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Key to families of Zygomycota

by Paul Kirk and Jerry Benny

1

Saprobic or, if parasitic or predacious, having mycelium immersed in the host tissue

(Zygomycetes) 2

 

Associated with arthropods and attached to the cuticle or digestive tract by a holdfast and not immersed in the host tissue

(Trichomycetes) 39

2 (1)

Mycelium ± regularly septate, septa with plugs; zygospores, where known, formed on undifferentiated hyphae

3

 

Mycelium, septa and zygospores not as above

4

3 (2)

Merosporangia bispored, septal plugs with polar protuberance, dissolving in 2-3% KOH

(Dimargaritales) Dimargaritaceae

 

Merosporangia unispored, septal plugs without polar protuberance, not dissolving in KOH

(Kickxellales) Kickxellaceae

4 (2)

Sporangia or 'conidia' forcibly released, or if not forcibly released then non-haustorial parasites of cicadas or nematodes with the latter producing two or more pedicellate, globose, spinose 'conidia' terminally and laterally from a coenocytic, erect pedicel

5

 

Sporangia not forcibly released; if non-haustorial parasites of nematodes spores formed in chains or if solitary, not both globose and spinose; fertile hyphae often septate at maturity

6

5 (4)

Sporangiophore arising from trophocyst; sporangia multispored, thallus mycelial

(Mucorales, p.p.) 11

 

Sporangiophore not arising from trophocyst; 'conidia' few- or unispored; thallus mycelial or consisting of hyphal bodies or protoplast-like cells

(Entomophthorales, p.p.) 28

6 (4)

Spores formed in sporocarps produced in soil or on organic material above ground, saprobes or ecto- or endomycorrhizal; or spores formed singly in soil, endomycorrhizal and forming arbuscules in roots of phycobiont

7

 

Spores not formed as above; saprobes or parasites

10

7 (6)

Sporocarps formed

8

 

Sporocarps not formed, spores formed singly in soil

(Glomales, p.p.) 33

8 (7)

Sporocarps containing only multispored sporangia

(Mucorales, p.p.) 11

 

Sporocarps containing zygospores, azygospores or chlamydospores

9

9 (8)

Only zygospores present in sporocarp; saprobes or ectomycorrhizal

(Endogonales) Endogonaceae

 

Sporocarps containing azygospores or chlamydospores; endomycorrhizal

(Glomales, p.p.) 33

10 (6)

Saprobes or facultative parasites; sporangiospores formed in sporangia, sporangiola, or less commonly merosporangia; arthrospores, chlamydospores and/or yeast-like cells sometimes present

(Mucorales, p.p.) 11

 

Obligate haustorial or non-haustorial parasites of fungi or small animals or their eggs; sporangiospores formed in multispored or unispored merosporangia; chlamydospores sometimes formed

(Zoopagales) 35

11 (5)

Sporangia ± lageniform, multispored; sporangiospores hyaline, smooth-walled, without appendages

12

 

Sporangia usually globose to obpyriform, never lageniform

13

12 (11)

Sporangia formed singly or in pairs; rhizoides present

Saksenaeaceae

 

Sporangia borne verticellately; rhizoids absent

Mucoraceae (p.p.)

13 (11)

Sporangia formed in a sporocarp

Mortierellaceae (p.p.)

 

Sporangia not formed in a sporocarp

14

14 (13)

Uni- or multispored merosporangia produced on a fertile vesicle

Syncephalastraceae

 

Merosporangia not produced; sporangia and/or sporangiola present

15

15 (14)

Sporangiospores fusiform to broadly fusiform, often with hyaline, hair-like, polar appendages; sporangia non-apophysate, wall persistent, fracturing regularly into 2 or irregularly into 3-4 segments by a preformed suture

16

 

Sporangiospores not with the above combination of characters, usually smooth-walled, rarely ornamented or appendaged; sporangium wall deliquescent' if persistent fracturing irregularly; sporangia sometimes absent and only sporangia produced

17

16 (15)

Sporangiospore wall pale brown to reddish-brown; sporangia only or sporangia and sporangiola (on separate sporangiophores) produced; orange pigment (carotene) usually produced in mycelium on rich media; zygosporangium smooth, thin, hyaline; zygospore reddish-brown, wall striate; suspensors apposed

Choanephoraceae (p.p.)

