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On Fungi in the Zoopagaceae and Cochlonemataceae

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On Fungi in the Zoopagaceae and Cochlonemataceae 日菌報 52: 19－27. 2011 総 説 ゾウパーゲ科およびコクロネマ科菌類について 犀 川 政 稔 東京学芸大学環境科学分野，〒184‒8501 東京都小金井市貫井北町 4－1－1 On fungi in the Zoopagaceae and Cochlonemataceae Masatoshi SAIKAWA Department of Environmental Sciences, Tokyo Gakugei University, Nukuikita-machi, Koganei-shi, Tokyo 184－8501, Japan （Accepted for publication September 10 2010） Morphological characteristics of fungi in the Zoopagaceae and Cochlonemataceae are outlined with a key for 11 genera, 99 species and 5 varieties in the two families. Observation techniques of these fungi are also shown briefl y. （Japanese Journal of Mycology 52: 19－27, 2011） Key Words―amoeba; conidium; key; morphology; Zoopagales とシグモイデオマイセス科（Sigmoideomycetaceae）と， 緒 言 それにピプトセファリス科（Piptocephalidaceae）の 3 ゾウパーゲ科（Zoopagaceae）とコクロネマ科（Co- 科が追加されている．追加の理由はこれらの菌類がどれ chlonemataceae）は接合菌門（Zygomycota）のゾウパー も分節胞子嚢（merosporangium）を生じるためという ゲ亜門（Zoopagomycotina），ゾウパーゲ目（Zoopagales） のである（Benjamin, 1979）．なお，現在ゾウパーゲ科 に所属する菌類である（Hibbett et al., 2007）．現在まで は Acaulopage，Cystopage，Stylopage，Zoopage と Zooph- に命名，記載されたすべての種はアメーバやセンチュウ agus の 5 属を，また，コクロネマ科は Amoebophilus， などの微小動物に寄生する．ゾウパーゲ科の菌はいずれ Aplectosoma，Bdellospora，Cochlonema，Endocochlus と も捕食性で，培地上に伸びる栄養菌糸が動物を捕え，吸 Euryancale の 6 属を含んでいる（Kirk et al., 2008）．こ 器を侵入して栄養を吸い取る（Fig. 1）．それに対してコ の 2 科の 11 属を合わせるとこれまで 99 種と 5 変種が記 クロネマ科の各種は吸器を生じることはなく，動物の体 載されている．ここではこのゾウパーゲ科とコクロネマ 表に付着した分生子が発芽して体内に栄養菌糸を伸長す 科の観察法，分生子および接合胞子の形成と発芽につい る．すなわち内部寄生性である（Fig. 2）．しかし，分生 て述べ，最後に 99 種と５変種を識別するための検索表 子が動物体表に付着した後，栄養菌糸ではなく吸器を生 を示す． じる着生性の数種（Fig. 3）も現在コクロネマ科に含ま れている． Ⅰ ゾウパーゲ科およびコクロネマ科菌類の観察 目の名前の「zoo」と「pagus」はそれぞれ「動物」 と「食う」という意味をもつ．かつてはこの動物寄生性 微小動物に寄生するこれら 2 科の菌類は，自然界では の 2 科だけを収容するゾウパーゲ目が存在したが（Dud- コケやよく湿った腐葉土に生息する．