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Stigmatomyces from New Zealand and New Caledonia: new records, new species and two new host families

     350 Illick Hall, 1 Forestry Drive, State University of New York College of Environmental Science and Forestry, Syracuse, New York 13210

Walter Rossi

     Dipartimento di Scienze Ambientali, Università dell’Aquila, 67010 Coppito, L’Aquila, Italy

	ABSTRACT

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS TAXONOMY KEY TO SPECIES LITERATURE CITED

 

Seven new records and three new species of Stigmatomyces (S. australis, S. baeopteri and S. clasiopellae), a genus of Laboulbeniales associated only with Diptera, are described from New Zealand and New Caledonia. Two new host families for Laboulbeniales are recorded, and a dichotomous key to Stigmatomyces species in New Zealand and New Caledonia is presented.

Key words: Insect-associated fungi, Laboulbeniales

	INTRODUCTION

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS TAXONOMY KEY TO SPECIES LITERATURE CITED

 
Despite the high endemism of New Zealand’s entomofauna and the marked host specificity of Laboulbeniales, only 18 species have been recorded from New Zealand (Thaxter 1931, Rossi 1984, Weir and Rossi 1997, Rossi and Weir 1998, Weir 2001). Recent and ongoing studies (Weir and Rossi 1997, Weir 2001) have nearly doubled the number of previously known species from this region, with the addition of a new genus, Triainomyces (Rossi and Weir 1998). Mouchacca (2000) lists no publications recording Laboulbeniales from New Caledonia; he also notes that mycological knowledge of this Gondwanan fragment, in general, is quite poor in comparison to the tropical flora.

As currently circumscribed, Stigmatomyces is a large genus comprising approximately 130 species and is associated exclusively with Diptera worldwide. Stigmatomyces species are quite variable but are characterized by Thaxter (1931) as having a two-celled receptacle, 2–4 ascogenic cells and four tiers of perithecial wall cells that correspond roughly to the venter, neck, tip and apex. The appendage is characteristic and consists of a stalk cell and a basal cell, the latter of which gives rise to a row of appendage cells and may bear antheridia (Thaxter 1908, Thaxter 1931). Antheridia are found singly or in groups of 2–3 (rarely 4), arising in a vertical series from the appendage cells or smaller flanking cells.

Two new host families and two new species of Stigmatomyces were discovered in New Zealand; one new species was discovered in New Caledonia. Seven new records for New Zealand and a comprehensive dichotomous key to Stigmatomyces in both countries are presented.

	MATERIALS AND METHODS

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS TAXONOMY KEY TO SPECIES LITERATURE CITED

 
Pinned specimens of flies carrying thalli of Laboulbeniales were obtained on loan from the New Zealand Arthropod Collection (NZAC) housed at Landcare Research in Auckland, New Zealand. Flies also were collected on the North Island by A. Weir and C. Carlton during Mar 2000, on the South Island by R. Meier and D. McAlpine in Jan 2000 and A. Weir and M. Hughes in Jan and Feb 2001, on Stewart Island by R. Didham and J. Ward in Jan 2000, and in Christchurch by M.H. Moore in Oct 2000. Flies were collected in New Caledonia by W.T. Starmer in Feb 2000. Fungi were mounted in glycerol or Amann’s lactophenol as outlined by Benjamin (1971) and were examined with a Nikon Eclipse E800 research microscope fitted with DIC optics. Photomicrographs were prepared with a Nikon Coolpix 995 digital camera. Holotypes of the newly described species are deposited in the New Zealand Fungal Herbarium (PDD) and the Mycological Herbarium at the SUNY College of Environmental Science and Forestry (SYRF). Isotypes and all other material are deposited in SYRF and in the collection of Walter Rossi. Localities are recorded as outlined in Crosby et al (1976), and all specimens examined are from New Zealand unless indicated otherwise.

	TAXONOMY

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS TAXONOMY KEY TO SPECIES LITERATURE CITED

 
Stigmatomyces australis sp. nov. M. B. Hughes, A. Weir, B. Gillen and W. Rossi. (fIGS. 1–3, 13)

View larger version (81K): [in this window] [in a new window]   FIGS. 1–6. Mature thalli of Stigmatomyces spp. All scale bars = 50 µm. FIGS. 1–3. Stigmatomyces australis. FIGS. 4, 5. Stigmatomyces baeopteri. FIG. 6. Stigmatomyces clasiopellae.

View larger version (59K): [in this window] [in a new window]   FIGS. 13–23. Appendage details of Stigmatomyces spp. Scale bar = 50 µm. FIG. 13. Stigmatomyces australis. FIG. 14. Stigmatomyces baeopteri. FIG. 15. Stigmatomyces clasiopellae. FIG. 16. Stigmatomyces ephydrae. FIG. 17. Stigmatomyces novozelandicus. FIG. 18. Stigmatomyces crassicollis. FIG. 19. Stigmatomyces limosinae. FIG. 20. Stigmatomyces hydrelliae. FIG. 21. Stigmatomyces spiralis. FIG. 22. Stigmatomyces purpureus. FIG. 23. Stigmatomyces rugosus.

