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Faculty of Environmental and Forest Biology, 350 Illick Hall, 1 Forestry Drive, State University of New York College of Environmental Science and Forestry, Syracuse, New York 13210
Richard Leschen
Landcare Research, Mount Albert Research Centre, Private Bag 92-170, Mount Albert, Auckland, New Zealand
Chris Judd
Ben Gillen
Faculty of Environmental and Forest Biology, 350 Illick Hall, 1 Forestry Drive, State University of New York College of Environmental Science and Forestry, Syracuse, New York 13210
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSTAXONOMYDISCUSSIONKEY TO SPECIES OF...LITERATURE CITED |
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Until now Rhachomyces kenodactyli Balazuc & W. Rossi has been the only species of Laboulbeniales known to occur on Coleoptera in the Bounty, Antipodes, Auckland, Campbell and Snares Islands, which lie 48° to 35° S. Four new species (Diphymyces depressus, Diphymyces leschenii, Laboulbenia subantarctica and Laboulbenia loxomeri) and five new records for the subantarctic (Cucujomyces phycophilus, Diphymyces penicillifer, Laboulbenia sp. 1, Rhachomyces sp. 1 and Teratomyces sp. 1) are reported, increasing the known number of taxa tenfold. An expanded geographic range for Rhachomyces kenodactyli is reported. A relatively high percentage (12%) of known beetle species in the subantarctic serve as hosts for Laboulbeniales. This host utilization rate is higher than that in tropical and north temperate regions. The high proportion of intertidal coleopteran taxa in the subantarctic fauna probably accounts for the greater number of host species utilized. Fungi on intertidal beetles (Omaliinae [Staphylinidae], Oopterus [Carabidae] and Kenodactylus audouini [Carabidae]) are known from many host individuals and collections, while those on terrestrial species are known from few, and in some cases, a single collection or host. The sporadic occurrence of some species encountered increases the likelihood that a few species of Laboulbeniales on Coleoptera probably remain undiscovered in the region.
Key words: biodiversity, biogeography, fungal distributions, insect-associated fungi, systematics
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYDISCUSSIONKEY TO SPECIES OF...LITERATURE CITED |
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). Of a total of 82 coleopteran families in New Zealand, 25 have been reported from one or more subantarctic islands (www.landcareresearch.co.nz/research/biodiversity/antarctica/html/beetlebase.html [Johnston et al 2002], Klimaszewski and Watt 1997, Leschen et al 2002). Fifteen of these 25 families have been recorded as hosts for Laboulbeniales in other parts of the world. Despite the diversity of potential hosts in the region and the worldwide occurrence of Laboulbeniales, only one species, Rhachomyces kenodactyli Balazuc & W. Rossi, has been reported from any group of subantarctic islands (Rossi 1984). Only two other taxa are known from the subantarctic region (Stigmatomyces australis M.B. Hughes et al and Stigmatomyces baeopteri M.B. Hughes et al). These are recently described and grow on Coelopidae (Diptera) (Hughes et al 2004). Both the endemic entomological fauna of the subantarctic region and the evident specificity of Laboulbeniales point toward the prospective fungal diversity in the region.
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYDISCUSSIONKEY TO SPECIES OF...LITERATURE CITED |
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, 1986). The fungi were observed on a Nikon Eclipse E-800 research microscope fitted with DIC optics. Determinations of freshly collected hosts were performed by R. Leschen. Holotypes, isotypes and paratypes of new species were deposited in the New Zealand Fungal Herbarium at Landcare Research (PDD). Some paratypes were deposited in the State University of New York College of Environmental Science and Forestry Mycological Herbarium (SYRF), with additional paratypes or isotypes at the Royal Botanic Gardens, Kew (K[M]). Using EstimateS version 6.01b (http://viceroy.eeb.uconn.edu/estimates [Colwell 2000]), a sample-based accumulation curve was constructed for the 2000 expedition. A curve was constructed for incidence data (fungal individuals of the same species on one or more hosts within the same collection considered as an encounter) using 100 randomizations of sample order without replacement.
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TAXONOMY |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYDISCUSSIONKEY TO SPECIES OF...LITERATURE CITED |
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Habit erect and robust. Receptacle and appendage hyaline. Perithecium tinged with brownish orange; olivaceous subterminally. Basal portion of appendage sometimes brownish yellow. Cell I obtriangular in optical section, the upper margin convex under the suprabasal cell (II) and the perithecial stalk cell (VI). Cells of main axis somewhat irregular in shape but equal in size, with rounded outer margins and thick walls. Cell II lying parallel to cell VI, about equal in size to cell I, with a straight inner and convex outer margin. Stalk cell of appendage isodiametric and superimposed by one or two pentagonal cells; the first pentagonal cell cut off obliquely on its upper margin by two or more smaller irregular corner cells, from which arise proliferating appendage branches. Appendage branches arising from cells IV, V and the accompanying smaller cells. Simple antheridia are borne on the appendage branchlets; seriate, intercalary and inconspicuous, lying parallel to the cells of the appendages. Perithecia arising from cells I and II. In the minority of cases, only one perithecium is formed, arising from cell I. Perithecial stalk cell (VI) about half again as long as cell I, slightly broader, and cut off obliquely on the upper margin by the basal cells. Cell VI obtriangular in optical section, the inner margin much more convex than the nearly straight outer margin. Basal cells of the perithecium subequal; of equal or slightly lesser height than the secondary stalk cell of the perithecium (VII). Perithecia narrowly subovoid, the inner margins less convex than the outer. Perithecial apices rounded, with one posterior lip cell slightly divergent.
Measurements. – Perithecium: 78–128 x 22–43 µm. Perithecial stalk cell: 12–44 x 11–29 µm. Length of main axis from foot to tip of appendage branches: 102–240 µm. Total length from foot to tip of perithecium: 124–210 µm.