 

Sporangiospore wall hyaline; sporangia only produced; mycelium hyaline; zygosporangium verruculose, blackish-brown; suspensors opposed

Gilbertellaceae

17 (15)

Sporangia uni- to multi-spored, acolumellate (columella-like structure septum-like or dome-shaped, never protruding into the sporangium); mycelium usually extremely fine

Mortierellaceae

 

Not with the above combination of characters; sporangia columellate; sporangiola columellate or acolumellate, sometimes arising from a vesicle; mycelium more robust

18

18 (17)

Sporangia never formed; sporangiola uni- or multispored, pedicellate, arising from a fertile vesicle

19

 

Sporangia formed; sometimes sporangiola also present

24

19 (18)

Fertile vesicles produced from primary vesicles present on uniseptate sporangiophore branch; uni- or multi-spored pedicellate or denticulate, acolumellate sporangiola present; zygospores where known smooth-walled, suspensors with appendages

Radiomycetaceae

 

Not with the above combination of characters; fertile vesicles produced terminally on the sporophore or its branches or on a lateral vesicle; sporangiola columellate or acolumellate; sterile spines sometimes present

20

20 (19)

Sporangiola columellate, uni- or multi-spored; saprophytic or gall-forming mycoparasites; sporophore sometimes bearing sterile spines

21

 

Sporangiola acolumellate, uni- or multi-spored; sterile spines on sporophore absent

22

21 (20)

Sporangiola uni-spored; sporophore bearing sterile spines

Chaetocladiaceae

 

Sporangiola multi-spored; sterile spines on sporophore absent

Thamnidiaceae (p.p.)

22 (20)

Sporangiospore wall pale brown to reddish-brown, striate; zygospores with apposed suspensors

Choanephoraceae (p.p.)

 

Sporangiospore wall hyaline, smooth or spinose; zygospore with opposed suspensors

23

23 (22)

Sporophore mostly branched; fertile vesicles terminal and lateral; sporangiola uni-spored, pedicel monomorphic, sporangiospores released by random fracture of the pedicel and/or dissolution of the sporangiolar wall; yeast-like cells absent

Cunninghamellaceae

 

Sporophore simple or rarely once or twice branched; fertile vesicles terminal; sporangiola uni- or multispored, pedicel mono- or dimorphic, sporangiospores released by fracture of the pedicel at preformed cicumscissile zone, sporangiolum wall persistent, separable from the sporangiospore wall; yeast-like cells produced after spore germination on rich media

Mycotyphaceae

24 (18)

Deliquescent sporangia and persistent-walled sporangiola present

Thamnidiaceae (p.p.)

 

Only sporangia produced, either with persistent or deliquescent wall

25

25 (24)

Sporangia ± apophysate

Mucoraceae (p.p.)

 

Sporangia non-apophysate; sporophore sometimes vesiculate or constricted below the sporangium

26

26 (25)

Sporangium wall blackish-brown to black, persistent; trophocyst sometimes produced; subsporangial vesicle sometimes produced, globose or broadly ellipsoid; sporangium liberated whole by collapse of the subsporangial vesicle or by cicumscissile rupture of the junction between the sporangium wall and the sporophore, or sometimes forcibly discharged; zygospore usually yellowish, suspensors apposed, zygosporangium thin, hyaline

Pilobolaceae

 

Sporangium liberation not as described above; subsporangial vesicles absent or small and hemispherical and then with a deliquescent sporangium; zygospore suspensors opposed or tongs-like

27

27 (26)

Sporophore unbranched, usually bluish-green; zygospore suspensors tongs-like, with branched appendages

Phycomycetaceae

 

Sporophore branched or unbranched, never bluish-green; zygospore suspensors opposed, without appendages

Mucoraceae (p.p.)

28 (5)

Nuclei large, >10 µm long, not staining with aceto-orcein, with a prominent nucleolus; sporophores with a vesicle subtending the spore ('conidium'); gametangia, if present, with small, beak-like accessory cells; contents of secondary spores and somatic hyphae in some species may divide ad produce small protoplast-like cells (appearing multispored); all cells uninucleate

Basidiobolaceae

 

Nuclei small, < 10 µm long, aceto-orcein nuclear staining and nucleolus visibility variable; sporophores without a subtending vesicle; gametangia, if present, without accessory cells; spores and mycelium not producing internal protoplast-like cells; cells not all uninucleate

29

29 (28)

Nuclei variable, 2-15 µm long during interphase, with much heterochromatin (staining readily in aceto-orcein and bismark brown); nucleolus not prominent; nucleus remaining visible during mitosis

Entomophthoraceae

 

Not as above

30

30 (29)

Nuclei relatively large (> 5 µm long), nucleoplasm appearing granular (chromatin condensed at interphase); sporophores short and unbranched, arising from sporogenous cells; spores forcibly discharged by papillar eversion; obligate intracellular parasites of fern gametophytes

