The Graminicolous Rust Fungi of New Zealand

R. F. R. McNabb

By

Plant Diseases Division, Auckland

[Received by the Editor, September 11, 1961.]

Abstract

Sixteen species of grass rusts are described and illustrated, including one new species and three species not previously recorded in New Zealand. Twenty-six new host records are given. The number of rusts recorded in New Zealand now totals 174.

Since the publication of Cunningham’s The Rust Fungi of New Zealand in 1931 there have been numerous alterations and additions to the graminicolous rust fungi of New Zealand.

Publications by overseas workers have necessitated nomenclatural changes affecting two rusts—viz., Puccinia hordei and P. recondita, previously recorded by Cunningham (1931) as P. anomala and P. elymi respectively.

Re-examination of New Zealand material of P. poarum shows it to agree with P. poae-nemoralis, a species not separated from P. poarum when Cunningham’s (1931) recording was made. P. poarum is now deleted from the list of New Zealand rusts.

Two species of Uromyces not previously recorded from this country have been collected, and teleutospores of Uredo crinitae have been found, which enables the rust to be placed in Puccinia as a new species. P. cynodontis recorded by Baker (1956) and P. flavescens (McNabb 1961) are also included, bringing the total number of grass rusts in New Zealand to 16. The 16 species occur on 60 hosts within the Gramineae, with 14 hosts carrying two rust species, thus giving a total of 74 rust-host combinations. Twenty-six new host records are given.

An interesting feature of the graminicolous rusts in New Zealand is the lack of host alternation. Of the eight known heteroecious species, in only one (Uromyces dactylidis) has the alternate host been found, and even in this species the alternation appears to be facultative rather than obligatory. The remaining seven heteroecious species appear to be non-obligatory and are able to overwinter by means of uredospores. Some of these rusts seldom or never produce teleutospores on certain hosts. It appears that host alternation in countries with mild winters is occasional or lacking, and that it is not necessary for the maintenance

of the rust. The absence of many alternate hosts in New Zealand probably accounts for the lack of host alternation in some species. The host alternation of the remaining eight species (with the possible exception of Uromyces danthoniae which may be autoecious) is not known.

Four rusts, Puccinia crinitae, Uromyces otakou, Uredo karetu and U. toetoe are according to the present knowledge endemic to New Zealand; a further three species, Puccinia flavescens, Uromyces danthoniae and U. ehrhartae are restricted to Australasia, while the remaining nine species are more or less cosmopolitan in distribution. Two cosmopolitan rusts ( Puccinia graminis and P. recondita) occur on hosts which are endemic or indigenous to New Zealand.

It is probable that some of the species at present considered endemic to New Zealand will be found in Australia when the rusts of that country are more thoroughly investigated. Hansford (pers. comm.) has found but not described, the uredo and teleuto stages of a rust on Dichelachne crinita from Australia, which seems to be identical with Puccinia crinitae. The writer wishes to thank Miss J. M. Dingley for her helpful advice, Mr V. D. Zotov, Botany Division, Lincoln, for identifying a number of the hosts and checking their nomenclature, and Professor L. W. A. Crawley, Classics Department, University of Auckland, for the Latin description of one species.

Materials and Methods For the examination of germ pores and spore markings, spores were heated in 0.2% lactophenol. Details of sori were obtained from hand-cut sections mounted in lactophenol-aniline blue (Cunningham 1931). Plant Diseases Division Herbarium accession numbers are given after each new host record and new rust recording; collections of recognised rust species and host recordings published prior to this paper are not cited. Conventional symbols for the various spore forms are placed after each host name and new host records are marked with an asterisk.

Key to the New Zealand Grass Rusts 1. Teleutospores predominantly 2-celled ( Puccinia) 2. Teleutospores coronate P. coronata Corda 2. Teleutospores not coronate . (3) 3. Uredospores with scattered germ pores ...... (4) 4. Uredosori with hyaline, capitate paraphyses (5) 5. Uredo paraphyses thick-walled P. poae-nemoralis Otth. 5. Uredo paraphyses thin-walled P. crinitae McNabb 4. Uredosori without hyaline, capitate paraphyses (6) 6. Uredospore germ pores 7-12 P. hordei Otth 6. Uredospore germ pores 5-8 ...... ...... P. recondita Rob. ex Desm. 6. Uredospore germ pores 4—7 P. flavescens McAlp. 3. Uredospores with equatorial germ pores ...... (7) 7. Uredospores echinulate ...... (8) 8. Teleutospores oblong-clavate, to 70m

long P. graminis Pers. 8. Teleutospores elliptical, to 40m long P. sorghi Schwein. 7. Uredospores verrucose P. cynodontis Lacroix in Desm. 1. Teleutospores 1-celled ( Uromyces) 9. Apex of teleutospores not or slightly thickened ; ...... (10) 10. Uredospore germ pores scattered U. dactylidis Otth 10. Uredospore germ pores equatorial U. tenuicutis McAlp. 9. Apex of teleutospores strongly thickened (11)

11. Apex acuminate U. danthoniae McAlp. 11. Apex rounded U. otakou G. H. Gunn. 11. Apex variable U. ehrhartae McAlp. 1, Teleutospores not known (Form Genus Uredo) 12. Epispore of uredospores hyaline to pallid yellow Uredo karetu G. H. Gunn. 12. Epispore of uredospores chestnut brown Uredo toetoe G. H. Gunn.

Puccinia coronata Gorda, Icon. Fung., 1: 6, 1837. (Text-fig. I, fig. 1.) Puccinia lolii Niels., Ugeskr. Landm. IV: 9, 549, 1875. P. rhamni Wettst., Verb. zool. bot. Ges. Wein, 35; 545, 1886. P. coronata var. himalensis Bard., Trans. Linn. Soc. Lend. (Bot.) 3: 227, 1891. P. coronifera Kleb., Z. PflKrankh. 4: 135, 1894. P. catharticae Lagerh., Tromso Mus. Aarsh. 17; 52, 1895. P. himalensis (Bard.) Diet., Nat. Pflanzenfam. 1: 63, 1900. P. beckmanniae McAlp., Rusts Austr. 116, 1906. P. pregracilis Arth., Bull. Torrey bot. Cl. 34; 585, 1907. P. erikssonii Bubak, Arch, naturw. Landes Forsch. Bohm. 13; 107, 1908. P. diarrhenae Miyabe & Ito, J. Coll. Agric. Sapporo, 3: 190, 1909. P. hierochloae Ito, loc. cit. 3: 193, 1909. P. alpinae-coronatae Muehlt., Zbl. Bakt. 11, 30: 418, 1911. P. mediterranea Trott., Ann. mycol. 10: 510, 1912. P. avenae-barbatae Frag., 801. Soc. Broter. 2: Ser. 2,7, 1923.

O, I. Unknown in New Zealand. 11. Uredosori amphigenous and on sheaths, at first scattered and elliptical, becoming aggregated and often linear, frequently reaching 2-3 mm long, bullate, brownish yellow, becoming pulverulent and naked; spores globose, elliptical or obovate, 20-34 x average 27 x 22ja; epispore pallid yellow, 1.0-1.5/x thick, finely and moderately echinulate; germ pores scattered, 6-8, indistinct; hyaline paraphyses present or absent. 111. Teleutosori amphigenous, scattered, up to 0.5 mm long, or confluent and linear when up to 2-3 mm long, black, bullate, compact, long covered; spores elongate-clavate, often cuneiform, 36-64 x 12-26 fi; apex truncate, thickened to 7/i, darker in colour, crowned with a variable number (5-9) of curved, digitate processes, variable in size and shape, frequently reaching 12ja long; base attenuate, basal cell longer, narrower, and paler in colour; not or slightly constricted at septum, septum often oblique; epispore smooth, pallid chestnut brown, 1.0-1.5/4 thick; pedicel persistent, coloured, to 20 x 10/i; mixed with and separated into compartments by coloured, subclavate paraphyses.