Slender, straight to slightly curved. Thallus hyaline to light amber yellow, except for basal cell of appendage, which is distinctly darker. Receptacle of nearly uniform width or slightly tapering to the foot, with the basal cell (I) almost as long as the suprabasal (II). Stalk cell (III) of the appendage rather narrow and elongate, slightly longer than the adjacent stalk cell of the perithecium. Free appendage outcurved or sigmoid, the axis composed of 5–9 relatively large obliquely superposed cells, the basal short, the others longer than broad, progressively smaller, slightly prominent externally. Each cell of the axis gives rise on the inner, upper margin to an elongate cell, which lies close to the adjacent distal cell of the axis and bears apically one antheridium externally and a smaller cell internally. The latter bears 1–2 antheridia. These are replaced in the upper portion of the appendage by long, sometimes curled, somewhat disorganized cells. Stalk cell of perithecium relatively large and elongate. Secondary stalk and basal cells relatively large, together forming a free stalk that is distally broader and hardly distinguished from the evenly oval venter; the latter tapers without abrupt constriction to the stout, slightly curved neck, which is about as long or longer than the venter, distally broadened at its junction with the tapering tip and apex, the latter ending in three rather well defined prominences of different heights.

Measurements. – Receptacle: 12.5–25 x 39–178 µm. Total length of appendage: 68–158 µm. Total length of perithecium: 165–380 µm. Venter 30–53 x 67–235 µm. Neck 10–15 x 47–200 µm. Total length of thallus: 200–600 µm. Antheridia 4–15 x 9–17 µm. Ascospores 3–4 x 15–30 µm.

Etymology. – Named for the southern hemisphere.

Specimens examined. – HOLOTYPE: STEWART ISLAND: Christmas Village Hut, on all parts of Icaridion debile (Coelopidae) collected at ultraviolet light, 17 Jan 2000, J. Ward (PDD 72041). ISOTY PE: STEWART ISLAND: Christmas Village Hut, on all parts of Icaridion debile collected at ultraviolet light, 17 Jan 2000, J. Ward, AW940. PARATYPES: STEWART ISLAND: Christmas Village Hut, on all parts of Icaridion nigrifrons (Lamb) collected at ultraviolet light, 17 Jan 2000, J. Ward, AW941 and WR2533. STEWART ISLAND: Smoky Beach, on the legs and thorax of I. nigrifrons collected at ultraviolet light, 14 Jan 2000, R.K. Didham, AW942. KAIKOURA: Paratal Point, on Australimyza? setigera Harrison (Australimyzidae), 26–27 Jan 2000, R. Meier and D.K. McAlpine, AW962, WR2543, WR2544, WR2545. KAIKOURA: on Icaridion debile, 26–28 Jan 2000, D.K. McAlpine and R. Meier, WR2575, WR2576, WR2616. CAMPBELL ISLAND: Lookout Bay, on legs of Australimyza anistomae Harrison, 14–23 Jan 1969, G. Kuschel, AW946. AUCKLAND ISLANDS: West Coast opposite Mount Story, on anterior left wing of A. anistomae collected on Hebe sp., 17 Jan 1963, J.L. Gressitt, AW947. AUCKLAND ISLANDS: Adams Island, Magnetic Cove, on legs, anterior tergites and left, mid-mesonotum of A. anistomae, 20 Jan 1966, K.A.J. Wise, AW948. AUCKLAND ISLANDS: Adams Island, Magnetic Cove, Malaise trap near shore, on anterior left wing, legs, anterior tergites and left, mid-mesonotum of A. anistomae, 29 Jan 1966, K.A.J. Wise, AW949. AUCKLAND ISLANDS: Adams Island, Magnetic Cove, on anterior left wing of A. anistomae, 3 Feb 1966, K.A.J. Wise, AW950. AUCK-LAND ISLANDS: Crozier Point, on thorax and legs of Icaridion nasutum Lamb on kelp, 28 Dec 1962, J.L. Gressitt, AW943.

