01_総説_入江 color.pdf

Size: px

Start display at page:

Download "01_総説_入江 color.pdf"

きのこけいれい
3 years ago
Views:

1 宮地賞受賞総説 1 * Studies of marine gastropods connecting natural history and evolutionary ecology 1. Intraspecific variation of shell characters and morphological taxonomy Takahiro Irie* Atmosphere and Ocean Research Institute, The University of Tokyo 1930 Monetaria 1930 Ernst Mayr Littorina Nucella G E Abstract: Within-species phenotypic variation has long attracted the attention of ecologists, taxonomists, geneticists, and evolutionary biologists. This article focuses on marine gastropods and reviews how scientists have addressed the problem of intraspecific variation in shell morphology since the 1930s. The molluscan classification at the species level relied mainly on conchological traits, even after important biological ideas were advocated such as the biological species concept, New Systematics, population genetics, and modern evolutionary synthesis. The first half of this article briefly explains the ontogeny and ecology of Monetaria species to examine the conceptual background of their inter- and intraspecific classification developed in Germany in the 1930s. This is to emphasize how phenotypically plastic traits are not desirable as taxonomic traits under the biological species concept. The last half summarizes studies primarily by North American and Scandinavian scientists of the relationship between morphological variation and inducible defense against predators, i.e., crabs. Their systems with Littorina and Nucella * irie@aori.u-tokyo.ac.jp 1

2 are an excellent example of how rearing experiments using marine gastropods successfully contributed to progress in ecology. A lesson from history is that to draw robust, appropriate conclusions from the observed phenotypic variation in both taxonomy and evolutionary ecology, it is critical to quantify the genetic and environmental components separately. Keywords: intraspecific taxonomy, ecotype, ecophenotype, phenotypic plasticity, inducible defense domestication Publius Vergilius Maro Gregor Mendel Monetaria genus Littorina genus Nucella 245 Moretzsohn 2014 Monetaria Meyer 2003 M. annulus M. moneta M. caputserpentis M. caputdraconis

自然史と進化生態学をつなぐ海産腹足類の研究 1 図 1 Monetaria 属各種の殻背面 a 沖縄 b 西サモア産のハナビラダカラ c ツアモツ諸島産のビャクレンダカラ d, f ツアモツ e ニューブリテン島産のキイロダカラ g セイシェル諸島 h 荒崎 i 沖縄 j ダンピア西オーストラリア k ロードハウ島 l パゴパゴアメリカ領サモア m オアフ島産のハナマルユキ n, o

annulus obvelata from Tuamotu, French Polynesia. M.

3 自然史と進化生態学をつなぐ海産腹足類の研究 1 図 1 Monetaria 属各種の殻背面 a 沖縄 b 西サモア産のハナビラダカラ c ツアモツ諸島産のビャクレンダカラ d, f ツアモツ e ニューブリテン島産のキイロダカラ g セイシェル諸島 h 荒崎 i 沖縄 j ダンピア西オーストラリア k ロードハウ島 l パゴパゴアメリカ領サモア m オアフ島産のハナマルユキ n, o イースター島産のリュウズダカラ個体群内での貝殻変異も著しいためこれらの標本はその産地で見られる個体の形態学的特徴を必ずしも代表するわけではない Figure 1. Dorsal views of Monetaria species. M. annulus from (a) Okinawa, Japan and (b) Western Samoa. (c) M. annulus obvelata from Tuamotu, French Polynesia. M. moneta from (d, f) Tuamotu, French Polynesia and (e) New Britain Island, Papua New Guinea. M. caputserpentis from (g) Seychelles, (h) Arasaki, Japan, (i) Okinawa, Japan, (j) Dampier, Western Australia, (k) Lord Howe Island, Australia, (l) Pago Pago, American Samoa, and (m) Oahu, Hawaii. (n, o) M. caputdraconis from Easter Island. These specimens do not necessarily represent the morphological characteristics of their localities, because variation within populations is also considerable. 図 2 ハナビラダカラの個体発生 a 母貝によって二枚貝死殻の窪みに生み付けられた卵塊 b 孵化直後のベリジャー幼生 c 孵化後約 1 ヶ月後のベリジャー幼生 d 着底後の貝殻成長 Figure 2. Ontogeny of Monetaria annulus. (a) An egg mass laid by a female. (b) A freshly hatched veliger. (c) A veliger ca. 1 month after hatching. (d) Shell ontogeny after the settlement. しているフランス領ポリネシア周辺に分布するビャクにベリジャー幼生と呼ばれる形態でプランクトン生活をレンダカラもまたかつては Monetaria 属の構成種であ送る時期を経験する図 2b, c 幼生期の長さを厳密にったが M. obvelata 現在はハナビラダカラの亜種とさ特定した研究はこれまで一例もないが孵化時の貝殻サれている M. annulus obvelata イズ殻幅約 100 μ m と幼生期の終了時点での貝殻サタカラガイ科には幼生期を持たない直達発生の種もいイズ殻幅約 1 mm から軟体動物の他の種の情報にるが Monetaria 属を含む浅海の種は卵から孵化した後基づいて推察するとどんなに短くとも 1 ヶ月はかかる 3

