwin considered to be of Euphrasia, seem to belong to Bartsia or at least to a genus not Euphrasia, on the basis of their serrate to crenate leaves, hairy anthers and entire corolla hoods (the first two characters are atypical of Sect. Trifidae and the last is a character separating the two genera). Contrary to Edwin I believe that E. tripartita, with its bilobed upper corolla lip and trifid leaves, belongs to Euphrasia Sect. Trifidae. Localities cited in the protologue were near Concepcion, Chile and at Cheuchin, Peru. In view of the several cases documented above, there seems no doubt that Sect. Trifidae occurs well to the north of its currently accepted distributional range. It appears that north of latitude 33°S the genus may be represented by a series of annuals occurring in the Andes as far as Peru. Note: The sectional description is based on material from NSW and MEL. The identity of the specimens was not checked. Perennials E. andicola Benth. E. debilis Wettst. E. chrysantha Phil. E. subexserta Benth. E. villaricensis Phil. E. aurea F, Phil. E. trifida Poepp. ex Benth. E. intricata Phil. E. muscosa Phil. E. flavicans Phil. Annuals ¥ E. antarctica Benth. (!,T) E. meiantha. Clos E. piossekii Desc. & Bors. E. pubescens Benth. E. philippii Wettst. E. tripartita Ruiz & Pavon E. perpusilla Phil. E. adenotata 1.M. Johnston 90 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia Vv. A TAXONOMIC REVISION OF EUPHRASIA IN AUSTRALIA A. INTRODUCTION The classification of Euphrasia in Australia has had to be completely restructured. Wettstein (1896), who published the last revision of the genus in Australia, recognized twelve species and proposed no formal infrageneric or infraspecific taxa. In the proposed classification, six of his twelve species are retained. Of the remainder five are reduced to subspecies and one is placed in synonymy. Euphrasia in Australia consists of six sections with eighteen species, of which eight are newly described and one is an upgrading of status. Four species, in particular E. gibbsiae with nine subspecies and E. collina with 14 subspecies, are polymorphic. In all, 42 taxa at the subspecies or species level have been described, of which 18 are new. Fifteen new combinations and/or changes in status have been made, but three were validated by Curtis in 1978 (Stones & Curtis 1967-78). The difficulties in the taxonomy of Euphrasia in Australia encountered by previous taxonomists have been caused mainly by the complexities of E. collina, which encompasses a large proportion of the Australian populations of Euphrasia. E. collina as defined in this revision is more or less equivalent to Mueller’s (1865) “E. brownii”. All taxa have limited geographical and ecological ranges. Clines and a limited amount of hybridization occur within the species. Interspecific hybrids are rare. B. TAXONOMIC HISTORY The first Australian collections of Euphrasia were apparently made by Labillardiére from southern Tasmania (see Nelson 1974; Apfelbaum 1977). He made at least three collections (verified in a note accompanying his handwritten description of the collection Labillardiére 43 in FI) all of which he considered to be E. cuneata Forst.f. of New Zealand, the sole known representative of the genus in the southern hemisphere at that time, but which belong to three taxa, ssp. psilantherea and ssp. kingii of E. gibbsiae and E. collina ssp. collina. It was Robert Brown who made the initial published contribution to the knowledge of Euphrasia in Australia. From perhaps no more than a dozen of his own collections, most of which are described exhaustively in his manuscript (Brown unpubl.), he described in his “Prodromus” (Brown 1810) eight species, all new, from localities throughout temperate Australia. Although not giving them any emphasis, he recognized the characters of duration and corolla coloration which have proved important in the proposed infrageneric revision. Subsequent works of Sprengel (1825) and Bartling (1845) were based largely on Brown’s work. Although produced over 35 years after Brown’s published treatment of Euphrasia and based on additional collections, Bentham’s (1846) large monograph of Scrophular- iaceae showed little modification of Brown’s concepts. He did not see Brown’s types and consequently applied Brown’s E. collina to the wrong taxon and erroneously erected a new species E. multicaulis. In addition he described four infraspecific taxa under Brown’s species. Two of these he later reduced to synonymy with other Brownian species (Bentham 1868). The third, E. paludosa f pedicularoides, was not referred to again; having seen no material, I cannot make any assessment of it. The fourth, E. scabra 8 arguta, he later reinstated as a species (see below, also p. 284). Subsequently, following explorations of montane to alpine areas of eastern temperate Australia, by Ferdinand Mueller, and Tasmania, by resident collectors such as Gunn and Archer sponsored by J.D. Hooker (see Burns & Skemp 1961), two