Type Locality. Europe, on “ Luzula albida” (= ? Bromus erectus Huds.) Hosts. Agrostis canina L.—II, III.* Auckland: Mt. Albert, December, 1960, R.F.R.Mc., 19534. Agrostis tenuis Sibth.—ll, 111. Agrostis stolonifera L.—II, III.* Auckland: Mt Albert, October, 1960, R.F.R.Mc., 19501; Mt Albert, December, 1960, R.F.R.Mc., 19700. Nelson: Karamea, February, 1961, R.F.R.Mc., 19604. Aira caryophyllea L.—ll.* Auckland: Karapiro, December, 1960, R.F.R.Mc., 19571. Arrhenatherum elatius (L.) J. & C. Presl.-—II, III.* Canterbury: Darfield, February 1961, R.F.R.Mc., 19616; Ladbrooks, February, 1961, R.F.R.Mc., 19623.

Wellington; Wanganui, February, 1961, R.F.R.Mc., 19578. Arrhenatherum elatius var. bulbosum (Willd.) Spenn. —11, lII.* Auckland: Mt Albert, December, 1960, R.F.R.Mc., 19541. Avena fatua L.—II, lII.* Auckland; Mt Albert, December, 1960, R.F.R.Mc., 19522; Owairaka, February, 1961. R.F.R.Mc., 19581.

Avena sativa L.—II, 111. Holcus lanatus L.—II, 111. Holcus mollis L.—II, 111. Lolium multiflorum Lam.—ll. Lolium perenne L.—II. Lolium temulentum L.—II. Phalaris minor Retz.—ll. Poa annua L.—ll.* Auckland: Mt Albert, December, 1960, R.F.R.Mc., 19885. Poa palustris L.—ll.* Marlborough: Koromiko, January, 1947, A. J. Healy, 5363. Poa pratensis L.—ll.* Auckland; Taupo, December, 1960, R.F.R.Mc., 19560; Karapiro, December, 1960, R.F.R.Mc., 19572. Canterbury: Lewis Pass, April, 1950, A. J. Healy, 7216; Waiau, December, 1954, A. J. Healy, 14206. Marlborough; Parikawa, December, 1946, A. J. Healy, 5370. Poa iridifolia Hauman —ll.* Wellington: Lower Hutt, October, 1953, A. J. Healy, 12691.

All of the hosts are introduced grasses.

Puccinia coronata s.lat. as here defined, has been shown to consist of numerous races separable by their reactions toward certain differential hosts. A number of these races have been considered separate species, but these cannot be maintained either on morphological or biological grounds. Cummins (1956) lists 21 synonymous species under the P. coronata complex and points out that amongst the synonyms of P, coronata s.lat. differences are mainly those of spore dimensions, length of crown digitations, presence or absence of uredo paraphyses and exposure of teleutosori. Cruickshank (1956, 1957) investigated the incidence of P. coronata on Avena sativa, Lolium multiflorum and L. perenne in this country and identified 13 physiological races of Puccinia coronata on oats in New Zealand.

Teleutospores have not been found on Lolium and Poa despite intensive searching. It seems probable that teleutospores are rarely produced on these hosts although uredospores are common on Lolium species during the winter. Jorstad (1950) comments that in Norway only the uredo stage has been found on L. perenne and L. multiflorum. Puccinia coronata is a heteroecious species with the aecidial hosts known to

be Rhamnus and Berchemia in the Rhamnaceae and Elaeagnus and Shepherdia in the Elaeagnaceae. The usual hosts are not naturalised in this country, and no alternate host has been established for the rust in New Zealand. The uredospores of Puccinia coronata and P. recondite are scarcely distinguishable on hosts such as Agrostis tenuis. Jorstad (1950) experienced the same difficulty. Uredospores of Puccinia coronata generally have thin, pale yellow walls and indistinct germ pores whereas those of P. recondite have thicker, yellow-brown walls with conspicuous germ pores. P. coronata is characterised by uredospores with pallid yellow walls and 6-8 indistinct germ pores, and teleutospores with coronate apices. Puccinia poae-nemoralis Otth, Mitt, naturf. Ges. Bern, 1870, 113, 1871. (Textfig. I, fig- 2.)

Puccinia poarum auct. non P. poarum Niels., Bot. Tidsskr. 111, 2: 34, 1877. P. perplexans Plowr. f. arrhenatheri Kleb., Abh. naturw. Ver. Bremen, 12: 366, 1892. P. milii Erikss., Fung. Paras. Scand. 450, 1895. P. arrhenatheri (Kleb.) Erikss., in Cohn, Beitr. Biol. Pfl. 8: 1, 1898. P. cognatella Bubak, Ann. mycol. 7: 378, 1909. P. poae-sudeticae (Westend.) Jorst., Nyt Mag. Naturv. 70; 325, 1932. P. anthoxanthina (Bubak) Gaum., Ber. schweiz. bot. Ges. 55: 74, 1945.

O, I. Unknown in New Zealand.

11. Uredosori mostly epiphyllous, scattered, orbicular when up to 0.5 mm diameter, or elliptical and up to 1 mm long, pulverulent, orange at first, becoming cinnamon brown; spores elliptical, 21-29 x 17-24//, average 25 x 21/t; epispore pallid yellow or almost hyaline, 1.5-2.0//, thick, densely and finely echinulate; germ pores scattered, 8-10; surrounded by and mixed with numerous clavate or capitate, hyaline, thick-walled paraphyses giving the sori a compact, cushion-like appearance.

111. Teleutosori hypophyllous, scattered, sparse, elliptical, to 2 mm long, pulvinate, deep chestnut brown, long covered; spores elliptical or subclavate, 35-53 x 15-22/x, average 48 x 20//; apex truncate or rounded, thickened to 6/t; base attenuate, basal cell narrower; slightly or not constricted at septum; epispore smooth, chestnut brown, 1,0-1.5// thick; pedicel persistent, short, tinted.

Type Locality. Switzerland, on Poa nemoralis L. Hosts. Anthoxanthum odoratum L. —ll.* Auckland: Mt Albert, October, 1960, R.F.R.Mc., 19506; Mt Albert, December, 1960, R.F.R.Mc., 19554. Canterbury: Stewart’s Gully, December, 1954, A. J. Healy, 14303. Wellington: Mangaroa, October, 1952, A. J. Healy, 11714, Poa annua L.—II. Poa pratensis L.—II.

All three hosts are common introduced grasses. The rust, which is an introduced species, was previously recorded in New Zealand under Puccinia poarum by Cunningham (1931). Jorstad’s (1932) interpretation of P. poae-nemoralis (P. poaesudeticae ) as a polymorphous species embracing a number of races showing only slight morphological differences, is generally acceptable to most workers (Wilson & Bisby 1954,

Cummins 1956). Gaumann (1941 b) criticises the erection of P. poae-nemoralis and considers the presence or absence of uredo paraphyses a labile character. Teleutospores of a rust on Anthoxanthum odoratum have been described by Gaumann (1945) under Puccinia anthoxanthina. The aecidial stage of this rust is not known. Viennot-Bourgin (1949) follows Gaumann (1945) and retains P. anthoxanthina for the rust with clavate uredo paraphyses on Anthoxanthum odoratum.

Little is known about the physiological specialisation of Puccinia poae-nemoralis and the alternate host is known only for the race on Arrhenatherum elatius which alternates with Herberts vulgaris (Gaumann 1934). According to Jorstad (1950) even on this host the alternation is non-obligatory. No alternate host has been established for Puccinia poae-nemoralis in New Zealand, but abundant uredosori produced in autumn and winter indicate that it is perpetuated by uredospores. Teleutospores have not been collected in this country and according to Jorstad (1950, 1958) the teleuto stage is rare. In Scandinavia, Hylander, Jorstad and Nannfeldt (1953) have been unable to find teleutospores on Anthoxanthum odoratum, Poa annua and other grasses, but report that they are fairly common on P. pratensis.

Puccinia poarum, with which this species has been confused, was originally described by Nielsen (1877) on Poa pratensis and proved by him to form aecidiosori on Tussilago farfara. Nielsen (1877) does not mention capitate, thick-walled uredo paraphyses in his original description, and Plowright (1889) who repeated Nielsen’s experiments specifically states that uredo paraphyses are absent. Jorstad (1932) repeated these experiments, and in no case did the resulting uredosori possess capitate paraphyses. Puccinia poarum appears to be an obligatory heteroecious species alternating with Tussilago farfara and species of Petasites. None of these hosts occur as grassland species in New Zealand.

