currently , 106 species are accepted in this genus ( denchev & denchev 2011a , 2011b , 2012 , vnky & abbasi 2011 , vnky 2012 , savchenko et al . 2013 ) , but this is certainly not a final number .
the magnitude of host plants reported in different publications for some putative species complexes suggests that more species are likely to exist , some of which may be well - delimited morphological species , while others are probably cryptic species that could be uncovered by molecular methods .
some species of anthracoidea were recently included in molecular systematic studies ( hendrichs et al .
distinct species could still be hidden under different generic names , especially under historical names that have not been reassessed in recent years ( pitek 2012 ) .
such historical names should be critically re - examined in addition to any comprehensive molecular studies directed to the description of novel anthracoidea species .
kenaica is such a neglected taxon name and a likely candidate to be a distinct member of anthracoidea .
micropoda collected in the kenai peninsula of alaska . that sedge is now accepted as a distinct species , carex micropoda , belonging to carex sect .
callistachys ) ( murray 2002a ) . savile ( 1952 ) provided the following description of cintractia carpophila var .
 zambettakis ( 1978 ) included it in anthracoidea , as  anthracoidea heterospora kukkonen var .
kenaica ( saville ) nov . comb .  , but without any indication of the basionym or a reference to the place of its valid publication , rendering the combination invalid ( art .
likewise , kukkonen ( 1963 ) and piepenbring ( 2000 ) considered this fungus to be a member of anthracoidea , but again without further treatment and any formal nomenclatural and taxonomic decisions .
vnky ( 2012 ) included this smut in two places in his monograph : first as a synonym of anthracoidea heterospora and later under excluded or invalidly published taxa , in both cases without detailed observations .
the aim of the present work is to clarify the nomenclatural and taxonomic status of cintractia carpophila var .
kenaica , and to provide a detailed characterization of this smut fungus as it lacks a detailed description and any iconography .
sori and spore characteristics were studied using dried herbarium material deposited in daom , s , and wrsl .
specimens were examined either by light microscopy ( lm ) and scanning electron microscopy ( sem ) or only by light microscopy ( lm ) . for light microscopy ( lm ) , hand - cut sections of sori or small pieces of sori were mounted in lactic acid , heated to boiling point and cooled , then examined under a nikon eclipse 80i light microscope .
spore size range , mean spore size , and standard deviation of 50 measured spores of each investigated specimen were calculated ( table 1 ) .
the spores were measured in plane view and measurements were adjusted to the nearest 0.5 m .
spore size ranges were assigned to one of the three groups distinguished by savile ( 1952 ) : ( 1 ) small - sized spores , 1321(23 )  917(20 ) m ; ( 2 ) medium - sized spores , 1525(27 )  1021 m ; and ( 3 ) large - sized spores , 1833  1328 m .
for scanning electron microscopy ( sem ) , spores taken directly from dried specimens were dusted onto carbon tabs and fixed to an aluminium stub with double - sided transparent tape .
the stubs were sputter - coated with carbon using a cressington sputter - coater and viewed under a hitachi s-4700 scanning electron microscope , with a working distance of ca .
sem micrographs were taken in the laboratory of field emission scanning electron microscopy and microanalysis at the institute of geological sciences of jagiellonian university ( krakw ) .
detailed morphological characteristics of the holotype , isotype , and two non - type specimens of cintractia carpophila var .
the internal soral structure in the holotype was typical of species of anthracoidea in that the spores were produced directly on the outer surface of the achene , and not within u - shaped cavities embedded in sterile stroma , a characteristic of cintractia ( kukkonen 1963 , kukkonen & vaissalo 1964 , piepenbring 2000 ) .
this indicated this smut fungus was better placed in anthracoidea , as was suggested in other studies ( kukkonen 1963 , zambettakis 1978 , piepenbring 2000 ) .
the spores were uniform in shape and size ranges between collections ( table 1 ) .
kenaica matched well the short description given by savile ( 1952 ) , although the spore surface was not smooth as stated in the protologue , but smooth or very finely punctate in lm , and very finely verruculose in sem .