distinctive new species were described, EF. alsa by Mueller (1855, 1856) and £. cuspidata by J.D. Hooker (1857). Nevertheless, while large collections were accumulating 91 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) from throughout the range of the genus in Australia encompassing all but a few localized species (e.g. E. phragmostoma, E. bella, E. bowdeniae) and the majority of infraspecific taxa, no other new taxa were recognized up to the end of the century. Publications utilized Brown’s (1810) taxa and Bentham’s (1846) taxonomy together with the two above additions. J.D. Hooker (1857, 1859a) agreed with Bentham that several species might be found to be forms of the one species. With his wide field experience of the genus, Mueller (1865) concurred and united five of Brown’s species and that of Bentham into a single species E. brownii. Despite an opinion to the contrary (van Royen 1972)*, the name £. brownii is illegitimate as the author clearly and deliberately based the new name upon seven validly published species names, most of which are legitimate (ICBN, Art. 63). E. striata was considered by Mueller (1865) to be distinct from E. brownii but he doubted its separation as a species. However, he had confused E. striata, a distinctive Tasmanian alpine species which he had not then seen in the field (but later saw in 1869), with an undescribed taxon which he had collected in the alpine regions of the “Munyang Mountains” and which indeed falls into the E. brownii complex, namely E. collina ssp. glacialis. Mueller erected a new variety psilantherea of E. brownii, and expressed some uncertainty as to the status of EF. arguta. In his second account of Euphrasia in Australia as part of “Flora Australiensis”, Bentham (1868) was able to refine his prior work. He recognized eight species. E. arguta was reinstated as a species following its misapplication in his earlier work. He reduced four species to synonymy under EF. collina, a name which he considered more appropriate than Mueller’s EF. brownii for the “collective species”. Bentham’s E. collina was thus narrower in circumscription than Mueller’s E. brownii. E. paludosa was segregated as a variety of that species. However, Bentham agreed with Mueller that E. striata was “not separated from E. collina by any marked characters” and that Brown’s E. speciosa and E. alpina may be further variants of the latter. He retained E. cuspidata and E. alsa, but considered the latter to be conspecific with the South American E. antarctica. . Following this productive half century in the taxonomy of Euphrasia in Australia came a period of almost a century in which Australian authors largely chose to follow the pre-existing concepts of the genus. Mueller (1882, 1885-88, 1889) retained his viewpoint of the taxonomy of the genus and the usage of E. brownii in his subsequent works on the Australian and Victorian flora, although he did follow Bentham in reducing E. a/sa to a synonym of E. antarctica. Early Australian botanists tended to favour his concepts in the genus, e.g. in the works of Spicer (1878) on the Tasmanian flora, Tate (1890) on the South Australian flora, and Moore (1884), Moore & Betche (1893) and Dixon (1906) on the New South Wales flora. Bentham’s ideas concerning Euphrasia, which differed from Mueller’s only in his recognition of four closely related species under E. brownii, were followed in the majority of other floristic works, such as those of Woolls (1891) on the plants of the Sydney region, Bailey (1883, 1890, 1901, 1913) on the Queensland flora, Gardner (1931) on Western Australian plants, Ewart (1931) and Galbraith (1955, 1967) on the Victorian flora, and Black (1926) and Robertson (1957) on the South Australian flora. Rodway (1903) in his “Tasmanian Flora”, Ewart (1908) in his “Contributions to the Flora of Australia” series, and Maiden & Betche (1916) in their “Census of New South Wales Plants” went a stage further with Mueller’s ideas by proposing as ee neene * Dr P. van Royen (pers. comm. 1972) has since expressed some doubts as to the correctness of his conclusions concerning the legitimacy of E. brownii. 92 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia varieties of E. brownii (Rodway, Maiden & Betche) or E. collina (Ewart), some or all of the four species and one variety which Bentham distinguished in the complex. Mueller at no stage proposed this idea as Maiden & Betche state, except with E. striata. In a comprehensive and detailed monograph of Euphrasia, Wettstein (1896) described 12 species from Australia. The total lack of published reference to his treatise until 1965 :n Australian works, other than by Du Rietz (1948a,b), is note- worthy, for the classification of the Australian species was dealt with in great detail and differed significantly from previous works. Even the hardly controversial re- instatement of E. alsa was not taken up to that time. Important German works in other genera, such as Plantago, Epilobium and Blennodia, were