Puccinia poae-nemoralis is characterised by abundant, thick-walled, capitate uredo paraphyses, a feature absent from other grass rusts in New Zealand, although P. crinitae possesses thin-walled, capitate paraphyses in the uredosori. Puccinia crinitae sp. nov. (Text-fig. 111, fig. 1.) Uredo crinitae G. H. Gunn., Trans. N.Z. Inst., 55: 41, 1924. O, I. Ignota.

11. Uredosori amphigeni, in maculis discoloribus sedentes, dispersi, non saepe confluentes, elliptici, ad 1 mm longi, rubro-aurantiaci, pulverulenti, epidermi rupta circumdati; sporae globosae vel subglobosae, 28-37 (31) x 24-32 (27) //,; epispora pallido-flava, subtiliter e.t presse echinulata, 2.0-2.5//, crassa; pori germinis dispersi, crebri, 10-14, magni, commixti paraphysibus tenuiparietibus, hyalinis, capitatis vel subclavatis.

111. Teleutosori sparsi, epiphyllii, nigri, elliptici, ad 0.5 mm longi, compressi, diu operti; sporae elongato-oblongae vel subclavatae, 36-50 (43) x 11-22 (18)/a; apex truncatus vel obtuse rotundatus, interdum obliquus, ad 4//, crassatus, colore fusciore; basis attenuata, cellula basali plerumque longiore, angustiore, colore clariore; ad septum leviter constricta vel non constricta; episporium leve, castaneo-fulvum, 1.5—2.0//, crassum in cellula superiore, 1.0-1.5//, in inferiore; pedicellus deciduus, coloratus, ad 15 x 6//,.

Mesosporae crebrae, clavatae vel inaequabiliter elongatae, 19-47 x 11-18//,. O, I. Unknown.

11. Uredosori amphigenous, seated on discoloured spots, scattered, seldom confluent, elliptical, to 1 mm long, reddish-orange, pulverulent, surrounded by ruptured epidermis; spores globose to subglobose, 28-37 x 24—32//,, average 31 x 27//,; epispore pallid yellow, 2.0-2.5//, thick, finely and closely echinulate; germ pores scattered, numerous, 10-14, large; mixed with thin-walled, hyaline, capitate or subclavate paraphyses.

111. Teleutosori sparse, epiphyllous, elliptical, to 0.5 mm long, black, compact, long covered; spores elongate-oblong to subclavate, 36-50 x 11-22//., average 43 x 18//,; apex truncate or bluntly rounded, sometimes oblique, thickened to 4//,, more deeply coloured; base attenuate, basal cell usually longer, narrower and lighter in colour; not or slightly constricted at septum; epispore smooth, chestnut brown, 1.5-2.0//, thick in upper cell, 1.0-1.5//, in lower; pedicel deciduous, coloured, to 15 x 6//,.

Mesospores numerous, clavate or irregularly elongate, 29-47 x 11-18//,. Type Locality. New Zealand, on Dichelachne crinita (Forst. f.) Hook. f. Host. Dichelachne crinita (Forst. f.) Hook. f. —II, lII.* Auckland: Karapiro, December, 1960, R.F.R.Mc., 19516; Moumoukai Valley, January, 1961, J. M. Dingley, 19690. Canterbury: Akaroa, January, 1922, W. D. Reid, 760, Type of Uredo crinitae; Porter’s Pass, February, 1961, R.F.R.Mc., 19630.

Marlborough: Pelorus Bridge, February, 1961, R.F.R.Mc., 19630. Wellington: Mt Victoria, February, 1961, R.F.R.Mc., Type Collection P.D.D. Herb. No. 19636. Dichelachne crinita is an indigenous species extending to Australia and Tasmania (Gheeseman, 1925, p. 163).

Puccinia crinitae is characterised by large globose or subglobose uredospores with numerous scattered germ pores, thin-walled, hyaline, capitate uredo paraphyses, and numerous mesospores in the teleutosori.

Puccinia hordei Otth, Mitt, naturf. Ges. Bern, 1870, 113, 1871. (Text-fig. I,

fig. 3.) Puccinia hordei Fckl., Jb. nassau Ver. Naturk. 27-28: 16, 1873. Uromyces hordei Niels., Ugeskr. Landm. 4: 9,1, 567, 1875. Puccinia anomala Rostrup., apud Thilm., Herb. Myc. Oecon. 451, 1878. P. rubigo-vera var. simplex Kcke. ex Wint., in Rabh. Krypt.-Fl. Ed. 2,1: 1, 218, 1882.

P. simplex (Kcke.) Erikss. & Henn., Landtbr. Akad. Handl. Tidsskr. 1894, 175, 1894; Z. Pfl. Krankh. 4: 260, 1894. (non P. simplex Peck, Rep. State Mus. N.Y. 34; 41, 1881.) P. hordei-murini Buchw., Ann. mycol. 41: 308, 1943. O, I. Unknown in New Zealand.

11. Uredosori amphigenous and on culms, scattered or confluent, to 2 mm long, cinnamon brown, pulverulent; spores elliptical or obovate, 23-34 x 17-27ju,, average 28 x 22/x; epispore pallid brown, 1.5-2.0/x thick, sparsely and finely echinulate; germ pores scattered, 8-12.

111, Teleutosori amphigenous and on culms, scattered or confluent, elliptical, to 1.5 mm long, cinnamon brown, pulvinate, compact, long covered; spores subclavate or cuneiform, 36-66 x 16-28/x; apex truncate, obtuse or bluntly acuminate, sometimes obscurely lobed, thickened to Bfi8 fi or not; base attenuate, basal cell cuneiform; slightly constricted at septum; epispore smooth, chestnut brown, 1.5-2.Oja thick; pedicel persistent, coloured, to 20 x Bft, continuous with spore; mixed with and separated into compartments by numerous brown clavate paraphyses.

Mesospores present, clavate or angularly obovate, 26-50 x 14-24ja. Type Locality. Switzerland, on Hordeum vulgare L. Hosts. Hordeum murinum L. —II, 111. Hordeum vulgare L. —II, 111.

Both hosts are introduced species. Puccinia hordei is an introduced rust previously recorded in New Zealand by Cunningham (1931) as P. anomala.

The rust on Hordeum murinum is sometimes recognised as a separate species, Puccinia hordei-murini (Buchwald 1943, Jorstad 1958). Buchwald (1943) maintains that P. hordei-murini differs from P. hordei Otth in possessing more germ pores in the uredospores (an average of 10-12 as against 8-9) ; a lower percentage of mesospores (usually around 28% and scarcely ever exceeding 40% as against 80% and never falling below 50%); and slightly larger teleutospores, ViennotBourgin (1941) also places emphasis on the percentage of mesospores to distinguish between the two species.

Oliveira (1937) maintains that the differences are not significant, while Jorstad (1958) believes that the distinguishing characters as outlined by Buchwald (1943) are relative, and states that separation on morphological grounds seems hardly possible. Despite this, Jorstad (1958) retains P. hordei-murini as a separate species because he is uncertain of its affinities. Inoculation experiments are not conclusive.

The rust on Hordeum murinum is classified here as a clearly defined race of Puccinia hordei Otth, the host alternation of which is unknown. Cummins (1956) places it in the compound species P. recondita, but both Oliveira (1937) and Jorstad (1958) point out that mesospores occasionally occur more abundantly than two-celled teleutospores, a feature which does not occur in P. recondita but which is the rule in P. hordei Otth.. The nomenclature of the barley rust has been clarified by Buchwald (1943).

P. hordei is a heteroecious species and aecidiosori shown by Tranzschel (1914) to occur on species of Ornithogalum, have not been collected in New Zealand.

The rust appears to be independent of its aecidial host in this country; Jorstad (1958) believes that probably it is everywhere independent of host alternation. Puccinia hordei is characterised by the numerous scattered germ pores of the uredospores and abundant mesospores in the teleutosori.

Puccinia recondita Roberge ex Desmazieres, Bull. Soc. bot. Fr. 4; 798, 1857. (Text-fig. I, fig. 4.)

