a taxonomic revision of the genus dioicomyces (laboulbeniales)

615

Sergio SANTAMARIA

Unitat de Bota[ nica. Departament de Biologia Animal, de Biologia Vegetal i d ’Ecologia. Facultat de Cie[ ncies. Universitat

Auto[ noma de Barcelona. 08193-Bellaterra (Barcelona), Spain.

E-mail : sergi.santamaria!uab.es

Received 21 September 2001; accepted 16 March 2002.

The genus Dioicomyces is reviewed on the basis of the examination of types, authentic material, or personal collections.

Three new species from Spanish anthicids are described: D. denticulatus, D. ladoi, and D. leptalei spp. nov. Eleven taxa

(including eight species, two forms, and one variety) are synonymized and included under D. anthici, which is

considered a very variable species. With additions and changes, Dioicomyces currently includes 23 species. In this

synopsis no taxa at infraspecific rank are recognised. For each accepted species synonyms are listed, mostly based on

the study of type material. Types for each taxon are included and lectotypes designated when necessary. This

monograph also includes descriptions, photographs and}or drawings for all species studied and a key for their

determination.

INTRODUCTION

The genus Dioicomyces was described by Thaxter

(1901) with three species : D. anthici (generitypus), D.

onchophorus, and D. spiniger (as D. spinigerus) from

thalli collected on specimens of Anthicus floralis

captured in the vicinity of Cambridge, MA, USA.

Subsequently, many other reports and the addition of

new taxa increased to 31 the number of taxa (including

one variety and one form) in the genus (Tavares 1985).

Only two species have been described that weren’t

included in Tavares’s monograph: D. africanus and D.

borneensis. Most of the known species of Dioicomyces

parasitize beetles of the family Anthicidae (Coleoptera),

with only one species on Tenebrionidae and two on

Staphylinidae. Species of Dioicomyces are dioecious. As

characterized by Thaxter (1901), Dioicomyces includes

Laboulbeniales having diminutive males consisting of

three superposed cells and a single terminal anther-

idium, which accompany the female perithecium-

bearing individuals.

Amorphomyces,Dioicomyces (Thaxter1901),Tetrand-

romyces (Thaxter 1912), Dicrandromyces, Nanomyces,

Rhizopodomyces, and Triandromyces (Thaxter 1931)

are closely related genera grouped all in subtribe

Amorphomycetinae (Tavares 1985).

The purposes of this study are : (1) to present an

illustrated taxonomic revision of Dioicomyces with a

key to the species ; and (2) to describe three new species.

Table 1 summarizes diagnostic information about the

23 species of this genus and may be useful for identifying

these species.

MATERIALS AND METHODS

Studies were made on specimens received from various

herbaria and from my own collections. The herbarium

material examined included type specimens from FH

(Thaxter’s collection), LPS (Spegazzini’s collection),

KRAM-F (Majewski’s collection), RO (Rossi’s col-

lection), Shizuoka University Herbarium (Sugiyama’s

collection), and specimens from BCB (author’s col-

lection). For each name the type is indicated as

mentioned in the original protologue. When any type

has been studied, ‘ ! ’ is added after the herbarium code.

When more than one locality was published in the

original description and no explicit indication of type

was included, a lectotype is designated from the slides

studied as a rule, choosing it, if available, from red-

labelled slides having the term ‘ type ’ written on the

label. In the paragraphs including specimens examined,

the original transcription of slide label is provided,

adding any necessary changes or corrections for better

understanding if appropriate.

Microscopic observations were made with the help of

differential interference contrast optics (DIC). In order

to follow a logical pattern some morphological terms

used in descriptions have been simplified and unified. In

descriptions, the black foot is not considered in the size

Mycol. Res. 106 (5) : 615–638 (May 2002). # The British Mycological Society

DOI: 10.1017}S0953756202005816 Printed in the United Kingdom.

A taxonomic revision of the genus Dioicomyces

(Laboulbeniales)

The

gen

us

Dio

icom

yces

616

Table 1. Diocomyces spp. : ecological, distributional, measurement, and morphological characteristics.

Species Hosts"

Known

distribution

Measurements (µm) Morphology

l m Outgrowths or evident swells on perithecium

Number of

wall cells$

Total

length

Total

length

Foot}appendage Perithecium Cell VI }® Position# Shape#

Origin

(wall cell)

africanus Formicomus (A) Sierra Leone 78–83 217–238 70–78 155–175¬45–53 30–38 ® 4, 5, 4, 4

anthici (A) Worldwide 36–68 140–365 39–69 78–210¬35–83 40–186 ® 5, 5, 4(–5), 4

borneensis Formicomus (A) Borneo 63–75 233–278 73–78 175–200¬50–53 48–65 apical finger- or tooth-like w%n

4, 4, 5, 4

denticulatus Cyclodinus (A) Spain 44–46 148–157 39–41 101–108¬40–45 25–33 apical tooth-like w%n

« 5, 5?, 4?, 4

floridanus Bledius (S) USA 68 193–229 54–62 146–168¬45–61 23–34 ® 5?, 5, 4(–5), 4

glossophorus Anthicus (A) Argentina 172–175 50–58 112–118¬35–42 30–33 subapical finger-like ? ?

inclinatus Anthicus (A) Philippines 186–193 43–45 143–146¬48–50 66–68 ® ?

indentatus Anthicus (A) Philippines 48–52 193–207 45–52 130–152¬36–43 57–66 ® 5, 5, 5, 4

italicus Formicomus (A) Philippines,

Japan, Spain,

Italy

68–79 242–371 75–83 158–250¬43–59 37–84 ® 4, 4, 4(–5), 4

ladoi Cyclodinus (A) Spain 54–58 181–190 48–53 119–135¬54–61 34–40 ® 5, 5, 4, 4

leptalei Leptaleus (A) Spain 43–47 131–154 42–45 88–127¬27–47 27–34 lateral spine-like w%m

5, 5, 4, 4

malleolaris Anthicus (A) Argentina 41–45 143–155 39–46 91–109¬50–59 80–91 ® 5, 4, 4, 4

myrmecophilus Myrmechixenus (T) Poland 54 146–160 47–54 85–110¬40–49 16–23 (2) apical &

subapical

(2) finger- &

tooth-like

w&n

w&m

5, 4, 5, 4

notoxi Notoxus (A) Guatemala 43–45 193–414 43–52 107–172¬32–52 71–228 ® 5, 4, 4, 4

obliqueseptatus (S) ‘Amazon ’ 200¬54–57 ® ?

onchophorus Anthicus (A) USA 48–57 154–200 43–54 128–143¬41–54 57–91 apical finger-like w&m

5, 4, 5, 4

proeminens Anthicus (A) Guatemala 45 150–196 46–50 59–77¬29–39 71–121 ® 5, 5, 4(–5), 4

rostellatus Anthicus (A) Argentina 183 55 130¬60 30 apical finger-like ? ?

spiniger Anthicus (A) USA, Spain 39–52 178–212 32–48 121–148¬37–63 36–43 lateral spine-like w%m

4–5(–6), 5, 4, 4

subtorulosus Anthicus (A) Argentina 228–229 52–53 128–134¬39–45 89 ® ?

torulosus Anthicus (A) Argentina 186¬43 143 ® ?

trinitatis Anthicus (A) Trinidad 41–52 342–550 52–59 143–200¬40–70 229–329 ® 4, 5, 4, 4,

umbonatus (A) Argentina,

Spain

57–62 230–254 69–72 117–146¬67–91 67–83 (2) lateral (2) swells n«w

"n«

5, 4, 5, 4

" (A) Anthicidae, (S) Staphylinidae, (T) Tenebrionidae.

# Number in parentheses when more than one outgrowth.

$ Arranged as : wm, w

n, w

n, and w

n«.

S. Santamaria 617

of the basal cell of the receptacle. A thallus is defined as

straight, arcuate or sigmoidal always in relation to the

main longitudinal axis, extending from the foot to the

top of the thallus. The efferent tube of the antheridium

has been defined as terminal (on the top of the

antheridial venter), subterminal (slightly below the top

of the venter) or lateral (located on one side). I have

used only the terms ovoidal and fusiform to define the

perithecial shape, avoiding similar and thus confusing

terms. For each description I have included (if observed,

thus depending on the quality of slide preservation) the

number of perithecial wall cells for each vertical row

with abbreviations such as wm, w

nor w

n«, representing

those rows formed, respectively, from perithecial basal

cells m, n, and n«, following the terminology used by

Tavares (1985). In several examples this number must

be considered as uncertain because of the difficulty in

seeing adequately the basal cells and thus, its cor-

responding wall cells. Other terms and abbreviations

are essentially those of Tavares (1985). Descriptions of

thalli are based on observations of specimens as seen

from one side to the other, in lateral view, because the

position of the fungi on the slide. In referring to some

elements of female thalli, anterior or ventral are in the

direction away from the appendage whereas posterior

or dorsal is in the direction toward the appendage.

Height precedes width in the measurements given.

When thalli are mounted on slides, the cover glass often

compresses perithecia and their outer wall cells may

separate, showing vertical gaps between vertical rows

(e.g. Figs 88, 95, 96). Of course, it should be considered

that mounting, including the type of medium used as

well as the mentioned pressure of cover glass, alterates

in a ³ degree some thallic features as cellular contents,

cellular orientations, etc. Efforts were made to allow for

such distortions in preparing descriptions and drawings.

TAXONOMY

Dioicomyces Thaxt., Proc. Amer. Acad. Arts 37 : 33(1901).

Dioecious. Male thallus consisting of four superposed

cells including a single, terminal, simple antheridium.

Female thallus consisting of a three-celled receptacle ;

the basal cell (I) obtriangular, longer than broad,

separated by an horizontal septum from the³ flattened,

suprabasal cell (II), which is triangular to trapezoidal in

section; the terminal cell (III) typically wedge-shaped,

triangular in section and inconspicuous, separated from

the cell II by an oblique septum. Primary appendage

free, subtended by cell III, consisting of a single cell.

Perithecium free, with well defined basal (m, n, n«) and

stalk cells (VI, VII) ; four vertical rows of outer wall

cells of 4–5(–6) cells each. Trichogyne downwardly

curved, consisting of two cells separated by an

horizontal septum, the proximal cell³basally con-

stricted, the distal cell forming terminal slender branch-

lets or papillae.

Type species : Dioicomyces anthici.

With this revision, the genus Dioicomyces includes

23 species. Species previously removed from the genus

were: D. bournieri, transferred to Dimeromyces

(Santamaria 1999) ; D. endogaeus, transferred to Picar-

della (Tavares 1985) ; D. mesoveliae, D. verruculosus,

and D. yongboi transferred to Triceromyces (Benjamin

1998).

Dioicomyces africanus W. Rossi, Accad. Naz. Lincei255 : 13 (1982). (Figs 25–26)

Type : ‘Parasitus Formicomi sp. in Africa occidentali.

Holotypus in herbario Instituti Botanici apud Universitatem

Studiorum Romae, n. 1191 ’ (Rossi 1982; RO; the number of

the type slide was erroneously published as 1190).

Male thallus 78–83 µm long from foot to antheridial tip,

hyaline to pale yellowish, slightly curved. Basal cell (I)

of receptacle two or more times longer than broad, ca

as long as the two cells above combined (excluding the

efferent tube). Suprabasal (II) and third cells similar,

1±5 times longer than broad, rectangular in section.

Antheridium with the venter shorter than the com-

bination of two cells below, one to two efferent tubes,

which are terminal and ³ laterally offset.

Female thallus 217–238 µm long from foot to peri-

thecial tip, brownish amber, slightly arcuate, 70–78 µm

long from foot to apex of primary appendage.

Suprabasal cell (II) nearly isodiametric. Terminal cell

(III)³isodiametric and slightly inflated dorsally. Pri-

mary appendage narrowly oblong, straight, slightly

constricted at base, sides slightly curved and narrowing

to a rounded apex. Cells I, II, III, and primary

appendage neither dark nor obscured, showing the

same depth of color as the remainder of the thallus.

Perithecium (with basal cells) 155–175¬45–53 µm,

asymmetric, fusiform, with one side strongly convex

and the opposite side straight to slightly concave,

broadest at or below the middle, gradually tapering

towards a short, broad neck and a broadly rounded

apex. The outer side of the third cell of the perithecial

wall cell row formed from cell m (w$m

) inflated.

Perithecial wall cell rows spirally arranged. The number

of perithecial wall cells for each row is : wm

¯ 4, wn¯

5 (including a small distal cell) and 4, wn« ¯ 4. Stalk cell

(VI) of perithecium 30–38 µm long, ca 1±5 times longer

than broad, distally broadened, strongly constricted at

the base. Trichogyne with the upper cell bearing

numerous short and rounded papillae. Free ascospores

not observed.

This species is only known from the type collection.