the very fine ornamentation of spores was probably outside the limits of resolution of savile s light microscope .
in general , the present examination confirms the decision of savile ( 1952 ) to consider this smut as distinct .
this is in line with the conclusion of kukkonen ( 1963 ) , who , however , did not formally make the transfer .
type : usa : alaska : kenai peninsula , head of palmer creek valley , 6049n , 14933w , on carex micropoda ( syn .
description : sori in all or single ovaries of the inflorescence , black , globose or ovoid , about 11.5 mm diam , at first covered by a silvery membrane and perigynium that later ruptures revealing agglutinated spores , powdery on the surface , the sori are partly hidden by the perigynium and scales .
sori develop around reduced achenes that are consecutively surrounded by a thin dark layer of the remnants of achene epidermis , a hyaline layer of sporogeneous hyphae with young spores , a layer of gradually maturing dark spores , and a thin membrane of host origin
. spores usually more or less flattened , chestnut brown , reddish brown to dark brown , quite regular in shape and size , globose , subglobose or broadly ellipsoidal , small , ( 14.0)15.020.5(22.0 )  ( 11.5)12.018.5(20.5 )
sd , 18.5  1.5  15.6  1.7 m , n = 200/4 ] , rarely enclosed by a thin , hyaline , mucilaginous sheath ; wall even , 1.01.5 m thick , somewhat darker than the rest of the spore , without protuberances and light - refractive spots , but with 25 distinct internal swellings ; surface smooth or very finely punctate in lm , spore profile smooth , surface very finely verruculose in sem .
( nannfeldt & lindeberg 1965 ) and is therefore illegitimate and to be rejected ( art .
also , liro s treatment can not constitute a valid description of a new species to be attributed to him alone due to the absence of a latin diagnosis ( nannfeldt & lindeberg 1965 ) , required in the period 1 january 1935 to 31 december 2011 ( art .
vnky ( 2012 ) considered uredo carpophila as an illegitimate name , which is correct , but also as a nomen nudum , which is not correct , since schumacher ( 1903 ) provided a short description of this species : 
kenaica is legitimate since an infraspecific name may be legitimate even if its final epithet was originally placed under an illegitimate species name .
verrucosa savile ( savile 1952 ) , was accepted as a legitimate varietal name that was elevated to the species rank as anthracoidea verrucosa ( savile ) nannf .
the type host of a. 
 kenaica is carex micropoda , but in addition savile ( 1952 ) assigned a single smut collection on carex deweyana ( in carex sect .
zambettakis ( 1978 ) included two sem pictures of spores from a specimen on c. deweyana and they are distinctly verrucose , unlike spores of specimens on c. micropoda .
anthracoidea kenaica was previously reported from the type locality on the kenai peninsula and two other collections from the same region ( savile 1952 ) . here
the smut is newly reported from pribilof island ( ak ) and from mt fougner in british columbia ; this last collection represents the first record of this species in canada .
kenaica can not be treated as a synonym or variety of anthracoidea heterospora since this species is different in having spores with better developed ornamentation , thicker walls ( 12.5 m ) , and occurs on host plants belonging to carex sect .
in contrast , the morphology of anthracoidea kenaica is characteristic of members of anthracoidea sect .
leiosporae , which includes species with smooth or very finely verruculose spores ( kukkonen 1963 ) . within this section ,
anthracoidea kenaica may be comparable only to five anthracoidea species having spores similar in size and with a smooth or very finely verruculose ( not papillate ) surface : a. elynae , a. externa , a. macranthae , a. nardinae , and a. scirpi .
the main morphological differences between these species include differences in wall thickness , the presence and the number of internal swellings , and the presence of a hyaline mucilaginous sheath enclosing the spores .
furthermore , all of them are restricted to host species belonging to different sections of carex , or to different genera ( kobresia , trichophorum ) , which could be used as supportive taxonomic characters .