Etymology.. Patronymic for one of the collectors, Rich Leschen, a specialist in beetle systematics.
HOLOTY PE: NEW ZEALAND. CAMPBELL ISLAND: Northwest Bay, 52°32'S, 169°8'E, 10–17 Mar 2000, on P. campbellicus, M. Bullians et al (PDD 72043).
PARATY PES: NEW ZEALAND. CAMPBELL ISLAND: Fence line from Tucker Cove, Col Ridge, 52°32'N, 169°8'E, 9 Mar 2000, on P. campbellicus, M. Bullians et al (SYRF AW972, PDD 72047); Beeman Hill, 52°32'S, 169°8'E, 10–17 Mar 2000, on P. campbellicus, M. Bullians, E. Edwards, R. Leschen (SYRF AW973, K[M]111876).
Known distribution and hosts. –
The only known host of Diphymyces leschenii is Paracatops campbellicus, which is restricted to Campbell Island (TABLE I). Diphymyces leschenii has been found only on the coxae of the host, and all other Diphymyces species we have discovered in New Zealand are specific to the elytra and mesonotum with the exception of D. curvatus (Thaxt.) I.I. Tav., which also occurs only on the coxae of its host.
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, 1918) later were segregated by Tavares (1985), who accommodated these by erecting a new genus, Diphymyces, characterized by a vertical septum between cells II and VI, which lie side by side above cell I (Tavares 1985, Santamaria 1993, Weir and Rossi 1997). Three new species of Diphymyces recently have been described: D. niger (T. Majewski) I.I. Tav. (as Corethromyces niger) (Majewski 1973), D. penicillifer A. Weir and W. Rossi (Weir and Rossi 1997) and D. urbasoli Santam. (Santamaria 1993). Diphymyces species grow on Leiodidae (Cholevidae or Colonidae) (Santamaria 1993), except for D. penicillifer from New Zealand, which grows on Stenomalium (Staphylinidae: Omaliinae) (Weir and Rossi 1997). Diphymyces species are common on Leiodidae in New Zealand and many undescribed species in our collection await study. The outer basal margin of the perithecium in Diphymyces leschenii is much less convex than that of Diphymyces depressus sp. nov., described below. The receptacle of D. leschenii also is less compact and the species has a deeper color than D. depressus. Diphymyces leschenii is distinguished from D. niger in its less compact habit, lighter color and perithecial apex, the lips of which are much smaller. The perithecial stalk and basal cells of D. leschenii are well distinguished but less slender than those of Diphymyces appendiculatus (Thaxt.) I.I. Tav., and the perithecium in D. leschenii is curved inward rather than outward. The lips of the perithecial apex in D. leschenii are less distinct than those of D. curvatus. In addition, the perithecium of the latter is curved outward rather than inward.
This is the first species of Diphymyces known to bear two perithecia regularly. Seventy-five percent of mature thalli (n = 12) in our collection bore two perithecia. Thaxter (1915, 1918, 1931) did not note the presence of multiple perithecia in any of his specimens now assigned to Diphymyces, and their presence also was unmentioned by Tavares (1985). Re-examination of the holotype material of Diphymyces bidentatus (Thaxt.) I.I. Tav. revealed that several thalli on Thaxter’s slides bear this arrangement. These specimens probably were assumed to be abnormal because many Laboulbeniales develop secondary perithecia when the primary one(s) become aborted or broken. However, we also have observed multiple perithecia in the aforementioned undescribed Diphymyces species and both perithecia observed in thalli of D. leschenii and D. bidentatus appear to be fully developed and functional. The frequency and abundance of multiple perithecia in D. leschenii, D. penicillifer (reported below), D. bidentatus and other undescribed species indicate that this is not an abnormal developmental phenomenon in the genus.
Diphymyces leschenii is remarkably similar in development and general habit to Asaphomyces tubanticus (Middelhoek & Boelens) Scheloske and A. cholevae Thaxt. Both species form multiple perithecia, mainly from the upper cells of the main axis, but sometimes the lowermost perithecial stalk cell arises from cell II. Majewski (1973) notes the presence of an "intermediate form" to A. cholevae and A. tubanticus, which has two perithecia, one each arising from the second and third cells of the main axis. In addition, the lowermost perithecia in A. tubanticus, may not develop fully. The perithecia in D. leschenii arise from cells I and II. Majewski (1994) observed single antheridia (not intercalary) in his specimens of Asaphomyces. Tavares (1985) also noted that the perithecia in Asaphomyces appear to form above a two-celled stalk, the upper cell of which is cell VI. Thaxter (1931), however, clearly illustrated intercalary antheridia in A. cholevae (Pl. 39, FIG. 14) and noted that the genus resembles Euphoriomyces and might be synonymous with it, the main difference being that A. cholevae is not symmetrically developed, its perithecia developed in a "unilateral series" (p. 311). However, Weir (1997) has noted that perithecia in Euphoriomyces are not always produced bilaterally. It is possible that members of Asaphomyces and Diphymyces are related closely, perhaps congeneric. Apart from one exception, the hosts of Diphymyces belong to Leiodidae, as in Asaphomyces and several members of Euphoriomyces, and multiple perithecia are present in at least three described members, D. leschenii, D. bidentatus and D. penicillifer. Our understanding of the generic limits of Diphymyces, Asaphomyces and Euphoriomyces, which are placed in three different tribes (Tavares 1985) would benefit from examination with molecular phylogenetic methods.