Described on Formicomus sp. from Sierra Leone, it

appears to be closely related to D. italicus. Dioicomyces

africanus differs from D. italicus by the strong inden-

tations at septa separating the different tiers of outer

perithecial wall cells (Rossi 1982). Septa in D. italicus

protrude (Rossi 1993). The type slide examined contains

12 mature females, 3 male thalli, several fragments as

well as 12 immature females bearing, some of them,

undamaged trichogynes. This species is evidently related

The genus Dioicomyces 618

Key to species of Dioicomyces

1 Female thalli with perithecia showing some swollen wall cells or bearing tooth-like, finger-like or ³ elongateprominences or outgrowths . . . . . . . . . . . . . . . . 2

Female thalli with perithecia not showing outgrowths or conspicuous swellings . . . . . . 11

2(1) Cell n« and contiguous upper wall cell strongly inflated and protruding on the side of the perithecium as twoevident swellings . . . . . . . . . . . . . . . . umbonatus

Without swellings on perithecial wall, but with outgrowths or prominences of different shape . . . . 3

3(2) Two outgrowths on and near the perithecial tip : a finger-like, sometimes uncinate process and a blunt, darkexcrescence. On Myrmechixenus . . . . . . . . . . . . .myrmecophilus

Only one outgrowth on perithecium . . . . . . . . . . . . . . . 4

4(3) Perithecial outgrowth lateral and always spine-like . . . . . . . . . . . . 5Perithecial outgrowth apical or subapical, tooth-like or finger-like . . . . . . . . . 6

5(4) The spine-like outgrowth projecting upward from the upper quarter of the side of the perithecium and extendingabove the perithecial apex. Female thalli 131–154 µm. On Leptaleus . . . . . . . leptalei

The finger-like outgrowth projecting upward from the middle of the perithecial side and not surpassing theperithecial apex. Female thalli 178–212 µm. On Anthicus . . . . . . . . . spiniger

6(4) Perithecial outgrowth tooth-like and apical . . . . . . . . . . . . . 7Perithecial outgrowth finger-like, apical or subapical . . . . . . . . . . . 8

7(6) Cells I, II, III, and primary appendage dark brown, becoming opaque towards the outer side. Male thallibrownish, 44–46 µm. Efferent tube of antheridium subterminal. On Cyclodinus . . . . denticulatus

Cells I, II, III, and primary appendage brownish amber, as in the remainder of the thallus. Male thalli hyaline topale yellowish, 63–75 µm. Efferent tube of antheridium terminal. On Formicomus. . . . . borneensis

8(6) Perithecial outgrowth showing a septum near the base. On Formicomus . . . . . . borneensis

Perithecial outgrowth without septum . . . . . . . . . . . . . . 9

9(8) Female thalli with a terminal cell (III) ca 1±5 times longer than broad. Primary appendage with a sharp pointedapex . . . . . . . . . . . . . . . . . . glossophorus

Female thalli with a terminal cell (III) typical, minute, wedge-shaped, and strongly flattened. Primary appendagenot sharp pointed . . . . . . . . . . . . . . . . . . 10

10(9) Perithecium ovoidal, broadest below the middle. Stalk cell (VI) of perithecium 57–91 µm long . . onchophorus

Perithecium broadly fusiform, broadest near the middle. Stalk cell (VI) of perithecium 30 µm long . rostellatus

11(1) Perithecial axis conspicuously perpendicular with relation to the main longitudinal axis of the thallus. Lower partof the perithecium strongly protuberant just above the basal cells, opposite the ostiolar tip, which is anteriorlydirected . . . . . . . . . . . . . . . . . . malleolaris

Perithecial axis never conspicuously perpendicular (but see in D. anthici the thalli formerly included in D. falcatusand D. uncinatus in which the perithecium does not protrude at the base) . . . . . . . 12

12(11) Perithecial sides marked with indentations or protrusions (corrugations) at the septa between wall cells . . 13Perithecial sides not indented or corrugated . . . . . . . . . . . . . 17

13(12) Perithecial sides with some indentations . . . . . . . . . . . . . . 14Perithecial sides with septal protrusions or corrugations . . . . . . . . . . . 15

14(13) Perithecium strongly arcuate, conspicuously indented at septa between the wall cells. Female thallus 193–207 µmlong. Male thallus 48–52 µm long. On Anthicus . . . . . . . . . . indentatus

Perithecium not strongly arcuate, with w$m

slightly inflated and with septa indented only above and below this cell,with slight corrugations at other septa. Female thallus 217–238 µm long. Male thallus 78–83 µm long. OnFormicomus . . . . . . . . . . . . . . . . . . africanus

15(13) Septal protrusions only at the base of the dorsal side of the perithecium. Terminal cell (III) squarish. Appendagenearly cylindrical . . . . . . . . . . . . . . . . subtorulosus

Septal protrusions present on both sides of the perithecium . . . . . . . . . . 16

16(15) Perithecium broadest near the basal third. Perithecial apex rounded. Stalk cell of perithecium up to 84 µm long.On Formicomus . . . . . . . . . . . . . . . . . italicus

Perithecium broadest above the middle. Perithecial apex truncate. Stalk cell of perithecium 143 µm long. OnAnthicus . . . . . . . . . . . . . . . . . . . torulosus

17(12) Perithecium small, short, 59–77 µm long, only slightly broader than stalk cell (VI) . . . . proeminens

Perithecium larger, longer than 78 µm, in most cases longer than 100 µm, definitely broader thanstalk cell (VI) . . . . . . . . . . . . . . . . . . . 18

18(17) Ascospores obliquely septate. On staphylinids . . . . . . . . . . obliqueseptatus

Ascospores not obliquely septate. On anthicids or staphylinids (see D. floridanus) . . . . . . 19

19(18) Primary appendage distinctively dark brown in comparison with pale yellowish cells I, II, and III.On Notoxus . . . . . . . . . . . . . . . . . . notoxi

Primary appendage with the same depth of colour as cells I, II, and III. On other anthicids . . . . 20

20(19) Perithecium with a strongly laterally offset apex; the ostiole is perpendicularly oriented in relation to the thalliallongitudinal axis. Cell w

%n« shows a distinctively thickened outer wall . . . . . . . ladoi

Perithecium not as above. No conspicuous thickenings in outer walls . . . . . . . . 21

21(20) Stalk cell of perithecium (VI) very elongated, 229–329 µm. Cells I, II, III, and primary appendage nearlyblackened . . . . . . . . . . . . . . . . . . trinitatis

S. Santamaria 619

Stalk cell of perithecium (VI) up to 186 µm long. Cells I, II, III, and primary appendage not blackened . . 22

22(21) Primary appendage with a broadly rounded apex . . . . . . . . . . . inclinatus

Primary appendage with a ³ pointed and inwardly curved apex . . . . . . . . . 23

23(22) Stalk cell of perithecium (VI) 23–34 µm long, 1±5 times longer than broad. Perithecium fusiform. On staphylinids(gen. Bledius) . . . . . . . . . . . . . . . . . floridanus

Stalk cell of perithecium (VI) (40–)75–186 µm long, many times longer than broad. Perithecium ovoidal.On anthicids . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . anthici

to D. italicus, not only because it parasitizes anthicids

of the genus Formicomus but also because of many

morphological traits of male (pale and with terminal

efferent tube) and female thalli (rather big and

isodiametric cell III ; cells I, II, III and primary

appendage with the same depth of color as the

remainder of the thallus ; similar shape of primary

appendage; perithecium with a broad and rounded

apex; etc). In addition to characteristics emphasized by

Rossi (1982), D. africanus may be separated from D.

italicus by the spiral arrangement of the perithecial wall

cell rows, by the slightly convex outer sides of the

primary appendage (which are concave in D. italicus)

and by the slightly inflated cell w$m

(Fig. 26, arrow).

Additionally, I have observed striation in cells w"m

and

w#m

, similar to that observed in other species of

Dioicomyces, such as D. ladoi (Fig. 88, see cells w"–$m

)

or D. leptalei (Fig. 89, see cells w"–#n

«).

Specimen examined : Sierra Leone : Southern Province : near

Zimi, on Formicomus sp., 11–13 Feb. 1980, W. Rossi (RO-

1191 – holotype).

Dioicomyces anthici Thaxt., Proc. Amer. Acad. Arts 37 :33 (1901). (Figs 1–24, 73–77)

Type : USA : Fresh Pond, Cambridge (Mass.), on Anthicus

floralis, Oct. 1900, R. Thaxter (FH-3488 – lectotypus hic

designatus).

Dioicomyces formicillae Thaxt., Proc. Amer. Acad. Arts48 : 169 (1912) ; as ‘ formicellae ’).

Type : Argentina : Parque 3 Feb., Buenos Aires, Palermo,

on Formicilla strangulata Pic, Oct. 1905, R. Thaxter 1692.

(FH-3476 – lectotypus hic designatus) (Figs 5–6)

Dioicomyces angularis Thaxt., Proc. Amer. Acad. Arts48 : 171 (1912).

Type : Argentina : Llavallol, on Anthicus parvus Pic, ‘no

date ’,R.Thaxter 1513A (FH-3468 – lectotypus hic designatus).

(Fig. 1)

Dioicomyces falcatus Speg., Anales Mus. Nac. Hist.Nat. Buenos Aires 29 : 522 (1917).

Type : Sobre el corselete y las elitras del Anthicus post-

maculatus en Sta Catalina, B. A., May 1916 [Spegazzini 1917;

LPS! – holotype]. (Fig. 4)

Dioicomyces formicillae f. anthicicola Speg., AnalesMus. Nac. Hist. Nat. Buenos Aires 29 : 523 (1917).

Type : Sobre las elitras en su borde posterior y sobre

las patas traseras del Anthicus floralis en La Plata, Jun.

1916 [Spegazzini 1917; LPS! – holotype]. (Fig. 7)

Dioicomyces formicillae f. brachygnathus Speg., AnalesMus. Nac. Hist. Nat. Buenos Aires 29 : 524 (1917).

Type : Sobre el corselete y las elitras de la Formicilla

Bruchi, en La Plata, May. 1916 [Spegazzini 1917; LPS! –

holotype]. (Fig. 8)

Dioicomyces infuscatus Speg., Anales Mus. Nac. Hist.Nat. Buenos Aires 29 : 526 (1917).

Type : Sobre las elitras del Anthicus pallidicolor

en La Plata, Ener. 1914 [Spegazzini 1917; LPS!®holotype].

(Fig. 10)

Dioicomyces pallidus Speg., Anales Mus. Nac. Hist.Nat. Buenos Aires 29 : 527 (1917).

Type : Sobre los genitales del Anthicus postmaculatus

en Sta Catalina, B. A., Feb. 1916 [Spegazzini 1917;

LPS! – holotype]. (Fig. 11)

Dioicomyces refractus Speg., Anales Mus. Nac. Hist.Nat. Buenos Aires 29 : 528 (1917).

Type : Sobre el corselete y las elitras del Anthicus

postsignatus, en la isla Santiago, La Plata, May. 1916

[Spegazzini 1917; LPS! – holotype]. (Fig. 12)

Dioicomyces uncinatus Speg., Anales Mus. Nac. Hist.Nat. Buenos Aires 29 : 532 (1917).

Type : Comu! n sobre el corselete y las elitras del

Anthicus parvus, en Sta Catalina, B. A., Oct. 1915

[Spegazzini 1917; LPS! – holotype]. (Fig. 13)

Dioicomyces anthici var. fuscescens Maire, Bull. Soc.Hist. Nat. Afrique Nord 8 : 135 (1920).

Type : Sur l’e! lytre droit d’ Anthicus Rodriguesi Latr. :

C. Bo# ne, embouchure de la Seybouse, mars 1918 (De

Borde) [Maire 1920; not seen].

Dioicomyces guatemalensis Thaxt., Mem. Amer. Acad.Arts 16 : 64 (1931).

Type : On the posterior leg of Anthicus vicinus Laf.,

No. 1638, Agua Caliente, Guatemala (Kellerman)

[Thaxter 1931; FH! – holotype]. (Fig. 9)


yellowish brown, nearly straight. Basal cell (I) of

receptacle two or more times longer than broad, as long

or longer than the three cells above combined (excluding

the efferent tube). Suprabasal cell (II) similar to or

longer than third cell, squarish to rectangular in section.

Antheridium with the venter shorter than the com-

bination of two cells below, the efferent tube nearly

lateral, straight to curved, ³ laterally offset.


thecial tip, brownish, sigmoid to ³ arcuate,

(39–)51–69 µm long from foot to apex of primary

appendage. Primary appendage conical, with a ³pointed and inwardly curved apex. Cells I, II, III, and

primary appendage dark brown, often delicately dotted,

becoming opaque towards the dorsal side. Perithecium


Figs 1–13. Dioicomyces anthici and some of its synonyms. Males (Fig. 2 and arrows in Figs 1, 5, and 6) and Females from

types. Fig. 1. D. angularis (FH3468). Figs 2–3. D. anthici (FH3488). Fig. 4. D. falcatus (LPS45120). Fig. 5. D. formicillae

(FH3477 – male–, FH3479 – female–). Fig. 6. D. formicillae (FH3476). Fig. 7. D. formicillae f. anthicicola (LPS45141). Fig.

8. D. formicillae f. brachygnathus (LPS45142). Fig. 9. D. guatemalensis (FH3526). Fig. 10. D. infuscatus (LPS45121). Fig.

11. D. pallidus (LPS45122). Fig. 12. D. refractus (LPS45123). Fig. 13. D. uncinatus (LPS45124). Bars¯ 50 µm.

S. Santamaria 621

Figs 14–36. For caption see p. 622.


(with basal cells) 78–210¬35–83 µm, asymmetric, ovoi-

dal, usually with one side strongly convex and the other

straight or even slightly concave, broadest below the

middle, gradually tapering towards a short, broad neck

and a broadly rounded apex. The number of perithecial

wall cells for each row is : wm

¯ 5 (including a small

distal cell), wn¯ 5 (including a small distal cell) and

4(–5), wn« ¯ 4. Stalk cell (VI) of perithecium

(40–)75–186 µm long, longer than broad, with nearly

parallel sides, sigmoid to slightly arcuate. Trichogyne

with the upper cell bearing few short, slender append-

ages. Ascospores dimorphic, 32–65 µm long (females),

26–50 µm long (males).

This is a widespread and very variable species

reported from all continents except Australia. It was

described by Thaxter (1901) on Anthicus floralis and A.

californicus from the USA. Subsequently it has been

reported worldwide on several species of Anthicus s.lato

(including Cyclodinus and Leptaleus) from Senegal

(Spegazzini 1914), Algeria (Maire 1916), Guatemala,

Jamaica, Cameroon (Thaxter 1931), Hungary (Ba! nhegi

1944), Libya (Maire 1948), Korea (Y. B. Lee & C. I.

