Backstory: is a high biomass ecosystem engineer that reworks and remineralizes estuary sediments (DeWitt et al. 2004, D’Andrea and DeWitt 2009)

Bopryid Isopod parasites infest decapod and effectively castrate their female hosts

Markham (2004) described , collected from the intertidal estuary mud shrimp, Upogebia pugettensis of Yaquina Bay OR in 2002

Orthione was introduced to North America from Asia in the 1980s (Chapman et al. 2012)

Orthione is associated with severe declines of Upogebia over its entire range (Chapman 2012, Dumbauld et al. 2011).

Interactions and mechanisms:

Variable host condition and possible vulnerability - Orthione restricted to reproductive sized Upogebia Highest prevalence among Upogebia females No relation between parasite load and host weight loss (Smith et al. 2008, Dumbauld et al. 2011, Griffen 2009, Repetto and Griffen 2012)

Could Orthione prevalence among Upogebia females result from sex change or greater male mortality? (Griffen 2009, Repetto and Griffen 2011) Asson

A relation between energetics and parasite load? (Repetto and Griffen 2011) Burton

Could trophic conditions for hosts control host vulnerability? ~Young Can an ecosystem engineer of Oregon estuaries, the blue mud shrimp, be spared from extinction? John Chapman Dept. Fisheries & Wildlife, HMSC Oregon State University [email protected] Acknowledgements: Amy Chapman, Ralph & Donelle Breitenstein, Brett Dumbauld, Nancy Edwards, MCWS, Lincoln County Community Consortium, and many, many others 2 Mar. 2017 MidCoast Watershed

Photo by Donna McCoy July 06 Council Geography of modern diversity http://sappingattention.blogspot.com.es/2012/11/reading-digital-sources-case- study-in.html

Main Shipping Routes in the Eastern Pacific: 1936 World Waterways Network: 2008 Exponentially increasing dominance of non-coevolved introduced in all marine ecosystems Ruiz et al. 2000 Invasions are Conservation’s Dark Side Introduced species are often not constrained by evolution

Introductions can become extinct

Introductions can cause extinction Upogebia pugettensis is an ecosystem engineer in estuaries due to its high biomass, and its reworking and remineralization of sediments (e.g. D’Andrea and DeWitt 2009) > 1 m depth burrows The trophic links of Upogebia to estuary sediment chemistry are intimate and significant. Journey Begins

6 60%

5 50%

4 40% density)

3 30% Prevalence

2 20%

Upogebia 1 10% Orthione ln( 0 0% 1988 1993 1998 2003

Year

Orthione griffenis Markham, 2004 (Published winter 2004 Orthione timing coincides with Upogebia 2008 Survey of U. declines Upogebia are possibly recruitment pugettensis limited (Dumbauld et al. 2011, Chapman et al. 2012) (Chapman et al. 2012)

. Reduced . Collapsed . Extinct Only known (first?) introduced bopyridan parasite

Bopyridan ecology, natural history and population biology unknown

Who? Previous records of bopryidans controlling host Me???? populations unknown Could effective castration limit Upogebia populations?

Could lost reproduction control Upogebia populations? 47 million 3 Benthic Yes. Upogebia 29 2010 are recruitment 24 abundances 2 19 increase with 14

limited Millions

PL Immigr. 2010 (millions) 2010 PL Immigr. 1 Neo Post Larvae 2010 9 immigration Upgebia 4

0 1/15 3/16 5/15 7/14 9/12 11/11 -1 1/15 3/16 5/15 7/14 9/12 11/11 UpoNight UpoDay Full New Neo Night Neo Day Full New Clouds 250 1,802 million 9 8

200 7 2011 2011 6 150 5

Post Larvae 2011 (millions) 4

100 Millions 3 Neo Post Larvae 2011 Upogebia 2 50 1

0 20100 and 2011 Upogebia 0-year benthic 1/15 3/16 5/15 7/14 9/12 11/11 density1/15 accumulation3/16 5/15 7/14 with9/12 immigration11/11 Flood-Night Flood-Day Ebb - Day Flood-Night Flood Day Ebb-Day Ebb - Night Full Moon New Moon Ebb-Night Full Moon New Moon 2010 Upogebia 3% of 2011 Upogebia in Yaquina Bay 2005-2014

250 2006 Yes. 200 Mature Upogebia 2011 2007

populations are 150 recruitment limited 100 2012 2013 2008

2005

Density of 1+ Shrimp in Following Year Following Shrimp1+ in Density of 2009 50 2010 0

0 50 100 150 200 250

Density of Recruits Can Orthione reach all hosts? Are Orthione host limited? Are there likely host refuges? Are the Orthione propagules limited? Who? Me????

Abundant, persistent Orthione populations? ~ 0.3 mm pelagic epicaridan

1 2

3 Brooding Orthione and its 3 larval dispersal stages Breitenstein et al. (unpublished MS) High estuary Cryptoniscan movements in flood and ebb abundance tide plankton samples: They wait.

600

500 Diel Equal emigration and abundance 400 immigration Number of Orthione larva 300 509 200

100 155

0 Day Night Only known (first?) introduced bopyridan parasite Bopyridan ecology, natural history and population biology

Who? No previous records of bopryidans controlling host Me???? populations First hypothesis – could effective castration limit reproduction?