入江貴博きくなることはなくまた軟体部の成長も起こらなくなるこれは完全変態の昆虫において羽化後は体サイズが増大しないことと同様の現象でそのような成長パターンは決定成長 determinate growth と呼ばれている海洋生物には成熟後も体サイズの増大が続く非決定成長 indeterminate growth の分類群が多いためタカラガ

4 入江貴博きくなることはなくまた軟体部の成長も起こらなくなるこれは完全変態の昆虫において羽化後は体サイズが増大しないことと同様の現象でそのような成長パターンは決定成長 determinate growth と呼ばれている海洋生物には成熟後も体サイズの増大が続く非決定成長 indeterminate growth の分類群が多いためタカラガイは海洋生物の中では少数派であると言える他にも後述するスイショウガイ科の腹足類は明らかな決定成長者であるまた最近の飼育実験でカルス形成期が終わっていったん成貝期に入ると殻の厚化は二度と起こらないことが確認された Irie and Morimoto 2016 タカラガイの個体発生種内分類貝殻種内変異については本稿で詳しく取り扱うがそれ以外の情報食性行動繁殖解剖学的特徴種より上位の分類を概説した記事は数年に一度の頻度で出版されているまず C. M. Burgess によって 1985 年に出版されたタカラガイの図 3 ハナビラダカラの生体貝殻が外套膜で覆われている様子 a 前方から撮影した幼貝 b 背面から撮影した幼貝 c 前方から撮影した成貝 d 背面から撮影した成貝 Figure 3. Living Monetaria annulus. Shells are covered by mantles. (a) Anterior and (b) dorsal views of juvenile individuals. (c) Anterior and (d) dorsal views of adult individuals. モノグラフにはハワイの貝類学者 E. Alison Kay によって執筆された本分類群の生物学に関する概論が 20 ページにわたって掲載されている Kay 1985 ドイツの Felix Lorenz Jr. と Alex Hubert によって 1993 年に出版され 2000 年に改訂されたモノグラフには軟体部の解剖学的特徴の概説が記されている Lorenz and Hubert 1993, 2000 豪州の多数の研究者によって執筆され 1998 年と考えられるベリジャー幼生が変態を経て海底に着底に出版された軟体動物学の大型専門書 The Southern すると底棲生活が始まる Synthesis には Barry Wilson によるタカラガイ科に関すいわゆる巻貝の殻として一般にイメージされるサザエる概説がある Wilson 1998 また Wilson は Peter やタニシといった貝類の殻は空間的な自己相似形を維 Clarkson との共著でオーストラリアの潮下帯に生息する持するような螺旋状の貝殻成長によって作られる本稿 Zoila 属と Umbilia 属のタカラガイに関するモノグラフをの後半で紹介するタマキビやチヂミボラの作る殻も同出版しておりここでも生物としてのタカラガイに関す様これに対してタカラガイの成貝が有する殻は自己る情報が豊富に記載されている Wilson and Clarkson 相似形とは完全にかけ離れているがこれは底棲生活が 2004 いわゆる生態的特徴を代表する食性について簡単開始した後の貝殻成長が次に示す 3 段階から構成されるに紹介するとタカラガイは種によって肉食藻食雑ためである図 2d まず幼貝期 juvenile stage には食に分かれ肉食の中でも別の貝類を捕食する種もいれ薄い殻が螺旋状に形成され主に軟体部が拡大されるば藻類を削って食べるのと同様の方式でカイメンやコ軟体部の増大が停止するとカルス形成期 callus-building ケムシを捕食する種もいる本稿で取り上げる Monetaria stage が始まり幼殻の主に外側にカルスとよばれる厚属の各種は主に藻類を好んで食べているとする研究がい殻が形成される幼貝期に見られる螺旋方向の殻の伸多い Kay 1960 Renaud 1976 Osorio et al 長やカルス形成期に見られる殻の厚化は過去に作られた殻の表面に炭酸カルシウムが結晶化することで進行種内分類の歴史するその際には石灰化の起こる箇所が軟体部の一部外套膜によって覆われる必要があるとされる図 3 最タカラガイの種内分類に初めて本格的に取り組んだの後にカルス形成期が終了すると成貝期に入り繁殖をはドイツの貝類学者 Franz Alfred Schilder と Maria Schilder である Schilder 夫妻は大航海時代に蒐集され行う reproductive stage 西欧の博物館に所蔵されていた世界各地の標本を比較し幼貝期が終了するとそれ以後は殻の内側の空間が大 4