2. Cucujomyces phycophilus A. Weir & W. Rossi, Can J Bot 75:793. 1997. FIG. 2B
Specimens examined. – NEW ZEALAND. AUCKLAND ISLANDS: Enderby I, Derry Castle Reef, 50°30'N, 166°19'W, 21 Mar 2000, on Omaliomimus venator (Broun), M. Bullians et al (SYRF AW693); Enderby I, East Bay, 50°30'S, 166°19'E, 21 Mar 2000, on Macralymma punctiventris Cameron, M. Bullians et al (SYRF AW965, PDD 77846–77848); Enderby I, 50°30'S, 166°19'E, 21 Mar 2000, on Nesomalium pacificum (Kiesenwetter), M. Bullians et al (SYRF AW919, PDD 92048). ANTIPODES ISLAND: 6 Feb 1971, on O. venator, Crymus kronei (Kiesenwetter), and M. punctiventris, G. Kuschel (SYRF AW964, PDD 77853–77876); Stella Bay, 19 Feb 1969, on undescribed species of Stenomalium, G. Kuschel (SYRF AW1013). CAMPBELL ISLAND: Beeman Point, 52°32'S, 169°8'E, 7 Mar 2000, on O. venator, M. Bullians et al (SYRF AW966, PDD 77851–77852); Beeman Hill, 52°32'S, 169°8'E, 7–10 Mar 2000, on Nesomalium campbellensis Steel, M. Bullians et al (SYRF AW967); Venus Cove, 52°32'S, 169°8'E, 14 Mar 2000, on O. venator, M. Bullians et al (SYRF AW968, PDD 77849–77850).
Known distribution and hosts. –
This fungus first was reported on Omaliomimus venator and Macralymma brevipenne Cameron (probably M. punctiventris Cameron) and was known only from Oreti Beach, Invercargill, New Zealand (Weir and Rossi 1997). Both the host and geographic range of the fungus have been expanded as outlined above (TABLE I).
Commentary. –
There is some morphological varation in new material as compared with the original species description (Weir and Rossi 1997), including shorter, less inflated perithecial stalk cells, more compact thalli and a greater variation in color. A representative of the shorter form is shown (FIG. 2B). Thalli may be nearly hyaline to violet brown, ruby or purple. Cucujomyces phycophilus commonly is encountered on intertidal Omaliinae (Staphylinidae). This is the first record of C. phycophilus occurring on other genera of Omaliinae in addition to Omaliomimus and Macralymma. Omaliinae are well represented in New Zealand, with 31 species in six genera (Klimaszewski et al 1996). At least 22 species in the four host genera Crymus, Stenomalium, Omaliomimus and Macralymma are known to occur under kelp and other seaweed in New Zealand (Klimazsewski et al 1996).
Laboulbeniales generally are restricted to hosts of the same species or genus. Host specificity in marine Laboulbeniales from New Zealand seems less stringent than that of terrestrial species. For example, Stigmatomyces australis is common on two genera of kelp flies, Icaridion and Australimyza (Hughes et al 2004). It might be that the marine environment affords greater ascospore viability and thus more possibilities for indirect transfer. Others (De Kesel 1996, Benjamin 1965) documented cases in which infection is simply a matter of spatial and temporal opportunity for the spore. Shared habitat between hosts of different genera might provide more possibilities for direct contact between unrelated hosts. Variation in integument characteristics of the host group also might play a role in specificity mechanisms (see commentary under Diphymyces penicillifer).
3. Rhachomyces kenodactyli Balazuc & W. Rossi, Kew Bull 39:754. 1984. FIG. 2C
Specimens examined. – NEW ZEALAND. ANTIPODES ISLANDS: Stella Bay, on K. audouini under kelp, 26–28 Feb 1969, G. Kuschel (SYRF AW1000, PDD 77900–77901). AUCKLAND ISLANDS: Camp Cove, on K. audouini under rocks on soft mudstone in intertidal zone, 14 Feb 1973, J.S. Dugdale (SYRF AW999, PDD 77898–77899). CAMPBELL ISLAND: Venus Cove, 52°32'S, 169°8'E, 14 Mar 2000, on Kenodactylus audouini (Guérin-Méneville), M. Bullians et al (SYRF AW995, PDD 77888); Tucker Cove, 52°32'S, 169°8'E, 16 Mar 2000, on K. audouini, M. Bullians et al (SYRF AW996, PDD 77889–77892); Venus Bay, 52°32'N, 169°8'W, 12 Mar 2000, on K. audouini, M. Bullians et al (SYRF AW997, PDD 77893–77896). SNARES ISLANDS: Gut, on K. audouini in sinkhole under stones in high supralittoral zone, 28 Dec 1976, J.W. Early (SYRF AW998, PDD 77897).
Known distribution and hosts. –
Previously known only from Antipodes Islands on Kenodactylus audouini, the geographic range now is expanded to the Campbell, Snares, and Auckland islands (TABLE I).
Commentary. –
No significant morphological variation from the type material described by Rossi (1984) is apparent.
4. Diphymyces depressus M.B. Hughes, A. Weir et C. Judd, sp. nov. FIG. 2D, E
Diphymyces bidentati Thaxt. habitu similis et probabiliter huic speciei proxima, sed perithecio edentato. Fungus hyalinis et labiis perithecii inaquaelibus parvis a Diphymycetibus ceteris statim distinguendus. Atque receptaculum atque cellula stirpis perithecii atque cellulae basales perithecii omnibus partibus verticaliter depressae. Margo basalis exterior perithecii satis convexa. HOLOTY PUS: PDD 72044.
Habit stout. Receptacle vertically compressed. Thallus hyaline to pale yellowish. Cell I stout, isodiametric. Appendage consisting of three or more seriate cells. Cell II broadly elliptical, cell III about twice as long and isodiametric. Cell IV subpentagonal in optical section, cut off on its upper margins by smaller cells which each form small appendage branches. Cell IV sometimes superimposed by another cell with a similar arrangement. Simple antheridia seriate, intercalary, small, curved upward, borne on variously branched and proliferating appendage branchlets, particularly in younger specimens. Perithecial stalk cell (VI) similar in size and shape to cell I. Perithecial basal cells inconspicuous and flattened, difficult to discern in mature specimens, subequal in size. Perithecium erect, its outer margin quite convex at the base, tapering gradually to the apex. The inner perithecial margin nearly straight. Perithecial tip distinctly asymmetrical, with two larger and two smaller lips.