also not followed during this period (Dr Hj. Eichler, pers. comm. 1973). Nevertheless, the taxonomic difficulties with Euphrasia, which continued to recent times (e.g. Willis 1967; Burbidge & Gray 1970), may have led to distrust of this revision produced by an overseas botanist who lacked recourse to field studies. Wettstein’s work was based entirely upon collections from European herbaria, but he failed to see those of the British Museum and Kew which housed most type specimens of the species described by Brown, Bentham and J.D. Hooker. As a result he misapplied the name E. arguta (see p. 284). E. glacialis and E. muelleri were described as new. No infraspecific taxa were recognized. Several species reduced by Bentham and Mueller and their followers under E. collina or E. brownii were reinstated. Substitute names were proposed for two species previously supplied with illegitimate names (later homonyms), the new name E. hookeri replacing E£. cuspidata Hook.f. non Host (1831), and E. diemenica Spreng. replacing EF. alpina R.Br. non Lamk. Mueller’s E. alsa was resurrected, E. antarctica being shown to belong to a separate section of the genus. In fact, Wettstein used characters identical to those proposed by Bentham (1846) to separate the South American species as a distinct section but which he (Bentham 1868) had subsequently ignored in his union of E. antarctica and E. alsa. Wettstein informally divided the Australian and New Zealand species on duration. Gandoger (1919) published seven new binomials in the “EZ. brownii complex” of Australia. Gandoger’s study was taxonomically superficial. Indeed, he had previously written a Flora of Europe which was highly unconventional (Stafleu 1967, 1972; McGillivray 1973). There is no evidence that he saw any material other than the single specimens upon which he based the names. Whereas such a course of action is non- scientific, his names are validly published with short diagnoses (in key form) and with specimens adequately cited. Until very recently when Briggs (in McGillivray 1973) supplied probable affinities of Gandoger’s species, workers in Euphrasia have rarely attempted to apply Gandoger’s names because of the brevity of the descriptive infor- mation and the lack of opportunity to see the pertinent types (cf. Du Rietz 1948b). Blake (1945) described and figured a distinctive new species E. bella from the mountains of southern Queensland. As a continuation of his earlier papers on Euphrasia in the southern hemisphere and Malesia (Du Rietz 1932a,b), Du Rietz (1948a,b) published two papers primarily to clarify the classification of the Tasmanian species. He went further than this, however, and discussed in some detail the taxonomy of the Australian species as a whole and their affinities in the genus elsewhere. His work was based mainly on material from European herbaria, including the important collections of the British Museum and Kew. A few specimens were sent to him from the herbarium of the Botany Department, University of Adelaide (now housed in AD). Throughout the work he mentioned possible distinct taxa which required further investigation. On such little material as he saw his predictions often proved remarkably good. Du Rietz described two new species, one of them, E. gibbsiae, with a number of 93 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) forms. Two of these forms he published formally, believing that, with further material and field studies, they might prove to be distinct species. He referred to several variants which he provisionally named as distinct species. While describing the second new species, E. gunnii, he rectified the misapplication of E. collina R.Br. by Bentham and subsequent authors. Like Wettstein he preferred to consider as distinct species the major variants in the “E. brownii complex”. Du Rietz was the only person to provide a formal infrageneric classification of the Australian species, with which the one proposed in the current infrageneric revision in many respects agrees. Finally, in view of the high polymorphy in E. gibbsiae, Du Rietz concluded that “possibly hybridism plays an important role in the Tasmanian Euphrasia-population”. Eichler (1965), on the basis of the Australian studies by Wettstein and Du Rietz, considered that several species were involved in South Australia under E. collina as circumscribed by Black (1926) and Robertson (1957). Curtis (1967) recognized eight species in her ‘Student’s Flora of Tasmania’. Her concepts also followed Wettstein and Du Rietz. She described one new species, FE. kingii, discussed the ecology and variability of the species, particularly of E. diemenica and E. gibbsiae, and noted Du Rietz’s comments on the role of hybridism in this variability. Willis (1967) published notes on Euphrasia in Australia, mainly on the taxa occurring in the Australian Alps. In acknowledging difficulty in’ taxonomic delimitation caused by intraspecific polymorphy and extensive hybridism, he preferred to