NEW ZEALAND. Northland: Trounson Park, trunk lesion of Agathis australis, 23 Mar. 2010, N. Waipara, culture REB 327-41 = ICMP 18404; Northland: Trounson Park, soil under A. australis, 23 Mar. 2010, A. Vannini, culture 327-60 = ICMP 18410; Northland: Raetea, trunk lesion of A. australis, 23 Mar. 2010, N. Waipara, culture 327-34 = ICMP 18401; Northland: Waipoua Forest, near Tane Mahuta, trunk lesion of A. australis, 23 Mar. 2010, N. Waipara, culture 327-47 = ICMP 18407; Northland: Waipoua Forest, trunk lesion of A. australis, 23 Mar. 2010, N. Waipara, culture 327-53 = ICMP 18408; Coromandel: Great Barrier Island, from wood of A. australis, Mar. 1972, P. Gadgil, culture ATCC 32256 = FRI 135 = ICMP 16471; Coromandel: Great Barrier Island, Kaiarara, from bleeding lesion on trunk of A. australis, 21 May 2009, N. Waipara, culture REB 326-154 = ICMP 18360; Auckland: Pakiri, trunk lesion of A. australis, 18 Nov. 2008, R.E. Beever, REB326-1 = ICMP 18244; Auckland: Waitakere Ranges, Cascades, from bleeding lesion on trunk of A. australis, 7 Oct. 2009, R.E. Beever, culture REB 326-221 = ICMP 18358; Auckland: Huia, soil under A. australis, 23 Mar. 2010, A. Vannini, culture 327-46 = ICMP 18406.

Differs from other Phytophthora Clade 5 species in its oogonium ornamentation with occasional and slightly raised protuberances, and its larger mean oospore diameter (31.9 µm). Found in association with Agathis australis. Differs from P. cocois and P. heveae in its DNA barcode sequence of ITS; differs from all other Phytophthora species in its DNA sequence of the ND1 gene. Description:—The species is homothallic, with isolates forming oogonia quickly (3–4 days) and abundantly on V8A. Oogonia are globose with a mean width of 31.9 μm, and ranging between (22.2–)29.7–32.2–32.7(–45) μm. Oogonium wall ornamentation is mildly stipulate. Oospores nearly fill the oogonia with a mean width of 27.7 μm, and ranging between (19.8–)25–27.5–29.7(–35) μm. Antheridia are amphigynous, globose some with knots at the base. Sporangia are globose to ovoid-ellipsoid, papillate, borne terminally from long thin branched sporangiophores and could be formed via internal proliferation. Sporangia are non-caducous (although some isolates have a somewhat defined septum near the base of the sporangium, see Fig. 5H). Sporangia have a mean width of 28.4 μm, and ranging between (12.4–)24.8–27.7–32.2(–50) μm, and a mean length of 39.6 μm, and ranging between (14.9–)32.4–37.5–47.5(–75) μm. Vegetative hyphae are simple, with slight swellings, and lacking chlamydospores in culture. Colony morphology after 7 days was very uniform across the isolates tested on most of the media examined. Colonies are loosely aerial. On 5% clarified V8-juice agar, there is a weakly stellate radial pattern. Minimum growth temperature 6°C; maximum 25°C; optimum 21.5°C. Complete morphometric statistics are presented in Table 3.

Known only from kauri (Agathis australis) trees and associated soil in northern (<38°S) New Zealand’s mixed podocarp broad-leaf forest.

Disease and management:—The root and collar rot of kauri (colloquially termed “kauri dieback”) is the subject of a long term management response led by the Ministry of Primary Industries, Department of Conservation, regional councils (Auckland, Bay of Plenty, Northland, Waikato), in partnership with the Tangata Whenua Roopu. Delimitation surveys have confirmed impacts upon kauri of all age classes, in forest remnants and plantations, throughout its geographic range. Phytosanitary measures have been put in place in high-access regional parks to control the spread of the pathogen by foot-traffic, and the use of phosphite to control the disease in planta has also commenced.

NEW ZEALAND. Coromandel: Great Barrier Island, from bleeding lesion on trunk of Agathis australis (D. Don) Lindl. ex Loudon, 23 Mar. 2006, R.E. Beever REB316-14, dried culture specimen, holotype PDD 91595; ex-holotype living culture preserved in a metabolically inactive state as ICMP 17027 = WPC P15175.