Commentary – The appendages of Stigmatomyces australis and Stigmatomyces baeopteri (described below) are unusual. The antheridia arise from distinct cells and their short stalks, adjacent to those of the main axis of the appendage. The appendages of both species are structurally similar to Stigmatomyces divaricatus Thaxt. from Cameroon on Borborus aptus Curran (Thaxter 1931) and, to a lesser extent, other species described by Thaxter on "Borboridae": Stigmatomyces borbori Thaxt., S. borboridinus Thaxt., S. divergatus Thaxt., and S. grenadinus Thaxt. (Thaxter 1931). Borborus aptus (now Gymnometopina apta (Curran)) is a member of the Sphaeroceridae. Most of the other species that Thaxter described from Sphaeroceridae, such as S. limosinae Thaxt. and S. crassicollis Thaxt., also have externally rounded cells of the main axis of the appendage, but in each series antheridia are only once removed from the cells of the main axis by a very short stalk cell. The variation in S. australis is notable. Thalli on Icaridion (fIGS. 1–3) are lighter in color and have more elongate appendages with more slender and more numerous cells in the main axis than those on Australimyza.

Approximately 90% (n = 122) of Coelopidae examined from Stewart Island were infected with S. australis or S. baeopteri. Other relatively high rates of Stigmatomyces infection (18%) on adult flies also have been observed on Amma blancheae McAlpine from Australia (McAlpine 1991). McAlpine (1991) suggests that the longevity of the adult form of this fly make it a particularly suitable host compared to other coelopids. Other factors of infection that have been implicated in northern temperate habitats include overlapping of mating generations, overwintering as adults and a moist climate (Weir and Hammond 1997, Hedström 1994, Hedström and Monge-Nájera 1998). Most species of both host genera are found near the ocean shore in New Zealand (Griffiths 1972).

Stigmatomyces australis and Stigmatomyces baeopteri are the first species of Laboulbeniales to be recorded from Coelopidae and Australimyzidae. Unidentified species of Stigmatomyces have been reported incidentally on five of the 28 known species of Coelopidae, all from Australia (McAlpine 1991): A. blancheae, Gluma keyzeri McAlpine, G. musgravei McAlpine, G. nitida McAlpine and Coelopella popeae McAlpine (McAlpine 1991).

Stigmatomyces baeopteri sp. nov. M. B. Hughes, A. Weir, and W. Rossi. (fIGS. 4, 5, 14)

Stigmatomyceti australi affinis, a quo differt validiore habitu, appendice longiore, antheridiis minoribus, perithecii collo breviore ac latiore, perithecii apice magis incurvato.

Slender and straight except for the perithecial neck, which often is curved. Thallus light amber yellow, except for the perithecial neck which is lighter or hyaline. Receptacle tapering gradually to the foot, of nearly uniform width. Cells I and II of receptacle subequal. Stalk cell of appendage (III) elongate, slightly longer than the adjacent stalk cell of the perithecium (VI). Free appendage crescent-shaped, sigmoid, or coiled distally, the axis composed of 6–10 relatively large obliquely superposed cells, the basal short, the others longer than broad, progressively smaller. Each cell gives rise on the inner margin to a cell that is isodiametric or slightly longer than broad and lies close to the adjacent distal cell of the axis, bearing apically one antheridium externally and a smaller cell internally. The latter bears two antheridia. The tip of the appendage may become disorganized, the lowermost 6–7 cells being fertile. Stalk cell of perithecium (VI) subrectangular. Secondary stalk cell (VII) and basal cells relatively large, forming a free stalk that is slightly less wide than the narrowly elliptical venter. The latter tapers gradually to the curved neck, which is distally and prominently broadened in several bulges at its junction with the tapering tip and apex. The apex is formed by one small lateral and two larger apical lips, all of different heights.

Measurements. – Receptacle: 17–24 x 82–120 µm. Total length of appendage: 70–180 µm. Total length of perithecium: 180–240 µm. Venter: 26–55 x 50–110 µm. Neck: 16–20 x 70–130 µm. Total length of thallus: 246–480 µm. Antheridia: 3–6 x 10–12 µm. Ascospores: 4 x 30–39 µm.

Etymology. – Named for the host genus, Baeopterus (Coelopidae).

Specimens examined. – HOLOTYPE: STEWART ISLAND: Christmas Village Hut, on legs of Baeopterus sp. collected at ultraviolet light, 17 Jan 2000, J. Ward, (PDD 72042). ISO-TYPE: same data as holotype except legs and mesonotum. AW944, WR2532. PARATYPE: STEWART ISLAND: Smoky Beach, on mesonotum of Baeopterus philpottii (Malloch), R.K. Didham, AW945.

Commentary. – This species differs from the allied S. australis in its more robust habit, the wider and generally shorter perithecial neck and the longer appendage with smaller antheridia. Ascospores are longer than, and lie outside the range of, those of S. australis. Moreover, the neck and perithecial tip region are curved inward toward the appendage. See further ecological remarks under S. australis.