そこで腐葉土など dington, 1973），最近広く認められているゾウパーゲ目 の 1 つまみを直径 9 cm のシャーレ内の水寒天培地（WA; はこの 2 科にヘリコセファリス科（Helicocephalidaceae） water agar）に載せ，湿度を保つためにシャーレ全体を ―19― 犀 川 ポリ袋などで包み，室内の直射日光の当たらないところ うな分節胞子嚢内にできる胞子嚢胞子ではなく真性の分 に 2～3 週間も放置すれば菌が WA 上に現れる．WA は 生子である．しかも多核であることが多い（Saikawa, 栄養分のない培地で，水道水に寒天粉末を 2% 入れ，煮 1986, fig. 10 ; Saikawa and Sato, 1991, fig. 10 ; Saikawa, てつくる．水道水培地（TWA; tap water agar）と呼ばれ 2011, fi gs. 5－8, 16, 17）． ゾウパーゲ科の分生子は発芽 ることもある．菌が出現したら次は培養である．培養と すると，発芽管はそのまま栄養菌糸となる（Fig. 1）． いっても菌だけを純粋培養できないので菌にアメーバな それに対して Euryancale を除くコクロネマ科では，分 どの微小動物を与えて培養する．だからこれらの菌類を 生子はまず宿主のアメーバまたは有殻アメーバの体表 培養するためには，まず微小動物を殖やすことが必要で （pellicle）に付着し，発芽して栄養菌糸を宿主の体内 ある．もし，WA にゾウパーゲ科またはコクロネマ科の に侵入させる（Fig. 2）．しかし，付着性の数種の分生子 菌が出現したら，その付近にいる未感染の動物がいる部 は付着後，栄養菌糸を生じる代わりに吸器を生じる（Fig. 分の寒天を 5 × 5 mm 程度の大きさに切り取って移植す 3）．なお，センチュウ内部寄生菌類の Euryancale では る．移植する増殖用の培地は SAA である（Saikawa and 分生子はセンチュウに飲み込まれ，体内に栄養菌糸を伸 Kadowaki, 2002）．ワムシの場合は SAA の平板上に培地 長する． とほぼ等量の蒸留水を入れ，5 × 5 mm 程度の大きさに 両科とも長紡錘形など長い形の分生子が多いが，発芽 切ったタマネギの表皮を２，３枚浮遊させる（Karling, 管は両端からではなく，ほとんどが一端または両端近く 1952）．1 週間もすればワムシは水底の寒天平板上や表 の分生子の側方に出る． 皮の直下で増えている． Ⅲ 接合胞子の形成と発芽 Ⅱ 分生子の形成と発芽 ゾウパーゲ科のAc. gomphoclada Drechsler と Zpa. ゾウパーゲ科とコクロネマ科の栄養胞子（無性胞子） cladosperma Drechsler，それにコクロネマ科の Am. dan- は原形質の分割によるのではなく，隔壁形成によって作 geardii Miura と Co. agamum Drechsler は単為生殖的に られる．よってそれは Benjamin（1979）が 主張するよ 接合胞子嚢を発生する．これら 4 種以外は 2 本の有性生 Figs. 1 － 3. ゾウパーゲ科およびコクロネマ科の形態を比較した模式図． Fig. 1, ゾウパーゲ科．培地上に伸びる栄養菌糸（VH）が動物（An） を捕え，吸器（H）を侵入して栄養を吸収する．分生子（Co）は発芽して栄養菌糸（VH）を生じる．Fig. 2, 内部寄生性の コクロネマ科．分生子（Co）が動物（An）の体表に付着した後，体内に栄養菌糸（VH）を伸ばす．吸器は生じない．Fig. 3, 着生性のコクロネマ科．分生子（Co）が動物（An）の体表に付着した後，吸器（H）を生じて栄養を吸収する．栄養菌糸 は生じない．CoG, 発芽した分生子 ; CP, 分生子柄 ; Z, 接合胞子． ―20― ゾウパーゲ科およびコクロネマ科菌類について 殖菌糸が別々の栄養菌糸に発生する．接合前，生殖菌糸 Peach, 1959; Park, 1971; Saikawa and Kadowaki, 2002）， が太くなり始めると太くなった部分の基部に隔壁が入 そのつのをもった分生子は水生不完全菌類の分生子とと り，隔壁よりも前方の太い部分は接合後の支持体となる． もに渓流で採取した水泡などからも確認される（Ingold, 2 本の有性生殖菌糸が多かれ少なかれ互いに絡み合って 1975）．同様に Ac. dichotoma Drechsler も水中からよく から接合することは 2 科に共通であるが，接合胞子嚢の 分離される菌である（Peach, 1948; Saikawa and Kadowa- 作られる場所はやや異なる．すなわち，ゾウパーゲ科の ki, 2002）． 