Etymology. – Named for the vertically depressed nature of cells I, II, VI, and the basal cells of the perithecium.
Measurements. – Basal cell (I): 10–14 x 6–10 µm. Stalk cell of appendage (II): 7–12 x 5–10 µm. Stalk cell of perithecium (VI): 7–15 x 4–10 µm. Perithecium (not including basal cells): 17.5–22.5 µm x 37.5–67.5 µm. Length from foot to tip of perithecium: 40–91 µm. Length of main axis (from base of cell II to tip of appendage): 35–106 µm.
HOLOTY PE: NEW ZEALAND. CAMPBELL ISLAND: Fence line from Tucker Cove, Col Ridge, 52°32'N, 169°8'E, 9 Mar 2000, on Paracatops campbellicus (Brookes), M. Bullians et al (PDD 72044, ISOTY PE K[M]111878).
PARATY PES: NEW ZEALAND. CAMPBELL ISLAND: Beeman Hill, 52°32'S, 169°8'E, 7–10 Mar 2000, on P. campbellicus, M. Bullians et al (SYRF AW974); fence line from Tucker Cove, Col Ridge, 52°32'N, 169°8'E, 9 Mar 2000, on P. campbellicus, M. Bullians et al (SYRF AW975, PDD 77877–77886); Beeman Hill, 52°32'S, 169°8'E, 10–17 Mar 2000, on P. campbellicus, M. Bullians et al (SYRF AW976, PDD 77887).
Known distribution and hosts. –
The only known host is Paracatops campbellicus from Campbell Island (TABLE I).
Commentary. –
Specimens were found growing on the elytra and were removed from the same beetles bearing individuals of Diphymyces leschenii. This species seems most closely allied with Diphymyces bidentatus (Thaxt.) I.I. Tav., sharing the same stout cell I and perithecial stalk. Both Diphymyces depressus and D. bidentatus are restricted to the dorsum of the host, but D. depressus lacks distinct "teeth" at the tip of the perithecium. Diphymyces depressus also differs from D. bidentatus, D. leschenii and D. penicillifer in its quite convex outer basal perithecial margin. The perithecial lips are of unequal height and not as divergent as those of D. penicillifer. The receptacle also is much more compact than that of either D. leschenii or D. penicillifer. The structure and branching of the appendage of D. depressus (FIG. 2E) closely resembles that of D. leschenii (FIG. 2A), yet the thalli bear only one perithecium without the orange and olivaceous coloration so characteristic of the latter species.
5. Diphymyces penicillifer A. Weir and W. Rossi, Can J Bot 75:793. 1997. FIG. 2F
Specimens examined. – NEW ZEALAND. unknown collection locality, on N. pacificum (SYRF AW979, PDD 72049). ANTIPODES ISLANDS: Central Valley, 25 Feb 1969, on Allodrepa decipiens Steel, G. Kuschel (SYRF AW977, PDD 72050–72051). AUCKLAND ISLANDS: Enderby Island, 21 Mar 2000, on unknown host, M. Bullians et al (SYRF AW919, PDD 72048). CAMPBELL ISLAND: Fence line from Tucker Cove, 52°32'N, 169°8'W, 21 Mar 2000, on "Stenomalium" sp., M. Bullians et al (SYRF AW978).
Known distribution and hosts. –
The host range is expanded to include Allodrepa decipiens and Nesomalium pacificum, and the geographic range is expanded to Campbell Island and the Antipodes and Auckland Islands (TABLE I).
Commentary. –
The newly discovered material fits the description of Weir and Rossi (1997), except that one thallus possesses two perithecia, one arising from cell I and the other from cell II. This ontogeny is identical to that observed in D. leschenii, D. bidentatus and other undescribed species of Diphymyces from New Zealand.
Allodrepa decipiens and N. pacificum both are terrestrial species, the former found in tussocks and the latter found commonly in bird nests and other habitats where there is decay (Steel 1964). Stenomalium species are found in similar terrestrial habitats. Reduced host specificity in D. penicillifer is reminiscent of Cucujomyces phycophilus, also inhabiting Omaliinae. Direct contact of hosts in different genera due to shared habitat might play a role in transmission. Integument chemistry is extremely variable (Blomquist and Dillwith 1985) and is considered to play a key part in host-specificity mechanisms among entomogenous fungi (Koidsumi 1957, Leucona et al 1997). It is possible that more uniformity than expected in the integument chemistry of Omaliinae could cause a decrease in host specificity of the fungi. The linkage of Laboulbeniales specificity to integument chemistry currently is speculative and not based on experimental studies with Laboulbeniales, but rather experimental studies of entomogenous Clavicipitaceae. This remains a difficult area of study in the Laboulbeniales due to the inability to culture the fungi or to rear many of the hosts in the laboratory for long enough periods of time.