CHINA. Hainan Island: forest soil, H. Ho, culture FFM 10-5C = ATCC MYA-3422 = ICMP 19635 = WPC P10661. TAIWAN. from soil, F. Panabieres culture F441= ICMP 16915 = SCRP 388; Nan Ton County: Lenhuachih (Lianhuachi), natural forest soil, 13 May 1988, H.S. Chang, Herb. IMI 325914, culture L-2A = ATCC 36818 = ICMP 19450 = CBS 587.85 = IMI 325914 = WPC P15598; Nan Ton County: soil under Castanopsis fargesii, 13 Nov 2010, S.E. Bellgard, culture REB326-308 = ICMP 18737. JAPAN. Ibaraki: from Castanea crenata trunk rot, 1971, K. Uchida, culture P1 = NBRC 30433 = ICMP 19435; Ibaraki: from C. crenata trunk rot, 1971, K. Uchida, culture P3 = NBRC 30434 = ICMP 19436; Ibaraki: from C. crenata shoot, 1971, K. Uchida, culture P12 = NBRC 30435 = ICMP 19437;

Description:—Katsura (1976) provided an initial description of this species and Ko & Chang (1979) described Taiwanese specimens. The more recent description of Erwin & Ribeiro (1996) is still useful, although the circumscription encompasses isolates we consider P. cocois. The primary morphological characters of isolates considered as P. castaneae examined in this work are presented in Table 3.

Known on trunk rot of Castanea crenata from Japan and Korea, on Castanopsis fargesii Franch. from Taiwan, and soil from China.

JAPAN. Ibaraki: from Castanea crenata Siebold & Zucc. trunk rot, 1971, K. Uchida, holotype specimen Herb. 1971-031 (Plant Pathology Herbarium, Kyoto Prefectural University, Japan); ex-holotype culture P8 = NBRC 9753 = ICMP 19434 = WPC P10187. Ko & Chang (1979) correctly cited the holotype of the species to be the Japanese specimen “No. 1971-031, Plant Pathology Herbarium, Kyoto Prefectural University, Japan”, but then described the morphology of an isolate from Taiwan (ATCC 36818); this isolate has subsequently been used as a reference strain for P. castaneae (ref Q-Bank). However, a culture of 1971-031 isolated by K. Uchida (P8) was deposited in the IFO (now NBRC) culture collection of Japan as NBRC 9753, and is thus an authentic ex-holotype culture.

USA. Hawaii: Kauai, from diseased fruit of Cocos nucifera, coll. J.Y. Uchida culture H1026 = ICMP 16949; CÔTE D’IVOIRE. Port Bouet, from nut of Cocos nucifera, De Franque-Ville, culture IMI 360596 = ICMP 19685.

Differs from other Phytophthora Clade 5 species in oogonium ornamentation with moderately bullate protuberances. Found in association with Cocos nucifera. The DNA barcode sequence of ITS distinguish P. cocois from all other Phytophthora species. Description:—The species is homothallic, with oogonia formed abundantly on V8A. Oogonia are globose with a mean width of 26.2 μm, and ranging between (22.3–)24.8–25–27.3(–35) μm. Oogonium wall ornamentation is mildly bullate, and often less ornamented isolated in host tissue. Oospores nearly fill the oogonia with a mean width of 24.2 μm, and ranging between (19.8–)22.3–24.3–25(–29.7). Antheridia are amphigynous, globose and often strongly reflexed (bent). Antheridial length in fresh host tissue is often longer than isolates in culture. Sporangia are globose to ovoid, papillate and non-caducous. Sporangia have a mean width of 25.4 μm, and ranging between (12.4–)22.4–27.2–29(–35) μm, and a mean length of 38.4 μm, and ranging between (18.6–)31.4–39.6–47.1(–50) μm. Colony morphology after 7 days was very uniform across the isolates tested on most of the media examined. Colonies are loosely aerial. Minimum growth temperature 10°C; maximum 30°C; optimum 22°C. Complete morphometric statistics are presented (Table 3), and Uchida et al. (1992) provide an original description

Known on diseased coconut (Cocos nucifera) from Hawaii, and Côte d’Ivoire. (Probably also on cocoa (Theobroma cacao L.) from Côte d’Ivoire.)

Latin genitive noun: cocois — of Cocos.