Stigmatomyces clasiopellae sp. nov. M.B. Hughes, A. Weir, and W. Rossi (fIGS. 6, 15)

Fungus brevis ac compactus, parum arcuatus aut sigmoideus, fuscoluteus, praeter receptaculum luteum et appendicis basalem cellulam cum exteriore cellulae appendicem sufferentis parte ferruginea. Receptaculum breve, ad basem valde angustatum. Basalis cellula quasi triangula, postice incurvata. Suprabasalis cellula trapezii instar, paulo longior quam basalis. Cellula appendicem sufferens cordi instar. Appendix brevis, 4 cellulis confecta, quarum basalis et apicalis bina, ceterae vero singula anteridia ferunt. Cellulae perithecium subiacentes suppares. Perithecii venter aeque ovatus, sine coarctatione producitur in validum collum, aeque longum ac venter, prae apicem inflatum. Apex directus aut ante versus, in parva supparia labia desinens.

Thalli relatively short and compact, slightly curved or sigmoid. Receptacle yellowish; basal cell of the free appendage and outer margin of its stalk cell rusty brown; the rest of the fungus is brownish yellow. Receptacle relatively short, strongly tapered below. Basal cell (I) subtriangular, more or less curved posteriorly. Suprabasal cell (II) subtrapezoidal, slightly longer than the basal. Stalk cell (III) of the appendage cordiform, its tapered base slightly overlapping the suprabasal cell of the receptacle. Appendage short, composed of four cells, the basal of which bears two antheridia, the others bearing two each, the series terminated by an eighth spinose antheridium. Perithecial stalk and basal cells subequal. Perithecial venter symmetrically ovoid, tapering without abrupt transition to the broad base of the neck, which is about as long as the venter, almost straight or curved slightly anteriorly, its margins very slightly concave, distally broader at its junction with the stout, tapering tip. The latter is straight to strongly bent anteriorly, ending in very small, subequal, hyaline lips.

Measurements. – Receptacle: 24–36 x 60–86 µm. Basal cell of appendage: 10 µm. Stalk cell of appendage: 20–23 µm. Total length of appendage: 48–53 µm. Total length of perithecium: 148–198 µm. Venter: 40–60 x 63–93 µm. Neck: 23–28 x 85–105 µm. Total length of thallus: 220–309 µm. Antheridia: 9 x 10 µm. Ascospores: 5 x 28 µm.

Etymology. – named for the host, Clasiopella.

Specimens examined. – HOLOTYPE: NEW CALEDONIA: near Nouméa: Mount Koghis, on Clasiopella austra Mathis (Ephydridae), 6 Feb 2000, W.T. Starmer, (AW873). ISO-TYPE: same data as holotype, in SYRF (AW873).

Commentary. – The only known species of Stigmatomyces with which this new species shares a similar habit is S. rampinii W. Rossi, described from Senegal by Rossi (1988). However, the lowermost and uppermost cells of the appendage in S. rampinii both have two antheridia, while the middle cell has only one.

Stigmatomyces rugosus Thaxt. (fIGS. 7, 23)

View larger version (83K): [in this window] [in a new window]   FIGS. 7–12. Mature thalli of Stigmatomyces spp. All scale bars = 50 µm. FIG. 7. Stigmatomyces rugosus. FIG. 8. Stigmatomyces purpureus. FIG. 9. Stigmatomyces ephydrae. FIG. 10. Stigmatomyces crassicollis. FIG. 11. Stigmatomyces hydrelliae. FIG. 12. Stigmatomyces limosinae.

(= S. harantii H.L. Dainat)

Specimens examined. – MID-CANTERBURY: Christchurch: Mona Vale, on tergites of female Psilopa metallica (Hutton) aspirated over Nymphea, 21 Feb 2001, A. Weir and M. Hughes, AW961. MID-CANTERBURY: Christchurch: Mona Vale, on midtergites of male Psilopa cf. flavimana (Hendel) (possibly a new species) aspirated over Nymphea, 21 Feb 2001, A. Weir and M. Hughes, AW925. MID-CANTERBURY: Christchurch: Hagley Park, Christchurch Botanic Garden, on mesonotum and midtergites of female P. metallica, sweeping plants, 5 Oct 2000, M.H. Moore, AW926. MID-CANTERBURY: Banks Peninsula: Duvauchelle, on ventral thorax just anterior to midlegs of male P. metallica, sweeping intertidal area at low tide, 21 Feb 2001, A. Weir and M. Hughes, AW927. TONGARIRO: Waikato River near Orakei-Korako Thermal Park, sweeping shoreline debris, on tergites of male P. metallica, 14 Mar 2000, A. Weir and C. Carlton, AW928.

Known distribution and hosts. – Stigmatomyces rugosus is a cosmopolitan fungus on members of the genus Psilopa. It has been reported in Europe (Santamaria et al 1991), Africa, Thailand and the Bismarck Archipelago (near New Guinea) (Rossi 1998). Rossi (1998) reported this fungus on Leptopsilopa nigrimana Williston in Bolivia. Stigmatomyces rugosus (as S. psilopae) originally was described from Jamaica and Grenada (Thaxter 1931, Rossi 1998).