Completoriaceae

 

Nuclei 2-5 µm long, with little heterochromatin (not staining in aceto-orcein or bismark brown), usually inconspicuous during mitosis; nucleolus prominent; spores discharged by papillar eversion, fluid discharge, or passively released; saprobes or parasites of insects, freshwater algae or soil invertebrates

31

31 (30)

Sporophores upright, simple, each bearing several spores; parasitic on soil invertebrates (nematodes and tardigrades)

Meristacraceae

 

Sporophores usually unbranched, one spore borne terminally; saprobes or parasites of insects, soil invertebrates, freshwater algae or causing mycotic infections of animals

32

32 (31)

Spores (asexual spores and zygospores) tending to be melanized (pale grey to black); zygospores budding from conjugation point of two hyphal bodies; obligate parasites of mites and insects (esp. Homoptera)

Neozygitaceae

 

Spores hyaline; zygospores formed in axis of parential hyphae as a result of conjugation between adjacent cells or scalariform conjugations between two hyphae; saprobes or obligate parasites of desmid algae, soil invertebrates, or facultative mycotic disease agents of animals

Ancylistaceae

33 (7)

Only arbuscules formed in mycorrhizal roots; azygospore-like bodies formed at the apex of sporogenous cells of fertile hyphae; auxiliary cells present

Gigasporaceae

 

Arbuscules and vesicles formed in mycorrhzal roots; chlamydospores formed terminally and laterally on or within fertile hyphae; auxiliary cells absent

34

34 (33)

Chlamydospores formed apically

Glomaceae

 

Chlamydospores formed from within the neck of a sporiferous saccule

Acaulosporaceae

35 (10)

Spores solitary, short pedicellate, formed on the surface of a vesicle or sessile and formed in pairs or chains from the coiled apex of a sporophore; spores of some taxa large, brown

36

 

Spores solitary (unispored) or in chains (multispored) and borne in merosporangia, long or short pedicellate, or sessile, or arising from a head-cell or a secondary branchlet, if arising from a vesicle usually multispored; unispored merosporangia not large and brown; sporophore apex not coiled

37

36 (35)

Fertile hyphae several times dichotomously branched, septate, terminating in sterile spines; fertile vesicles pedicellate, arising in pairs from the cell at the branching point of the fertile hyphae; spores ± hyaline, relatively small; parasites of fungi

Sigmoideomycetaceae

 

Fertile hyphae simple, coenocytic, without sterile spines; sporophore apex forming a fertile vesicle or not swollen and coiled; spores brown, large; parasites of microfauna or their eggs

Helicocephalidaceae

37 (35)

Haustorial mycoparasites, especially of Mucorales

Piptocephalidaceae

 

Haustorial ectoparasites or non-haustorial endoparasites of soil microfauna

38

38 (37)

Predacious, with adhesive material on hyphae to trap prey; vegetative mycelium formed outside host

Zoopagaceae

 

Ecto- or endoparasitic; vegetative mycelium within host, only sporophores formed outside host

Cochlonemataceae

39 (1)

Spores (trichospores) produced exogenously, usually bearing one or more basal appendages; zygospores produced in most genera

(Harpellales) 40

 

Spores (sporangiospores) or amoeboid cells produced endogenously, or thallus breaking up into arthrospores; zygospores not present

41

40 (39)

Thallus unbranched, usually attached to the peritrophic membrane of the midgut (less common to the hindgut cuticle)

Harpellaceae

 

Thallus branched, attached to the hindgut cuticle

Legeriomycetaceae

41 (39)

Thallus branched and septate, producing arthrospores

(Asellariales) Asellariaceae

 

Thallus unbranched, or branched only at the base; non-septate; producing sporangiospores or amoeboid cells

42

42 (41)

No amoeboid cells; sporangiospores usually produced singly in basipetal series of terminal sporangia

(Eccrinales) 43

 

Amoeboid cells produced at some stage; entire thallus functioning as sporangium, releasing spores or amoeboid cells more or less simultaneously

(Amoebidiales) Amoebidiaceae

43 (42)

Thalli producing directly only one type of spore; sporangia sometimes germinating in situ

Palavasciaceae

 

Thalli usually producing at least two types of spores; sporangia not germinating in situ

44

44 (43)

Primary infestation spores produced in thalli that become converted entirely or partly into multispored sporangia

Parataeniellaceae

 

Primary infestation spores produced singly in series of terminal sporangia

Eccrinaceae