use varietal rank in describing distinctive taxa, and questioned the status of several Tasmanian species recognized by Du Rietz. Under E. scabra he described a new alpine and subalpine variety caudata from the Australian Alps. This variety seemed to pass gradually into E. alsa, which he showed to be quite different from £. antarctica and which he reduced to a further variety of E. scabra. To Wettstein’s E. glacialis of the Kosciusko and adjacent regions, he added the new variety eglandulosa, abundant in the higher central parts of the Victorian Alps, but with outliers as far away as the Brindabella Range, Australian Capital Territory. Burbidge & Gray (1970) could provide only a tentative treatment of Euphrasia in the Australian Capital Territory. Of the five taxa recognized, only two were attributed published names. In extending his concepts in the ‘Handbook to Plants in Victoria’, Willis (1973) re- iterated doubts of past authors concerning the separation at the species level of E. collina and E. speciosa, which had been reinstated by Wettstein (1896) and Du Rietz (1948b). Six taxa were described and illustrated in Stones & Curtis’s (1967-78) ‘The Endemic Flora of Tasmania’. In the final part, following an exchange of correspondence, Curtis published lists including all taxa recognized by me as endemic to Tasmania. Most unpublished names and combinations were correctly cited as unpublished, but three (comb. & stat. nov.) were validated. In conclusion, throughout the various taxonomic works dealing with Euphrasia in Australia, doubts have been frequently expressed as to the status of most species recognized to this time. Thus, species placed in separate infrageneric taxa by one worker (e.g. Du Rietz 1948a,b) have been considered distinguishable only at the infraspecific level in a later publication (e.g. Willis 1967). In addition much difficulty has been found in applying correct names to the taxa (e.g. Burbidge & Gray 1970), and consequently names have sometimes been wrongly applied. As Willis (1967) stated, these deficiencies have obviously arisen from a lack of detailed taxonomic study of Euphrasia throughout Australia based on extensive field experience. Indicative of the inadequacy of previous concepts of the genus is the fact that almost half of the taxa recognized in the proposed classification are new; the proportion would be increased even further if Gandoger’s arbitrarily designated and never utilized new species were discounted. 94 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia C. TAXONOMIC PHILOSOPHY AND PRESENTATION The aim of the work has been to produce a classification of the genus in Australia reflecting the natural situation, thus providing a relatively stable framework into which any future discoveries can be fitted. Field work was undertaken in two of the main centres of diversity in Australia, encompassing two months in late 1970-early 1971 in Tasmania, and a year later a brief visit to the Grampians and six weeks in the Australian Alps. It has not been possible to visit the third important region of northern and central eastern New South Wales. In the course of the revision many problems have been encountered which, through paucity of material or need of field study, are unresolvable. In cases where distinct taxa are undoubtedly involved I have given them their postulated status and treated them informally as if species or subspecies, including distinguishing them in the keys. Already, early recognition of E. bowdeniae, E. phragmostoma and E. semipicta as distinct taxa on the basis of a few specimens has been supported by subsequent collections. It is suggested that in future ssp. diemenica (p. 185) and the ssp. paludosa -ssp. speciosa-ssp. diversicolor complex (p. 168) of E. collina may be usefully divided into varieties. Odd variants of more doubtful status and origin or apparently discordant localities are treated in notes under the appropriate taxon. Where such a variant might apply to several taxa, it is noted under the appropriate species or section to which they all belong. Determination of taxa and rank As far as possible in this work the basic unit for the natural delimitation of taxa has been the population. My collections have enabled characteristics of the population to be studied in the herbarium, for they are largely population samples made up of single specimens of each of the several to many plants collected. Variation in taxa of which I have little or no field experience, has been assessed on the basis of experience with other taxa. In determining rank, a combination of the number of characters, the extent of overlap in their variation and the degree of intergradation with allied taxa has formed the primary basis. It became clear that there were two or possibly three main levels in the natural grouping of taxa near species level. A formal species-subspecies hierarchy (with possible potential extension to variety) was eventually decided upon. The view that in large complexes