Commentary. – The material described here differs from that of Thaxter (1908) in the lack of rugosity in the venter of the perithecium and in the length of the specimens, which range from 215 µm to 600 µm. The synonymy of S. psilopae and S. rugosus probably accounts for these differences. S. psilopae was described to be much more variable in length, with a maximum total length of 540 µm and with an often indistinct verrucosity of the perithecium.

Rossi (1988) notes that the number of cells in the appendage is variable (4–6) and that only specimens growing on the wings of insects are relatively uniform in having a four-celled appendage and smooth perithecium. We did not observe any specimens on the wings but did notice variation in the number of both appendage cells and antheridia. Rugosity varied more with position of growth than with geographic locality. Santamaria and Rossi (1998) record specimens up to 540 µm in length from the coxae and indicate that shorter specimens were found growing on the wings and mesonotum. The variability in thallus length for our specimens was not linked to specific positions of growth on the host.

Stigmatomyces purpureus Thaxt. (fIGS. 8, 22)

(= S. scatellae Batra)

(= S. purpureus f. scatellae (Batra) Balazuc)

Specimens examined. – GISBORNE: Urewera National Park, Black Beech Track at Lake Waikaremoana, on posterior tergites of female Scatella subvittata Tonnoir & Malloch, 18 Mar 2000, sweeping bank of lake in late afternoon, A. Weir and C. Carlton, AW960. GISBORNE: Urewera National Park, Black Beech Track at Lake Waikaremoana, on sternites of S. nitidithorax Malloch, 18 Mar 2000, sweeping bank of lake in late afternoon, A. Weir and C. Carlton, AW937. TAUPO: Lake Taupo, Waihi Beach, on legs, sternites, and behind the right eye of male, sternites of female S. nitidithorax, 13 Mar 2000, sweeping beach debris, A. Weir and C. Carlton, AW938. GISBORNE: Urewera National Park, Lake Waikaremoana, sweeping shoreline between Waikaremoana and Aniwaniwa, on leg of male S. nitidithorax, and tergites of female S. nitidithorax and S. subvittata, 7 Mar 2000, A. Weir and C. Carlton, AW939. AUCKLAND: Lynfield, surface of swimming pool, on abdomen of S. nitidithorax, 11 Dec 1983, B.A. Holloway, AW686.

Known distribution and hosts. – This species generally is found on the abdomen of members of the genus Scatella. It has been reported throughout Europe (Weir and Rossi 1995, Majewski 1994, Santamaria et al 1991, Santamaria and Rossi 1993), North America (Santamaria et al 1991), Asia (Santamaria et al 1991) and Algeria (Rossi 1988).

Commentary. – The measurements and description of the New Zealand material correspond well to descriptions of the fungus in both Majewski (1994) and Thaxter (1908). This is the first record of S. purpureus on S. subvittata. Some thalli were found growing behind the right eye of a male. Infections on the face also were found in a closely related fungus, S. hydrelliae, in Britain (Weir and Rossi 1995). Other positions of growth are consistent with the hypothesis of sexual transmission.

Stigmatomyces ephydrae Mercier & Poiss. (fIGS. 9, 16)

(= S. bottnicus Huldén)

(= S. setacerae Huldén)

Specimens examined. – AUCKLAND: Miranda, on mesonotum and legs of male and female Ephydrella novaezelandiae (Tonnoir & Malloch) sweeping muddy margins of salt marsh, 27 Mar 2000, A. Weir and C. Carlton, AW934. GIS- BORNE: Urewera National Park, on Black Beech Track at Lake Waikaremoana, on legs, tergites and mesonotum of male and female Ephydrella aquaria (Hutton) sweeping banks of the lake in late afternoon, 18 Mar 2000, A. Weir and C. Carlton, AW935. GISBORNE: Urewera National Park, Lake Waikaremoana between Waikaremoana and Aniwaniwa, on mesonotum, sternites, tergites, hind legs of male and female E. aquaria, 18 Mar 2000, A. Weir and C. Carlton, AW936.

Known distribution and hosts. – This species has been reported in France, Finland and Russia (Santamaria et al 1991) on Ephydra and Setacera. S. ephydrae has been reported more recently from Italy and the U.K. (Rossi 1993, Weir and Rossi 1995).

Commentary. – Mercier and Poisson (1927) described this species as having a total length of 240–300 µm, while British specimens are 200–235 µm. The New Zealand specimens are up to 400 µm in length. Finnish specimens (recorded as the two synonyms listed above) differ only in the tip of the perithecium, which is variable, and not necessarily variable by host as Rossi (1993) has indicated. Observations of the New Zealand material support the synonymy.