Disease and management:— In Hawaii the disease has killed hundreds of coconut trees. Areas with highly conducive environments have lost most of their trees, thus the disease is now less common than in previous decades. Infected trees, which are identified by dead young leaves, cannot be saved by the application of systemic fungicides. Rapid removal and destruction of infected trees and removal of old leaves and petioles is needed to prevent infection of healthy trees (Uchida 2004). No other palm in Hawaii has been found to be affected by P. cocois and thus other ornamental palms are planted where coconut fruit is not required.

USA. Hawaii: Kauai, from diseased fruit (husk) of Cocos nucifera L., J.Y. Uchida H1024, May 1990, dried culture specimen, holotype PDD 103199; ex-holotype living culture preserved in a metabolically inactive state as ICMP 16948.

AUSTRALIA. New South Wales: Wyong, soil under Eucalyptus pilularis Sm., 1976, L. Gerrettson-Cornell, culture ATCC 64863 = DAR 27023 = ICMP 16691 = IMI 208224= WPC P3475. CHINA. Hainan Island: forest soil, Hon Ho, culture LMM6 = ICMP 17964 = WPC P10660. GUATEMALA. Los Aposentos: canker on trunk of Persea americana Mill., 1975, G.A. Zentmyer, culture ATCC 38692 = CBS 954.87 = ICMP 19452 = WPC P1000. MALAYSIA. Trengganu: Jerangau Estate, Theobroma cacao, 14 Jan 1968, P.D. Turner, specimen Herb. IMI 131093, culture ICMP 16914 = IMI 131093 = WPC P0578. USA. North Carolina: dieback of Rhododendron, 1976, DM Benson, culture 1400 = ATCC 46299 = CBS 958.87 = ICMP 19453 = WPC P1717.

MALAYSIA. “P. heveae Thompson, showing sporangia and oospore formation”, Plate 1D (line drawing) in A.W. Thompson, Malayan Agricultural Journal 17(3–4): 53–100 (1929), lectotype designated here [IF550520]. MALAYSIA. Selangor: pod rot of Hevea brasiliensis (Willd. ex A. Juss.) Müll. Arg., Dec. 1927, A.W. Thompson, culture preserved in a metabolically inactive state, CBS 296.29 (= IMI 180616 = WPC P3428 = ATCC 58815 = ICMP 19451 = KACC 44943), epitype designated here [IF550521]. Thompson (1929) did not designate a type specimen in his manuscript (not a requirement for valid publication prior to 1958). However, in the same year he deposited a culture of P. heveae as CBS 296.29. In 1974 this culture was deposited as IMI 180616, and in 1980 a dried culture of CBS 296.29 was deposited in the CBS herbarium as CBS H-7641. These cultures have the note “isotype” in the collection databases, but we have not been able to find any publication mentioning the typification of P. heveae, and the dates of deposition are too far apart for the cultures to be accepted as isotypes. Thus, to clarify the typification of P. heveae we designate Thompson’s original protologue line drawing of sporangia and oogonia (reproduced here as Fig. 7) as the lectotype, and his authentic culture as the epitype. Description:—A description of this species is provided by Thompson (1929), and more recently by Erwin & Ribeiro (1996). Descriptions of sporangial caducity are inconsistent in the secondary literature. Caducous sporangia were not noted by Thompson despite extensive description of the sporangia on different media; if free sporangia had been observed he would very likely have described them, as he did for another species (P. palmivora) in the same article. The epitype culture CBS 296.29 = ICMP 19451 is described by the fungal database at Q-bank as non-caducous (www.q-bank.org). A P. heveae isolated from Rhododendron in North Carolina, USA (1400 = ICMP 19453) had persistent sporangia (Benson & Jones 1980). However, Albuquerque et al. (1974) reported caducous sporangia on P. heveae isolated from leaf blight of Brazil nut (Bertholletia excelsa Bonpl.), but also reported some paragynous antheridia, a character not observed in P. heveae in our research, nor reported by others. We examined sporangial caducity for an additional two isolates (ICMP 16691 and ICMP 19452) and found only persistent sporangia, even with very vigorous shaking in an attempt to dislodge the sporangium. Therefore we emend the description of P. heveae to include persistent (non-caducous) sporangia, the same characteristic as all other Phytophthora Clade 5 species (Table 3).