Lee 1982), Poland (Majewski 1986), Finland (Hulde!n1983), Bali (Lee & Sugiyama 1984), Spain (Santamaria

1989), France (Balazuc 1990), India (Kaur & Mukerji

1996), and the United Kingdom (Weir 1996).

I have studied several hundreds of fungal thalli

collected on anthicids from the Iberian Peninsula that

may be forms or varieties included in D. anthici. I have

borrowed the type of D. anthici from FH and after the

study of this material I decided to define the character-

istics of D. anthici in a broad sense. It seems more

appropriate to regard D. anthici as a species complex or

collective species because its forms or varieties are

difficult to separate on the basis of morphological

characters. I have hesitated to maintain many of the

synonyms proposed here as infraspecific taxa (varieties

or forms might be adequate) because I doubt the

taxonomical value of such names.

The type slide of D. anthici consists of several mature

females and few males. The females have a ovoidal,

rather symmetric perithecium, which is apparently not

the commonest perithecial form in D. anthici as

indicated by illustrations subsequently published by

other authors. This adds support to the concept of wide

variation in D. anthici. Only molecular studies might

Figs 14–36. Dioicomyces spp. Figs 14–24. D. anthici. Variability of forms among Iberian samples. Arrow in Fig. 22

indicate the ‘protuberant basal cell ’ of a form similar to ‘D. refractus ’. Figs 15 and 21 represent males, others are females

(Fig. 14: BCB-SS1063; Fig. 15: BCB-SS0843b; Fig. 16: BCB-SS1432; Fig. 17: BCB-SS1462; Fig. 18: BCB-SS1087; Fig.

19: BCB-SS1069; Figs 20–21: BCB-SS0883b; Fig. 22: BCB-SS0890; Fig. 23: BCB-SS0965b; Fig. 24: BCB-SS1062). Figs

25–26. D. africanus. Male (Fig. 25) and female (Fig. 26) thalli from type slide. In Fig. 25 arrows indicate the efferent

necks of two antheridia. In Fig. 26 arrow indicates the inflated w$m

(RO1191). Figs 27–30. D. borneensis. Male (Fig. 28)

and female thalli from type slide. In Fig. 27 a detail of perithecial apex showing the digitate outgrowth with a basal

septum (arrow). In Fig. 29 a female thallus with perithecial digitate outgrowth (arrow). In Fig. 30 a female thallus with

tooth-like outgrowth (arrow) on perithecial tip (Fig. 28: K-S-3228d; Figs 27, 29–30: K-S-3228a). Figs 31–34. D.

denticulatus. Male (Fig. 33) and female thalli from type. Arrows in Figs 31, 32, and 34 indicate the perithecial tooth-like

outgrowth (Fig. 31: BCB-SS1061; Fig. 32: BCB-SS1059; Figs 33–34: BCB-SS1064a). Figs 35–36. D. floridanus. Male (Fig.

35) and female thalli from type slide (Fig. 35: FH3474; Fig. 36: FH3473). Bars¯ 50 µm except in Fig. 31 which is 25 µm.

add new features to an understanding of the taxonomic

value of such variations. A discussion follows of each of

the synonymized taxa.

Dioicomyces angularis was described by Thaxter

(1912) on Anthicus parvus Pic from Argentina. Later,

Spegazzini (1917) reported the species on the same host

from Argentina. Thaxter (1931) stated that this species

was very closely allied to D. anthici, differing from this

by the ‘angular ’ outer side of its perithecium, although

it ‘ is not invariably as striking as in the two individuals

represented in figures 41 and 42 ’. Thaxter also

questioned the identification of Argentinian thalli by

Spegazzini (1917). After examination of types, it seems

preferable to synonymize this form with D. anthici

because ‘ the angular outer margin of the perithecium ’

is not clearly identifiable among the thalli examined

(Fig. 1). Other characters seem of minor value.

Dioicomyces anthici var. fuscescens was described by

Maire (1920) on Leptaleus rodriguesi (as Anthicus

rodriguesi) from Algeria. This variety should be included

among the wide variation of the collective species D.

anthici. There is no reason to maintain this name.

Dioicomyces falcatus was described by Spegazzini

(1917) on the elytra and superior surface of the thorax

of Anthicus postmaculatus from Argentina. Later,

Thaxter (1931) mentioned ‘ the unusual development of

the very broad, coarse lips of its upcurved tip and apex ’

of the perithecium as the main diagnostic characteristic

rather than the ‘ falcate ’ habit. Later, Majewski (1988)

reported the species on legs of Pseudoleptaleus valgipes

from Japan. The Polish author mentioned some doubts

about the determination of this Japanese material. Fig.

11 in Majewski’s paper illustrated a female thallus with

‘ falcate ’ habit but without the so-called ‘ typical ’

perithecial tip described by Spegazzini. Terada (1991)

photographed two females and one male of this species

collected on the same Japanese host ; these photographs

show a broad variation with one falcate thallus (Terada

1991: fig. 2) and one sigmoid thallus (Terada 1991: fig.

1). The type slide received from LPS included three

females and no males. Spegazzini’s drawing of the

perithecial tip shows broad terminal lobes. The exam-

ination of these thalli demonstrated that these lobes are

not visible, as shown in my photograph of the type (Fig.

4). The tip is broad and blunt, as Spegazzini indicated

it, but he showed a more slender primary appendage. I

have examined many similar thalli among the Iberian

S. Santamaria 623

Figs 37–59. Dioicomyces spp. Figs 37–38. D. glossophorus (LPS45143). Fig. 37 represents a detail of the perithecial apex.

Arrows indicate the perithecial outgrowth, which in Fig. 38 is out of focus. Fig. 39. D. inclinatus (FH3528). Female

thallus. Arrow indicates the rounded apex of the primary appendage. Figs 40–42. D. indentatus. Male (Fig. 40: FH3530)

and females (Fig. 41: FH3530; Fig. 42: FH3531). Figs 43–46. D. italicus. Males (Fig. 43: BCB-SS417a; Fig. 45: FH3485)

and females (Fig. 44: BCB-SS417d; Fig. 46: FH3485). Arrows in Fig. 45 indicate the efferent necks of two antheridia.

Figs 45 and 46 were from type slide of D. formicomi. Figs 47–49. D. ladoi (BCB-2261a). Female (Fig. 47), male (Fig. 48)

and detail of perithecium (Fig. 49). Arrows in Figs 47 and 49 indicate the thickened outer wall of cell w%n

«. Figs 50–52. D.

leptalei. Male (Fig. 50: BCB-SS661), female (Fig. 51: BCB-SS674a), and detail of perithecium (Fig. 52: BCB-SS1520).

Arrows in Figs 51 and 52 indicate the perithecial outgrowth. Fig. 53. D. malleolaris (FH3537). Two females and one male

(arrow). Figs 54–56. D. myrmecophilus. Male (Fig. 54: TM-912), female (Fig. 55: TM-912), and detail of perithecium (Fig.

56: TM-908). Arrow in Fig. 54 indicates the sharp apex of the antheridium; arrows in Figs 55 and 56 indicate perithecial

outgrowths. Figs 57–59. D. notoxi. Male (Fig. 58: FH3541) and females (Fig. 57: FH3540; Fig. 59: FH3542). Bars¯50 µm except in Figs 37 and 45 which is 25 µm.


material (Figs 20, 74), most growing on the abdomen

tip of their hosts, with a very large range of variation;

therefore I have decided to include this species among

the synonyms of D. anthici.

Dioicomyces formicillae was described (as ‘D.

formicellae ’) by Thaxter (1912) on Formicilla strangu-

lata (host as Formicella) from Argentina. As mentioned

by Thaxter himself, this species was the largest species

(by size) at the time of its description and had ‘no

striking peculiarities ’. Spegazzini (1917) reported and

illustrated this species on Formicilla leporina from

Argentina and described two additional forms, D.

formicillae f. anthicicola on Anthicus floralis and

D. formicillae f. brachygnatus on Formicilla bruchi, both

from Argentina. The former of these two forms is

identical to D. anthici (Fig. 7) and the second shows a

more abruptly narrowed apex than the typical form

and demonstrates another variation (Fig. 8). Thaxter

(1931) reported D. formicillae as allied to D. anthici but

‘easily ’ distinguished by its large size, long, dark,

coarse-tipped perithecium, and the rather gradual

transition from the basal cell region to that of the

venter. After examination of types, this ‘species ’ is

being included among the synonyms of D. anthici

because it is too difficult to define adequately any

diagnostic characteristic.

Dioicomyces guatemalensis is only known from the

type described by Thaxter (1931) on the posterior legs

of Anthicus vicinus from Guatemala. It is hardly

separable from D. anthici. According to Thaxter (1931)

it ‘recalls ’ D. falcatus in general habit. The type slide

received included one female (Fig. 9) and two male

thalli, which seem to be unique and the same as

described and illustrated by Thaxter. This material is

very scarce to be used to describe a species. However,

the female primary appendage is broad and abruptly

narrowed to the apex, like that of D. falcatus. This

fungus also resembles D. refractus (see below) by the

shape of the primary axis of females. I don’t see an

adequate reason for maintaining this as a separate

species.

Dioicomyces infuscatus was described by Spegazzini

(1917) on elytra of Anthicus pallidicolor from Argentina.

According to the author, this species is very near D.

anthici but may be distinguished by its short, stout

habit and deeper coloration. Thaxter (1931) pointed

out the absence of any striking peculiarities. The male

was not described. The single slide received from LPS

and labelled as the type included only one female

thallus (Fig. 10) what seems to be the same drawn by

Spegazzini (1917). I have collected several specimens of

D. anthici similar to this form (Fig. 17) with inter-

mediates between this and other forms.

Dioicomyces pallidus which was only known from the

type (Spegazzini 1917) on Anthicus postmaculatus from

Argentina, lacks distinctive characters (Fig. 11). It was

described by Spegazzini as rather similar to D. anthici.

Spegazzini’s drawings magnify several characters that I

regards irrelevant. For example, what is indicated by

Spegazzini as a ‘pale ’ thallus is indeed a yellowish

colour, common for many other species of Dioicomyces.

Moreover, colour alone, is not a good character on

which to base a species.

Dioicomyces refractus, described by Spegazzini (1917)

on Anthicus postsignatus from Argentina, has never

been reported again. Spegazzini mentioned that this

form was originally considered teratological, but later,

when it was found in relative abundance it was described

as a separate species. Thaxter (1931) compared this

species with D. inclinatus because of the angle of

inclination of perithecium at the level of the basal cells.

As stated by Thaxter, the orientation of this incline is

opposite in the two species. The type slide received from

LPS contains two females and no males (which had not

been described originally). I considered this species

distinct from D. inclinatus especially because of the

rounded primary appendage of the latter (Fig. 39),

which represents a distinctive character in relation to D.

anthici. Spegazzini’s drawing (1917: 529 nu! m. 71)

represents a thallus with very exaggerated character-

istics, specially that of the protuberant basal cell (Fig.

12). Among Iberian specimens I have found several

thalli with similar ‘ inclinations ’ (Fig. 22), that seem to

be related to unfavourable growth positions on the host

body (e.g. under the abdomen, legs, and tarsi).

Dioicomyces uncinatus was described by Spegazzini

(1917) on the pronotum and elytra of Anthicus parvus

from Argentina. In the type slide there are thalli which

represent transitional forms to ‘more typical ’ D. anthici

as well as thalli which are totally typical D. anthici. The

author compared his species with D. malleolaris, but I

think D. uncinatus is more similar to D. falcatus from

which it only clearly differs by a longer cell VI (Fig. 13).

Spegazzini’s description of males seems erroneous

because his drawings are imprecise and do not

accurately represent the thalli studied. I have found

many similar forms among Iberian thalli (Fig. 18),

mostly growing near the elytral tips of their hosts.

Specimens examined : Argentina : Llavallol, on Anthicus

parvus [no date], R. Thaxter 1513A (FH-3468, lectotype of D.

angularis) ; ibidem, Apr. 1906, R. Thaxter 1513A (FH-3469,

isolectotype of D. angularis) ; idem, Temperley, Apr. 1906, R.

Thaxter 1513-1513A (FH-3470, FH-3471 – paratypes of D.

angularis). Santa Catalina, on Anthicus postmaculatus, 9 May

1916, C. Spegazzini 477-1916 (LPS-45120, D. falcatus). Parque

3 Feb., Buenos Aires, Palermo, on Formicilla strangulata,

Oct. 1905, R. Thaxter 1692 (FH-3476 – lectotype of D.

formicillae ; FH-3477, FH-3478, FH-3480, FH-3481, FH-

3482, FH-3483, FH-3484 – isolectotypes of D. formicillae) ;

ibidem, Feb. 1905, R. Thaxter 1692 (FH-3479 – paratype of

D. formicillae). La Plata [‘LP ’ in the slide label], on Anthicus

floralis, 1 June 1916, C. Spegazzini 482-1916 (LPS-45141, D.

formicillae f. anthicicola). La Plata [‘LP ’ in the slide label], on

Formicella [sic], 15 Jan. 1914, C. Spegazzini 136b (LPS-45142,

D. formicillae f. brachygnathus). ‘Without locality on the slide

label ’, on Anthicus sp., 15 Jan. 1914, C. Spegazzini 400b-1914

(LPS-45121, D. infuscatus). Santa Catalina, on ?, 1 to 3 Feb.