6. Laboulbenia subantarctica M.B. Hughes, A.Weir et C. Judd, sp. nov. FIG. 3A, C
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Habit erect; some thalli slender and elongate, others short and squat. Variably olive brown, except for the lighter colored receptacle, stalk cell of the perithecium and inner appendage. Cell I usually curved at the base. Cell II broader than cell I and of equal length or up to twice as long; cut off obliquely on its upper margin by cell VI. Cell III twice as long as broad, subrectangular to subtrapezoidal, the inner margin convex, the outer straight. Cell IV subrectangular, the outer margin straight. Cell V distinctly flattened and displaced between cell IV, the insertion cell, and the perithecium, twice as long as broad, oblong to subtriangular, the IV–V septum oblique and curved. Basal cell of outer appendage rectangular, twice as long as broad. Outer appendage simple, variably olive brown, much darker distally; composed of up to 10 seriate thick-walled cells slightly constricted at the septa. The outer appendage occasionally splitting above the third cell, giving rise to a thinner and shorter inner branch. Basal cell of inner appendage shorter than that of the outer appendage, slightly longer than broad to twice as long as broad. Inner appendage consisting of two small superposed cells, the upper of the two bearing paired antheridia. On the other hand, the inner appendage dichotomously branched, each of the four terminal cells bearing 2–4 ventricose antheridia. The inner appendage rarely giving rise to a sterile branch up to five cells long. Stalk cell of perithecium (VI) slightly longer than broad; rhomboidal, subtriangular or elliptical. Basal cells of perithecium irregularly shaped and subequal, slightly smaller than secondary stalk cell of perithecium. Perithecium with a convex outer margin, the upper third free; broadly subovoid, the lower two-thirds olive brown. The perithecium somewhat paler below the deep olive brown preostiolar spots. The apex of the perithecium nearly hyaline, consisting of two rounded asymmetric, outwardly facing lips.
Measurements. – Length of appendage (from insertion cell to tip): 128–504 µm. Length from foot to insertion cell: 140–448 µm. Perithecium: 72–154 x 38–96 µm. Total length: 196–502 µm.
Etymology. – Named for the subantarctic region. HOLOTY PE: NEW ZEALAND: AUCKLAND ISLANDS: Port Ross, on Oopterus plicaticollis Blanchard, Mar 2000, M. Bullians et al (PDD 72045, ISO-TY PES SYRF AW981, PDD 77828–77829).
PARATY PES: NEW ZEALAND. AUCKLAND ISLANDS: Enderby I, Derry Castle Reef, 50°30'N, 166°19'W, 21 Mar 2000, on O. plicaticollis and O. marrineri, M. Bullians et al (SYRF AW983, PDD 72054, 77841–77844, K[M]111884); Port Ross, Erebus and Terror Cove, 50°30'S, 166°19'E, 22 Mar 2000, on O. plicaticollis, M. Bullians et al (SYRF AW984); Ender-by I, Spar Cr Area, 50°30'S, 166°19'E, 21 Mar 2000, on O. plicaticollis, M. Bullians et al (SYRF AW988, K[M]111887); Port Ross, Erebus and Terror Cove, 50°30'S, 166°19'E, 12 Mar 2000, on O. marrineri and O. clivinoides, M. Bullians et al (SYRF AW989); Enderby I, Northeast Cape, 50°30'S, 169°19'E, 21 Mar 2000, on O. clivinoides, M. Bullians et al (SYRF AW990); Enderby I, 50°30'S, 166°19'E, 30 Mar 2000, on O. plicaticollis, M. Bullians et al (SYRF AW991, PDD 77845); Port Ross, Erebus and Terror Cove, 50°30'S, 166°19'W, 22 Mar 2000, on O. plicaticollis, M. Bullians et al (SYRF AW992); Enderby I, Derry Castle Reef, 50°30'N, 166°19'W, 21 Mar 2000, on O. plicaticollis, M. Bullians et al (SYRF AW993, PDD 77830–77833); Enderby I, Stella Hut, 50°30'N, 166°16'W, 21 Mar 2000, on O. plicaticollis, M. Bullians et al (SYRF AW994, PDD 77834–77840); Ewing I, 7 Feb 1973, on O. clivinoides on rotten Olearia lyall Hook. f. and live Colobanthus muscoides Hook. f. mats, C.J. Horning (SYRF AW1001, PDD 77910–77915); Enderby Island, Sandy Bay, 26 Feb 1973, on Oopterus sp. under rocks at stream edge, D.S. Horning (SYRF AW1002, PDD 77909); Adams I, Fairchild’s garden, 20 Jan 1966, on O. plicaticollis in litter, G. Kuschel (SYRF AW1003); Adams I, Fairchild’s garden, 2 Feb 1973, on O. plicaticollis in peat among roots and stems, D.S. Horning (SYRF AW1004, PDD 77907–77908). CAMPBELL ISLAND: Northwest Bay, 100m from shore, 13 Mar 2000, on Oopterus marrineri Broun, E. Edwards, P.J. Johnston (SYRF AW982, PDD 72055–72057); Mount Yuan, Villarceau, 3 Dec 1975, on Oopterus clivinoides Guérin-Méneville under rock-hopper penguins 20 m above high water, B.M. May (SYRF AW1014, PDD 72058–72059); Camp Stream, 52°32'S, 169°8'E, 11 Mar 2000, M. Bullians et al (SYRF AW985); Beeman Point, 52°32'S, 169°8'E, 7 Mar 2000, on O. marrineri, M. Bullians et al (SYRF AW986, PDD 72052); Northwest Bay, 52°32'S, 169°8'E, 13 Mar 2000, on O. marineri, M. Bullians et al (SYRF AW987, PDD 72053).
Known distribution and hosts. –
Oopterus clivinoides and O. plicaticollis in the Auckland Islands and on O. marrineri in Campbell Island (TABLE I).
Commentary. –
Abundant collections of this fungus were made in spring 2000. It also was encountered numerous times on museum specimens of Oopterus clivinoides and O. plicaticollis collected in the 1960s and 1970s in the Auckland Islands. This is a variable fungus in total size, thickness and length of the outer appendage, degree of branching of the inner appendage and number of antheridia per branchlet (FIG. 2A, C). In immature specimens, cell V generally is obtriangular in optical section with an oblique IV–V septum, but as the perithecial venter expands cell V is compressed between cell IV, the insertion cell, and the perithecial venter. Apart from Laboulbenia sp. 1. (below), also on Zolini (Carabidae: Trechinae), few other species share this arrangement.