characters used to discriminate these five species of anthracoidea from a. kenaica ( table 2 ) are contrasted and discussed below .
anthracoidea elynae is distinguished from a. kenaica by the mostly smooth spores with a thicker wall , fewer internal swellings , a more or less evident mucilaginous sheath , and occurrence on kobresia .
the internal swellings in a. elynae are weakly visible in lm ( savile 1952 , kukkonen 1961 , vnky 1994 , 2012 ) , and recent tem studies of spores from a romanian collection did not report internal swellings ( parvu et al .
in fact , the spore presented in figure 3 of the latter study has a shallow thickening on the lower flattened side , which may be interpreted as a weak internal swelling .
by contrast , internal swellings of anthracoidea kenaica are prominent and clearly visible in lm .
anthracoidea externa is morphologically distinct in having absolutely smooth spores surrounded by a thick mucilaginous sheath , a thicker spore wall without internal swellings , and occurrence on species of carex sect .
anthracoidea macranthae differs from a. kenaica as it has completely smooth spores with prominent and common hyaline caps ( a mucilaginous sheath ) on the flattened sides , an absence of internal swellings , a somewhat thinner spore wall , and in occurring on kobresia ( guo & wang 2005 ) .
although not discussed in the protologue , the occurrence of a mucilaginous sheath in the form of hyaline caps is the most valuable diagnostic character of anthracoidea macranthae .
it is distinguished by nearly smooth spores , a thicker spore wall , fewer internal swellings , and occurrence on carex sect .
the type host of a. nardinae is c. nardina , but kukkonen ( 1963 ) assigned this smut to two collections on c. elynoides , which belongs to a different carex section ( sect .
the examination of one collection of anthracoidea on c. elynoides [  plants of southern colorado , carex elynoides holm n. sp . , near pagosa peak , aug .
extracted from the isotype of c. elynoides in wrsl ( phanerogamic herbarium ) ] , had globose , subglobose to broadly ellipsoidal spores , ( 14.5)15.019.5  ( 11.0 ) 12.017.5 m , av .
sd , 17.2  1.2  14.8  1.7 m , n = 50 , wall even , 1.01.5 m , without internal swellings , surface smooth without mucilaginous sheath ] , though different from those studied by kukkonen , revealed a complete absence of internal swellings typical of a. nardinae .
it could be yet another distinct species or a form of a. externa without a hyaline sheath .
anthracoidea scirpi is distinguished from a. kenaica by the minutely punctate spores , which are usually surrounded by hyaline , mucilaginous sheaths on the flattened sides , a thicker spore wall , the absence of internal swellings , and occcurrence on trichophorum species ( vnky 1994 , 2012 ) .
differences between smut specimens in the ovaries of carex micropoda and all aforementioned anthracoidea species from sect .
leiosporae support a. kenaica as a distinct species specialised to a host in carex sect .
host plants of these anthracoidea species are placed in one of the four / five major clades of the tribe cariceae , the so called  core unispicate clade  , which includes carex subgen .
leiosporae is uncertain , and the problem remains open for future studies using molecular methods . in a recent molecular phylogenetic study , by hendrichs et al .
( 2005 ) , the only accessioned smooth - spored species , a. elynae , was recovered as sister to the verrucose - spored species a. curvulae on carex curvula , which is also a member of the  core unispicate clade  .
 cintractia carpophila var . 
 kenaica , a neglected taxon described from alaska more than half a century ago , is re - described and illustrated . its nomenclature and taxonomic status are discussed . 
 this smut species is characterised by small spores with a very finely verruculose surface rarely enclosed by a thin , hyaline , mucilaginous sheath , a wall with 25 distinct internal swellings , and parasitism on carex micropoda ( carex sect . 
 dornera ) . 
 it is reallocated to the genus anthracoidea as a distinct species , anthracoidea kenaica comb . 
 nov . , and assigned to anthracoidea section leiosporae which includes species having smooth or very finely verruculose spores . 
 morphological and biological characteristics of the five most similar anthracoidea species are contrasted and discussed . 