Uredo rubigo-vera DC. FI. Fr. 5: 83, 1815. Puccinia rubigo-vera (DC.) Wint., in Rabh. Krypt.-Fl. Ed. 2, I, 1: 217, 1882. P. perplexans Plowr., Quart. J. micr. Sci. 25: 164, 1885. P. persistens Plowr., Brit. Ured. 180, 1889. P. apocrypta Ell. & Tracy, J. Mycol. 6: 76, 1890. P. agrostidis Plowr., Gdnr’s Ghron. Ser. 3,8: 139, 1890. P. agropyri Ell. & Everh., J. Mycol. 7: 131, 1892. P. borealis Juel, Ofvers Vetensk Akad. Forh. Stockh. 1894, 411, 1894. P. dispersa Erikss. & Henn., Z. Pfl. Krankh. 4: 141, 1894. P. aquilegiae Lagerh., Tromso Mus. Aarsh. 17: 47, 1895. P. clematidis Lagerh., loc. cit. 17: 54, 1895. P. adspersa Diet. & Holw., Erythea, 3: 81, 1895. P. dispersa Erikss., Ann. Sci. nat. Ser. 8,9: 268, 1898. P. bromina Erikss., loc. cit. 9: 268, 1898. P. triticina Erikss., loc. cit. 9: 270, 1898. P. agropyrina Erikss., loc. cit. 9: 273, 1898. P. holcina Erikss., loc. cit. 9: 274, 1898. P. symphyti-bromorum F. Muell., Beih. bot. Zbl. 10: 201, 1901.

P. actaeae-agropyri Ed. Fisch., Ber. schweiz. bot. Ges. 11: 8, 1901. P. brachypus Speg., An. Mus. nac. B. Aires, Ser. 3,1: 61, 1902. P. triticorum Speg., loc. cit. 1: 65, 1902. P. impatientis Arth., Bot. Gaz. 35: 19, 1903. P. cinerea Arth., Bull. Torrey bot. Cl. 34: 583, 1907. P. perminuta Arth., loc. cit. 34: 584, 1907. P. glyceriae Ito, J. Coll. Agric. Sapporo, 3: 200, 1909. P. elymi-sibericae Ito, loc. cit. 3: 202, 1909. P. bromi-japonicae Ito, loc. cit. 3: 205, 1909, P. fujiensis Ito, loc. cit. 3: 210, 1909. P. alternans Arth., Mycologia, 1: 248, 1909. P. obliterata Arth., loc. cit. 1: 250, 1909. j P. actaeae-elymi Eug. Mayor, Ann. mycol. 9; 361, 1911. P. secalina Grove, Brit. Rust. Fungi, 261, 1913. P. madritensis Maire, Bull. Soc. Hist. nat. Afr. N. 10: 145, 1919. P. bromi-maximi Guyot, Ann. Ec. Agric. Grignon, Ser. 3,2: 105, 1940-41. P. clematidis-secalis Dupais, Bull. Soc. mycol. Fr. 64: 182, 1948.

O, I. Unknown in New Zealand.

11. Uredosori amphigenous, scattered or crowded, elliptical or linear, 0.5-2.0 mm long, rarely confluent, light ferruginous brown, pulvinate, becoming pulverulent; spores subglobose, ovate or elliptical, 23-36 x 17-28/a; epispore finely and closely echinulate, 1.5-2.0/* thick, yellowish-brown; germ pores scattered, 6-8, conspicuous.

111. Teleutosori mostly hypophyllous and on culms, elliptical or more frequently linear when up to 3 mm long, black, compact, long covered; spores elongate -oblong to subclavate, 34-65 x 14-22 /*; apex truncate, rounded or bluntly acuminate, often oblique, thickened to 10/*, more deeply coloured; base attenuate, basal cell longer, paler in colour; slightly constricted at septum; epispore smooth, chestnut brown, 1.5-2.0/* thick in upper cell, 1.0-1.5/* in lower; pedicel persistent, coloured, stout, continuous with spore, to 20 x 7/*, divided into compartments and surrounded by dark brown, stout, clavate naraphyses.

Type Locality. France, on Secale cereale L. Hosts. Agropyron repens (L.) Beauv.—ll, III.* Auckland: Mt Albert, August, 1956, J. M. Dingley, 14320. Canterbury: Islington, November, 1954, A. J. Healy, 14232; Riccarton, April, 1954, A. J. Healy, 14239; Aranui, January, 1955, A. J. Healy, 14235; Lincoln, June, 1959, H. C. Smith, 18969; TaiTapu, February, 1961, R.F.R.Mc., 19601; Ladbrooks, February, 1961, R.F.R.Mc., 19602. Marlborough: Havelock, May 1959, A. J. Healy, 19070.

Agropyron scabrum (Labill.) Beauv.—ll, lII.* Auckland: Taupo, December, 1960, R.F.R.Mc., 19600. Marlborough: Pelorus Bridge, February, 1961, R.F.R.Mc.. 19592; Havelock, February, 1961, R.F.R.Mc., 19590.

Nelson: Kikawa, January, 1925, H. H. Allan, 3432. Bromus gussonii Pari.—ll, 111. Bromus hordeaceus L.—II, 111. Bromus mollis L.—II, III.* Auckland; Mt Albert, October, 1960, R.F.R.Mc., 19521; Dargaville, November, 1960, J. M. Dingley, 19705; Taupo, December, 1960, R.F.R.Mc., 19564. Canterbury: Waiau, December, 1954, A. J. Healy, 14209; Springfield, February, 1961, R.F.R.Mc., 19621.

Bromus sterilis L. —11, 111. Deyeuxia forsteri Kunth—ll, 111. Secale cereale L.—II, 111. Triticum aestivum L.—II, 111.

All hosts except Agropyron scabrum and Deyeuxia forsteri are introduced species. Puccinia recondita has been recorded on Agropyron scabrum in Australia as Puccinia agropyri by Me Alpine (1906). The rust is an introduced heteroecious species previously recorded in New Zealand by Cunningham (1924, 1931) as P. elymi.

P. recondita is regarded as a polymorphous species embracing a large number of races previously recognised as distinct species—e.g., leaf rust of wheat, P. triticina, is classified as a race of P. recondita restricted primarily to the genus Triticum but which may infect certain members of other genera. Although small differences may occur in spore dimensions and alternate hosts, these races have the same general form and are morphologically indistinguishable. Hosts such as Bromus and Secale cereale are susceptible to a number of races or race groups of Puccinia recondita (Gaumann, 1936, Viennot-Bourgin 1941, Guyot 1941, Jorstad 1958).

Considerable confusion exists in the literature concerning the name of this rust, due mainly to the recognition of species on the basis of host specialisation alone. To overcome this confusion, Cummins & Caldwell (1956) propose that P. elymi be considered a separate species, related to but morphologically distinct from the leaf rust of cereals. This recommendation follows suggestions by Mains (1932), Hylander, Jorstad & Nannfeldt (1953) and Wilson & Bisby (1954) (as Rostrupia elymi ), and has met with general approval. The acceptance of Puccinia elymi as a separate species makes P. recondita the oldest valid name for the species complex.

Although the rust is heteroecious, aecidiosori which occur on many hosts within the Ranunculaceae, Boraginaceae and Hydrophyllaceae have not been collected in New Zealand, and the rust is perpetuated by uredospores.

P. recondita is characterised by echinulate uredospores with 6-8 scattered germ pores and yellow brown walls, and the usually truncate but occasionally rounded or bluntly acuminate apices of the teleutospores.

Puccinia flavescens McAlpine, Proc. Linn. Soc. N.S.W. 28: 558, 1903. (Text-fig. 11, fig. i.) ..... Puccinia flavescentis McAlp., Rusts Austr. 119, 1906 O, I. Unknown.

11. Uredosori epiphyllous, linear, up to 1,5 mm long, often confluent, chestnut brown, pulverulent, arranged along furrows of leaf; spores subglobose or obovate, 22-30 x 19-25//, average 25 x 22//.; epispore pallid yellow, 1.5-2.0//, thick, closely echinulate; germ pores scattered, 4-7.

111. Teleutosori epiphyllous, scattered, elliptical, to 0.5 mm long, occasionally confluent, dark chestnut brown; spores elliptical, 33-48 x 18-26//, average 44 x 24//; apex rounded, thickened to 8//; base rounded, sometimes attenuate; constricted at septum; epispore smooth, dark chestnut brown, 1.0-1.5// thick; pedicel persistent, tinted, to 70 x 7//,.