Stigmatomyces crassicollis Thaxt. (fIGS. 10, 18)

(= S. papuanus Thaxt. var. leiostoma Maire)

(= S. longicollis Thaxt.)

(= S. hackmanii Huldán)

Specimens examined. – GISBORNE: Urewera National Park, Lake Waikaremoana, on mesonotum of Pseudocollinella maculifrons (Becker) collected by sweeping shoreline between Lake Waikaremoana and Aniwaniwa, 7 Mar 2000, A. Weir and C. Carlton, AW929. GISBORNE: Urewera National Park, Black Beech Track at Lake Waikaremoana, on unknown host, and on head, anterior mesonotum and right mid-tergites of P. maculifrons collected by sweeping bank of lake, 18 Mar 2000, A. Weir and C. Carlton, AW930. TAUPO: Lake Taupo, Waihi Beach, 38°57'S, 175°45'E, behind right eye of P. maculifrons collected by sweeping over beach debris, 13 Mar 2000, A. Weir and C. Carlton, AW931.

Known distribution and hosts. – This fungus has been recorded in many European countries and in Jamaica and North Africa on Limosina, Leptocera, Spelobia, Pseudocollinella (formerly Opacifrons), Opalimosina, Kimosina and Terrilimosina (Sphaeroceridae) (Thaxter 1931, Majewski 1994, Santamaria et al 1991, Santamaria and Rossi 1993, Weir and Rossi 1995) and in North Africa. The fungus recently was collected in Bolivia on Rachispoda merga Wheeler (Rossi 1998).

Commentary. – Thaxter (1931) notes that this species is related to S. papuanus but is distinguished by its truncate, unmodified apex with no conspicuous protrusion. Majewski (1994) reports this as a variable species with a thick, stout receptacle, the longest Polish specimens occurring on Leptocera (290–430 µm) and the appendage consisting of 10–13 superposed cells, with thalli on Spelobia possessing eight slightly flattened cells. The New Zealand material displays 9–15 cells, but these are not as flattened as the British material described by Weir and Rossi (1995). Thaxter noted great variation in the length of both the appendage and thallus, the latter being dependent on position of growth, with those on the legs being stouter. We also found similar variation, although this could not be associated with position of occurrence. Overall, the size of the New Zealand material is consistent with other records but the thalli are darker. The major difference from Thaxter’s description of the fungus is in the number of appendage cells, which originally were described as 9–10.

The paired sessile and stalked antheridia in each series were illustrated by Thaxter but not explicitly mentioned in the description. Weir and Rossi (1995) use this character in their key to British species of the genus. Further collections of Stigmatomyces species possessing paired antheridia arising putatively directly from the appendage cells should be examined in detail for the presence of short antheridial stalk cells described above. The nature of the antheridial arrangement is clear in appendages separated from the thallus and viewed under high magnification.

Stigmatomyces hydrelliae Thaxt. (fIGS. 11, 20)

Specimens examined. – TARANAKI: Waitaanga Forest, sweeping the bankside at confluence of Morgan Stream and Tangarakan River, on midsternites of unknown host, 11 Mar 2000, A. Weir and C. Carlton, AW953. GISBORNE: Urewera National Park, Black Beech Track at Lake Waikaremoana, sweeping bank of lake in late afternoon, on sternites and tergites of female Hydrellia enderbii (Hutton), 18 Mar 2000, A. Weir and C. Carlton, AW954. GISBORNE: Urewera National Park, Black Beech Track at Lake Waikaremoana, sweeping bank of lake in late afternoon, on trochanter and sternites of male and sternites of female Hydrellia velutinifrons Tonnoir and Malloch, 18 Mar 2000, A. Weir and C. Carlton, AW955. GISBORNE: Urewera National Park, Lake Waikaremoana, sweeping shoreline between Waikaremoana and Aniwaniwa, on tergites of male and female H. velutinifrons, coxae of male H. velutinifrons, 7 Mar 2000, A. Weir and C. Carlton, AW957. TONGARIRO: Waikato River near Orakei-Korako Thermal Park, sweeping shoreline debris, on tergites of male H. enderbii, 14 Mar 2000, A. Weir and C. Carlton, AW958. MID-CANTERBURY: Christchurch, Mona Vale, on midtergites of H. enderbii, 21 Feb 2001, A. Weir and M. Hughes, AW959. GISBORNE: Urewera National Park, Black Beech Track at Lake Waikaremoana, near motor camp, on posterior sternites of Hydrellia velutinifrons Tonnoir & Malloch, 18 Mar 2000, A. Weir and C. Carlton, AW956.

Known distribution and hosts. – This fungus has been recorded throughout Europe (Santamaria et al 1991, Majewski 1994, Weir and Rossi 1995, Santamaria and Rossi 1993) and the U.S.A. (Thaxter 1901) on several members of Hydrellia.