1916, C. Spegazzini 424-1916 (LPS-45122, D. pallidus). Isla

Santiago, on Anthicus postsignatus, 5 May 1916, C. Spegazzini

S. Santamaria 625

462-1916 (LPS-45123, D. refractus). Santa Catalina, on

Anthicus postmaculatus, 1 to 3 Feb 1916, C. Spegazzini 420-

1916 (LPS-45124, D. uncinatus). Guatemala : Agua Caliente,

on Anthicus ?, Feb. 1908, R. Thaxter 1637 (FH-3526 – type of

D. guatemalensis). – Portugal : Beira Alta : Vouzela ; Perimetro

Florestal da Penoita, Serra do Caramulo, on Anthicus tristis,

6 Nov. 1996, S. Santamaria (BCB-SS1887). – Senegal :

Diake' ne, on Anthicidae indet., 20 Jul. 1968, MZB (BCB-

SSE201). – Spain : Barcelona : Cerdanyola del Valle' s ;Bellaterra, on Leptaleus rodriguesi, 4 Apr. 2000, S. Santamaria

(BCB-SS2342). Gualba; Gualba de baix, Gualba stream, on

Hirticomus hispidus, 9 Jul. 1991, 21 Jul. 1991, 9 Aug. 1991, 11

Aug. 1991, S. Santamaria (BCB-SS1069, SS1083a-b, SS1087) ;

ibidem, on Leptaleus rodriguesi, 20 Aug. 1995, S. Santamaria

(BCB-SS1873). El Prat de Llobregat, on Cyclodinus minutus,

4 Oct. 1986, S. Santamaria (BCB-SS0417}1). Mura; Nespres

stream, on Leptaleus rodriguesi, 9 May 1987, 23 Jul. 1990, S.

Santamaria (BCB-SS0701, SS0965a-d). Viladrau; El

Montseny, Sant Marc: al, on Cyclodinus humilis, 10 Jul. 1988,

21 Jul. 1988, 7 Aug. 1988, 29 Sep. 1988, 11 May 1989, 20 June

1989, 7 Jul. 1989, Zoologia-UAB (BCB-SS1057, SS1058b,

SS1059}2, SS1060, SS1061}2, SS1062, SS1063, SS1064b).

Cantabria : Santon4 a, marshlands, on Anthicidae indet., 19

June 2000, S. Santamaria (BCB-SS2313a-e). Ciudad Real :

Pedro Mun4 oz; Alcahozo lagoon, on Cyclodinus constrictus,

24 June 1998, S. Santamaria (BCB-SS2008a-c). Villamayor de

Calatrava; Cucharas lagoon, on Cyclodinus constrictus, 25

June 1998, S. Santamaria (BCB-SS2014a-b). Villamayor de

Calatrava; los Almeros lagoon, Idem (BCB-SS2031). Cuenca :

Las Mesas ; Manjavacas lagoon, on Cyclodinus constrictus, 24

June 1998, S. Santamaria (BCB-SS2007). Gerona [Girona] :

Riells i Viabrea; Ju! nior Parc, on Leptaleus rodriguesi, 20 Apr.

1987, 24 May 1987, 2 Aug. 1987, 2 Jul. 1989, 2 Sep. 1989, 1

Oct. 1989, 20 Apr. 1992, 2 Aug. 1992, 2 Aug. 1993, 20 May

1995, S. Santamaria (BCB-0674a, SS0702, SS0740a, SS0858,

SS0883a-b, SS0890, SS0891, SS1462, SS1544, SS1716a,c,

SS1717, SS1718, SS1719, SS1720, SS1863c,e). Gran Canaria

(Canary Islands) : Isla de la Montan4 a Clara, on Anthicus sp.,

20 Aug. 1989, MZB (BCB-SSE90). Guadalajara : Puebla de

Belen4 a; lagoons, on Anthicus antherinus, 25 June 1997, S.

Santamaria (BCB-SS1896b, SS1896d-h). Huesca : Aisa, on

Anthicus laeviceps, Aug. 1986, J. Franc (BCB-SS0852a). Ibiza

[Eivissa] : Sant Antoni Abad, on Anthicus tristis, Nov. 1935,

MZB (BCB-SS1432). LeU rida [Lleida] : Baix Pallars ; Gerri de

la Sal, on Cyclodinus coniceps, 15 Apr. 1918, MZB (BCB-

SS1181). Madrid : Madrid city ; Isabel II channel, on Anthicus

sp., no date, MNCN (BCB-SS1345, SS1347). Mallorca :

Bunyola; Biniatzar, on Anthicus sp., Oct. 1942, MZB (BCB-

SS1424). Cabrera Island, on Cordicomus instabilis ?, Jul. 1935,

MZB (BCB-SS1421). Palma; El Molinar, on Anthicus sp.,

June 1936, MZB (BCB-SS1425).Navarra : Pitillas, on Anthicus

sp., 7 Jul. 1996, I. Ribera (BCB-SS1945). Zamora : Revellinos

de Campo, Villafa! fila lagoons, Las Salinas lagoon, on

Cyclodinus constrictus, 21 June 1999, 23 June 2000,

S. Santamaria (BCB-SS2261c-h, BCB-SS2333a-b). – USA :

Massachusetts : Cambridge, Fresh Pond, on Anthicus floralis,

Oct. 1900 (FH-3488 – lectotype of D. anthici).

Dioicomyces borneensis T. Majewski & K. Sugiy.,Trans. mycol. Soc. Japan 27 : 428 (1986).

(Figs 27–30)

Type : ‘In Formicomo sp. (Coleoptera : Anthicidae), Kota

Kinabalu, Saba, Insulae Borneo, Malaysiae Orientalis, die 28

Martii 1981, K. Sugiyama leg., K-S-3228 in Herbario K.

Sugiyamae in Universitate Shizuokae ’ [Majewski & Sugiyama

1986; K. Sugiyama Herbarium!].


hyaline to pale yellowish, straight to slightly curved.

Basal cell (I) of receptacle two or more times longer

than broad, longer than the two cells above combined

(excluding the efferent tube). Suprabasal cell (II) 1±5times longer than broad, rectangular in section. Third

cell flattened, rectangular in section. Antheridium with

the venter shorter than the combination of two cells

below, the efferent tube terminal and ³ laterally offset.


thecial tip (not including outgrowth), brownish amber,

variably arcuate, 73–78 µm long from foot to apex of

primary appendage. Terminal cell (III)³isodiametric

and slightly inflated dorsally. Primary appendage

dolabriform, with a rounded and nearly straight apex.

Cells I, II, III, and primary appendage neither dark nor

obscured, showing the same color depth as the

remainder of the thallus. Perithecium (with basal cells

and without outgrowth) 175–200¬50–53 µm, asym-

metric, fusiform, with one side strongly convex and the

opposite side straight, broadest just below the middle,

gradually tapering towards a short, broad neck and a

broadly rounded apex, which extends into a blunt,

small tooth-like outgrowth or into a finger-like,

subacute, basally septated, 65–67 µm long outgrowth.

The number of perithecial wall cells for each row is : wm

¯ 4, wn¯ 5 and 4 (including outgrowth), w

n« ¯ 4. Stalk

cell (VI) of perithecium 48–65 µm long, ca twice longer

than broad, distally broadened, strongly constricted at

the base. Free ascospores not observed.

Only known from the type. According to its authors,

D. borneensis is well-characterized by having a pro-

jection from the perithecial apex. The authors compared

their species with D. myrmecophilus which also has a

perithecial projection. Nevertheless, according to de-

scription and drawings, D. borneensis should be

compared with D. italicus and D. africanus, which also

parasitize anthicids of the genus Formicomus. Majewski

& Sugiyama (1986: fig. 13) illustrated a specimen

without the finger-like process which seems identical to

D. italicus. A study of the type slides shows that there

are two characters overlooked by Majewski and

Sugiyama that should be emphasized: (1) the thalli

without the finger-like process show a tooth-like

projection formed from cell w%m

(Fig. 30, arrow), and

(2) the finger-like outgrowth has a septum near its base

(Fig. 27, arrow). The sharp projection, blunter apex,

and more gradual narrowing to apex distinguish those

thalli without the elongate excrescence from D. italicus.

Majewski & Sugiyama (1986) only mentioned an

elongate anterior lip-cell curved over the ostiole, but

their drawing (fig. 13) diminishes this characteristic.

Specimens examined : East Malaysia (Sabah) : Borneo : Kota

Kinabalu, on Formicomus sp., 28 March 1981 (K-S-

3228a – holotype; K-S-3228b, K-S-3228c, K-S-3228d –

isotypes).


Dioicomyces denticulatus Santam., sp. nov.(Figs 31–34, 78–81)

Etym. : denticulatus-based on the small tooth-like

outgrowth on perithecial apex.

Thallus mas 44–46 µm longus ab pede ad antheridii apicem,

brunneolus, in totum subtiliter punctatus, rectus. Basalis

cellula (I) receptaculi bis vel pluries longior quam latior, tam

longa quam tres cellulae superiores coniunctae (collo excluso).

Suprabasalis (II) et tertia cellula similes, quadratae in sectione.

Antheridium cum ventre tam longo quam duae cellulae

inferiores coniunctae ; collum subterminale et rectum.

Thallus femineus 148–157 µm longus ab pede ad perithecii

apicem, succineus-brunneus, rectus, 39–41 µm longus ab pede

ad primariae appendicis apicem. Primaria appendix conica,

cum ³ acuto et leviter introrsum curvo apice. Cellulae I, II,

III et primaria appendix atrobrunneae, subtiliter punctatae,

opacescentes in dorsalem marginem versum. Perithecium

(cum basalibus cellulis) 101–108¬40–45 µm, parum asi-

metricum, ovoidale, latus circa medium, paulatim angustatum

in breve, latum collum et obtusum apicem, qui in denti-

formem, rotundatam, parvam prominentiam protenditur.

Pedicellaris cellula (VI) perithecii 25–33 µm longa, leviter

longior quam latior, distaliter lata facta. Ascosporae parvum

dimorphae, 34–36 µm longae (femineae), 28–30 µm longae

(mares).

Typus : Hispania orientalis ; ‘Barcelona: Viladrau; El

Montseny, Sant Marc: al ’, super Cyclodinus humilis, 7 Jul.

1989, Zoologia-UAB (BCB SS1064a – holotypus).


brownish, entirely and delicately dotted, straight. Basal

cell (I) of receptacle two or more times longer than

broad, as long as the three cells above combined

(excluding the efferent tube). Suprabasal (II) and third

cells similar, squarish in section. Antheridium with the

venter as long as the combination of two cells below,

the efferent tube subterminal and straight.


thecial tip, amber-brown, straight, 39–41 µm long from

foot to apex of primary appendage. Primary appendage

conical, with a ³ pointed and slightly inwardly curved

apex. Cells I, II, III, and primary appendage dark

brown, delicately dotted, becoming opaque towards

the dorsal side. Perithecium (with basal cells)

101–108¬40–45 µm, slightly asymmetric, ovoidal,

broadest near the middle, gradually tapering towards a

short, broad neck and a blunt apex, which extends into

a tooth-like, rounded, small outgrowth. The number of

perithecial wall cells for each row is : wm

¯ 5, wn¯ 5?

and 4?, wn« ¯ 4 (forming the outgrowth). Stalk cell (VI)

of perithecium 25–33 µm long, slightly longer than

broad, distally broadened. Ascospores slightly dimor-

phic, 34–36 µm long (females), 28–30 µm long (males).

Dioicomyces denticulatus may be compared with

other species having perithecial outgrowths. Dioi-

comyces onchophorus may be considered the closest

species because of several morphological traits, in-

cluding that of the perithecial outgrowth. Nevertheless,

D. denticulatus may be distinguished from D. oncho-

phorus by: (1) the shorter outgrowth, tooth-like in the

former, more elongate, finger-like in the later, also

(although this is difficult to see) outgrowths were

formed from different rows in the two species, from wm

in D. onchophorus and from wn« in D. denticulatus ; (2)

all the measurements are smaller in D. denticulatus ; and

(3) cell VI is arcuate and elongate in D. onchophorus,

whereas is stout and short in D. denticulatus.

Specimens examined : Spain : Barcelona : Viladrau; El

Montseny, Sant Marc: al, on Cyclodinus humilis, 21 Jul. 1988,

7 Aug. 1988, 29 Sep. 1988, 7 Jul. 1989, 20 Jul. 1989, Zoologia-

UAB (SS1064a – holotype; BCB-SS1058a, SS1059,

SS1061 – paratypes).

Dioicomyces floridanus (Thaxt.) Thaxt., Proc. Amer.Acad. Arts 37 : 33 (1901). (Figs 35–36)

Amorphomyces floridanus Thaxt., Proc. Amer. Acad.Arts 28 : 159 (1893).

Type : ‘On the abdomen of Bledius basalis Lec., Florida ’

[Thaxter 1893; FH!].

Male thallus 68 µm long from foot to antheridial tip,

brownish, slightly sigmoid. Basal cell (I) of receptacle

two or more times longer than broad, slightly shorter

than the three cells above combined (excluding the

efferent tube). Suprabasal cell (II) twice as long as third

cell, rectangular in section. Third cell flattened. An-

theridium with the venter as long as the suprabasal cell

below, the efferent tube nearly lateral, curved.


thecial tip, yellowish brown, sigmoid to arcuate,

54–62 µm long from foot to apex of primary appendage.

Primary appendage conical, with a ³ pointed and

inwardly curved apex. Cells I, II, III, and primary

appendage slightly darker than the remainder of the

thallus. Perithecium (with basal cells) 146–168¬45–

61 µm, asymmetric, ³ elongate fusiform, broadest

below the middle, gradually tapering towards a short,

broad neck and a broadly rounded, laterally offset

apex. The number of perithecial wall cells for each

row is : wm

¯ 5? (including a small distal cell ?), wn¯ 5

and 4(–5), wn« ¯ 4. Stalk cell (VI) of perithecium

23–34 µm long, 1±5 times longer than broad, constricted

at the base, distally broadened. Free ascospores not

observed.