Mesospores present, clavate or elliptical, 28-40 x 14-21/r. Type Locality. Australia, on Stipa flavescens Labill. Host. Stipa variabilis Hughes—ll, 111. S. variabilis is an introduced grass of Australian origin first appearing in New Zealand in the Christchurch area (Allan, 1940), and the rust has probably been introduced with its host. Greene & Cummins (1958) reviewed the rusts occurring on Stipa and Nasella throughout the world but make no comment on the presence of mesospores in Puccinia flavescens. Since the rust was first recorded in New Zealand (McNabb, 1961) teleutospores have been found.

P. flavescens is characterised by uredospores with 4-7 scattered germ pores and elliptical teleutospores with thickened apices. Puccinia graminis Persoon, Syn. Meth, Fung. 228, 1801. (Text-fig. 11, fig. 2.)

Puccinia culmorum Schum., Enum. PI. Saell. 2: 233, 1801. P. cerealis Mart., FI. mosq. 183, 1812. P. anthoxanthi Fckl., Jb. nassau Ver. Naturk. 27: 15, 1873. P. poculiformis Wettst., Verb, zool-bot. Ges. Wein, 35: 544, 1886. P. phlei-pratensis Erikss. & Henn., Z. Pfl. Krankh, 4: 140, 1894. P. jubata Ell. & Barth., Erythea, 4: 2, 1896. P. megalopotamica Speg., An. Mus. nac. B. Aires, 6: 224, 1898. P. anthistiriae Bard., List Ured. Simla, 2: 246, 1899. P. vilis Arth., Bull. Torrey bot. Cl. 28: 663, 1901. P. sesleriae-coerulae Ed. Fisch., Beitr. Kryptogamenfl. Schweiz. 2: 259, 1904. P. culmicola Diet., Engl. Jb. 37: 100, 1905. P. avenae-pubescentis Bubak, Ann. mycol. 4: 107, 1906. P. dactylidis Gaum., Ber. schweiz. bot. Ges. 55: 79, 1945

O, I. Unknown in New Zealand. 11. Uredosori amphigenous and on stems, sheaths and inflorescences, scattered or more frequently confluent and linear, up to 2-3 mm long or more, reddishbrown, bullate, pulverulent; spores elliptical, obovate, sometimes subglobose, often laterally compressed, 24-34 x 16-20/x; epispore golden brown, 1.5-2.0/t thick, coarsely and moderately echinulate; germ pores equatorial, 4-5.

111. Teleutosori on sheaths, stems and inflorescences, occasionally on leaves, at first scattered, becoming confluent and linear, up to 15 mm or more, pulvinate, compact, black, becoming naked and conspicuous; spores elliptical, oblong-clavate or subclavate, 29-65 x 15-25//; apex acuminate, sometimes oblique, thickened to 12//. deeply coloured; base attenuate, basal cell longer and narrower; constricted at septum; epispore smooth, chestnut brown, to 2.5// thick; pedicel persistent, coloured, to 70 x 10//, continuous with spore.

Type Locality. Europe, on Triticum aestivum L. (Lectotype) Hosts. Agropyron re pens (L.) Beauv.—ll, lII.* Canterbury: Lincoln, March, 1960, J. M. Dingley, 19486. Agropyron scabrum (Labill.) Beauv. —11, 111. Anthoxanthum odoratum L.—II, 111. Anthoxanthum pueli Lee. & Lamb.—ll.* Canterbury: Paparua, January, 1955, A. J. Healy, 14302. Avena fatua L. —11, 111. Avena saliva L.—II, 111. Avena strigosa Schreb.—ll, 111. Briza minor L.—ll.*

Auckland; Mt Albert, November, 1960, R.F.R.Mc., 19517; Te Aroha, December, 1960, R.F.R.Mc., 19533. Dactylis glomerata L.—II, 111. Deschampsia caespitosa (L.) Beauv.—ll, 111. Deyeuxia avenoides (Hook, f.) Buch.—ll, III.* Canterbury: Porter’s Pass, February, 1961, R.F.R.Mc., 19628. Echinopogon ovatus (Forst. f.) Beauv.—ll, 111. Festuca pratensis Huds.—ll, 111. Glycerin maxima (Hartm.) Holmb.—ll, 111. Hierochloe redolens (Vahl) Roem. & Schult.—ll, III.*

Wellington: Rangipo Desert, March, 1961, R.F.R.Mc., 19599. Lolium multiflorum Lam.—ll, 111. Lolium perenne L.—II, 111. Lolium temulentum L.—II, 111. Poa anceps Forst. f.—II, 111. Poa trivialis L.—II, lII.* Wellington; Waimarino, May, 1930, H. H. Allan, 10045. Phalaris tuberose L.—II, 111. Triticum aestivum L.—II, 111.

Among the hosts listed above, Deyeuxia avenoides and Poa anceps are endemic grasses, while Agropyron scahrum, Echinopogon ovatus and Hierochloe redolens are indigenous species. Puccinia graminis is sporadic in its occurrence on these grasses and has been recorded from Australia on Agropyron scahrum and Echinopogon ovatus by McAlpine (1906).

The rust on Anthoxanthum odoratum and A. pueli is commonly considered a distinct species, Puccinia anthoxanthi. Jorstad (1950) includes P. anthoxanthi as a race of P. graminis s. lat., differing in that sori are soon naked and in the comparatively small teleutospores. (In Norwegian material, teleutospores measure 29-41 x 14.4-2lju). New Zealand specimens yield the following measurements: teleutospores 29-45 x 16-22 /a, average 37 x 19/a, apex thickened to Sjx, three-celled teleutospores and mesospores occasionally present. These differences are insufficient to warrant the acceptance of P. anthoxanthi as a valid species and the rust is retained within P. graminis s.lat, as is done by Viennot-Bourgin (1949) and Cummins (1956). Teleutospores are rare on the specimens examined. Both Me Alpine (1906) and Jorstad (1950) comment that they are not common on Anthoxanthum.

There is some evidence that races of Puccinia graminis on wheat in New Zealand are of Australian origin (Smith, 1960). Smith (1960) suggests that the rust overwinters in Queensland and spreads to New South Wales and across the Tasman Sea to New Zealand in spring and early summer, appearing in New Zealand about November of each year. Despite the fact that Berberis vulgaris is common in New Zealand, there is no evidence that it acts as an alternate host for Puccinia graminis (Smith, 1960).

P. graminis is characterised by usually laterally compressed, echinulate uredospores with 4-5 equatorial germ pores, and teleutospores with strongly thickened apices. Other grass rusts in New Zealand possessing echinulate uredospores with

equatorial germ pores are P. sorghi and Uromyces tenuicutis, but neither of these have laterally compressed uredospores. Puccinia sorghi Schweinitz. Trans. Am. Phil. Soc. Ser. 11. 4: 295, 1832. (Textfig. 11, fig. 4.) Puccinia maydis Bereng., Atti. Sci. Ital. 6: 475, 1845. P. arundinacea var. maydis Cast., Catal. 1: 199, 1845. P. zeae Bereng., in Klotzsch, Herb. Viv. Myc. Suppl. 18, 1851. O, I. Unknown in New Zealand.

11. Uredosori amphigenous, scattered, elliptical, to 1 mm long, or confluent forming lines to 5 mm long, cinnamon brown, bullate, pulverulent; spores subglobose or occasionally obovate, 26-32 x 23-29/*., average 28 x 25/*,; epispore cinnamon brown, 1.5-2.0/*, thick, finely and moderately echinulate; germ pores approximately equatorial, 3-4. 111. Teleutosori amphigenous, scattered, elliptical or occasionally linear, rarely confluent, to 3 mm long, pulvinate, compact, brown to black, soon naked; spores elliptical, 30-46 x 16-24/*,, average 40 x 21/*.; apex rounded, often bluntly acuminate, thickened to 8/*,; base attenuate, frequently rounded; both cells same size and colour, slightly constricted at septum; epispore smooth, chestnut brown, 1.5-2.0/*, thick; pedicel persistent, coloured, to 80 x 8/*,.