this leads to a system which is cumbersome to use (Yeo 1978b) is disputed, as in the proposed formal framework only a trinomial has to be used, identical to that widely accepted in species with few infraspecific taxa. Such a system gives the user not only the benefit of a natural system, but also the practical advantage of being able to avert the unavoidable difficulties of subspecific determination in a complex where the species name would amply suit his needs. Furthermore, it is believed that recognition of Australian subspecies as “microspecies”, which has been done elsewhere in the genus (Sell & Yeo 1970; Yeo 1978b), would camouflage the well- defined species in Australia. From autecological, breeding and herbarium studies Karlsson (1974, 1976) believes that the Swedish taxa, traditionally recognized principally as species, can be more naturally treated under a species-subspecies system, although his conclusion suffers from coverage of only a portion of the range of the complexes involved. In his treatment of Euphrasia in central Europe, Hartl (1972) has also used a broader species concept in association with an infraspecific hierarchy incorporating subspecies, varieties, subvarieties and forms. The species in this revision are distinct on a number of characters, some of which show little overlap. They show no evidence of complete clinal intergradation with their closest allies, although there is occasionally limited hybridization between species in 95 W. R. Barker J. Adelaide Bot. Gard. 5 (1982) different sections (p. 287 ), often involving an increase in pollen sterility in the hybrids. Subspecies have been designated because of their substantially fewer character differences and/or greater degree of overlap of variation compared with the species. They often intergrade morphologically with their closest relatives on an ecotonal or geographical basis. In such cases tests of pollen sterility (p. 40) show that the intergrades may differ little in sterility from parent taxa. Because of the long history of controversy about the rank of the taxa of Euphrasia in Australia and elsewhere, it is essential to highlight the degree of distinction of taxa from their closest allies by demonstrating the amount of clinal intergradation between taxa, particularly using studies in the field (treated under the observations on the nature of polymorphism in relevant species and E. collina ssp. diemenica), and by indicating all important diagnostic characters. Descriptions and keys In relation to the content of descriptions and keys, Yeo’s (1978b) comments on his own experience with the genus are pertinent and echo my own views: “Bearing in mind the minor characters that have to be used to distinguish the species [his “microspecies”: equivalent to subspecies here], it should be obvious that here is a group where it is especially important to hold fast to the principle that in classification as many characters as possible should be taken into account. There are many groups in which this principle can be ignored at the species level, since it is possible to separate absolutely the members of any pair of taxa by the use of one or two ‘key characters’.” He then refers to the breakdown of such weighted characters in the European species. : Detail in the keys is particularly important in this revision, as it provides a more complete summary of the differences between related taxa than is often found in discussion of relationships traditionally placed under each taxon. The keys provide a ready summary of the number and degree of overlap of characters used in assessing the rank at which taxa are separated. In using keys in this complex group, all characters must be treated equally, and ideally the variation of several plants from a population should be determined. At any point in a key where character states overlap, the greater the number of diagnostically important characters there are, the more certain will be the identification. This also makes for more reliable identification of individual specimens. I can see no better way of providing a simple and as reliable Australia-wide key to taxa. Simplification should be possible, however, in regional accounts. Intergradation between taxa As already intimated, two types of intergradation between Australian taxa of Euphrasia are apparent. The first involves intergradation of a primary nature between closely-related (apparently sister) taxa, e.g. ssp. comberi and ssp. kingii of E. gibbsiae, ssp. collina and ssp. diemenica of E. collina, and ssp. diversicolor and ssp. glacialis of E. collina. These intergrades occur on narrow to broad ecotones. As there is little or no evidence of increased pollen sterility in the intermediates, it is believed that these represent a stage in a process of vicarious or parapatric speciation, which seems to have been the principal method of diversification in Australia. The clines evident within taxa such as ssp. collina and ssp. tetragona of E. collina are considered to represent incipient phases of this process, while the vicarious-parapatric pattern of distribution in Sect. Lasiantherae and its closer allies in Sect. Scabrae, and vicarious pattern in the &. bella-E. sp. ‘Tamworth’-E. bowdeniae lineage are a logical end- point of the process following isolation of parts of clines and subsequent divergence of the isolated populations. 