This species was originally described in the genus

Amorphomyces (Thaxter 1893), but later was transferred

to Dioicomyces (Thaxter 1901). According to Thaxter

(1908) the female thalli of this species resembles that of

D. anthici. After examination of Thaxter’s slides I

concluded that D. floridanus should be regarded as

distinct although it lacks any striking characteristic. It

may be distinguished from D. anthici by the unusually

shortened cell VI (as compared with the average size in

D. anthici), by the ³ elongate fusiform shape of

perithecium and by the absence of darkening on the

dorsal side of cells I, II, and III. Dioicomyces floridanus

and D. obliqueseptatus are the only species parasitizing

Staphylinidae beetles, and with D. myrmecophilus the

only species of the genus not affecting Anthicidae

beetles.

S. Santamaria 627

Specimens examined : USA : Florida: on Bledius basalis, R.

Thaxter (FH-3473 – holotype). Ipswich, on small Bledius, R.

Thaxter (FH-3474).

Dioicomyces glossophorus Speg., Anales Mus. Nac.Hist. Nat. Buenos Aires 29 : 525 (1917).

(Figs 37–38)

Type : ‘Sobre las tibias y los lados del abdomen del Anthicus

postmaculatus en Sta Catalina, B. A., Febr. 1916 ’ [Spegazzini

1917; LPS!].

Male thallus not observed.


thecial tip (excluding the outgrowth), yellowish, ³straight except at the strongly arcuate perithecial tip,


Terminal cell (III) ca 1±5 times longer than broad, with

inflated sides. Primary appendage conical, with a sharp-

pointed and inwardly curved apex. Cells I, II, III, and

primary appendage dark brown. Perithecium (with

basal cells) 112–118¬35–642 µm, asymmetric, fusiform,

broadest below the middle, gradually tapering towards

a short neck and a strongly laterally offset and blunt

apex. A subterminal, finger-like outgrowth borne near

the perithecial apex. The outer side marked with two

slight prominences at those septa between cells VII and

n«, and between cells w"n

« and w#n

«. The perithecial wall

cells poorly defined and their number no easy to

determine in the material studied. Stalk cell (VI) of

perithecium 30–33 µm long, twice longer than broad,

distally broadened, constricted at the base; its distal

septum located above the upper level of cell VII. Free

ascospores not observed.

This is a well-distinguished species only known from

the original description based on thalli collected on legs

and abdominal margins of Anthicus postmaculatus from

Argentina (Spegazzini 1917). The single slide received

on loan includes four females (two of them broken) and

no males. Female thalli show very peculiar characters :

a finger-like, sigmoid outgrowth near the bent peri-

thecial apex (Fig. 37), a cell III longer than broad and

a sharp-pointed, spiny primary appendage (not shown

in the photographs here included). Description of males

by Spegazzini (1917) may be based on misinterpreted

thalli, as may occur with other descriptions of males of

Dioicomyces species given by this author.

Specimen examined : Argentina : Santa Catalina, ‘pedibus ’

Anthicus postmaculatus, 1}3 Feb. 1916, C. Spegazzini 422-

1916 (LPS-45143).

Dioicomyces inclinatus Thaxt., Mem. Amer. Acad. Arts16 : 64 (1931). (Fig. 39)

Type : On legs of Anthicus setosus Laf. No. 2554, Manila,

P. I. [Thaxter 1931; FH!].

Male unknown.


thecial tip, dark brown, arcuate, 43–45 µm long from


cylindrical to conical, with a broadly rounded apex.

Cells I, II, III, and primary appendage dark brown

towards the dorsal side. Perithecium (with basal cells)

143–146¬48–50 µm, asymmetric, fusiform, broadest

below the middle, gradually tapering towards a short,

broad neck and a broadly rounded, laterally offset

apex. Cells VII and n distinctly inflated. The perithecial

wall cells poorly defined and their number no easy to


perithecium 66–68 µm long, three or more times longer

than broad, slightly broadened distally, arcuate. Free


This species, which was only known from the type on

Anthicus setosus from the Philippines (Thaxter 1931)

was regarded as similar to D. guatemalensis by its

author. Certainly it resembles this species, but also it

resembles D. refractus and therefore D. anthici. The

type slide received from FH included four females and

no males. I distinguish Dioicomyces inclinatus from the

closely allied D. anthici by the rounded, almost

cylindrical and straight primary appendage of the

former (Fig. 39, arrow) whereas it is pointed, more

conical, and inwardly curved in D. anthici.

Specimens examined : Philippines : Manila, on Anthicus

setosus, June 1912, R. Thaxter 2554 (FH-3528 – holotype).

Dioicomyces indentatus Thaxt., Mem. Amer. Acad. Arts16 : 65 (1931). (Figs 40–42)

Type : ‘On the mid left elytron of Anthicus setosus Laf., No.

2553, Manila, P. I. ’ [Thaxter 1931; FH!].


pale yellowish, arcuate. Basal cell (I) of receptacle 1±5times longer than broad, slightly longer than the

suprabasal cell. Suprabasal cell (II) ca 1±5 times longer

than broad, rectangular in section. Third cell minute,

shorter, rectangular in section. Antheridium with the

venter as long as the suprabasal cell, the efferent tube

terminal, curved, and slightly laterally offset.


thecial tip, pale yellowish, strongly arcuate, 45–52 µm

long from foot to apex of primary appendage. Primary

appendage subconical, with a blunt and straight apex.

Cells I, II, III, and primary appendage neither dark nor

obscured, showing the same colour depth as the

remainder of the thallus. Perithecium (with basal cells)

130–152¬36–43 µm, asymmetric, fusiform, with one

side strongly convex and the opposite side concave,

broadest near the base, ³ conspicuous indentations

marking the septa between the wall cells, gradually

tapering towards a long, broad neck and a broad,

truncate apex. Basal cells (VII, m, n, n«) inflated,

especially the cells VII and n«. The number of perithecial



distal cell), wn¯ 5 and 5, w

n« ¯ 4. Stalk cell (VI) of the

perithecium 57–66 µm long, three times longer than

broad, distally broadened, constricted at the base.


Ascospores dimorphic, 34 µm long (females), 28 µm

long (males).

This species is clearly distinguished by the

‘ indentations ’ observed at the septa separating peri-

thecial wall cells (Figs 41–42). It is only known from the

type from the Philippines.

Specimens examined : Philippines : Manila, on Anthicidae,

June 1912, R. Thaxter 2553 (FH-3530 – holotype; FH-3531).

Dioicomyces italicus Speg., Anales Mus. Nac. Hist. Nat.Buenos Aires 27 : 51 (1915).

(Figs 43–46, 82–84, 97)

Type : ‘118–1915}Dioicomyces}italicus}typus}Conegliano,

VII-1914 ’ [according to R 1993: 125, caption of figs

1–3–; LPS, not seen].

Dioicomyces formicomi Thaxt., Mem. Amer. Acad. Arts16 : 63 (1931).

Type : ‘In various positions on Formicomus inhumeralis Pic,

No. 3142 (Type), Los Ban4 os, Luzon, P. I. (Weston) ’ [Thaxter

1931; FH!]. (Figs 45–46)


pale yellowish, straight to slightly curved. Basal cell (I)

of receptacle 1±5–2 or more times longer than broad, as

long as the combination of two cells above or ca as long

as the combination of three cells above (excluding the

efferent tube). Suprabasal cell (II) 1±5 times longer than

broad, rectangular in section. Third cell similar to

suprabasal cell or ³ isodiametric, squarish to rec-

tangular in section. Antheridium with the venter as long

as any of the two cells below, the efferent tube terminal,

curved, and ³ laterally offset.


thecial tip, pale yellowish to amber, sigmoid to slightly

arcuate, 75–83 µm long from foot to apex of primary

appendage. Terminal cell (III) ³ isodiametric, slightly

inflated dorsally. Primary appendage subconical, with a

rounded and straight apex. Cells I, II, III, and primary


same depth of colour as the remainder of the thallus.


asymmetric, elongate-fusiform, with one side strongly

convex and the opposite side concave, broadest near

the basal third, gradually tapering towards a short,

broad neck and a broadly rounded apex, the septa

between the wall cells protruding to form ³ con-

spicuous corrugations. The number of perithecial wall

cells for each row is : wm

¯ 4, wn¯ 4 and 4(–5), w

n« ¯4. Stalk cell (VI) of perithecium 37–84 µm long, slender

or stout, variably longer than broad, either with nearly

parallel sides and no basal constriction or distally

broadened and strongly constricted at the base.

Trichogyne with the upper cell bearing numerous short,

irregular rounded papillae and few slender appendages.

Ascospores not dimorphic, 36–40 µm long.

Spegazzini (1915) described D. italicus on the basis of

material collected on Anthicus (hispidus?) from Italy. D.

formicomi was described by Thaxter (1931) on Formi-

comus inhumeralis from the Philippines. Subsequently,

D. formicomi has been reported on Formicomus bra-

minus from Japan (Sugiyama 1973) and on Formicomus

pedestris from Spain (Santamaria 1989). Rossi (1993)

found more fungi on F. pedestris from Italy, identical

with those reported by Sugiyama and Santamaria; he

studied the type of D. italicus and concluded that

D. formicomi was probably synonymous; moreover,

Anthicus hispidus is now called Hirticomus hispidus,

which is an unlikely host of D. italicus. I borrowed the

type of D. formicomi from FH (Figs 45–46) and agree

with Rossi’s opinion about this synonymy.

Specimens examined : Philippines : Los Ban4 os, P. I., on base

left … [illegible script label] anthicid, ‘no date ’, 3105,

W.H.W. (FH-3485 – holotype of D. formicomi) – Spain :

Barcelona : El Prat de Llobregat, on Formicomus pedestris

(Rossi), 4 Oct. 1986, S. Santamaria (BCB-SS0417a-e).

Dioicomyces ladoi Santam., sp. nov.(Figs 47–49, 85–88)

Etym. : ladoi – named for Carlos Lado, Spanish

mycologist.


brunneolus, parum arcuatus. Basalis cellula (I) receptaculi bis

vel pluries longior quam latior, longior quam tres cellulae

superiores coniunctae (collo excluso). Suprabasalis (II) et

tertia cellula similes, rectangulares in sectione, leviter com-

planatae. Antheridium cum ventre tam longo quam duae

cellulae inferiores coniunctae ; collum subterminale, rectum et

parum lateraliter inclinatum.


apicem, succineus-brunneus, sigmoideus ad parum arcuatum,

48–53 µm longus ab pede ad primariae appendicis apicem.

Primaria appendix conica, cum ³ rotundato et recto apice.

Cellulae I, II, III et primaria appendix atrobrunneae, subtiliter

punctatae, opacescentes in dorsalem marginem versum.

Perithecium (cum basalibus cellulis) 119–135¬54–61 µm,

asimetricum, ovoidale, latus circa tertiam partem basalem,

paulatim angustatum in breve, latum collum et rotundatum,

valde lateraliter inclinatum apicem; ostiolum ad perpen-

diculum cum longitudinali thalli axe. Cellula w%n

cum externa

pariete conspicue incrassata. Pedicellaris cellula (VI) perithecii

34–40 µm longa, crassa, inflata, leviter longior quam latior,

valde constricta circa basem, distaliter latior. Ascosporae

dimorphae, 46–48 µm longae (femineae), 34–36 µm longae

(mares).

Typus : Hispania centralis ; ‘Zamora: Revellinos de Campo ’

in lacubus ‘Villafa! fila ’, super Cyclodinus constrictus, 21 June

1999, S. Santamaria (BCB SS2261a – holotypus).


brownish, slightly arcuate. Basal cell (I) of receptacle

two or more times longer than broad, longer than the

three cells above combined (excluding the efferent

tube). Suprabasal (II) and third cells similar, rectangular

in section, slightly flattened. Antheridium with the

venter as long as the two cells below combined, the

efferent tube subterminal, straight, and slightly laterally

offset.


thecial tip, amber-brown, sigmoid to slightly arcuate,


S. Santamaria 629

Figs 60–72. Dioicomyces spp. Fig. 60. D. obliqueseptatus (FH3550). Broken female thallus ; arrow indicates the oblique

septum of an ascospore. Figs 61–62. D. onchophorus (FH3545). Female thalli showing the perithecial outgrowth (arrows).

Figs 63–64. D. proeminens (FH3511). Female thalli (that of Fig. 64 broken). Fig. 65. D. rostellatus (LPS45144). Female

thallus showing the perithecial outgrowth (arrow). Fig. 66. D. spiniger (BCB-SS851). Female thallus showing the

perithecial outgrowth (arrow). Fig. 67. D. subtorulosus (LPS45145). Female thallus showing perithecial corrugations

(arrows). Fig. 68. D. torulosus (LPS45146). Upper part of a broken female thallus showing perithecial corrugations

(arrows). Figs 69–70. D. trinitatis (FH3562). Male (Fig. 69) and female thallus (Fig. 70). Figs 71–72. D. umbonatus.

Female (Fig. 71: BCB-SS740b) and male thallus (Fig. 72: BCB-SS744a). Some perithecial basal cells (n« and VII) are

labelled as well as inflated cell w"n

« (arrow). Bars¯ 50 µm.

Primary appendage conical, with a rounded and straight


brown, delicately dotted, becoming opaque towards the

dorsal side. Perithecium (with basal cells)

119–135¬54–61 µm, asymmetric, ovoidal, broadest

near the basal third, gradually tapering towards a short,

broad neck and a rounded, strongly laterally offset

apex; the ostiole is perpendiculary oriented in relation

to the thallial longitudinal axis. The number of


¯ 5, wn¯ 5

(including a small distal cell) and 4, wn« ¯ 4. The cell

w%n

« shows a distinctively thickened outer wall. Stalk

cell (VI) of perithecium 34–40 µm long, stout, inflated,

slightly longer than broad, strongly constricted at the

base, distally broadened. Ascospores dimorphic,

46–48 µm long (females), 34–36 µm long (males).