5 1 Schilder 1927 Schilder and Schilder Schilder and Schilder Schilder 400 Monetaria Carolus Linnaeus 1758 Schilder Monetaria 4a Schilder and Schilder Monetaria caputserpentis Schilder and Schilder caputserpentis 7 mikado reticulum argentata 3 Schilder caputdraconis Monetaria Erosaria Ravitrona Meyer 2003 Monetaria Schilder and Schilder Griffiths 1958 Griffiths 1958 caputophidii caputserpentis 4c Schilder caputserpentis, caputophidii, kenyonae, caputanguis b Schilder and Schilder 1971 Brian N. Tissot 1984 Tissot 1984 Griffiths c Tissot 1984 mtdna Meyer subspecies race variety morph form Schilder and Schilder geographic race Ernst Mayr 1942 Ernst Mayr p.193 p.106 Bernhard Rensch Rensch

4 a Schilder and Schilder 1938-39 7 b Schilder and Schilder 1971 4 c Figure 4. Historical transition of the taxonomy of Monetaria caputserpentis and caputdraconis.

6 4 a Schilder and Schilder b Schilder and Schilder c Figure 4. Historical transition of the taxonomy of Monetaria caputserpentis and caputdraconis. (a) Distributions of caputserpentis consisting of seven subspecies and caputdraconis, proposed by Schilder and Schilder ( ). (b) Distributions of caputserpentis consisting of four subspecies and caputdraconis, proposed by Schilder and Schilder (1971). (c) Currently accepted taxonomy of caputserpentis subdivided into two subspecies and caputdraconis. 6

7 1 Mayr 1942 ecotypeecophenotype ecological race ecological subspecies Martin and Hine 2016 p.27 phenotypical plasticity p.28 Schilder Schilder and Schilder a mikado kenyonae caputanguis 1 h, j, k 3 capuserpentis, reticulum, argentata 1 g, i, l 2 3 Schilder Mayr 1942 Mayr 1942 ecotypeschilder and Schilder 1936 Ernst Mayr Schilder Mayr p.27 p.143 Mayr Schilder and Schilder R. J. Griffiths Schilder and Schilder Griffiths 1958 Tissot caputserpentis caputophidii 4c Tissot Irie