96 J. Adelaide Bot. Gard. 5 (1982) Studies in Euphrasia The secondary type of intergradation involves hybridization or introgression of distantly related taxa. On the available data most examples evident today exhibit high pollen sterility and occur in disturbed sites rather than natural ecotones. An exception would be possible introgression in E. gibbsiae ssp. pulvinestris (p. 132). Furthermore, it is proposed that in the evolution of the genus a few taxa, namely E. orthocheila ssp. peraspera, E. semipicta and possibly E. crassiuscula, may have originated through hybridization or have been modified through introgression. .All of these extant and historic cases involve parent taxa which are apparently not sister taxa (i.e. which are not derived directly by divergence from the same ancestral taxon). Most of the proposed parent taxa occupy different sections of the genus, but E. orthocheila ssp. peraspera and E. collina ssp. paludosa x ssp. diversicolor are exceptions. The relative importance of hybridization in intergrade situations in Australian Euphrasia at first seems to contrast greatly with the substantial hybridization and introgression within the ecotypically differentiated microspecies of Europe (Yeo 1978b). However, the “hybrid swarms” of Yeo (1978b) occur in disturbed or intermediate habitats and are most frequent between closely related species growing nearby. Since in these instances intermediates are highly fertile, some of these may be analogues to the primary type of intergradation described above. Order of presentation of the taxa Taxa are ordered in a sequence which as far as possible places closest allies adjacent. Species begin with that postulated to be closest to the progenitors of the section or lineage. Subspecies begin with the autonymous taxon. Distribution and conservation status Many of the Australian taxa of Euphrasia are geographically and ecologically restricted. Some taxa are confined to rare localized habitats, e.g. in alpine habitats, E. collina ssp. lapidosa of fjaeldmark, E. collina ssp. glacialis of moist communities with short herbage, and FE. gibbsiae ssp. pulvinestris of blanket bog, and at lower elevation probably the cliff face home of E. bella, E. bowdeniae and E. phragmostoma. Other taxa, particularly of montane and lowland regions, occur in a range of often more widespread communities, but even some of these taxa, e.g. ssp. fetragona and ssp. osbornii of E. collina, are rare, if occasionally forming large local populations. A number of taxa, i.e. E. gibbsiae ssp. psilantherea, E. arguta, E. collina ssp. muelleri and unnamed taxa such as an annual in the Blue Mountains, New South Wales (p. 284) and E. sp. ‘Tamworth’, are known only from old specimens. The lack of collections may reflect extinction, but cases such as the recent discovery of E. phrag- mostoma after 80 years’ absence from collections, and single recent collections of each subspecies of E. orthocheila, previously not collected for decades, indicate that some of these taxa may still survive. Conservation status of each taxon is provided using the formulae of Leigh, Briggs & Hartley (1981). The numerical rarity code relates to distributional range, 2 being very restricted, e.g. less than 100km, and 3 being for wider ranges of taxa with small local populations. The letter risk codes are: X, not collected in recent years, believed extinct; E, endangered, in risk of disappearing in 1-2 decades of continued present pressure; V, vulnerable, not endangered but at risk through depletion or if potential changes to land use threaten survival; R, rare but not endangered. The addition of a letter C indicates occurrence in a National Park or proclaimed reserve. 97 ; W. R. Barker J. Adelaide Bot. Gard. 5 (1982) EUPHRASIA L. For synonymy, description, typification and distribution, see p. 73. KEY TO THE SECTIONS OF EUPHRASIA IN AUSTRALIA la. Anthers glabrous about connective. [ Perennial.] 2a. Rearmost pair of anther awns (1.5)2.5-3.2 mm long, as long as or longer than anthers, needle-shaped or sometimes distally twisted or dilated and erosulate. Corolla lacking striations. [Branches or shoots on main inflorescence-bearing axes occurring at widely spaced groups of consecutive WeulHbastecoqsogcsanrc Il. Sect. Phragmostomae (p. 102) 2b. Rearmost pair of anther awns 0.2-0.8 mm long, shorter than anthers, needle-shaped, entire. Corolla partially to extensively striated. 3a. Branches or shoots on main inflorescence-bearing axis(es) developing in basi- petal sequence in consecutive axils from I-few nodes below inflorescence. Aah ern danepGnadonsbucwnaie onided