At first glance, D. ladoi does not show very striking

characters, but careful observation and comparison

with other species reveals a very peculiar morphology.

The perithecial shape and its apex structure may be its

more distinctive traits (Figs 47 and 49, arrows),

combined with the stout cell VI. This species may be

compared with D. spiniger, if its perithecial outgrowth

is disregarded.


Specimens examined : Spain : Zamora : Revellinos de

Campo; Villafa! fila lagoons, Las Salinas lagoon, on Cyclodinus

constrictus (Curtis), 21 June 1999, S. Santamaria (BCB-

SS2261a – holotype; SS2261b, SS2261e – isotypes).

Dioicomyces leptalei Santam., sp. nov.(Figs 50–52, 89–92)

Etym. : leptalei – referring to the host genus name

Leptaleus.


brunneolus, sparsim et subtiliter punctatus, rectus. Basalis

cellula (I) receptaculi pallidior quam reliquus thallus, bis vel

pluries longior quam latior, leviter brevior quam tres cellulae

superiores coniunctae (collo excluso). Suprabasalis (II) et

tertia cellula similes, quadratae in sectione. Antheridium cum

ventre parum breviore quam duae cellulae inferiores coni-

unctae; collum subterminale, breve, rectum.


apicem, rubellus-brunneus, fere rectus, 42–45 µm longus ab

pede ad primariae appendicis apicem. Primaria appendix

conica, cum ³ obtuso et recto apice. Cellulae I, II, III et

primaria appendix atrobrunneae, subtiliter punctatae, opaces-

centes in dorsalem marginem versum. Perithecium (cum

basalibus cellulis) 88–127¬27–47 µm, fusiforme, latus circa

medium, paulatim angustatum in breve, latum collum et late

rotundatum et valde lateraliter inclinatum apicem. Spini-

formis prominentia, 37–46 µm longa, eminens sursum ab

supero quadrante marginis perithecii, cellulis w%m

formata.

Haec prominentia valde arcuata est, basaliter dilatata,

paulatim angustata in rotundatum sursum versum et ali-

quantum clavatum apicem. Pedicellaris cellula (VI) perithecii

27–34 µm longa, parum longior quam latior, basaliter

constricta, distaliter latior. Ascosporae dimorphae, 34–38 µm

longae (femineae), 26–31 µm longae (mares).

Typus : Hispania orientalis ; ‘Girona: Riells i Viabrea ’,

super Leptaleus rodriguesi, 9 Aug. 1987, S. Santamaria (BCB

SS0744b – holotypus).


brownish, sparsely and delicately dotted, straight. Basal

cell (I) of receptacle paler than the remainder of the

thallus, two or more times longer than broad, slightly

shorter than the three cells above combined (excluding

the efferent tube). Suprabasal (II) and third cells similar,

squarish in section. Antheridium with the venter slightly

shorter than the two cells below combined, the efferent

tube subterminal, short, and straight.


thecial tip, reddish brown, nearly straight, 42–45 µm


appendage conical, with a blunt and straight apex. Cells

I, II, III, and primary appendage dark brown, delicately

dotted, becoming opaque towards the dorsal side.

Perithecium (with basal cells) 88–127¬27–47 µm, fusi-

form, broadest near the middle, gradually tapering

towards a short, broad neck and a broadly rounded and

strongly laterally offset apex. A spine-like process,

37–46 µm long, projects upward from the upper quarter

of the side of the perithecium, formed by cell w%m

. This

outgrowth is strongly arcuated, basally broad at point

of origin, gradually tapering towards a rounded,

upwardly turned, and somewhat clavate tip, which

extends beyond the perithecial apex. The number of


¯ 5 (including

the outgrowth), wn¯ 5 (including a small distal cell)

and 4, wn« ¯ 4. Stalk cell (VI) of perithecium 27–34 µm

long, slightly longer than broad, constricted at the base,

distally broadened. Ascospores dimorphic, 34–38 µm

long (females), 26–31 µm long (males).

This species should be carefully compared with D.

spiniger. The material now referred to here as D. leptalei

was determined earlier as D. spiniger (Santamaria

1989), at a time when I had only collected thalli on

Leptaleus rodriguesi. Later, when additional material

was collected from other hosts and the type of D.

spiniger was studied, several invariable differences were

observed. The position and shape of the perithecial

outgrowths are clearly different : in D. leptalei the

outgrowths is formed from the upper quarter of the

perithecium or higher (Figs 51–52, 89–90), it extends

beyond the perithecial apex (Figs 51–52, 89–90), is

clavate at the tip (Fig. 51, arrow), shorter, and more

upwardly turned than the outgrowth of D. spiniger,

which is formed laterally at the middle of the

perithecium (Figs 66, 95–96), does not reach the

perithecial apex and is slender, more acute, and longer

than the outgrowth of D. leptalei. Moreover, both

species may be separated by differences in the shape of

the perithecial apex as well as by other minor characters,

such as the measurements, which are a bit smaller in D.

leptalei, and the reddish color of the female thallus in D.

leptalei.

Specimens examined : Spain : Barcelona : Cerdanyola del

Valle' s ; Bellaterra, on Leptaleus rodriguesi, 10 Apr. 1987, 14

Apr. 1987, S. Santamaria (BCB-SS0659, SS0661). Cerdanyola

del Valle' s ; Can Borrell damp, on L. rodriguesi, 14 Aug. 1989,

S. Santamaria (BCB-SS0867). Gualba; Gualba de baix,

Gualba stream, on L. rodriguesi, 20 June 1992, S. Santamaria

(BCB-SS1520). Gerona [Girona] : Riells i Viabrea; Ju! nior

Parc, on L. rodriguesi, 20 Apr. 1987, 2 Aug. 1987, 9 Aug.

1987, 2 Aug. 1992, 20 May 1995, S. Santamaria (BCB-

SS0674a-b, SS0741, SS0744b – holotype – , SS883d, SS1544,

SS1863b, d).

Dioicomyces malleolaris Thaxt., Proc. Amer. Acad. Arts48 : 170 (1912). (Fig. 53)

Type : Argentina : Llavallol (?), on Anthicus parvus, Apr.

1906, R. Thaxter 1513 (FH-3535 – lectotypus hic designatus)

[The label script referring to locality was illegible.]


brownish, ³ straight. Basal cell (I) of receptacle 1±5times longer than broad, longer than the suprabasal

cell. Suprabasal cell (II) slightly longer than broad,

rectangular in section. Third cell minute, flattened,

rectangular in section. Antheridium with the venter as

long as the two cells below combined, the efferent tube

terminal, curved, and slightly laterally offset.


thecial tip, yellowish brown, strongly arcuate, 39–46 µm

S. Santamaria 631

n!

73

82

74

n

m

w5n

II

I IIIpa

75

76

A B

77

VII

n!

n

w5n

78

7980

VII

w4n!

81

n!

VII

nm

n!

n!

VIIVIpa

IIIIII

83 84

8587

w5n

VII

n m 86

ts

m

VII

n!

w4m

n!

VII

n m

w5n

w4n

88

9189

90

92

98100

1021019997

969594

93

w4m

n!

m

VII

nn

m

w5n

t

t

n

VII

n!

w1n!

VI

t

n

m

Figs 73–102. Iberian species of Dioicomyces, with designation of some cells. Abbreviations not used in the text are : pa,

primary appendage; t, trichogyne; ts, trichogyne scar. Figs 73, 75, 78, 82, 86, 91, 94, 101 represent pairs of ascospores.

Figs 74, 77, 80, 81, 83, 85, 88, 89, 90, 95, 96, 102 represent mature female thalli. Figs 76, 79, 84, 87, 92, 93, 100 represent

male thalli. Figs 97–99 represent immature female thalli with trichogynes. Figs 73–77, 98. D. anthici (Figs 73–74: BCB-

SS1716a; Figs 75–77: BCB-SS1896d; Fig. 98: BCB-SS1997b). Figs 78–81. D. denticulatus (Figs 78–79: BCB-SS1058a; Fig.

80: BCB-SS1059; Fig. 81: BCB-SS1064a). Figs 82–84, 97. D. italicus (Figs 82–83: BCB-SS0417b; Fig. 84: BCB-SS0417a;

Fig. 97: BCB-SS0417c). Figs 85–88. D. ladoi (BCB-SS2261a). Figs 89–92. D. leptalei (Figs 89 and 92: BCB-SS0867; Fig.

90: BCB-SS0674b; Fig. 91: BCB-SS1520). Figs 93–96. D. spiniger (Fig. 93: BCB-SS0843c; Fig. 94: BCB-SS0843a; Figs

95–96: BCB-SS0843a). Figs 99–102. D. umbonatus (Figs 99, 101: BCB-SS1863a; Figs 100, 102: BCB-SS0744a). Bars¯50 µm (Bar B for Figs 74, 82–84, and 98–102: Bar A for all others).



appendage subconical, with a broadly rounded and

straight apex. Cells I, II, III, and primary appendage

slightly darker than the remainder of the thallus.


strongly asymmetric, ovoidal, with its main axis

perpendicular to the main longitudinal axis of the

thallus, the dorsal side strongly convex, strongly

protuberant basally, on the end opposite the rounded,

ostiolar tip, which is anteriorly directed. The number of


¯ 5 (including

a small distal cell), wn¯ 4 and 4, w

n« ¯ 4. Stalk cell (VI)

of perithecium 80–91 µm long, three or more times

longer than broad, with nearly parallel sides, arcuate,

slightly constricted at the base. Ascospores dimorphic,

34–39 µm long (females), 25–34 µm long (males).

This species was described by Thaxter (1912) on the

elytral tips of Anthicus parvus from Palermo and

Llavallol (Argentina). Later, Spegazzini (1917) added

several collections on the same host from Argentina.

This may be the most striking species of the genus

Dioicomyces, showing a very peculiar perithecial shape

and position on the axis of the thallus. This abnormal

position also influences the location and form of

perithecial outer wall and basal cells (Fig. 53).

Specimens examined : Argentina : Llavallol (?) [the label

script referring to locality was illegible], on Anthicus parvus

Apr. 1906,R.Thaxter 1513 (FH-3535 – lectotype). Temperley,

idem, Apr. 1906, R. Thaxter 1513 (FH-3536, FH-3537, FH-

3538 – paratypes).

Dioicomyces myrmecophilus T. Majewski, Acta Mycol.9 : 115 (1973). (Figs 54–56)

Type : Poland : Sado! wka, Nowy Dwo! r Mazowiecki county,

on Myrmecoxenus subterraneus Chevrolat (Coleoptera,

Colydiidae) in anthill of Formica rufa at edge of wood, 8 Jan.

1972 (TM-913 – holotype) [Majewski 1973; KRAM-F!].

Male thallus 54 µm long from foot to antheridial tip,

amber, straight. Basal cell (I) of receptacle two or more

times longer than broad, slightly longer than the three

cells above combined (excluding the efferent tube).

Suprabasal (II) and third cells similar, flattened,


long or longer as suprabasal and third cells below

combined, antheridial apex sharp pointed, the efferent

tube lateral, straight, laterally offset.


thecial tip (including the outgrowth) brownish, ³arcuate, 47–54 µm long from foot to apex of primary

appendage. Primary appendage conical, with a pointed

and inwardly curved apex. Cells I, II, III, and primary


same depth of color as the remainder of the thallus.

Perithecium (with basal cells and without outgrowth)

85–110¬40–49 µm, asymmetric, ovoidal, broadest be-

low the middle, gradually tapering towards a short,

undistinguished neck and a broad, blunt, and truncated

apex, which bears a finger-like, apically curved, some-

times uncinate outgrowth and a blunt, dark prominence

near the perithecial tip. The number of perithecial wall

cells for each row is : wm

¯ 5 (with short prominence),

wn¯ 4 and 5 (with finger-like outgrowth), w

n« ¯ 4.

Stalk cell (VI) of perithecium 16–23 µm long, slightly

longer than broad, distally broadened, constricted at

the base. Free ascospores not observed.

This species is only known from Polish localities

(Majewski 1973a, 1994). It represents a well-dis-

tinguished and characteristic species. The host was

originally called Myrmecoxenus subterraneus and was

reported to belong to the family Colydiidae of the

Coleoptera. According to Majewski (1994) the correct

name of the host is Myrmechixenus subterraneus and it

belongs to the Tenebrionidae. The host is a myrmeco-

philous beetle. I have studied four slides from

Majewski’s collection including the holotype and my

measurements are a bit greater than those published by

Majewski (1994).

Specimens examined : Poland : Sadowka pow. Novy Dwo! rM., on Myrmecoxenus subterraneus, 8 Jan. 1972, T. Majewski

(TM-913 – holotype; TM-908, TM-912, TM-914).

Dioicomyces notoxi Thaxt., Mem. Amer. Acad. Arts 16 :66 (1931) ; as ‘notosci ’, which was based on theincorrect spelling of the beetle genus Notoscus, nowNotoxus. (Figs 57–59)

Type : ‘On the elytra of Notoscus eximius Camp. No. 1596,

El Rancho, Guatemala (Kellerman) ’ [Thaxter 1931; FH!].


yellowish brown, straight to slightly arcuate. Basal cell

(I) of receptacle two or more times longer than broad,

as long or longer than the three cells above combined

(excluding the efferent tube). Suprabasal cell (II)

isodiametric, square in section. Third cell flattened,

rectangular in section. Antheridium with the venter


tube terminal to subterminal, ³ laterally offset.


thecial tip, yellowish brown, sigmoid to slightly arcuate,


Primary appendage subconical, with a rounded and

straight apex. Cells I, II, III, and primary appendage

darker than the remainder of the thallus. Perithecium

(with basal cells) 107–172¬32–52 µm, asymmetric,

fusiform, typically arcuate, with one side convex and

the other straighter, broadest below the middle,

gradually tapering towards a well-distinguished, narrow

neck and a thin, subacute apex. The number of


¯ 5 (including

a small distal cell), wn¯ 4 and 4, w

n« ¯ 4. Stalk cell (VI)

of perithecium 71–228 µm long, four to ten times longer

than broad, with nearly parallel sides, sigmoid to

slightly arcuate. Ascospores dimorphic, 32 µm long

(females), 27 µm long (males).

