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s a p a h u a i s l d d a l r i s u e r C e e e u c c l s i n a r a r o t l I u l c s a m s e l s d s s d i g s i n s i y m u l s e e o o u h r s l u i a l m a n a c i i i n l s o s s g l i i a a n r c u a l i a i p e l s g r a y l s l n b a i i c v a s o h m o m s d q a s a u a o r s a a b b a l i s r l e u s i u s s u a s s s s a w l n c s s i c u m c p u C r m o u s o d b a y s a o u e s r l l s u o m s s y g l d o s e l s a m i u l l o i a r u y s i l t a i a z s r n o o o y a x a d i h u n y r c h a e r i s a l u ( l e a a n s i r m p u b p s t l l l i o a l r h i u i i s p a i u r u y o r s s a n r l u o i i n a u p p u a o t a u i a l e o o i l e o i e s i o s u m g n l r t u a e i t a e l l e l n h a u a o i m u s d o l i a u l r u s h e i i l d s a e i p n u e p m i i l i l i r r e h s l p t a c O a o d l s u l r l m i l d m h h s i a l l o o o i s i s b e m m l u l l e i s a s a l a i a o o l p s e e g a c f l s o l s p l i r n h h l r a l r z h a o r r u o a r a h n r c o p 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l r i m p b e m r i c i e o e r n l d i r e l r l e h a r b c p t e c z r r t o t h r s s d m p s d t u i t i l e e o u p l i e m l d a c c i i t o l d u b b h t p c c p t o o a t p i a o c y u p o o m u o e o o e i e o o o o o r a o z o a a a n a i a i e e e i m r b l e e d a y e u o c l l e o m g r e l o i y l m n e o a r y a p e l t c o m l l p p p n r b i l a s t r o y y y y a l a r y l a a s e o l m h u a o r h m o r r o o n r e n t a l a o r i l x l r y i n r n l a a h a o h p h s r u s e x u e n n a e c s a y o m a r r a p g t e l h t a i l p m m h o r r r u r i r t m c p p c c c n e c o h l u e o c a i e o m c t o r r e v i i a i a t u i s l b o s a e a y m y a o y y l a a r l o y h o o o o o r e y y h a h u e e y e l r y c e r a a n s g s o s t h g l u c i o a o e t h i t r o o l u r e o p h g g i l r a y o o r e y y e a G P e a e M P T S H T P H P T H T G S H G R C G P P P A P S T C C H C T D C T F U E P S W E N P S O P M G S G O C A G T P C U M L C C X D C C S G R S B p a a T a u e n y y l l m l e h y R S F F X A M B M C O G M P C L T E N A L C A C L L C L C C P B C C C I H P G S H P A P T H S G R S C G P B P G B X S T L C P P J S R L A B B H H A A L A P C G N P G F S F P C T L a b c 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 P A P X P T H H C 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 4 4 5 6 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 5 6 7 9 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3 3 3 3 4 4 4 4 4 4 4 4 4 4 5 5 5 5 5 5 5 5 5 5 6 6 6 6 6 6 6 6 6 6 7 4 8 1 2 3 4 5 6 7 8 9 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3 3 3 3 4 4 4 4 4 4 4 4 4 4 4 4 4 5 5 5 5 5 5 5 5 5 5 6 6 6 6 6 6 6 6 6 6 7 7 7 7 7 7 7 7 7 7 8 8 8 8 8 8 8 8 8 8 9 9 9 9 9 9 9 9 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 9 9 e e a Cortinar- e 2 3 3 2 2 2 2 3 3 3 3 a 2

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ewiaceae T b e o c 3 2 T r a L r l c H n r 5 5 Tricholomataceae Suillineae Sclerodermatineae GOMPHALES E A c E a Psathyrellaceae e Auriscalpiaceae 1 i a A e E N

1 O o Hygrophoraceae a LECANOROMYCETES t l

A 3 o l a a E l C C b I h b C

o Hymenogastraceae h o e A e 1 1 e PHALLALES

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Mycenaceae n i T c e T A i o M a M o p t i M p Strophariaceae LEOTIOMYCETES SORDARIOMYCETES o r p S n N L SPHAERO- O o O O n e S Physalacriaceae- POLYPORALES THELEPHORALES Y I O n a g a I S S r I S I

n Russulaceae o O 1 2 E

y 2 T BOLALES 2 N P T C E

Schizophyllaceae O E C D m C H T I O E T M L O y C H H E S G R H G 2 Y T T T E 1 2 GEASTRALES C 1 O U

1 D O O 2 S D Y

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PLUTEOID CLADE HYGROPHOROID CLADE MARASMIOID CLADE TRICHOLOMATOID CLADE E E A D GLOEOPHYLLALES M M C E G 1 R C M A O C

5 Y L L Y O A 1 1 I M

2 2 2 2 1 5 O B Y A O R M R H L Z 2 Agaricaceae- I O E T M E BOLETALES Z L C S Nidulariaceae O E

CORTICIALES S S E

RUSSULALES S S A P 1 O M E E C N I E E E E I A D T X S I O A L L M L L H R 4 E T E L K S E A A A ANOMOPORIALES A C H I L E C C G L T L R 2 I P O R Y A C PEZIZOMYCOTINA L L E K O A N A