Commentary. – This represents the first record of the fungus for the Southern Hemisphere on two new hosts of Hydrellia, suggesting a worldwide, or at least temperate, distribution. These specimens are different from previous records in that the tips of the perithecia are rounded, with no apical projections. Also, the perithecial wall cells are not spiralled as are most of the European and American collections. Weir and Rossi (1995) indicate that the position of growth on the British hosts (on the face) supported the hypothesis of spread other than sexual means; infection patterns here are consistent with both sexual contact of the hosts, or indirect infection and autoinfection.

Stigmatomyces limosinae Thaxt. (fIGS. 12, 19)

(= Stigmatomyces pedunculatus T. Majewski)

Specimens examined. – AUCKLAND: Miranda, on posterior tergites and sternites, midcoxae, mid-left tibia, and left hind femur of Rachispoda fuscipennis (Haliday). Collected by sweeping muddy margins of salt marsh, 27 Mar 2000, A. Weir and C. Carlton, AW932. UNITED KINGDOM: DEVON: Slapton Ley, on leg of Leptocera sp. (Sphaeroceridae), 17 Aug 1995, A. Weir, AW718.

Known distribution and hosts. – Thaxter’s specimens were collected in the eastern and western United States, Mexico and Jamaica (Thaxter 1931). This species also occurs in Poland (Majewski 1994), England (Weir and Rossi 1995), Spain (Santamaria and Rossi 1993) and Italy (Rossi 1993). Hosts are members of Rachispoda.

Commentary. – Thaxter (1931) notes that the Californian material differs constantly in the perithecium, which is longer and less distinctly inflated, its apex showing no modification of the lip cells. The New Zealand material shares these characteristics. The distinctly inflated perithecial venter as seen in the British specimens (Weir and Rossi 1995; FIG. 12) was not observed. In Poland the species appears to be quite variable (Majewski 1994, Pl. 80; FIGS. 1–16) in size, morphology of the perithecial apex and shape of the perithecium and perithecial stalk cell (VI). The vast majority of the New Zealand material, although collected at one locality, showed immense variability in these characters. The main difference between S. limosinae and S. crassicollis traditionally has been regarded as the shape of the perithecial stalk cell (VI), being distinctly oblique in the latter and subrectangular in the former. Observations of the New Zealand material and illustrations of Polish specimens (Majewski 1994, Pl. 80; FIGS. 1–16) do not support this as a defining character, and even Thaxter (1908) noted that specimens of S. limosinae from California had very oblique stalk cells. Although not regarded as taxonomic characters in the genus, the most consistent differences between the two species in New Zealand are in the color of the thallus, shape of the perithecial apex and the length of the appendage with respect to the perithecium. In New Zealand, the specimens of S. limosinae consistently are smaller than S. crassicollis in all characters except antheridial length.

Stigmatomyces novozelandicus A. Weir & W. Rossi (FIG. 17)

Commentary. – This fungus has not been collected since the initial report by Weir and Rossi (1997). It is the tallest known species of Stigmatomyces, ranging in length from 1000 µm to nearly 1400 µm.

Stigmatomyces spiralis Thaxt. (FIG. 21)

Known distribution and hosts. – Hosts include members of Hyadina and Philygria. S. spiralis has been found in Italy (Rossi 1993), Spain (Santamaria and Rossi 1993), France (Rossi 1993, Santamaria et al 1991) and the U.S.A. (Santamaria et al 1991, Rossi 1993).

Commentary. – No additional collections have been made since the initial report by Weir and Rossi (1997).

Stigmatomyces ceratophorus Whisler

(= Fanniomyces ceratophorus [Whisler] T. Majewski)

Specimens examined. – TAUPO: on mid- and posterior tergites, mesonotum and head of Muscidae indet., on window-sill, 21 Feb 1935, J.S. Armstrong, AW951.

Known distribution and hosts. – Known on Fannia canicularis (L.) from California. A similar form was found by I.I. Tavares in Thaxter’s collection of undescribed Stigmatomyces in the Farlow Herbarium, Cambridge, Massachusetts, and is from an anthomyid fly collected in Guatemala (Whisler 1968). Individuals also have been found in Poland on the legs of males and abdomen of male and female flies in urban houses (Majewski 1994).

Commentary. – Although the material is in somewhat poor condition, it is easily recognizable as S. ceratophorus by the distinct branching of the appendage. The presence of this fungus in New Zealand suggests introduction into the country on its host. "Fannia" spp. are found in farm habitats where dung or urine is present.

Stigmatomyces ilytheae Thaxt.

Specimens examined. – AUCKLAND: Huia, side of Whatipu Road, on female Zeros invenatus Lamb, 26 Apr 1970, H.A. Oliver, AW952.

