The Futurability of Biodiversity Chapter 2 Why have

ØIn review Biodiversity＝Gathering of organisms with interactions Ecosystems whose biodiversity is high enrich human life - for example, providing various foods or medicines, keeping diseases in check, affecting local culture, and so on. Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

ØToday’s Topics 1. Evolution of biodiversity How has the present biodiversity of the earth evolved? 2. Maintaining mechanisms of biodiversity How is the present biodiversity of the earth maintained? Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

1. Evolution of biodiversity n How has the present biodiversity of the earth evolved? 1. Biological evolution 2. Environmental factors of evolution 　　Isolation & Niche 3. Ecological factors of evolution 　　 Coevolution & Cospeciation Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

Ø 1. Evolution of biodiversity 1) Biological evolution What is the ancestor of dogs?

Ø 1. Evolution of biodiversity 1) Biological evolution Evolutionary history Copyright 2010 Research Institute

Ø 1. Evolution of biodiversity 2) Environmental factors of evolution Isolation Bat Lion Elephant Mole Koala Reindeer Primitive mammal Camel Deer Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved. Whale

Ø 1. Evolution of biodiversity 2) Environmental factors of evolution Niche arboreal terrestrial hypogeal aquatic herbivorous Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved. carnivorous

Ø 1. Evolution of biodiversity 2) Environmental factors of evolution Herbivorous and terrestrial mammals Desert Foraging method Temperate zone Big body Arctic zone There are few animals which live in the same environmental conditions with the same habitat. Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

Ø 1. Evolution of biodiversity 3) Ecological factors of evolution Coevolution Herbivores Mustard oil

Ø 1. Evolution of biodiversity 3) Ecological factors of evolution These new Insects eat the plant. new Some insects have evolutionarily acquired the metabolic mechanism which detoxicates the chemical. new The plant synthesizes a poisonous chemical. Both plants and animals continue to evolve, affected by a vicious circle (arm’s race). Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

Ø 1. Evolution of biodiversity 3) Ecological factors of evolution Mutualism means … close interaction between organisms which help each other. This is not the truth! Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

Ø 1. Evolution of biodiversity 3) Ecological factors of evolution 　 The process of establishing pollinative mutualism ―The true aspect of mutualism― 1. Insects were initially herbivores (enemies of plants). Oh no! Our flowers will be eaten! I’m hungry. Oh! I found food! Food Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

Ø 1. Evolution of biodiversity 3) Ecological factors of evolution The process of establishing pollinative mutualism ―The true aspect of mutualism― 2. Some flowers have evolved to secrete nectar. Continuously produced nectar is better food than flowers, isn’t it? えさ Nectar Pollination Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved. That’s true. Oh! It’s so sweet! I will come here again.

Ø 1. Evolution of biodiversity 3) Ecological factors of evolution The process of establishing pollinative mutualism ―The true aspect of mutualism― 3. Plants make the insects pollinate. Yes, of course. (We did it ! She will pollinate. ) Nectar Give me nectar. I have collected nectar from other flowers like you, but I need more! Pollination Mutualism is the result of evolution whereby a species has tried for a long time to take advantage of the other species, which was originally hostile. Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

Ø 1. Evolution of biodiversity 3) Ecological factors of evolution Partnerships between flowers and pollinators Characteristics of flower ? Color Uniqueness Time zone of providing nectar White Strong smell Night Yellow Large number of flowers Daytime Red Tubulous flower Daytime Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved. Pollinator

Summary 1. Evolution of biodiversity n Biodiversity is the result of evolution. l. The present biodiversity on the earth is the evolutionary state still in progress. l. Evolution is promoted by isolation, such as continental drift. l. Life history varies from species to species, caused by adaptations to different niches. l. Interspecific interaction also promotes evolution. Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

2. Maintaining mechanisms of biodiversity n How is the present biodiversity of the earth maintained? 1. Segregation of niche 　　space, time and food 2. Fixation of interspecific interaction 　　Cospeciation and host-shift 3. Keystone species Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

Ø 2. Maintaining mechanisms of biodiversity　1) Segregation of niche Spatial segregation of ant species inhabiting tropical rainforest, Southeast Asia Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

Ø 2. Maintaining mechanisms of biodiversity　1) Segregation of niche Temporal segregation Diurnal species Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved. Nocturnal species

Ø 2. Maintaining mechanisms of biodiversity　1) Segregation of niche Food segregation (South America) Copyright

Ø 2. Maintaining mechanisms of biodiversity　1) Segregation of niche in ant community Carnivorous Herbivorous Several species which inhabit the same place coexist by segregation of niche. Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

Ø 2. Maintaining mechanisms of biodiversity 2) Cospeciation and host-shift Mutualism between Macaranga species and Crematogaster ants Nesting space (hollow stem) 　Food bodies (food) Protection against herbivores (bodyguard) Macaranga sp. Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved. Crematogaster ants Photo: (right) Tamiji Inoue

Ø 2. Maintaining mechanisms of biodiversity 2) Cospeciation and host-shift When mutualistic ants are experimentally removed… Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

Ø 2. Maintaining mechanisms of biodiversity 2) Cospeciation and host-shift Sharing the same fate started diversification Molecular phylogeny of Crematogaster ant and Macaranga spp. Crematogaster ant C. borneensis 1 M. trachyphylla Macaranga spp. M. bancana C. borneensis 2 C. decamera 1 Host-shift C. decamera 2 C. sp. 4 M. hullettii M. havilandii M. lamellata M. beccariana M. hypoleuca M. hosei C. sp. 2 M. Winkler The above partnerships are species-specific. Fixation of the partnerships = survival of each species Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved. (Itino et al. 2001)

Ø 2. Maintaining mechanisms of biodiversity 3) Keystone species Is any species indispensable to maintaining ecosystem? Keystone species Excessive hunting of sea otters in 1900 s for their fur Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved. Species which greatly influences an ecosystem. If the species decreases or eradicates, the ecosystem changes and its balance breaks down.