5 Y Jaapia clade N M A E C 5 AGARICOID CLADE I P T L E M O R 2 Y C W

D S

5 5 U I 5 M

2 L O A M AGARICALES A C Y L H L I H Y A A D2 B E L H C T R T R E N A I E S D A M N Evolutionary relationships of FUNGI U S C 2 O T A

5 N E R H E A A I A C

What does this diagram represent? This is a phylogenetic tree. It shows the pattern of evolutionary relationships that connect extant (living) species of Fungi. T N T O P R T I C Y What do the names on the tips and branches of the tree mean? The labeled tips and branches of the tree are the names of taxa (sing. taxon), which are groups of organisms that have been given scientific names. D

AGARICOMYCETIDAE E C T

To be recognized as a formal taxon, a group should be monophyletic, which means that it is a single and complete branch of the tree (i.e., an ancestor and all of its descendants). Another word for a monophyletic group is a clade. T O M 5 C Some of the taxa at the tips are species, recognized by their binomial names, consisting of a generic name and specific “epithet”, e.g. Typhula phacorrhiza. Other taxa listed here are more inclusive groups that contain multiple species. 5 5 5 Y O The branching points on the tree are speciation events. C

5 Y O M I C Is every species of Fungi present in this phylogenetic tree? Absolutely not! There are roughly 100,000 described species of Fungi, but the total number of extant species is surely much greater. One popular estimate suggests that there are 1,500,000 5 5 5 Y

extant species of Fungi. This tree emphasizes the fleshy mushroom-forming Fungi that are most often collected on forays, including agarics, boletes, polypores, coral fungi, cup fungi, etc. However, much of the diversity of Fungi is composed of M O Y D M

microscopic or inconspicuous forms, such as yeasts (single-celled Fungi), molds (filamentous asexual Fungi), corticioid fungi (crustlike resupinate forms), and diverse symbionts (Fungi living in or on other organisms, including plants, animals, I O R

and other Fungi). Only a few exemplars of these cryptic forms are included here. 4 M O R How was the phylogenetic tree constructed (what is the evidence?)? To estimate relationships, fungal systematists use both morphological characters (physical features) and molecular characters (DNA or amino acid sequences of genes and proteins Z I O O T shared among diverse taxa). Molecular and/or morphological characters are scored from each species and then encoded in a data matrix, which is analyzed using numerical methods that are implemented by computer algorithms. N I I

4 H C

This tree is a composite based on many independent phylogenetic analyses that were conducted using molecular characters. Y N G

How do we know the tree is correct? We can never “know” with absolute certainty that our reconstruction of evolutionary relationships is correct. However, using statistical analyses we can assess confidence in individual branches. We can also test R I U hypotheses using independent analyses of different genes. When multiple genes support the existence of a particular clade, then we can have strong confidence in that result. Sometimes the data are not decisive, and a particular region of the H A R C C

tree must be left unresolved. For example, the branching order among major groups of Agaricales is not well understood, which is why the “backbone” of the Agaricales is drawn as an unresolved “polytomy”. M L

Why should we construct phylogenetic trees? Phylogenies are central in biology. Important uses of phylogenies include the following: O C 4 I To structure biological classifications (taxonomy). Taxonomy, which provides the names used for communication about all living organisms, is based on phylogeny. Taxonomy provides a core element in the language of biology, which is essential T T to basic fields, such as ecology, as well as applied disciplines, such as agricultural science and medicine. U 5 5 5 5 N S

To create tools for identification (molecular ecology). The molecular sequences used for phylogeny reconstruction can also be used to identify unknowns (as long as there is a well-sampled database of reference sequences). For example, molecular 4 P

approaches have been used to show that the symbionts of non-photosynthetic “indian pipes” plants (Monotropa uniflora) are members of the Russulaceae. E 5 U To understand the evolution of organismal features (character evolution). Phylogenetic trees provide a framework for reconstructing the changes in organismal features that have occurred throughout evolution. For example, phylogenetic trees of 4 Basidiomycota show that the puffball type fruiting body has evolved repeatedly in clades such as Agaricales (e.g., Lycoperdon), Boletales (Scleroderma), and Phallomycetidae (Geastrum). 5 5 5 5 To reconstruct the past movements of organisms (historical biogeography). By mapping the distribution of organisms onto the tips of a phylogeny it is possible to infer the geographic origins of groups of organisms. For example, historical “BASAL FUNGAL biogeographic methods were used to locate the origins of the West Nile Virus in Europe. AGARICOMYCOTINA ASCOMYCOTA To understand the history of life. LINEAGES” Where can I go to learn more about Fungi and phylogenetics? The following web sites all contain many links to other mycology resources on the internet. 4 The Tree of Life Web Project (http://tolweb.org/tree/) provides an excellent, user-friendly overview of the higher-level relationships of Fungi and other organisms. MykoWeb (http://www.mykoweb.com/), which is run by the Mycological Society of San Francisco, includes images, keys, and other information. Click on the “systematics” link. Tom Volk’s Fungi pages (http://botit.botany.wisc.edu/Toms_fungi/) provides detailed, accessible information about diverse fungi through the “fungus of the month pages” (and much more). BASIDIOMYCOTA 4 The Mycological Society of America (http://www.msafungi.org/) is the major professional society of mycologists in the USA. 5 Electronic copies of this poster are available from David Hibbett, Biology Department Clark University (http://www.clarku.edu/faculty/dhibbett/) DIKARYA