Known distribution and hosts. – This fungus has been reported from Jamaica, Grenada, Maine, Borneo and Cameroon (Thaxter 1931). The only record after 1931 is from Bolivia on Zeros fenestralis (Cresson) (Rossi 1998).

Commentary. – Although this fungus is variable, the single specimen from New Zealand is remarkably similar to Thaxter’s original description and illustration (Thaxter 1931, Pl. XXVII; FIG. 4). The discovery of this fungus in New Zealand suggests it might be quite cosmopolitan.

	KEY TO SPECIES

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS TAXONOMY KEY TO SPECIES LITERATURE CITED

 

1. Extremely long and slender, the suprabasal cell of the receptacle (II) contributing more than half the length of the thallus; more than 1000 µm in length (on Lauxanidae). S. novozelandicus 1'. Not extremely slender, the suprabasal cell (II) not contributing more than half the length of the thallus; less than 1000 µm in length (not on Lauxanidae). 2     2. Perithecial wall cells distinctly spiraled 3     2'. Perithecial wall cells not distinctly spiraled 5 3. Appendage with dichotomous branching near the base; antheridia pedunculate, borne on tips of the appendage branchlets (on Muscidae) S. ceratophorus 3'. Appendage not branching dichotomously near the base; antheridia sessile, borne on an axis of superposed cells (not on Muscidae) 4     4.Perithecial apex consisting of four papillae arranged around a somewhat apical projection; cell II of receptacle nearly twice as long as cell I S. spiralis     4'. Perithecial apex consisting of two elongate, erect external lips and two smaller internal ones; cells I and II of receptacle sub equal S. purpureus 5.Venter of perithecium with distal rounded bulges 6 5'. Venter of perithecium without distal rounded bulges 7     6. Appendage and uppermost antheridium erect; appendage 3-(rarely 4-)celled S. hydrelliae     6'. Appendage with curved outer margin, uppermost antheridium facing slightly inward; appendage usually 5-celled S. ephydrae 7. Appendage cells of main axis numbering 4–5 (rarely 6), their outer margin almost straight; antheridia sessile, borne directly on the cells of the axis (on Ephydridae) 8 7'. Appendage cells of main axis numbering 6 or more, their outer margin convex; antheridia borne directly on the cells of the axis and on stalks (on Sphaeroceridae, Australimyzidae or Coelopidae) 10     8. Surface of perithecial venter more or less rugose S. rugosus     8'. Surface of perithecial venter not rugose 9 9. Receptacle elongate. Perithecial neck tapering gradually to the tip, its apex bearing four small lips, one distinctly longer and subtriangular. Uppermost appendage cell bearing one terminal antheridium S. ilytheae 9'. Receptacle relatively short and strongly tapered below, the basal cell subtriangular. Perithecial neck tapered but distally broader at its junction with the tip, which is stout. Perithecial apex consisting of small subequal lips. Terminal appendage cell bearing two antheridia S. clasiopellae     10. Antheridia on lowermost cells of the axis borne in pairs; one sessile, the other on a very short stalk 11     10'. Antheridia in each series sometimes two, but usually three; always borne not on cells of the main axis but on relatively large adjacent superposed cells and small stalk cells 12 11. Apex of perithecium blunt, tip of appendage reaching above the middle of the venter. Stalk and basal cells of perithecium compact S. crassicollis 11'. Apex of perithecium consisting of small, rounded lips, tip of the appendage reaching the lower to middle portion of the venter. Stalk and basal cells of perithecium elongate S. limosinae     12. Perithecial neck wider (16 µm–20 µm) and generally shorter. Neck curved inward toward appendage. Appendage up to 180 µm with 6–10 cells in the main axis; sigmoid or coiled distally S. baeopteri     12'. Perithecial neck narrower (10 µm–15 µm) and generally longer. Neck straight and erect. Appendage up to 160 µm with 5–9 cells in the main axis; outcurved S. australis

	ACKNOWLEDGMENTS

 
We gratefully acknowledge J. Ward, M.H. Moore, D. Mc-Alpine, W.T. Starmer, and R.K. Didham for collections of host insects, S. Marshall, D. McAlpine, and P. Clausen for determination of hosts, C. Judd for curatorial assistance, and the New Zealand Arthropod Collection based in Auckland, New Zealand, for the loan of pinned specimens. We also would like to thank L. Cajani for reviewing the Latin diagnoses. This research was supported by the National Science Foundation Biotic Surveys and Inventories Program (NSF DEB 9972083 awarded to A. Weir, C. Carlton and R. Leschen).

	FOOTNOTES

 
Accepted for publication July 19, 2003.

¹ Corresponding author. E-mail: aweir{at}syr.edu

	LITERATURE CITED

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS TAXONOMY KEY TO SPECIES LITERATURE CITED

 
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