Ø 2. Maintaining mechanisms of biodiversity 3) Keystone species Fluctuations of population Sea otter Fish Sea urchin Kelp Prey-predator interaction Indirect interaction Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

Ø 2. Maintaining mechanisms of biodiversity Copyright 2010 Research Institute for Humanity and Nature.

Ø 2. Maintaining mechanisms of biodiversity 3) Keystone species Photo: Tamiji Inoue Copyright 2010

Ø 2. Maintaining mechanisms of biodiversity 3) Keystone species Which is the keystone species? Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved. Photo: Yasunori Maezawa & Echigo-Matsunoyama Museum of Natural Science, ‘Kyororo’

Summary 2. Maintaining mechanisms of biodiversity l Organisms which are seen inhabiting the same environment actually differentiate their living space, time of activity, food, and so on. l In mutualistic and parasitic relationships, fixation of species-specific partnerships enables many species to survive. l Keystone species is an especially important species for maintaining an ecosystem. Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

ØSummary of Today’s Topics Why have the earth’s organisms become so diverse? 1. Biodiversity has evolved through long-time association between organisms and environment, and interspecific interaction. 2. The present biodiversity is maintained by segregation of niche and fixation of interspecific interaction. 3. We often recognize the whole web of an ecosystem by the loss of keystone species. However, there are many ecosystems whose keystone species has not been identified yet. Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

ØExercises Let’s do the exercises below: 1. Among the following three phenomena, mention all examples which you consider to be the process of evolution. In those which you do not choose, explain the reason why you think so. a) A cabbageworm grows into a pupa, then becomes a butterfly. b) A mule is produced by crossing a male donkey and a female horse. c) A plague of harmful insects occurs in paddies. Pesticide which had effectively controlled the insects until last year is not effective against the plague. 　 Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

ØExercises Let’s do the exercises below: 2. Mention an example of mutualism and explain each profit which is brought by the partner. Then think about how the mutualism has evolved. 3. Biological evolution is proceeding now. What kind of species may appear in the future, evolved from the present species? Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

ØColumn 1: Cospeciation and host-shift Parasitism sometimes brings a number of new species. -parasitism between stag beetles and mites- Parasitic mite Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved. Photo: National Institute for Environmental Studies, Koichi Goka

ØColumn 1: Cospeciation and host-shift Copyright 2010 Research Institute for Humanity and Nature. All

ØColumn 2: Keystone species General flowering period in tropical rainforest in Southeast Asia Fig

ØColumn 2: Keystone species No general flowering period in tropical rainforest in Southeast Asia

ØGlossary (1/3) Arm’s race Phenomena in which prey and predator species develop their attack strategy and defense strategy, respectively, and evolve together. Coevolution Phenomena in which several species have influenced each other’s survival and reproduction in the process of evolution. Cospeciation In the process of evolution, several species have influenced each other’s survival and reproduction, then synchronously produce new species. General flowering Flowering phenomena which are observed in tropical rainforests in Southeast Asia. Many canopy tree species synchronously flower at intervals of several years. Triggers of flowering are thought to be severe drought or low temperature. Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

ØGlossary (2/3) Interspecific interaction Several kinds of interactions in which organisms influence on each other, such as competition, predation, mutualism, and so on. Isolation Preventing gene exchange among populations by geographical barriers. Molecular phylogeny Phylogeny is a tree diagram which expresses interspecific relationships and evolutionary history. In molecular phylogeny, the degree of similarity of base sequences is a measure of interspecific affinity. Mutualism A kind of interspecific interaction in which all organisms within the interaction increase their fitness. Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

ØGlossary (3/3) Pollinative mutualism Mutualism between angiosperms and pollinators (insects, birds, mammals, etc). Angiosperms have the profit of carrying pollens to conspecifics, and pollinators have the profit of getting food (nectar and pollens). Species-specific relationship Relationships in which specified species are closely tied in with each other, and do not accept other species. Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

ØReferences Itino T. , Davies S. J. , Tada H. , Hieda Y. , Inoguchi M. , Itioka T. , Yamane S. and Inoue T. (2001) Cospeciation of ants and plants. Ecological Research 16: 787 -793 Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.

ØAuthors & Credits The Futurability of Biodiversity Chapter 2 Why have the earth’s organisms become so diverse? Authors Aya Hatada Takao Itioka Masahiro Ichikawa Stewart Wachs Martin Piddington Applications Cut. PRO 3 Real tough. Animal and Insect version (Design Office Kyowa) Microsoft Power. Point® Sugoneta illustration package. Animal encyclopedia (Grapac Japan) Illustration & design Be 4°TECH Koubou Yecoruka Photos Aya Hatada Biodiversity Photos Chihiro Handa Daisuke Fukuda Echigo-Matsunoyama Science of Natural Museum, ‘Kyororo’ Hiroshi Tanaka Koichi Goka National Institute for Environmental Studies Shoko Sakai　　　　　 Takao Itioka Takashi Matsumoto 　　 Tamiji Inoue Yasunori Maezono Data provider Koichi Goka Copyright 2010 Research Institute for Humanity and Nature. All Rights Reserved.