Type Locality. North America, on Zea mays L. Host. Zea mays L.— II, 111. Both host and rust are introduced. Arthur (1904) first proved that Puccinia sorghi is a heteroecious species by inoculating aecidiospores from Oxalis cymosa on to maize leaves. Smith (1926) obtained aecidiosori by inoculating with teleutospores, on O. corniculata, O. europaea and O. tropaeoloides, while Rice (1933) reports successful inoculations with O. stricta and O. repens. Although O. corniculata, O. latifolia and O. stricta are common in New Zealand (Healy 1958, Allan 1961), the aecidial stage has not been found.

Since Cunningham (4931) first recorded the rust in New Zealand, teleutospores have been found. Puccinia sorghi is characterised by echinulate uredospores with 3-4 approximately equatorial germ pores, and elliptical teleutospores. Puccinia cynodontis Lacroix in Desmazieres PL Crypt. II: no. 655, 1859. (Textfig. 11, fig. 3.) Puccinia varians Diet., Ann. mycol. 6: 224, 1908. O, I. Unknown in New Zealand.

11. Uredosori mainly hypophyllous, sometimes on purple lesions, scattered or gregarious, oval to elliptical, to 1 mm long, cinnamon brown, pulverulent; spores globose, subglobose, or obovate, 21-28 x 19-26/x, average 23-21 ft; epispore pallid cinnamon brown, 2-3 /x, thick, finely and closely verruculose; germ pores equatorial, 2-3.

111. Teleutosori mainly hypophyllous, scattered, occasionally arranged in lines, elliptical, to 0.5 mm long, compact, chocolate brown, soon naked; spores elliptical or oblong-elliptical, 30-45 x 17-22/x, average 37 x 19//,; apex obtuse or acuminate, thickened to 12//,; base attenuate, frequently rounded; both cells approximately the same size or basal cell slightly smaller, slightly constricted at septum; epispore smooth, chestnut brown, 1.5-2.5//, thick; pedicel persistent, coloured, to 60 x 5//.

Type Locality. France, on Cynodon dactylon (L.) Pers. Host, Cynodon dactylon (L.) Pers.—ll, 111.

Both host and rust are introduced species. Puccinia cynodontis is a heteroecious rust shown by inoculation experiments to possess a plurivorous haploid phase. Aecidial hosts may belong to the genera Delphinium, Mercurialis, Adonis, Plantago, Valerianella, Veronica and Viola (Guyot & Viennot-Bourgin, 1943), but Jorstad (1958) comments that in nature other genera may also carry the aecidial stage. No alternate host has been established for Puccinia cynodontis in New Zealand, and it appears to be independent of host alternation in this country.

Since the rust was first recorded (Baker, 1956), teleutospores have been found. P. cynodontis is characterised by the closely and finely verruculose uredospores with 2-3 equatorial germ pores. It is the only grass rust in New Zealand with verruculose uredospores.

Uromyces dactylidis Otth, Mitth. naturf. Ges. Bern, 1861, 85, 1861. (Text-fig. 111, fig. 2.)

Uromyces poae Rabenh., in Marcucci, Unio. Itin. 38, 1866. U. graminum Gke., Handbk. Br. Fungi, 520, 1871. U. alopecuri Seym. Proc. Bost. nat. Hist. Soc. 24: 186, 1889. U. festucae Syd., Hedwigia, 39: 117, 1900. U. phyllachoroid.es P. Henn., Hedwigia, 40: 129, 1901. U. ranunculi-festucae Jaap, Verb. bot. Ver. Brandenb. 47: 90, 1905. U. ranunculi-distichophylli Semad., Zbl. Bakt. 16: 385, 1906. U. schlerochloae Tranz., Ann. mycol. 5: 550, 1907. U. atropodis Tranz., loc. cit. 5: 550, 1907.

U. poae-alpinae Rytz, Mitt, naturf. Ges. Bern, 67: 6, 1910. U. lygei Syd., Monogr. Ured. 2: 331, 1910. U. festucae-nigricantis Frag., Gontr. fl. mic. Guad. 33, 1914. U. phlei-michelii Cruch., Bull. Soc. vaud. Sci. nat. 51: 75, 1916. U. scleropoae Baud. & Picb., Bull. Inst. bot. Univ. Belgrade, 1: 62, 1928. U. agrostidis (Frag.) Guyot, Les Ured., Uromyces, Encycl. Myc. 8: 73, 1938. U. vulpiae Losa Espana, Ann. Jard. bot. Madr. 7: 422, 1945. U. vulpiae Camara, Agron. Lusit. 11: 166, 1949. U. volkartii Gaum. & Terrier, Ber. schweiz. bot. Ges. 62: 299, 1952.

O. Pycniosori epiphyllous, scattered or in small groups accompanying the aecidiosori, immersed, minute, honey-coloured.

I. Aecidiosori hypophyllous and on petioles, seated on yellow spots, crowded in small groups, orange; peridia cupulate, margins recurved, lacerate, white; spores polygonal, 17-25/t, average 19 ( u; epispore pallid yellow, I.o[jl thick, finely and closely verruculose.

11. Uredosori mostly epiphyllous, scattered or arranged in lines, to 2mm long, pallid brown, pulverulent; spores subglobose or obovate, 22-32 x 17-26/a, average 28 x 24/x; epispore golden brown, 1. thick, finely and sparsely echinulate; germ pores scattered, 6-8, conspicuous.

111. Teleutosori mostly hypophyllous, scattered or arranged in lines, to 2mm long, compact, black, bullate, long covered; spores obovate, often pyriform, seldom elliptical, 20-32 x 14-20 jx, average 22 x 16/*; apex bluntly rounded, seldom acuminate, sometimes truncate, not or slightly thickened (to 4/i) , darker in colour; base sub-attenuate; epispore smooth, golden brown, 1.0-1 thick; pedicel persistent, tinted, to 30 x sft; surrounded and divided into compartments by numerous clavate, thick-walled, brown paraphyses.

Type Locality. Switzerland, on Dactylis glomerata L. Hosts. Ranunculus repens L.—O, I. Dactylis glomerata L.—-11, 111.

Both hosts are introduced and the rust is an introduced, heteroecious species. The interpretation of Grove (1934), Jorstad (1950) and Cummins (1956) has been followed. They consider that Uromyces dactylidis is a polymorphous species consisting of a number of races infecting mainly Dactylis, Festuca and Poa. Races

of Uromyces dactylidis s.lat. were previously separated into species (Gaumann, 1941 a) , but both Grove (1934) and Jorstad (1950) did not find the races sufficiently distinct to warrant specific rank. Cummins (1956) states that the differences involved between synonymous species are almost exclusively minor variations in spore dimensions. Where known, the host alternation of these races is with species of Ranunculus.

Uromyces dactylidis is characterised by small teleutospores with unthickened or slightly thickened apices, and brown, clavate subepidermal paraphyses in the teleutosori. It is the only heteroecious grass rust in New Zealand of which the aecidial stage has been found. Uromyces tenuicutis McAlpine, Rusts Austr., 87, 1906. (Text-fig. 111, fig. 3.) O, I. Unknown.

11. Uredosori mainly epiphyllous and on culms and stems, scattered, rarely confluent, elliptical, to 0.5 mm long, chestnut brown, becoming pulverulent; spores elliptical to ovate, 26-36 x 18-24/*, average 31 x 22/*; epispore cinnamon brown, 1.5-2.0/* thick, moderately and coarsely echinulate; germ pores equatorial or nearly so, 3-5, conspicuous.

111. Teleutosori hypophyllous, scattered, elliptical, to 0.5 mm long, long covered, compact; spores angular, obovate to oblong, or pyriform, 21-32 x 15-22/*, average 27 x 18ft; apex variable, truncate, irregularly rounded or bluntly acuminate, thickened to 4/*, base attenuate; epispore smooth, chestnut brown, I.sft thick; pedicel mostly deciduous, tinted, to 40 x 7/*.

Type Locality. Australia, on Sporobolus capensis Kunth. Host. Sporobolus capensis Kunth.—ll, lII.* Auckland: Onerahi, October, 1960, R.F.R.Mc., 19577; Karekare, November, 1960, R.F.R.Mc., 19576; Mt Albert, January, 1961, R.F.R.Mc., 19619; Cascades, February, 1961, R.F.R.Mc., 19617; Dargaville, March, 1961, J. M. Dingley, 19629; Waikaraka, April, 1961, R.F.R.Mc., 19779.