This species is only known from the type, described

by Thaxter (1931) on Notoxus eximius from Guatemala.

It is very similar to D. anthici. The slides studied include

short (Fig. 59) and long thalli (Fig. 57). The longer and

more slender thalli (Fig. 57) may be easily distinguished

S. Santamaria 633

from D. anthici by the long fusiform perithecia, with

narrow neck and almost pointed apex. The receptacle

(cells I, II, and III) is pale yellowish, whereas the

primary appendage is distinctively dark brown; this

character helps to separate this species from the wide

range of forms of D. anthici.

Specimens examined : Guatemala : El Rancho, on Notoxus

eximius Champ., Feb. 1908, R. Thaxter 1596 (FH-3540,

Female Type; FH-3541, Male Type; FH-3542).

Dioicomyces obliqueseptatus (Thaxt.) Thaxt., Proc.Amer. Acad. Arts 37 : 33 (1901). (Fig. 60)

Amorphomyces obliqueseptatus Thaxt., Proc. Amer.Acad. Arts 35 : 431 (1900).

Type : ‘On the antennae of an undetermined staphilinid,

British Museum, No. 398, Ega, Amazon River ’ [Thaxter

1900; FH!].

Male unknown.

Female thallus amber brown. Receptacle and primary

appendage unknown. Perithecium (with basal cells)

200¬54–57 µm, asymmetric, fusiform, strongly arcu-

ate, broadest near the base, gradually tapering towards

a short, broad neck and a broadly rounded apex. The

perithecial wall cells poorly defined and their number

no easy to determine in the material studied. Stalk cell

(VI) of perithecium broken. Ascospores dimorphic,

45 µm (females), 41 µm (males).

This species was described as Amorphomyces obli-

queseptatus on a staphylinid beetle allied to Myrmedonia

from Ega, Amazon River (Thaxter 1900). Later

(Thaxter 1901) the taxon was transferred to Dioi-

comyces. This new condition was assessed by Thaxter

(1908) who mentioned that this species is undoubtedly

a Dioicomyces because its obliquely septate spores (Fig.

60, arrow). At that time Thaxter did not realize that

Amorphomyces spores are once-septate ; he had not

noticed any septation. An oblique septum seems to be

exclusive to D. obliqueseptatus in the genus. Some

species of Amorphomyces (e.g. A. italicus) have asco-

spores with an oblique septum (Santamaria 2000).

The type slide borrowed included only two perithecia

each borne on a broken cell VI. This material seems

inadequate for description of a species, and although

the structure of the perithecial base indicates that this

thallus does not belongs to the genus Amorphomyces,

the exact identity of the genus is questionable. There

are two sizes among ascospores measured, so dioecy is

likely. Therefore, this species should be placed in

Dioicomyces, although with reservations.

Specimen examined : Amazon [see above], on Myrmedonia?,

R. Thaxter 398 (FH-3550 – holotype).

Dioicomyces onchophorus Thaxt., Proc. Amer. Acad.Arts 37 : 34 (1901). (Figs 61–62)

Type : ‘Usually on the basal half or at the base of the left

elytron of Anthicus floralis Linn. Fresh Pond, Cambridge ’

[Thaxter 1901; FH!].


yellowish, arcuate. Basal cell (I) of receptacle two or

more times longer than broad, slightly longer than the

three cells above combined (excluding the efferent

tube). Suprabasal (II) and third cells similar, flattened,


long as or shorter than suprabasal and third cells below

combined, the efferent tube subterminal, ³ laterally

offset.


thecial tip (including the outgrowth), brownish, strongly

arcuate, 43–54 µm long from foot to apex of primary

appendage. Primary appendage conical, with a pointed

and inwardly curved apex. Cells I, II, III, and primary

appendage dark brown. Perithecium (with basal cells

and outgrowth) 128–145¬41–54 µm, asymmetric, ovoi-

dal, broadest below the middle, gradually tapering

towards a short, undistinguished neck and a broad,

rounded apex, which bears a finger-like, erect out-

growth. The number of perithecial wall cells for each

row is : wm

¯ 5 (including outgrowth), wn¯ 4 and 5, w

n«

¯ 4. Stalk cell (VI) of perithecium 57–91 µm long, three

or more times longer than broad, distally broadened,

constricted at the base. Free ascospores not observed.

Only known from original description (Thaxter

1901), it was described on Anthicus floralis Linne! from

USA. This species is readily distinguished by the

terminal prominence of the perithecium (Figs 61–62,

arrows). Although at the time of its decription it was

the only species of the genus known to bear such a

perithecial outgrowth (as mentioned by Thaxter 1908) ;

many species are now known to have similar processes

(Table 1). This character is invariable among the 14

mature females included in the slides received from FH.

Specimens examined : USA : Fresh Pond, Cambridge, on

Anthicus floralis, Oct. 1900,R.Thaxter? (FH-3544 – holotype;

FH-3545, FH-3546, FH-3547, FH-3548).

Dioicomyces proeminens Thaxt., Mem. Amer. Acad.Arts 16 : 67 (1931) ; as ‘prominens ’. (Figs 63–64)

Type : Guatemala : Agua Caliente, on Anthicus sp., Feb.

1908, R. Thaxter 1583 (FH-3511 – lectotypus hic designatus).


yellowish brown, ³ straight. Basal cell (I) of receptacle

1±5 times longer than broad, as long as the two cells

above combined. Suprabasal (II) and third cells similar,

squarish in section. Antheridium with the venter longer

than the two cells below combined, the efferent tube

terminal or subterminal, ³ laterally offset.


thecial tip, brownish, ³ straight, 46–50 µm long from


conical, with a rounded and straight apex. Cells I, II,

III, and primary appendage dark brown. Perithecium

(with basal cells) 59–77¬29–39 µm, asymmetric, ovoi-

dal, broadest below the middle, gradually tapering

towards a short, undistinguished neck and a broadly


rounded, laterally offset apex. The number of perithecial



distal cell), wn¯ 5 (including a small distal cell) and

4(–5), wn« ¯ 4. Stalk cell (VI) of perithecium 71–121 µm

long, three or more times longer than broad, widening

toward the upper end, straight. Ascospores dimorphic,

37 µm long (females), 30 µm long (males).

This species is only known from the type described

by Thaxter (1931) on Anthicus sp. from Guatemala.

The two slides borrowed from FH include four mature

females, three mature males, and two broken female

thalli. On each slide label the name ‘anthici ’ was

written with soft pencil and later erased; therefore

Thaxter had questioned of the distinctiveness of this

species in relation to D. anthici. The identity of the host

is not written on the labels. Although it is closely

related to D. anthici, this species is clearly distinguished

by the small size and the shape of the perithecium which

appears rather reduced (Figs 63–64) ; by the elongated

and straight stalk cell, the broadened apex of which is

nearly as broad as the maximum breadth of the

perithecium. Also, the males are very small with an

almost terminal efferent tube.

Specimens examined : Guatemala : Agua Caliente, on ?,

Feb. 1908, R. Thaxter 1583 (FH-3511 – lectotype; FH-

3512 – isolectotype).

Dioicomyces rostellatus Speg., Anales Mus. Nac. Hist.Nat. Buenos Aires 29 : 529 (1917). (Fig. 65)

Type : ‘Al a! pice de las elitras del Anthicus floralis en La

Plata, Dic. 1915 ’ [Spegazzini 1917; LPS!].

Male unknown.

Female thallus 183 µm long from foot to perithecial

tip (including the outgrowth), dark brown, arcuate,

55 µm long from foot to apex of primary appendage.

Primary appendage narrowly oblong, with a pointed

and straight apex.Cells I, II, III, and primary appendage

dark brown, becoming opaque towards the dorsal side.

Perithecium (with basal cells and without outgrowth)

130 x 60 µm, asymmetric, broadly fusiform, with both

sides convex, broadest near the middle, abruptly

tapering towards a short, undistinguished neck and a

broadly rounded apex, which bears a finger-like,

outwardly directed outgrowth. The perithecial wall



perithecium 30 µm long, slightly longer than broad,

distally broadened, constricted at the base. Free


This species is only known from the original

description by Spegazzini (1917) on the elytral apices of

Anthicus floralis from Argentina. It is a well-dis-

tinguished species which is evidently related to D.

onchophorus. The unique slide received from LPS bears

a single female thallus (Fig. 65).

Specimen examined : Argentina : ? [only ‘L ’ in slide label

script], on Anthicus floralis, 23 Dec. 1915, C. Spegazzini 390-

1915 (LPS-45144 – holotype).

Dioicomyces spiniger Thaxt., Proc. Amer. Acad. Arts37 : 34 (1901) ; ‘as spinigerus ’. (Figs 66, 93–96)

Type : ‘On Anthicus floralis Linn., with the last two species,

more commonly on the inferior surface of the abdomen.

Fresh Pond, Cambridge ’ [Thaxter 1901; FH!].


pale brown, straight to slightly arcuate. Basal cell (I) of

receptacle two or more times longer than broad, longer

than the three cells above combined (excluding the

efferent tube). Suprabasal (II) and third cells similar,

usually flattened. Antheridium with the venter slightly


tube subterminal, long, and straight, ³ laterally offset.


thecial tip, brown, straight to slightly arcuate, 32–48 µm


appendage narrowly oblong, with a ³ pointed and

inwardly curved apex. Cells I, II, III, and primary

appendage dark brown, often delicately dotted, be-

coming opaque towards the dorsal side. Perithecium

(with basal cells) 121–148¬37–63 µm, asymmetric,

ovoidal, broadest near the basal third, gradually

tapering towards a short, broad neck and a rounded,

strongly laterally offset apex; the ostiole is perpendicu-

larly oriented in relation to the thallial longitudinal

axis. A spine-like process, 41–66 µm long, projects

upward from the middle of the perithecium, formed by

cell w%m

. This outgrowth is straight to slightly arcuated,

wide at the base, gradually tapering towards a rounded,

upwardly turned tip, which does not reach the

perithecial apex. The number of perithecial wall cells

for each row is : wm

¯ 4–5(–6) (including the out-

growth), wn¯ 5 (including a small distal cell) and 4, w

n«

¯ 4. The cell w%n

« shows a distinctively thickened outer

wall. Stalk cell (VI) of perithecium 36–43 µm long, 1±5times longer than broad, constricted at the base, distally

broadened. Ascospores slightly dimorphic, 34–43 µm

long (females), 32–38 µm long (males).

Described by Thaxter (1901) from Anthicus floralis

from the USA; Benjamin (1970: Fig. 3) illustrated one

female. The records from Spain on Leptaleus rodriguesi

by Santamaria (1989) belong in D. leptalei.

Specimens examined : Spain : Huesca : Aisa, on Anthicus

bifasciatus (Rossi), Aug.– Sep. 1986, J. Franc (BCB-SS0842a,

SS0843a, SS0843c-d, SS0851). – USA : Fresh Pond,

Cambridge, on Anthicus floralis, Oct. 1900 and 1902, R.

Thaxter? (FH-3552 – holotype; FH-3553, FH-3554, FH-

3555, FH-3556, FH-3557).

Dioicomyces subtorulosus Speg., Anales Mus. Nac. Hist.Nat. Buenos Aires 29 : 530 (1917). (Fig. 67)

Type : ‘Sobre el vientre del Anthicus decerptus en la isla

Santiago, La Plata, May 1916 ’ [Spegazzini 1917; LPS!].

Male unknown.


thecial tip, yellowish, sigmoid, 52–53 µm long from foot

to apex of primary appendage. Terminal cell (III) ³isodiametric.Primary appendagenearly cylindrical, with

S. Santamaria 635

a rounded and straight apex. Cells I, II, III, and

primary appendage yellowish like the remainder of

the thallus. Perithecium (with basal cells) 128–

134¬39–45 µm, asymmetric, fusiform, with one side

strongly convex and with some septal protrusions

near the base, the opposite side ³ straight, broadest

below the middle, gradually tapering towards a short

neck and a broadly rounded apex. The perithecial wall



perithecium 89 µm long, slender, strongly arcuate, four

or more times longer than broad, constricted at the

base, distally broadened. Free ascospores not observed.

Only known from the type described by Spegazzini

(1917) on the basis of specimens found on the ‘venter ’

of Anthicus decerptus collected in Argentina. It repre-

sents a well defined species showing characteristic

septal protrusions at the base of the dorsal side of the

perithecium (Fig. 67, arrows), a large, squarish cell III,

and a cylindrical, blunt appendage. The male thallus is

unknown. The unique slide received from LPS was in

rather poor condition and included only two females.

Specimens examined : Argentina : Isla Santiago, ‘venter ’

Anthicus decerptus, 1 May 1916, C. Spegazzini 461-1916

(LPS-45145 – holotype).

Dioicomyces torulosus Speg., Anales Mus. Nac. Hist.Nat. Buenos Aires 29 : 531 (1917). (Fig. 68)

Type : ‘Al a! pice de las elitras y del abdomen del Anthicus

decerptus en la isla Santiago, La Plata, May 1916 ’ [Spegazzini

1917; LPS!].

Male not observed.

Female thallus 307 µm long from base of cell VI to

perithecial tip, yellowish, arcuate. Receptacle and

primary appendage not observed in studied material.

Perithecium (with basal cells) 186¬43 µm, strongly

asymmetric and arcuate, fusiform, with both sides

marked by protrusions at septa between cells in vertical

outer wall cell rows; broadest above the middle,

abruptly tapering towards a short neck and a broad,

blunt, and truncate apex. The perithecial wall cells

poorly defined and their number no easy to determine

in the material studied. Stalk cell (VI) of perithecium

143 µm long, straight, seven or more times longer than

broad, with nearly parallel sides. Free ascospores not

observed.