A large group of Laboulbenia species parasitic on pterostichid Carabidae have simple outer appendages (Santamaria 1998). Many species resemble L. subantarctica: L. oedodactyli Thaxt., L. madeirae Thaxt., and L. erecta Thaxt., L. aubryi Balazuc, L. vulgaris Peyr., and L. benjaminii Balazuc ex Santam. Hosts for these species are scattered within the Carabidae (Platynini, Bembidiinae, Trechinae, Licininae). The host for L. oedodactyli is a staphylinid beetle. Laboulbenia oopteri Thaxt., described from New Zealand, occurs not on Oopterus but on Mecyclothorax rotundicollis White. The usually simple inner appendage, the shape and displacement of cell V, and the shape of the perithecium and its relation to the upper cells of the receptacle serve in combination as diagnostic characters of L. subantarctica.
7. Laboulbenia loxomeri M.B. Hughes et A. Weir, sp. nov. FIGS. 3B, D
Perithecium angustum, minus quam dimidie liberum. Margo perithecii exterior fere recta. Perithecium infra apicem abrupte decrescens. Cellulae labiales perithecii hyalinae, atque supra infuscatam preapicatem zonam parum in-flatae. Cellula suprabasalis (II) latior quam cellula I, atque in latitudine omnino subuniformis. HOLOTY PUS: PDD 72046.
Thalli erect and elongate; dirty yellow. Perithecium and outer appendage olivaceous brown. Cell I usually curved at the base. Cell II wider than cell I throughout and of approximately equal width along its length; about twice as long as cell I and cut off obliquely on its upper margin by cell VI. Cell III about twice as long as broad, with a straight outer and convex inner margin. Cell IV slightly longer than broad, its upper margin cut off obliquely by cell V. Cell V obtriangular in optical section, the upper part somewhat free. Outer appendage generally forming 2–3 branches above the basal cell; basal cell about as long as broad, dirty yellow. Distally, cells of the outer appendage darker, becoming olivaceous brown, slightly longer than broad, thick-walled, constricted at the septa; septa may be blackened in age. Inner appendage variously dichotomously branched, each terminal cell bearing 2–4 antheridia. Perithecial stalk cell stout, about as long as broad, about twice as large as the basal cells. Perithecium slender, the inner margin coincident with that of the receptacle, about one-third free, somewhat paler below the abruptly narrowed tip. Tip deep olive-brown. Perithecial apex hyaline, slightly broadened above the tip; tip consisting of two rounded asymmetric, outwardly facing lips.
Measurements. – Length of appendage (from insertion cell to tip): up to 240 µm (immature specimens). Length from foot to insertion cell: 360–470 µm. Perithecium: 55–84 x 168–240 µm. Total length: 420–580 µm.
Etymology. – Named for the host genus, Loxomerus.
HOLOTY PE: NEW ZEALAND. AUCKLAND ISLANDS: Adams I, Fairchild’s garden, 20 Jan 1966, on Loxomerus sp., G. Kuschel (PDD 72046, ISOTY PE K[M]111883).
PARATY PES: NEW ZEALAND. AUCKLAND ISLANDS: Raoul I, Jan 1963, on Loxomerus nebrioides Guerin-Meneville, L.J. Dumbleton (SYRF AW1005); Adams I, Northeast ridge of Mount Dick, 1200–1800 feet, 17 Jan 1966, on Loxomerus sp., G. Kuschel (SYRF AW1006, PDD 77906); Ewing I, 15 Sep 1943, on L. nebrioides, R.A. Falla (SYRF AW1007, K[M]111880); Ranui Cove, 22 Feb. 1973, on L. nebrioides, on peat under old boards, C. Horning (SYRF AW 1008, PDD 77903–77905).
Known distribution and hosts. –
Known only from Loxomerus nebrioides and Loxomerus sp. in the Auckland Islands (TABLE I).
Commentary. – This species is somewhat similar in inner appendage characteristics to Laboulbenia subantarctica and in outer appendage characteristics and general habit to L. pterostichi Thaxt. The narrow perithecium and the distinct shape of the tip, the size and shape of cell II, and the branching pattern of the outer and inner appendage all are characters that distinguish it from other species. Loxomerus species are of conservation concern in New Zealand because they are preyed upon by mice, an introduced pest.
8. Teratomyces sp. 1 FIG. 4A, D
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Measurements. – Perithecium: 28 x 132–145 µm (52 x 136 µm for short-stalked specimen) Stalk cell of perithecium (VI): 90–160 µm (65 µm long for short-stalked specimen) Appendage length: 75–85 µm. Receptacle (first three cells only): 28 x 75 µm (40 x 80 µm in short-stalked specimen) Total length: 320–380 µm (396 µm in short-stalked specimen).
Specimens examined. – NEW ZEALAND: AUCKLAND ISLANDS: Port Ross, Erebus Cove, 50°32'S, 166°12'E, on Quedius secretus Cameron in leaf litter, 20 Mar 2000, M. Bullians et al (SYRF AW1011); Enderby I., Stella Hut, 50°30'S, 166°16'E, on Q. secretus in leaf litter, 21 Mar 2000, M. Bullians et al (SYRF AW1012).
Known distribution and hosts. –
Known only from Auckland Islands on Quedius secretus (TABLE I).
Commentary. –
Specific delineation in Teratomyces remains difficult. Species are distinguished by such characters as receptacle structure, number of perithecia and length of the appendages. Unlike the vast majority of Laboulbeniales, these characters often are variable in fungi removed from one host individual. Defining a species in this genus is complicated further by the fact that the different morphological forms do not always cluster in discrete positions on the host, as do many other species of Laboulbeniales. Weir and Rossi (2001) note that further collections are required to clarify relationships within the genus.
Thaxter (1915) illustrated what he regarded to be two forms of Teratomyces insignis Thaxt., each with a much different receptacle, one wholly melanized and the other with all three cells translucent. He considered them the same species due to differences in age of the specimens or in position of growth. Both forms, however, were removed from the abdomen. In other cases Thaxter considered similar morphological differences sufficient to distinguish between species, even when those fungi were removed from the same beetles. Teratomyces actobii Thaxt. and T. brevicaulis Thaxt. both were removed by Thaxter from the same host species, Actobius nanus Horn, and they are distinguished only by the size of the perithecial stalk-cell and the clavate shape of the larger appendage cells in T. brevicaulis. Both occur on the legs and abdomen.