接合胞子嚢は接合部付近の生殖菌糸がそのまま側方に偏 りながら膨大して作られるが，コクロネマ科のものは偏 Ⅴ ゾウパーゲ科およびコクロネマ科菌類の検索表 りが極端で，接合部付近の生殖菌糸に芽が生じ，その芽 が膨大して作られることが多い．接合胞子はいずれの場 今までに記載された 11 属 99 種と 5 変種を識別するた 合も接合胞子嚢の中にただ 1 つ作られ，しかも発芽する めの検索表を作成した．以下のとおりである．種名の次 まで胞子嚢壁の内側にある．そのため一般的には接合胞 の星印のついた数字は検索表に付記した文献番号を示し 子嚢の細胞壁も含めた全体を「接合胞子」と呼んでいる． ている．なお，2 種間の識別が困難なところには括弧内 発芽は Hirotani-Akabane and Saikawa（2010 ）が発見し にコメントした． た．それによると，ゾウパーゲ科のAc. lophospora Drechsler の接合胞子は 1 か月間の休眠後，突然 24 時 1 捕食性．栄養菌糸は宿主の外に伸びる 間前から原形質流動が激しくなって発芽し，その発芽管 ………………………………………… 2（ゾウパーゲ科） はそのまま栄養菌糸となって寒天培地上を伸び直ちにア 1 内部寄生性または付着性 ……… 6（コクロネマ科） メーバを捉えたという．一方，コクロネマ科のCo. 2 動物を捕捉するための捕食器官（捕食短枝）をもつ cerasphorum Drechsler と Co. megalosomum Drechsler の …………………………………………… 11（ Zoophagus） 場合には，接合胞子は長期間（それぞれ6 か月と 2 か月） 2 動物を捕捉するための特別な構造を生じない…… 3 の休眠後，発芽の 24 時間前にやはり急に原形質流動が 3 分生子ではなく球形の厚壁胞子を作る 活発となり，発芽管は分生子柄となって空気中に分生子 …………………………………………… 14（ Cystopage） 鎖を生じたという． 3 分生子を作る………………………………………… 4 4 分生子は鎖状に連なる………………… 20（ Zoopage） 4 分生子は鎖状に連ならない………………………… 5 Ⅳ ゾウパーゲ科およびコクロネマ科菌類の生態 5 分生子は無柄，または極めて短い柄の上に 1 つ作ら 最近 Zygomycota は Glomeromycota（ Hibbett, et al., れる…………………………………… 30（ Acaulopage） 2007）と呼ばれるようになったが，この接合菌類のうち， 5 分生子は立った分生子柄上に作られる ハルペラ目（Harpellales）とアセラリア目（Asellariales） …………………………………………… 58（ Stylopage） のほとんどの種は水生昆虫の幼虫の腸に着生しているの 6 付着性で，栄養菌糸を生じない．分生子は宿主の体 で，幼虫とともに水生といえるかもしれない．しかし， 表に着生する…………………………………………… 7 これら旧トリコミケス綱以外のもう１つの綱であった旧 6 内部寄生性で，分生子は発芽して体内に栄養菌糸を 接合菌綱は本質的に陸生で，水生の Aquamorteirella ele- 生じる…………………………………………………… 8 gans Embree & Indoh などは例外であった（Embree and 7 付着した分生子は大きく育ち，宿主内に吸器を伸ば Indoh, 1967）．ゾウパーゲ目のゾウパーゲ科とコクロネ す…………………………… Bdellospora helicoides（*1） マ科も最近までは陸生と考えられており，両科のほとん 7 付着した分生子は大きくならず，吸器も小さい どの属と種を命名，記載したDrechsler（1935 a, b, ……………………………………… 74（ Amoebophillus） 1936b, 1947a, 1959）も，論文中「陸生のアメーバ（terri- 8 分生子は鎖状に連なる……………………………… 9 colous amoebae）」という言葉をしばしば用いていた． 8 分生子は鎖状に連ならない…………………………10 しかし，少なくともゾウパーゲ科には Zph. insidians 9 栄養菌糸はねじれて伸び，とぐろを巻く Sommerstorff などもっぱら水中で生活している種を含 ………………………………………… 77（ Cochlonema） め水陸両生の種がかなり多いようである．