Could lost reproduction control host populations? Unexpected Infest > 12 mm shrimp and ~2 females for each male infestation Dumbauld et al. 2011 Smith et al. 2011 patterns No Evidence of Energetic Cost

No apparent relation between lipid No apparent loss of host weight No apparent association of weight concentrations and relative size of associated with Orthione (Burton loss and parasite size (Smith et al. parasite (Repetto and Griffen 2012) et al. In Prep.) 2008) Do Orthione limit host reproduction or otherwise stress their hosts?

Who? 1) Do they change host sex? Me???? 2) Do they kill host males?

3) Do they take enough host resources for effective castration? 400

350

300

250

200

Frequency 150

100

50

0 0 5 10 15 20 25 30 35 Carapace Length (mm)

All Infested No evidence that the introduced parasite, Orthione griffenis, causes sex change or differential mortality in the native mud shrimp, Upogebia pugettensis Danielle Asson, John W. Chapman, Brett R. Dumbauld Do Orthione limit host reproduction or otherwise stress their hosts?

Who? 1) Do they change host sex? NO Me???? 2) Do they kill host males? NO

And there seems to be an energetic cutoff at around 12 mm Remember But more questions remain this decline Do Orthione limit host reproduction or otherwise stress their hosts?

Who? 1) Do they change host sex? NO Me???? 2) Do they kill host males? NO

3) Do they take enough host resources for effective castration? Energetic losses to the introduced bopyrid isopod parasite, Orthione griffenis Andrea R. Burton, John W. Chapman, Brett R. Dumbauld, Richard Caldwell, and Louise A. Copeman

Infested shrimp lose lipids! Energetic losses to the introduced bopyrid isopod parasite, Orthione griffenis Andrea R. Burton, John W. Chapman, Brett R. Dumbauld, Richard Caldwell, and Louise A. Copeman

They lose lipids a lot of lipids!3 Growth adjustments by Upogebia pugettensis hide energetic losses to the introduced bopyrid isopod parasite, Orthione griffenis Andrea R. Burton*1, John W. Chapman2, Brett R. Dumbauld1, Richard Caldwell3, and Louise A. Copeman4

Crustacea adjust their sizes to energy/biomass Do Orthione limit host reproduction or otherwise stress their hosts?

Who? 1) Do they change host sex? NO Me???? 2) Do they kill host males? NO 3) Do they take enough host resources for effective castration? Yes Do Orthione limit host reproduction or otherwise stress their hosts? Yes. YOU! End of Who? Me???? story? No. Invasion of San Francisco Bay by Not Upogebia major A newly recognized non-native end of species with potentially large ecosystem consequences story Michael McGowan Maristics John Chapman Oregon State University Ralph Breitenstein Oregon State University Andrew Cohen C.R.A.B. Kathy Hieb California Dept. Fish and Game Pacific Ocean 244 213 323 215 345 322 214 110 325 109 321 106 216 317 103 243 211 USACE2011 July 320 108 142 105 212 318 Central Bay 319 346 107 San Pablo Bay San Pablo 104 101 447 427 Suisun Bay 102 South Bay 428 429 432 430 140 431 433 Upogebia 534 West Delta 535 736 837 San Francisco Bay Study Bay Francisco San Boat Sampling Stations Sampling Boat 750 sp. sp. benthic in mud 751 Channel Station Channel Shoal Station 760 853 761 752 863 762 864 865 Pacific Ocean 244 213 323 215 345 322 214 DFG larval Upogebia larval DFG 110 325 109 321 106 216 317 103 243 211 320 108 142 105 212 318 Central Bay 319 346 107 San Pablo Bay San Pablo 104 101 447 427 Suisun Bay 102 South Bay 428 429 432 430 140 431 433 534 West Delta 535 736 837 San Francisco Bay Study Bay Francisco San Boat Sampling Stations Sampling Boat 750 751 Channel Station Channel Shoal Station 760 853 761 752 863 June 2010 June MLMLbenthic in mud 762 864 865 The Asian mud shrimp, Upogebia major invasion of the northeast Pacific: a coevolved host of the introduced Asian bopyrid isopod, Orthione griffenis John W. Chapman1, Michael F. McGowan2, Ralph A. Breitenstein3, Andy Cohen4 Kathy Hieb5 and Ralph Appy5 How come so late a discovery?

San Francisco Bay 2010 Drakes Estero 2015 Bodega Harbor 2016 The Asian mud shrimp, Upogebia major invasion of the northeast Pacific: a coevolved host of the introduced Asian bopyrid isopod, Orthione griffenis John W. Chapman1, Michael F. McGowan2, Ralph A. Breitenstein3, Andy Cohen4 Kathy Hieb5 and Ralph Appy5

Upogebia San Francisco Bay 2010 major is a Drakes Estero 2015 Bodega Harbor 2016 native host What’s that got to do with of Orthione me???? griffenis Did you More questions remember?

2008 Core and yabby pump samples

Possibly more to this than “time at size” A B C

0.4 m

2 m

Cores miss the biggest and deepest shrimp Deep Core Project 2017

Team 1: Omar M. Alkhaldi, Eric M. Beebe, Cade T. Burch. Team 2: Connor Churchill, Patrick L. Finn, Jacob A. Garrison. Team 3: Zachary M. Gerard, Evan J. Leal, Derrick E. Purcell, Derrick E Yamada et al. 2015

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