Several forms of the fungus described above were removed from the host. One form (FIG. 4A) removed from the legs differs most notably in the size, shape and color of receptacular cells, all of them being more elongate than in the form described above. The perithecia in this type are also more ovoid, with a short, inconspicuous stalk cell. However, another form from the legs had a very narrow, long-stalked perithecium but shared the same receptacular structure. Two other mature individuals were removed from the abdomen, which had long-stalked, narrow perithecia and compact receptacles (FIG. 4D). These variations are reminiscent of Thaxter’s observations (1931) of different types of thalli removed from the same hosts and indicate the confused systematic state of Teratomyces. Rather than contribute to the confusion, we prefer to reveal the presence of the genus in the subantarctic without describing this (or these) species formally. Molecular investigation of genetic differences between morphotypes in the genus is needed.
9. Rhachomyces sp. 1 FIG. 4B
Thallus slender, erect, dirty yellow to brownish yellow, apart from the opaque appendages. Basal cell (I) of primary receptacle subrectangular, and longer than broad. Secondary receptacle composed of 11–12 superposed cells, which are slightly broader than long and successively larger. Appendages opaque, long, stiff, nearly straight, curved slightly upward, becoming slightly broader distally, each composed of up to nine cells, (perhaps more); appendage cells about twice as long as broad, with blackened septa. Perithecium ellipsoid, free from appendages, the outer margin more convex than the inner margin, the four tiers of wall cells unequal in size.
Measurements. – Perithecium: 36–45 x 93–110 µm. Appendages: 10–18 x up to 146 µm. Receptacle: 117–130 µm. Length from foot to tip of perithecium: 210–240 µm.
Specimens examined. – NEW ZEALAND. AUCKLAND ISLANDS: Port Ross, Erebus and Terror Cove, 50°30'S, 166°19'S, on O. plicaticollis in leaf litter and under rotting wood, 22 Mar 2000, M. Bullians et al (SYRF AW923).
Commentary. – This species was recovered from one host individual. If the specimens examined represent an undescribed species, its closest morphological allies are Rhachomyces speluncalis Thaxt. from West Virginia, USA, and Rhachomyces anophthalmi Thaxt. and Rhachomyces capucinus Thaxt., both from Europe. All three species are found on Trechinae. Rhachomyces sp. 1 differs from the others in its short stalk cell and basal cells of the perithecium, as well as the perithecial shape.
10. Laboulbenia sp. 1 FIG. 4C
Thallus robust and erect; brownish yellow, except black perithecium and base of appendages. Cell I small, about twice as long as broad, wider at its apex. Cell II much wider than cell I, thick-walled, about twice as long as broad, cut off obliquely on its upper margin by cell VI. Cell III isodiametric, slightly longer than broad, the inner margin slightly convex. Cell IV subrectangular, slightly larger than cell III, its inner margin juxtaposed with the lower, outer curved margin of cell V. Cell V distinctly displaced, appearing enclosed by cell IV, the perithecium, and the insertion cell, its upper margin black and indistinguishable from the blackened insertion cell and the base of the appendages. Cell VI ellipsoid and equal in size to the other basal cells. Outer appendage broken distally, olivaceous with blackened and slightly constricted septa, the cells slightly longer than broad and the basal cell stout and obscured by melanization. Inner appendage consisting of two branches, the outer of about equal width to the outer appendage and broken distally; dirty yellow, the septa unconstricted and not blackened, the inner branch much smaller, olivaceous brown with a darkened septum, broken off close to the base. Perithecium ovoid, nearly black excepting the basal cells, the tip slightly constricted below the apex, which consists of two outwardly facing lips.
Measurements. – Length of appendage (from insertion cell to tip): 100–160 µm. Length from foot to insertion cell: 280–300 µm. Perithecium: 112 x 180–192 µm. Total length: 375 µm.
Specimens examined. – NEW ZEALAND. SNARES ISLANDS: Penguin Creek, 15 Dec 1974, on Synteratus ovalis Broun under dead branches of O. lyalli on peat, D.S. Horning (SYRF AW1010).
Known distribution and hosts. – Known only from Synteratus ovalis, which occurs in New Zealand only on Snares Islands.
Commentary. – This species shares the peculiar displacement of cell V found in L. subantarctica. Both species grow on Zolini (Carabidae: Trechinae). This species is undoubtedly new, but because of the broken appendages it is not formally described. Fresh collections in good condition are needed.
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DISCUSSION |
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). The uncommon, and sometimes very rare, occurrence of Laboulbeniales species makes it likely that the species accumulation curve is only an indication of a lower bound to the total number of species in the subantarctic; there are at least a few more species left to find on Coleoptera alone. The most likely host groups are Ptiliidae, Pselaphinae (Staphylinidae), Aleocharinae (Staphylinidae), Byrrhidae, Corylophidae, Tenebrionidae and Diglymma (Carabidae); these all are terrestrial taxa that have been found serving as hosts for Laboulbeniales in mainland New Zealand. They also are commonly infected elsewhere in the world.