Nelson: Karamea, February, 1961, R.F.R.Mc., 19575. The host is an introduced grass common on poor land throughout the country. Uromyces tenuicutis has not previously been recorded in New Zealand. U. tenuicutis is characterised by large uredospores with 3-5 conspicuous, equatorial germ pores, and irregularly-shaped teleutospores with slightly thickened apices.

Uromyces danthoniae McAlpine, Rusts Austr. 85, 1906. (Text-fig, 111, fig. 4.) 0. Unknown. 1. Unknown in New Zealand. 11. Uredosori mostly epiphyllous and on culms, scattered, elliptical, to 3 mm long, or linear when up to 1 cm long, pulverulent, pallid chestnut brown; spores subglobose to shortly elliptical, 25-44 x 22-36//,; epispore pallid chestnut brown, 2.0/X-3.5//, thick, finely and moderately echinulate; germ pores scattered, conspicuous, 7-10; mixed with hyaline, cylindrical to subclavate paraphyses.

111. Teleutosori epiphyllous, scattered, elliptical, to 1 mm long, or linear when up to 1 cm long, chocolate brown; spores ovate, pyriform or fusoid, 28-43 x 18-25/*; apex acuminate, occasionally rounded, thickened to 12/*; base rounded, often subattenuate to attenuate; epispore smooth, chestnut brown, 1.5-2.5/* thick; pedicel persistent, tinted, to 200 x 12/*; hyaline, cylindrical to subclavate paraphyses occasionally present.

Type Locality. Australia, on Danthonia semi-annularis (Labill.) R. Br. (lectotype). Hosts. Danthonia cunninghami Hook. f. —II, 111, Danthoria flavescens Hook. f.—II. Danthonia pilosa R. Br.—ll, 111. Danthonia raoulii var. rubra Ckne.—ll, lII.* Canterbury: Lewis Pass, September, 1960, L. J. Metcalf, 19487; Maruia, February, 1961, R.F.R.Mc., 19573.

Wellington: Desert Road, December, 1960, R.F.R.Mc., 19565; Rangipo Desert, March, 1961, R.F.R.Mc., 19614. Danthonia semi-annular is (Labill.) R.Br.—II, 111. Danthoria setifolia (Hook.) Ckne.—ll, lII* Auckland: Turangi, December 1960, R.F.R.Mc., 19557. Wellington: Rangipo Desert, December, 1960, R.F.R.Mc., 19545. Among the species of Danthonia listed above, D. pilosa and D. semiannularis are indigenous grasses which occur in Australia (Cheeseman, 1925, p. 178). The remainder are endemic.

Aecidiosori tentatively placed with Uromyces danthoniae have been described by McAlpine (1906) on a species of Danthonia, but as pointed out, this was done as a matter of convenience. There is some doubt as to whether the aecidosori are biologically connected with the uredo and teleuto stages for they were not found on the same plant. If a biological connexion does exist, Uromyces danthoniae will be one of the few known autoecious grass rusts.

Uredo and teleutospores on endemic hosts are consistently larger than those on the two indigenous species; uredospores average 36 x 32 /t (as against 30 x 26/t) and teleutospores 36 x 19ft (as against 32 x 21ft). Small differences occur in the density of uredospore echinulations and in teleutosori and teleutospore shape, but these are considered insufficient to warrant the erection of a separate species.

McAlpine (1906) does not designate a type specimen. An examination of Australian material from McAlpine’s herbarium, Department of Agriculture, Melbourne, shows that the rust on Danthonia semi-annularis agrees closely with New Zealand specimens on this host. It is proposed that the collection on D. semi-annularis (Killara, 9 Oct., 1902, Coll. G. H. Robinson) be accepted as the lectotype.

The rust is characterised by thick-walled uredospores with scattered, conspicuous germ pores, and the long, persistent pedicels and thickened apices of the teleutospores.

Uromyces otakou G. H. Cunningham, Trans. N.Z. Inst. 54; 627, 1923. (Textfig. IV, fig. 1.) O, I. Unknown. 11. Uredosori epiphyllous, scattered, rarely arranged in lines, to 2 mm long, orange yellow, pulverulent; spores subglobose, occasionally shortly elliptical, 21-23 x 18-24ju,, average 25 x 21/x; epispore pallid yellow, 1.5-2.0//, thick, moderately and finely echinulate; germ pores scattered, 6-10.

111. Teleutosori epiphyllous, scattered, seldom confluent, to 3 mm long, dark chestnut brown, pulvinate, compact; spores ovate, 24-30 x 14-24ju,, average 25 x 18/x; apex rounded, seldom bluntly acuminate, thickened to 12/x; base subattenuate, frequently rounded; epispore smooth, chestnut brown, 2.0-2.5/a thick; germ pore apical; pedicel persistent, tinted, to 60 x Bju,.

Type Locality, New Zealand, on Poa caespitosa Sprengel. Hosts. Poa anceps Forst. f.—II, 111. Poa caespitosa Sprengel.—ll, 111. Poa colensoi Hook. f.—ll.* Wellington: Rangipo Desert, December, 1960, R.F.R.Mc., 19528, Poa litorosa Gheesem. —11.

Of the hosts listed above, Poa caespitosa is an indigenous grass extending to Australia and Tasmania (Gheeseman, 1925, p. 190) : the remainder are endemic species.

Cunningham (1923, 1931) states that uredospore walls are hyaline with densely and minutely verruculose ornamentation; Cummins (1956) describes them as

echinulate with hyaline-golden walls. An examination of type material shows that mature uredospores are moderately and finely echinulate with pallid yellow walls. Uromyces otakou is an endemic rust characterised by small teleutospores with thickened apices and persistent pedicels, and small, moderately echinulate uredospores. Uromyces ehrhartae McAlpine, Agr. Gaz. N.S.W. 6: 855, 1895. (Text-fig. IV, fig. 2.)

O, I. Unknown

11. Uredosori amphigenous and on sheaths, scattered, occasionally confluent and linear, orbicular, to 0.5 mm long, yellow brown, pulverulent; spores subglobose to obovate, 21-27 x 17-20/*, average 24 x 19/*; epispore cinnamon brown, 1.0-1.5/* thick, finely and moderately echinulate; germ pores scattered, 6-8, conspicuous.

111. Teleutosori sparse, amphigenous and on sheaths, mainly linear, to 1 mm long, chocolate brown; spores variable in shape, sub-globose or elliptical, fusiform or oblong, 19-33 x 13-19/*, average 25 x 15/*; apex conical or truncate, thickened to 9/*; epispore smooth, chestnut brown, .5-2.0//, thick; pedicel persistent, tinted, to 50 x 7/*.

Type Locality. Australia, on Microlaena stipoides (Labill.) R. Br. Host. Microlaena stipoides (Labill.) R.Br.—II, lII.* Auckland: Taupo, December, 1960, R.F.R.Mc., 19618; Kamo, April, 1961, R.F.R.Mc., 19780; Kirikopuni, April, 1961, R.F.R.Mc., 19846. Marlborough; Pelorus Bridge, February, 1961, R.F.R.Mc., 19597; Goose Bay, March, 1961, R.F.R.Mc., 19610; Oaro, March, 1961, R.F.R.Mc., 19624. Nelson: Karamea, February, 1961, R.F.R.Mc., 19598. Taranaki; New Plymouth, February, 1961, R.F.R.Mc., 19586; Uriti, February, 1961, R.F.R.Mc., 19594. Wellington: Upper Hutt, March, 1952, A. J. Healy, 11469; Wallaceville, January, 1953, A. J. Healy, 11743; Greytown, March, 1953, A. J. Healy, 12054.

M. stipoides is an indigenous grass also occurring in Australia and Tasmania (Cheeseman, 1925, p. 144). The rust has not previously been recorded in New Zealand, but it appears to be widespread throughout the country. Teleutospores are scarce and usually found on the dry, lower leaves. Uromyces ehrhartae is characterised by uredospores with cinnamon brown walls and 6-8 scattered germ pores, and the thickened apices and variable shape of the teleutospores. Uredo karetu G. H. Cunningham, Trans. N.Z, Inst. 55: 41, 1924. ( Text-fig. IV, fig. 3.)