8 Schilder and Schilder mikado reticulum 4a Irie 1997 reciprocal transplant experiment CG common-garden experiment 1900 Frederic E. Clements Harvey M. Hall Hagen 1984 CG Göte Turesson ecotype CG John E. Fauth CG Fauth 1998 CG genus Littorina David G. Reid 1996 Reid 1996 Littorina saxatilis 5a-l Kerstin Johannesson 1980 Johannesson Reid 1996 Littorina arcana, L. saxatilis, L. compressa 3 arcana saxatilis 2 arcana Reid 1996 A4 9 crabwavesoft sediment barnacle 4 ecotype Johannesson 2016 Johannesson 2 8

1 5 a-fg-l m n o-q r Figure 5. Littorina saxatilis from (a-f) Trebeurden, Brittany, France and (g-l) Varangerfjord, Norway. Littorina obtusata from (m) Bretagne, France, (n) St.

9 1 5 a-fg-l m n o-q r Figure 5. Littorina saxatilis from (a-f) Trebeurden, Brittany, France and (g-l) Varangerfjord, Norway. Littorina obtusata from (m) Bretagne, France, (n) St. Jacut de la Mer, Brittany, France, and (o-q) Nahant, Boston, USA. Littorina littorea from (r) Solstrand, Norway. Johannesson Littorina sitkana Yamazaki and Goshima NPE = nonlethal predator effect NCE = nonconsumptive effect 1988 Robert D. Appleton A. Richard Palmer family Muricidae Nucella lamellosa 6a-dCancer productus Appleton and Palmer 1988 e.g., Kincaid 1957 Nucella 2014 Marko et al. 9

10 入江貴博図 6 a-c ブレマートンアメリカ d ステイラコームアメリカ産のヒレチヂミボラ e イロデュヴェルドレフランス f ウェールズイギリス g ソルストランドノルウェー h プゲットサウンドアメリカ i, j ナハントアメリカ産のヨーロッパチヂミボラ Figure 6. Nucella lamellosa from (a-c) Bremerton, Washington, USA and (d) Steilacoom, Washington, USA. Nucella lapillus from (e) Ilot du Verdelet, France, (f) Wales, England, (g) Solstrand, Norway, (h) Puget Sound, Washington, USA, and (i, j) Nahant, Boston, USA て染色体数の変異 Robertsonian variation が報告され同じく Nucella 属のヨーロッパチヂミボラ Nucella 分布の大半は 2n = 26 の個体が占めるのに対して一部の lapillus 図 6e-j は大西洋北部の潮間帯に生息する直個体群では最大で 2n = 36 の個体が優占するようだこの達発生の種で貝殻形態の種内変異に関する研究は種染色体数の地理的変異は当初貝殻種内変異の原因で内分類の観点から 1910 年代 40 年代に繰り返し取り組あることが疑われたが現在では両者間の因果関係は否まれた Largen 1971 さらに年代には Roger N. 定されている Galante-Oliveira et al Hughes などの英国の潮間帯生態学者によって詳しい研究実は本種で観察される貝殻種内変異は捕食者からが進められていたが本種は現在貝殻誘導防御の代表のシグナルを含むその個体にとっての環境条件に基づ的な研究対象となっている本種が捕食者のヨーロッパく表現型可塑性だけでは説明できないそのひとつの証イチョウガニ Cancer pagurus に対して殻厚に誘導防御拠として殻形の変異は孵化前の胚の段階で既に存在しを示すことはヒレチヂミボラとほぼ同時に報告されたており母貝の殻形態と対応関係にあることが挙げられ Palmer 1990 図 6 に示したように本種も著しい貝殻る Guerra-Varela et al 年以降この問題の種内変異があり波浪の影響が少ない生息地では捕食者解明には大きな進展があり大規模な交換移植実験と CG が多いために厚い殻を作る個体が多く反対に波あたり実験の結果殻形変異には表現型可塑性 E と遺伝 G の強い海岸では捕食リスクが低いために薄い殻を有するの影響に加えて母貝が経験した環境捕食者からシグとされている本種からは 1950 年に H. Staiger によっナルが子の貝殻形態を変化させる母性効果までもが関 10