たとえば Ac. 9 栄養菌糸は座布団形 tetraceros Drechsler は「陸生のアメーバ」に寄生する種 ………………………… Aplectosoma microsporum（*2） として記載されたが（Drechsler, 1935b），水に沈んだ落 10 分生子は横たわった分生子柄上に間隔をおいて 1 つ ち葉などからよく出現し（Tubaki, 1958; Jones and ずつ立つ……………………………… 95（ Endocochlus） ―21― 犀 川 10 分生子は横たわった分生子柄の分枝先端部にのみ作 20 宿主は有殻アメーバである…………………………21 られる………………………………… 98（ Euryancale） 20 宿主はアメーバである………………………………22 21 分生子は両端に原形質が抜けた突起をもつ Zoophagus Sommerst.（1911） 栄養菌糸は直線的に伸び， …………………………………… Zpa. toechospora（*13） 分枝は少ない．菌糸には短い捕食分枝が密生する． 21 分生子は原形質が抜けた突起をもたない ワムシやセンチュウを捕え，体内に吸器を伸ばす． ……………………………………… Zpa. tryphera（*14） 11 休眠胞子は長紡錘形で，鹿角状の捕食分枝をもつ 22 分生子は 5 個以上が空気中で鎖状に連なる………23 ……………………………………………Zph. cornus（*3） 22 分生子鎖は 4 個以下の分生子で構成される 11 休眠胞子は長紡錘形であるが，それが捕食分枝をも …………………………………… Zpa. tetraspora（*15） つことはない……………………………………………12 23 分生子の長いものは数回湾曲する 12 捕食分枝は先端に多数の触手のような分枝を持つ… ………………………………… Zpa. nematospora（*16） ……………………………………… Zph. tentaclum（*4） 23 分生子は湾曲しない…………………………………24 12 捕食分枝の先端に分枝を持たない…………………13 24 分生子の長いものは 150 µm を超える 13 捕食分枝は長円筒形である…… Zph. insidians（*5） …………………………………… Zpa. mitospora（*17） 13 捕食分枝の先端は球形をしたノブ状突起をもつ 24 分生子の長さは 10－60 µm の範囲である ………25 …………… Zph. pectosporus（*6） と Zph. tylopagus（*7） 25 分生子の発芽管は異様に太い （Zph. tylopagus はノブ状突起から出た粘液の塊が突起 …………………………………… Zpa. pachyblasta（*13） の一部とみなされて命名されたが，Zph. pectosporus 25 分生子の発芽管は細く，異様に太くなることはな と同一種である） ……………………………………………………………26 26 吸器の分枝は宿主内をカーブして伸びる Cystopage Drechsler（1941） 栄養菌糸は直線的に伸び， ………………………………… Zpa. thamnospira（*17） 分枝は少ない．菌糸のいたるところでセンチュウ， 26 吸器の分枝はカーブしない…………………………27 またはアメーバを捕え，体内に吸器を伸ばす．分生 27 吸器の分枝はすべて 5 µm 以上である ……………28 子の代わりに厚壁胞子を作る． 27 吸器の分枝は短く，長さは 2-6 µm である ………29 14 宿主はセンチュウである……………………………15 28 接合胞子は単為生殖的によって作られる 14 宿主はアメーバなどの原生動物である……………17 ………………………………… Zpa. cladosperma（*16） 15 厚壁胞子は培地上に伸びる菌糸の途中に作られる 28 接合胞子は単為生殖的によって作られない ……………………………………………………………16 ………………………………… Zpa. atractospora（*16） 15 厚壁胞子は培地上に伸びる菌糸の先端に作られる 29 分生子は円筒形である………… Zpa. virgispora（*13） …………………………………………Cy. cladospora（*8） 29 分生子は長紡錘形である………… Zpa. phanera（*1） 16 厚壁胞子は菌糸の中央に作られる ……………………………………… Cy. intercalaris（*9） Acaulopage Drechsler（1935） 栄養菌糸のいたるところ 16 厚壁胞子は細胞が側方に肥大して作られる でアメーバを捕え，吸器で栄養を吸収する．