Very closely allied to the previous species, it is only

known from the type described on the same host and

locality (and probably on the same individual) as D.

subtorulosus. I hesitated to synonymize these two species

because of the inadequate material in the borrowed

slide. The single slide received from LPS includes only

one broken female (Fig. 68), no males, and one mature

female thallus of D. subtorulosus. The perithecium

shows a rough surface with corrugations at almost all

the septa in the vertical rows of outer wall cells (Fig. 68,

arrows). Unfortunately the absence of the receptacle in

the single thallus examined prevented the observation

of a fundamental character, which is very distinctive in

D. subtorulosus : the shape of the appendage. The

drawing of Spegazzini (1917: nu! m. 74) of this character

is indeed very different from that of D. subtorulosus, but

in our experience the accuracy of Spegazzini drawings

is suspect. In the same way the male is misdrawn by

Spegazzini, according to Thaxter (1931).

Specimen examined : Argentina : Isla Santiago, ‘venter ’

Anthicus decerptus, 1 May 1916, C. Spegazzini 460-1916

(LPS-45146 – holotype).

Dioicomyces trinitatis Thaxt., Mem. Amer. Acad. Arts16 : 69 (1931). (Figs 69–70)

Type : Trinidad : Maraval Brook, on ‘Anthicidae ’, ‘no

date ’, R. Thaxter 2877 (FH-3562 – lectotypus hic designatus).


brownish, ³ straight. Basal cell (I) of receptacle 1±5times longer than broad, as long or slightly longer than

suprabasal cell. Second cell longer than third cell,

squarish in section. Third cell flattened, rectangular in

section. Antheridium with the venter shorter than the

two cells below combined, the efferent tube nearly

terminal, straight to curved, ³ laterally offset.


thecial tip, yellowish to dark-brown, sigmoid, 52–59 µm


appendage conical, with a broadly rounded and straight


brown, nearly blackened. Perithecium (with basal cells)

143–200¬40–70 µm, asymmetric, fusiform, sigmoid to

arcuate, broadest near the base, gradually tapering

towards a short neck and a thin and rounded apex. The

number of perithecial wall cells for each row is : wm

¯4, w

n¯ 5 (including a small distal cell) and 4, w

n« ¯ 4.

Stalk cell (VI) of perithecium 229–329 µm, four to eight

times longer than broad, with nearly parallel sides,

sigmoid. Ascospores dimorphic, 54 µm long (females),

39 µm long (males).

Dioicomyces trinitatis was only known from type on

Anthicus sp. from Trinidad (Thaxter 1931). According

to Thaxter this species is very similar to D. formicillae

‘of which it may be merely a form ’. I included D.

formicillae in D. anthici because of its wide variation

and the inconsistent characters (‘ large size, dark and

coarse-tipped perithecium, and the rather gradual

transition from the basal cell region to that of the

venter ’). Dioicomyces trinitatis may be separated from

D. anthici by the unusually elongated cell VI, by the

slender, curved perithecium and by blackening of the

receptacle and primary appendage (Fig. 70).

Specimens examined : Trinidad : Maraval Valley, on

‘Anthicidae ’, R. Thaxter 2877 (FH-3559, FH-3560 – para-

types). Maraval, Port of Spain, idem (FH-3561 – paratype).

Maraval Brook, idem (FH-3562 – lectotype).

Dioicomyces umbonatus Thaxt., Proc. Amer. Acad. Arts48 : 170 (1912). (Figs 71–72, 99–102)

Type : ‘At the base of the elytra near the inner margin of

several specimens of Anthicus parvus Pic. ; Temperley, No.

1513C ’ [Thaxter 1912; not seen].



brownish, straight to slightly arcuate. Basal cell (I) of

receptacle two or more times longer than broad, slightly

longer than the three cells above combined (excluding

the efferent tube). Suprabasal (II) and third cells similar,

³ isodiametric, squarish in section. Antheridium with

the venter shorter than the two cells below combined,

the efferent tube terminal, very short, and slightly

laterally offset.


thecial tip, brownish, sigmoid, 69–72 µm long from foot

to apex of primary appendage. Primary appendage

conical, with a rounded and inwardly curved apex.

Cells I, II, III, and primary appendage dark brown,

delicately dotted, becoming opaque towards the dorsal

side. Perithecium (with basal cells) 117–146¬67–91 µm,

asymmetric, ovoidal, with one side strongly convex,

broadest below the middle part, gradually tapering

towards a short neck and a broadly rounded and

laterally offset apex. Cell n« and contiguous upper wall

cell strongly inflated and protruding in the opposite

side of perithecium; especially remarkable is the wall

cell (w"n

«) which forms a prominent swell. The number

of perithecial wall cells for each row is : wm

¯ 5, wn¯

5 (including a small distal cell) and 4, wn« ¯ 4 (including

swell). Stalk cell (VI) of perithecium 67–83 µm long,

two or more times longer than broad, broadened

distally. Trichogyne with the upper cell bearing few

short slender appendages. Ascospores arcuate, dimor-

phic, 57–59 µm long (females), 42–44 µm long (males).

This species was described on Anthicus parvus from

Argentina (Thaxter 1912) and found there again on the

same host by Spegazzini (1917). Later, Santamaria

(1989) reported it from Spain on Leptaleus rodriguesi. It

is very well distinguished by the two inflated lower cells

of the ventral row of perithecial outer wall cells (Figs

71, 102) which seem correspond to the basal cell n« and

first wall cell of the row (w"n

«). The position of cell n«might seem to be too high, but lateral position of early

asci (Fig. 71) supports this idea, because of ascogenic

and n« cells are formed from cell VII.

Specimens examined : Spain : Barcelona : Mura; Nespres

stream, on Leptaleus rodriguesi, 23 July 1990, S. Santamaria

(BCB-SS0965d). Gerona [Girona] : Riells i Viabrea; Ju! nior

Parc, on L. rodriguesi, 2 Aug. 1987, 9 Aug. 1987, 2 Sep. 1989,

1 Oct. 1989, 2 Aug. 1992, 2 Aug. 1993, 20 May 1995, S.

Santamaria (BCB-SS0740b, SS0744a, SS0883c, SS0891,

SS1544, SS1716b, SS1863a).

DISCUSSION

The male in Dioicomyces species is very simple and in

all instances consists of four cells, the upper functioning

as a simple antheridium. The basal cell usually

represents nearly half of the total length of the male

thallus and develops from the lower cell of the original

spore; the remaining three cells, including the an-

theridium, develop from the short, upper cell of the

spore. The comparative length of cells is a good

character to separate some species. In most of the

species of the genus, the efferent tube of the antheridium

diverges slightly and is subterminal or else lateral (D.

myrmecophilus, Fig. 54). Only in few species this

discharge tube is strictly terminal (D. africanus, Fig. 25;

D. borneensis, Fig. 28; D. indentatus, Fig. 40; D.

italicus , Figs 43, 45, 84; D. notoxi, Fig. 58) or nearly so

(D. trinitatis, Fig. 69). The original spore apex is blunt

and does not persist as a spine-like process as in many

other genera of Laboulbeniales. Only in D. myrmeco-

philus (Fig. 54) is the antheridial apex sharp-pointed. In

most species there is a more or less evident dimorphism

in ascospores (Figs 73, 75, 78, 86, 91, 94, 101), those

that will develop males being smaller than those that

will develop into females ; only in D. italicus (Fig. 82) is

this dimorphism not evident. Often, the ascospores

germinate inside the perithecium, before release, each

showing the dark foot (Figs 1, 4, 5–6, 8, 12, 23, 52, 53).

Typically, males of Dioicomyces species have only one

antheridium, but sometimes a second one may be

present (D. africanus, Fig. 25, arrows; D. italicus, Fig.

45, arrows) as well a third antheridium (in D. anthici,

unpubl. obs.) ; it is not unusual for two antheridia to be

present, only one remaining functional (Benjamin 1970).

Thaxter described three additional genera: Dicran-

dromyces (Thaxter 1931), Triandromyces (Thaxter

1931), and Tetrandromyces (Thaxter 1912, 1931),

basically separated by the characters of the males,

which have, respectively, two, three or four functional

antheridia.

The receptacle of female thalli in Dioicomyces is also

simple, consisting in mature specimens of only three

cells (I, II, and III). Most species are very constant in

shapes of these cells ; cell III is usually small and

triangular in section or wedge-shaped in three

dimensions. Typically the dorsal side of the receptacle

appears darkened and sometimes finely dotted. The

appendage is always unicellular and typically conical,

with a more or less acute apex, although there are

exceptions showing a rounded (D. inclinatus, Fig. 39,

arrow) or else a sharp-pointed or spiny apex (D.

glossophorus). This appendage represents the undivided

upper cell of the original spore. The stalk cell (cell VI)

of perithecium is very variable in shape and size, and its

variability is often related with the position of growth

of thalli. Most of the characteristics separating the

species of Dioicomyces are based on the perithecium.

Species with perithecial outgrowths, prominences,

excrescences, swellings or other similar processes may

be easily separated, and such processes are invariably

observed in all thalli for every collection with the only

exception being D. borneensis (Figs 27, 29–30) which

was reported to have two different types of outgrowths

in different thalli. The number of perithecial wall cells

in each vertical row ranges from four to five (or else 6)

depending on the row. Their number seem to vary in

some degree and it is not uncommon to found variability

in this character among the thalli of the same species.

Tavares (1985) mentioned five cells in each vertical row

S. Santamaria 637

but also recognizes the possibility of a variation in this

point. Only the vertical row developed from n« shows

invariably four cells in all species (Table 1).

Benjamin (1995, 1998) makes extensive analyses of

dioecism in the order Laboulbeniales. Fourteen genera

of Laboulbeniales are strictly dioecious (Amorphomyces,

Apatomyces,Corylophomyces,Dimeromyces,Dimorpho-

myces, Dioicomyces, Herpomyces, Nycteromyces, Par-

vomyces, Picardella, Polyandromyces, Rhizopodomyces,

Tetrandromyces – incl. Dicrandromyces and Triandro-

myces – and Trenomyces), including only dioecious

species. Five genera (Aporomyces, Cryptandromyces,

Euphoriomyces, Laboulbenia, and Nanomyces) include

both dioecious and monoecious species. Only one

genus (Triceromyces) includes monoecious, dioecious,

and trioecious species (see Benjamin, 1998: Table 2

page 14). The subtribe Amorphomycetinae (tribe Laboul-

benieae) includes six genera according to the usage of

Tavares (1985) : Amorphomyces, Corylophomyces, Dioi-

comyces, Rhizopodomyces, Tetrandromyces (incl. Di-

crandromyces and Triandromyces), and Nanomyces. All

genera are strictly dioecious excluding Nanomyces that

has monoecious and dioecious species (Tavares 1985).

Only Amorphomyces and Rhizopodomyces have female

receptacles with only two cells, whereas in the other

genera of the subtribe they consist of three cells.

Nanomyces and Rhizopodomyces seem to be not closely

related to other genera (Benjamin 1995). Amorphomyces

seems also an isolated genus, whereas the three

remaining genera have manymorphological similarities.

I have studied also the types of the species of

Tetrandromyces, Dicrandromyces, and Triandromyces ;

these observations will be published separately but here

it can be said that unique differences between these

genera and Dioicomyces are found in features of the

male thalli. Although this may be a poor basis for

separating four genera, the male characteristics ob-

served are very consistent and they are supported by

ontogeny. In a recent work (Rossi & Santamaria 2000)

a new species of Tetrandromyces is described (T.

weirianus) and the synonymy of Tetrandromyces,

Dicrandromyces, and Triandromyces is questioned.

Nevertheless, it is evident that Dioicomyces, Tetrand-

romyces, Dicrandromyces and Triandromyces are very

closely related. Lastly, Corylophomyces (Benjamin

1995) is related to Dioicomyces in several characteristics

such as receptacle and trichogyne features (Figs 97–99)

but differs in the perithecial wall cell rows.

ACKNOWLEDGEMENTS

I express my gratitude to the curatorial staff of herbaria for the loan

of specimens: Donald H. Pfister, G. Cacavio, and E. W. Wood

(Farlow Herbarium, Harvard University, Cambridge, MA), A.

Arambarri, H. Spinedi, and J. Chayle (‘Instituto de Bota! nica C.

Spegazzini ’, La Plata, Argentina). Also, I wish to express my

gratitude to T. Majewski (Warsaw Agricultural University), Walter

Rossi (Universiti degli studi dell’ Aquila), and K. Sugiyama (Shizuoka

University) for the loan of some type slides ; to Isabelle I. Tavares

(University of California at Berkeley) and W. Rossi for critically

reading the manuscript ; to J. Fortes (Universitat Auto' noma de

Barcelona) for translating the Latin diagnoses ; to O. Escola' (Museu

de Zoologia de Barcelona), I. Izquierdo (Museo Nacional de Ciencias

Naturales, Madrid), and I. Ribera (Barcelona) for supplying

parasitized insects ; to G. Nardi (Italy) for determining the identity of

some Iberian anthicids. I am also indebted to C. Lado (Real Jardı!nBota! nico, Madrid) and Victor Jime!nez Rico (Universidad Com-

plutense, Madrid) for their company in the field. I also thank Ll. Sa! ez(Universitat Auto' noma de Barcelona, Bellaterra) for help with

nomenclatural problems. This work was supported by the DGES

project no. PB-98-0538-C04-04 (‘Flora Micolo! gica Ibe! rica IV ’).

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Corresponding Editor: D. L. Hawksworth

a taxonomic revision of the genus dioicomyces (laboulbeniales)

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