11 1 Pascoal et al transgenerational phenotypic plasticity Donelan and Trussell Geerat J. Vermeij Carcinus maenas 19 Cape Cod Boston Massachusetts 1982 Halifax Nova Scotia Dartmouth Geerat J. Vermeij 1982 Vermeij 1982 Vermeij Vermeij 1996 Large and Smee 2010 evolutionary arms race ecological arms race Vermeij 1982 Robin Hadlock Seeley Littorina obtusata 5m-q New England Seeley 1986 Geoffrey C. Trussell Trussell 1996 New England CG Evolution Trussell 2000 Maine Lubec Massachusetts Manchester Lubec Manchester Conover and Schultz 1995 cogradient variation CoGV countergradient variation CnGV CoGV CnGV CoGV Trussell PNAS L. David Smith Trussell 2000 Trussell and 11

12 Smith Matthew O. Nicklin 2 Massachusetts Nahant Manchester Trussell and Nicklin 2002 Cancer productus crusher claw Smith and Palmer 1994 L. David Smith 10 Maine crusher claw Smith 2004 Geerat J. Vermeij Littorina littorea 5r 2 Vermeij 1982 Vermeij 1982 Vermeij 1982 Timothy C. Edgell Rémy Rochette Edgell and Rochette 2008 Fundy Nova Scotia Maine Paul Kemp Mark D. Bertness New England Kemp and Bertness Kemp and Bertness

1 Paul E. Bourdeau family StrombidaeLobatus gigas 7 CITES II Caribbean Marine Research Center Allan W. Stoner Megan Davis 1 8 Martín-Mora et al. 1995 Panulirus argus Delgado et al.

13 1 Paul E. Bourdeau family StrombidaeLobatus gigas 7 CITES II Caribbean Marine Research Center Allan W. Stoner Megan Davis 1 8 Martín-Mora et al Panulirus argus Delgado et al Monetaria Figure 7. A juvenile specimen of Lobatus gigas from South Florida. G E Vermeij Monetaria Schilder Christian Lindemann Philippe Poppe JSPS JP16K