菌糸の ………………………………………… Cy. lateralis（*10） ところどころに分生子を 1 つずつ生じるが，分生子 17 厚壁胞子は菌糸の中央に作られる…………………18 柄はないか，極めて短い．なお，Ac. tigrina（ *18） 17 厚壁胞子は菌糸の側方に肥大して作られる………19 は菌類とは思えないので表からは除外した． 18 宿主は原生動物の Lecythium mutabile である 30 分生子は針形である…………………………………31 ………………………………………… Cy. subtilis（*10） 30 分生子は針形ではない………………………………39 18 宿主はアメーバである………… Cy. sacciformis（*11） 31 分生子本体の先端に突起がある……………………37 19 厚壁胞子は球形である……… Cy. sphaerospora（*12） 31 分生子本体の先端に突起はない……………………32 19 厚壁胞子は楕円形である…… Cy. ellipsospora（*12） 32 分生子は先細りでなく，いきなり丸くなっている ……………………………………………………………33 Zoopage Drechsler（1935） 栄養菌糸のいたるところでア 32 分生子は先細りである………………………………34 メーバ，または有殻アメーバを捕え，吸器で栄養を 33 分生子は基部に向かって細い……… Ac. retusa（*19） 吸収する．菌糸からは分生子柄が立ち上がり，分生 33 分生子は円筒形である………… Ac. baculispora（*20） 子鎖を形成する． 34 分生子の多くは長さが 45 µm 以下である ………35 ―22― ゾウパーゲ科およびコクロネマ科菌類について 34 分生子は長さが 46 µm 以上である ……………………………………… Ac. longicornis（*12） ……………………… Ac. ischnospora var. pleacra（*11） 50 分生子本体の長さは 16－24 µm である 35 分生子は上部が二又となることがある ……………………………………… Ac. tetraceros（*21） …………………………………… Ac. macrospora（*21） 50 分生子本体の長さは 25－41 µm である 35 分生子の上部が二又となることはない……………36 …………………………… Ac. tetraceros var. longa（*25） 36 分生子は長さが 25－45 µm で接合胞子は 5－7 µm 51 分生子は倒卵形である………… Ac. lophospora（*26） である………………………… Ac. rhaphidospora（*21） 51 分生子は球形または，かぶら形である……………52 36 分生子は長さが 25－60 µm で接合胞子は 6－8 µm 52 分生子はしばしば１～数個が密着し，融合する である……………………………… Ac. stenospora（*10） …………………………………… Ac. dactylophora（*12） （両種の識別は困難である． Ac. stenospora は Ac. 52 分生子密着，融合することはない…………………53 rhaphidospora とすべきである） 53 分生子の突起は 18 本以下である …………………54 37 分生子の長さは 55 µm 以下である ………………38 53 分生子の突起は 20 本以上である …………………55 37 分生子の長さは 56 µm 以上である 54 分 生 子 は 10－12 × 9－10.5 µm， 突 起 は 3－6 × …………………………………… Ac. ischnospora（*22） 1－2 µm である ………………… Ac. acanthospora（*17） 38 接合胞子は単為生殖によって作られる 54 分生子は 12－19 × 8－15 µm，突起は 5－8 × 1.5－2 …………………………………… Ac. gomphoclada（*23） µm である ………… Ac. acanthospora var. magna（*27） 38 接合胞子は単為生殖によって作られない
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