Uredosori hypophyllous, seated on discoloured spots visible above, linear, to 1 mm long, seldom confluent, pulverulent, orange yellow; spores subglobose or obovate, 26-37 x 24-31/*, average 32 x 28/*; epispore hyaline or tinted yellow, 2.0-2.5/* thick, finely and densely echinulate; germ pores scattered, numerous, 6-10, indistinct; mixed with a few hyaline paraphyses.

Type Locality. New Zealand, on Hierochloe redolens (Vahl) Roem. & Schult. Host. Hierochloe redolens (Vahl) Roem. & Schult.—ll. The host is an indigenous grass extending to Australia, Tasmania and South America (Cheeseman, 1925, p. 147). Cunningham (1924) in his original description, states that uredospores are finely and moderately echinulate, but later (1931) describes them as verruculose, as does Cummins (1956). An examination of the type specimen shows that uredospores are finely and densely echinulate,

Uredo karetu is characterised by finely and densely echinulate uredospores with numerous scattered germ pores.

Uredo toetoe G. H. Cunningham, Trans. N.Z. Inst. 55: 41, 1924. (Text-fig. IV, fig. 4.)

Uredosori hypophyllous, scattered, elliptical, 0.5-1.0 mm long, or confluent when up to 4 mm long, bright reddish-brown, pulverulent; spores elliptical or subglobose, 22-35 x 21-30/t, average 30 x 28/x; epispore pallid chestnut brown, 2.5-3.0/t thick, finely and closely echinulate; germ pores scattered, numerous, 7-12, conspicuous.

Type Locality. New Zealand, on Arundo richardii Endl. Host. Arundo richardii Endl.—ll. The host is an endemic species abundant throughout the country. Uredo toetoe is characterised by reddish-brown sori and thick-walled uredospores with numerous, scattered, conspicuous germ pores.

Host Index

Agropyron repens Puccinia graminis Agropyron repens Puccinia recondita Agropyron scabrum Puccinia graminis Agropyron scabrum Puccinia recondita Agrostis canina Puccinia coronata Agrostis stolonifera Puccinia coronata Agrostis tenuis Puccinia coronata Air a caryophyllea Puccinia coronata Anthoxanthum odoratufn Puccinia graminis Anthoranthum odoratum Puccinia poae-nemoralis Anthoxanthum pueli Puccinia graminis Arrhenatherum elatius Puccinia coronata Arrhenatherum elatius var. bulbosum Puccinia coronata Arundo richardii Uredo toetoe Avena fatua Puccinia coronata Avena fatua Puccinia graminis Avena sativa Puccinia coronata Avena sativa Puccinia graminis Avena strigosa * Puccinia graminis Briza minor ~ Puccinia graminis Bromus gussonii ...... Puccinia recondita Bromus hordeaceus Puccinia recondita Bromus mollis . Puccinia recondita Bromus sterilis , Puccinia recondita Cynodon dactylon ...... Puccinia cynodontis Dactylis glomerata Puccinia graminis Dactylis glomerata Uromyces dactylidis Danthonia cunninghamii Uromyces danthoniae Danthonia flavescens Uromyces danthoniae Danthonia raoulii var. rubra Uromyces danthoniae Danthonia pilosa Uromyces danthoniae Danthonia semi-annularis ...... ...... Uromyces danthoniae Danthonia setifolia ...... Uromyces danthoniae Deschampsia caespitosa Puccinia graminis Deyeuxia avenoides ~... Puccinia graminis Deyeuxia forsteri Puccinia recondita Dichelachne crinita Puccinia crinitae Echinopogon ovatus Puccinia graminis Festuca pratensis Puccinia graminis Glyceria maxima Puccinia graminis Hierochloe redolens Puccinia graminis Hierochloe redolens Uredo karetu Holcus lanatus Puccinia coronata Holcus mollis ...... ...... Puccinia coronata Hordeum murinum Puccinia hordei Hordeum vulgare Puccinia hordei Lolium multiflorum Puccinia coronata Lolium multiflorum Puccinia graminis Lolium perenne Puccinia coronata Lolium perenne Puccinia graminis Lolium temulentum Puccinia coronata Lolium temulentum ...... Puccinia graminis Microlaena slip aides Uromyces ehrhartae Phalaris minor Puccinia coronata Phalaris tuberosa Puccinia graminis

Poa anceps ...... '.. Puccinia graminis Poa anceps Uromyces otakou Poa annua Puccinia coronata Poa annua Puccinia poae-nemoralis Poa caespitosa ...... ....; ...... Uromyces otakou Poa colensoi ...... - ...... ...... . Uromyces otakou Poa iridifolia Puccinia coronata Poa litorosa Uromyces otakou Poa palustris ...... Puccinia coronata Poa pratensis Puccinia coronata Poa pratensis Puccinia poae-nemoralis Poa trivialis ...... Puccinia graminis Ranunculus repens Uromyces dactylidis Secale cereale Puccinia recondita Sporobolus capensis Uromyces tenuicutis Stipa variabilis Puccinia flavescens Triticum aestivum Puccinia graminis Triticum aestivum Puccinia recondita Zea mays Puccinia sorghi

Literature Cited

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Cruickshankj I. A. M., 1956. Physiological specialization of Oat Crown Rust {Puccinia coronata f. sp. avenae) in New Zealand. N.Z.].Sci.Tech. 38A: 359-362. Cummins, G. 8., 1956. Host index and morphological characterization of the grass rusts of the world. Plant Dis. Reptr, Suppl. 237, 1-52. Cummins, G. 8., and Caldwell, R. M., 1956. The validity of binomials in the leal rust fungus complex of cereals and grasses. Phytopathology, 46: 81-82. Cunningham, G. H., 1923. The Uredinales or rust-fungi of New Zealand: Part 1— Pucciniaceae, Tribe Puccineae. Trans. Proc. N.Z. Inst. 54: 619—704. Part 11. Trans. Proc. N.Z. Inst. 55: 1-58. Gaumann, E., 1934. Mykologische notizen. Ann. mycol. 32: 300-301. Ges. 46: 229-247.

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Valerianella en rapport avec Puccinia cynodontis Desm. Ann. Ec. Agric. Grignon, Ser. 3,3: 100-105. Healy, A. J., 1958. Contributions to a knowledge of the adventive flora of New Zealand, No. 7. Trans, roy. Soc. N.Z. 86: 113-118. Hylander, N., Jorstad, 1., & Nannfeldt, J. A., 1953. Enumeratio Uredinearum Scandinavicarum. Opera Bot. 1: 1, I—lo2.

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McAlpine, D., 1906. The Rusts of Australia. Government Printer, Melbourne. McNabb, R. F. R., 1961. Additions to the Rust Fungi of New Zealand, 111. Trans, roy. Soc. N.Z. (in press). Mains, E. 8., 1932. Host specialization in the leaf rust of grasses, Puccinia rubigo-vera. Mich. Acad. Sci. Arts, Letters, Papers 17, 289-394. Nielsen, P., 1877. Bemaerkninger om nogle rustarter. Bot. Tidsskr. 3: Ser. 2, 26. Oliveira, B. d’., 1937. Brown rust of the wild species of Hordeum. Rev. agron., Lisboa, 25: 230-234.

Plowright, G. 8., 1889. British Uredineae and Ustilagineae. Kegan Paul, Trench & Co., London. Rice, Mabel A., 1933. Reproduction in rusts. Bull. Torrey hot. Cl. 60: 23-54. Smith, H. C., 1960. Notes. Commonwealth Phytopath. News. 6: 59-60. Smith, M. A., 1926. Infection and spore germination studies with Puccinia sorghi. Phytopathology, 16: 69. Tranzschel, W. A., 1914. Culturversuche mit Uredineen in den Jahren. Mykol. Zbl. 4: 70-71.

Viennot-Bourgin, G., 1941. La rouille jauve des Graminees, Ann. Ec. Agric. Grignon, Sen 3,2: 129-217. Wilson, M., and Bisby, G. R., 1954. List of British Uredinales. Trans. Brit, mycol. Soc. 37: 61-86.

Mr R. F. R. McNabb, M.Sc., Plant Diseases Division, D.5.1.R., Private Bag, Auckland.