Based upon the list of taxa provided by Johnston et al (2002) (www.landcareresearch.co.nz/research/biodiversity/antarctica/html/beetlebase.html), the percentage of beetle species infected with Laboulbeniales (12%) is higher than that found at tropical (6–7%) and north temperate (9.5%) sites, as reported by Weir and Hammond (1997). All areas investigated are relatively well-inventoried, so the figure is probably significant but not necessarily a function of latitude. In the subantarctic investigation, the majority of taxa infected with Laboubeniales are intertidal, and the intertidal beetles have more frequent and heavier infections than terrestrial beetles. Frequency and abundance of infection in kelp flies is also high (Hughes et al 2004, McAlpine 1991). The higher percentage of infected taxa in the subantarctic probably is due to the propensity for Laboulbeniales infection in moist habitats (Weir and Hammond 1997, Huldén 1983) and shared host habitat of diverse hosts, facilitating greater opportunities for direct or indirect transfer of spores. The broad host range of Cucujomyces phycophilus and Diphymyces penicillifer also contribute to the higher percentage of host utilization; at least one host species, however, is shared between these two fungi (TABLE I).
The subantarctic Laboulbeniales are a mixture of putative endemics (Rhachomyces kenodactyli, Laboulbenia subantarcticus, Laboulbenia loxomeri, Laboulbenia sp. 1, Diphymyces leschenii, and Diphymyces depressus), and other species found more widely in mainland New Zealand (Cucujomyces phycophilus and Diphymyces penicillifer). It is possible that Rhachomyces kenodactyli is of Gondwanan origin. This would be confirmed by the presence of the species on Kenodactylus audouini in other areas such as the Falkland Islands or Patagonia. Rhachomyces kenodactyli has not been found in mainland New Zealand.
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KEY TO SPECIES OF SUBANTARCTIC LABOULBENIALES |
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ACKNOWLEDGMENTS |
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FOOTNOTES |
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1 Corresponding author. E-mail: mohughes{at}mailbox.syr.edu
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———. 1971. Introduction and supplement to Roland Thaxter’s contribution towards a monograph of the Laboulbeniaceae. Bibl Mycol 30:1–155.
———. 1986. Laboulbeniales on semiaquatic Hemiptera. V. Triceromyces: with a description of monoeciousdioecious dimorphism in the genus. Aliso 11:245–278.
Blomquist GJ, Dillwith JW. 1985. Cuticular lipids. In: Kerkut GA, Gilbert LI, eds. Comprehensive Insect Physiology, Biochemistry, and Pharmacology. Vol. 3: Integument, Respiration, and Circulation. New York: Pergamon Press. p 117–154.
De Kesel A. 1996. Host specificity and habitat preference of Laboulbenia slackensis. Mycologia 88:565–573.
Gotelli NJ, Colwell RK. 2001. Quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species richness. Ecol Lett 4:379–391.
Hughes M, Weir A, Gillen B, Rossi W. 2004. Stigmatomyces from New Zealand and New Caledonia: new records, new species, and two new host families. Mycologia 96: 834–844.
Huldén L. 1983. Laboulbeniales (Ascomycetes) of Finland and adjacent parts of the U.S.S.R. Karstenia 23:31–136.
Klimaszewski J, Newton AF, Thayer MK. 1996. A review of the New Zealand rove beetles (Coleoptera: Staphylinidae). N Z J Zool 23:143–160.
———, Watt JC. 1997. Fauna of New Zealand (Ko te Aitanga Pepeke o Aotearoa). Coleoptera: family-group review and keys to identification. Lincoln, New Zealand: Manaaki Whenua Press. 199 p.
Koidsumi K. 1957. Antifungal action of cuticular lipids in insects. J Ins Physiol 1:40–51.
Leschen RAB, Lawrence JF, Kuschel G, Thorpe S, Wang Q. 2002. Insecta: Coleoptera Genera of New Zealand. Gordon DP, ed. The New Zealand Inventory of Biodiversity: A Species 2000 Symposium Review. Christchurch, New Zealand: Canterbury Press (In press).
Leucona R, Clement J-L, Riba G, Joulie C, Juarez P. 1997. Spore germination and hyphal growth of Beauveria sp. on insect lipids. J Econ Ento 90:119–123.
Majewski T. 1973. Rare and new Laboulbeniales from Poland. III. Acta Mycol 9:111–124.
———. 1994. The Laboulbeniales of Poland. Polish Botanical Studies 7:466.
McAlpine DK. 1991. Review of the Australian kelp flies (Diptera: Coelopidae). Syst Entomol 16:29–84.
O’Connor T, ed. 1999. New Zealand’s Subantarctic Islands. Auckland, New Zealand: Reed Books. 104 p.
Rossi W. 1984. New Laboulbeniales from Australia and New Zealand. Kew Bull 39:753–757.
Santamaria S. 1993. A new species of Diphymyces from Spain. Mycol Res 97:791–794.
———. 1998. Flora Mycologica Iberica Vol. 4: Laboulbeniales, I. Laboulbenia. Real Jardín Botanico Madrid. 185 p.
Steel WO. 1964. Insects of Campbell Island. Coleoptera: Staphylinidae. Pacific Insects Monographs 7:340–375.
Tavares II. 1985. Laboulbeniales (Fungi, Ascomycetes). Mycol Mem 9:1–627 p.
Thaxter R. 1915. New Indo-Malayan Laboulbeniales. Proc Amer Acad Arts Sci 51:1–51.
———. 1918. New Laboulbeniales from Chile and New Zealand. Proc Amer Acad Arts Sci 54:205–232.
———. 1931. Contribution toward a monograph of the Laboulbeniaceae. Part V. Mem Amer Acad Arts Sci 16:1–435.
Weir A. 1997. Comparative biological and taxonomic studies of tropical and temperate Laboulbeniales (Fungi; Ascomycota) [Doctoral thesis]. United Kingdom: University of Newcastle upon Tyne. p 1–106, 1–429.
———, Hammond P. 1997. Laboulbeniales on beetles: host utilization patterns and species richess of the parasites. Biodiv and Cons 6:701–719.
———, Rossi W. 1997. New and interesting Laboulbeniales (Ascomycetes) from New Zealand. Can J Bot 75:79–798.
———, ———. 2001. New and interesting Bolivian Laboulbeniales. Mycologia 93:171–180.
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