14 Appleton RD, Palmer AR (1988) Water-borne stimuli released by predatory crabs and damaged prey induce more predatorresistant shells in a marine gastropod. Proceedings of the National Academy of Sciences of the United States of America, 85: (2017)., Conover DO, Schultz ET (1995) Phenotypic similarity and the evolutionary significance of countergradient variation. Trends in Ecology & Evolution, 10: Delgado GA, Glazer RA, Stewart NJ (2002) Predator-induced behavioral and morphological plasticity in the tropical marine gastropod Strombus gigas. Biological Bulletin, 203: Donelan SC, Trussell GC (2015) Parental effects enhance risk tolerance and performance in offspring. Ecology, 96: Edgell TC, Rochette R (2008) Differential snail predation by an exotic crab and the geography of shell-claw covariance in the northwest Atlantic. Evolution, 62: Fauth JE (1998) Investigating geographic variation in interspecific interactions using common garden experiments. In ResetaridisWJ Jr, Bernardo J (eds), Experimental Ecology, Issues and Perspectives, Oxford University Press, New York Galante-Oliveira S, Marçal R, Pacheco M, Barroso CM (2012) Nucella lapillus ecotypes at the southern distributional limit in Europe: variation in shell morphology is not correlated with chromosome counts on the Portuguese Atlantic coast. Journal of Molluscan Studies, 78: Griffiths RJ (1958) Subspeciation in two species of Cypraeidae. Proceedings of the Malacological Society of London, 33: Guerra-Varela J, Colson I, Backeljau T, Breugelmans K, Hughes RN, Rolán-Alvarez E (2009) The evolutionary mechanism maintaining shell shape and molecular differentiation between two ecotypes of the dogwhelk Nucella lapillus. Evolutionary Ecology, 23: Hagen JB (1984) Experimentalists and naturalists in twentiethcentury botany: experimental taxonomy, Journal of the History of Biology, 17: Irie T (1997) Relationships between geographic variation of shell structure and water temperature in Cypraea caputserpentis (Gastropoda: Cypraeidae). Yuriyagai: Journal of Malacozoological Association of Yamaguchi, 5:17-29 Irie T, Morimoto N (2016) Intraspecific variations in shell calcification across thermal window and within constant temperatures: Experimental study on an intertidal gastropod Monetaria annulus. Journal of Experimental Marine Biology and Ecology, 483: (2001)., 51: Johannesson K (2016) What can be learnt from a snail? Evolutionary Applications, 9: Kay EA (1960) The functional morphology of Cypraea caputserpentis L. and an interpretation of relationships among the Cypraeacea. Internationale Revue der gesamten Hydrobiologie und Hydrographie, 45: Kay EA (1985) About the cowries. In: Burgess CM, Burgess cowries of the world, Gordon Verhoef Seacomber Publications, South Africa Kemp P, Bertness M (1984) Snail shape and growth rates: evidence for plastic shell allometry in Littorina littorea. Proceedings of the National Academy of Sciences of the United States of America, 81: Kincaid T (1957) Local races and clines in the marine gastropod Thais lamellosa Gmelin - A population study. The Calliostoma Company, Washington Large SI, Smee DL (2010) Type and nature of cues used by Nucella lapillus to evaluate predation risk. Journal of Experimental Marine Biology and Ecology, 396:10-17 Largen MJ (1971) Genetic and environmental influences upon the expression of shell sculpture in the dog whelk (Nucella lapillus). Proceedings of the Malacological Society of London, 39: Lorenz F Jr., Hubert A (1993) A guide to worldwide cowries. Christa Hemmen, Wiesbaden Lorenz F Jr., Hubert A (2000) A guide to worldwide cowries (2nd revised ed.). ConchBooks, Hackenheim Marko PB, Moran AL, Kolotuchina NK, Zaslavskaya NI (2014) Phylogenetics of the gastropod genus Nucella (Neogastropoda: Muricidae): species identities, timing of diversification and correlated patterns of life-history evolution. Journal of Molluscan Studies, 80: Martin E, Hine R (2016) A dictionary of biology (7th ed). Oxford University Press, Oxford Martín-Mora E, James FC, Stoner AW (1995) Developmental plasticity in the shell of the queen conch, Strombus gigas. Ecology, 76: Mayr E (1942) Systematics and the Origin of Species. Columbia University Press, New York Meyer CP (2003) Molecular systematics of cowries (Gastropoda: Cypraeidae) and diversification patterns in the tropics. Biological Journal of the Linnean Society, 79: Moretzsohn F (2014) Cypraeidae: how well-inventoried is the best-known seashell family? American Malacological Bulletin, 32: (2004). Argonauta, 10:19-41 Osorio C, Jara F, Ramirez ME (1993) Diet of Cypraea caputdraconis (Mollusca: Gastropoda) as it relates to food availability in Easter Island. Pacific Science, 47:34-42 Palmer AR (1990) Effect of crab effluent and scent of damaged conspecifics on feeding, growth, and shell morphology of 14

15 1 the Atlantic dogwhelk Nucella lapillus (L). Hydrobiologia, 193: Pascoal S, Carvalho G, Creer S, Rock J, Kawaii K, Mendo S, Hughes R (2012) Plastic and heritable components of phenotypic variation in Nucella lapillus: an assessment using reciprocal transplant and common garden experiments. PLoS ONE, 7:e30289 Reid DG (1996) Systematics and evolution of Littorina. Ray Society, London Renaud ML (1976) Observations on the behavior and shell types of Cypraea moneta (Mollusca: Gastropoda) at Enewetak, Marshall Islands, Pacific Science, 30: Rensch B (1934) Kurze Anweisung für zoologischsystematische Studien. Akademische Verlagsgesellschaft, Leipzig Schilder FA (1927) Revision der Cypraeacea (Moll. Gastr.). Archiv für Naturgeschichte, 91A:1-171 Schilder M, Schilder FA (1936) Revision of the genus Monetaria (Cypraeidae). Proceedings of the Zoological Society of London, 106: Schilder FA, Schilder M ( ) Prodrome of a monograph on living Cypraeidae. Proceedings of the Malacological Society of London, 23: Schilder FA, Schilder M (1971) A catalogue of living and fossil cowries: Taxonomy and bibliography of Triviacea and Cypraeacea, (Gastropeda Prosebranehia). Institut royal des sciences naturelles de Belgique Mémoires, (2) 85:1-246 Seeley RH (1986) Intense natural selection caused a rapid morphological transition in a living marine snail. Proceedings of the National Academy of Sciences of the United States of America, 83: Smith LD (2004) Biogeographic differences in claw size and performance in an introduced crab predator Carcinus maenas. Marine Ecology Progress Series, 276: Smith LD, Palmer AR (1994) Effects of manipulated diet on size and performance of brachyuran crab claws. Science, 264: Tissot BN (1984) Multivariate analysis of geographic variation in Cypraea caputserpentis (Gastropoda: Cypraeidae). The Veliger, 27: Trussell GC (1996) Phenotypic plasticity in an intertidal snail: The role of a common crab predator. Evolution, 50: Trussell GC (2000) Phenotypic clines, plasticity, and morphological trade-offs in an intertidal snail. Evolution, 54: Trussell GC, Smith LD (2000) Induced defenses in response to an invading crab predator: An explanation of historical and geographic phenotypic change. Proceedings of the National Academy of Sciences of the United States of America, 97: Trussell GC, Nicklin MO (2002) Cue sensitivity, inducible defense, and trade-offs in a marine snail. Ecology, 83: Vermeij GJ (1996) Privileged hands: a remarkable scientific life. W.H. Freeman & Company, New York. (2000)., Vermeij GJ (1982) Phenotypic evolution in a poorly dispersing snail after arrival of a predator. Nature, 299: Wilson B (1998) Family Cypraeidae. In: Beesley PL, Ross GJB, Wells A, Mollusca: The Southern Synthesis. Part B. CSIRO Publishing, Melbourne. Wilson B, Clarkson P (2004) Australia s spectacular cowries: A review and field study of two endemic genera Zoila and Umbilia. Odyssey Publishing, El Cajon (CA). (2001) G E, 51: Yamazaki T, Goshima S (2012) Adaptive differences in morphology of the periwinkle Littorina sitkana between two habitats separated by an artificial structure. VENUS, 70:

68 7 2013 2 Hall (1992) 20 Arthur 2011 Hall 1992 evo-devo 20 Arthur evo-devo evo-devo p. x Hall 1992 Arthur 2011 2.1 Arthur 2011 developmental repatte

68 7 2013 2 Hall (1992) 20 Arthur 2011 Hall 1992 evo-devo 20 Arthur evo-devo evo-devo p. x Hall 1992 Arthur 2011 2.1 Arthur 2011 developmental repatte Evo-devo eco-evo-devo Evo-devo and eco-evo-devo: Two extended syntheses in evolutionary biology Yoshinari YOSHIDA, Katsuya TAKAO, Hisashi NAKAO 1 30 80 e.g., Amundson 2005 90 evolutionary developmental