Towards trait-based modeling for predator-prey interactions under climate change in the NE Pacific

Dr. Natasha Hardy Postdoc in Biological Sciences @ U Alberta, AB, Canada & Hopkins Marine Station @ Stanford, CA, USA Collaboration with NOAA Environmental Research Division @ Monterey, CA, USA Within a context of increasing climatic variability and warming mean climate states…

NOAA > Global climate series > Climate at a glance > Jul–Sep …and shifts in thermal habitat causing redistributions RCP 2.6 RCP 8.5

Projected shifts in distributions for 303 west coast species by 2100 (Jul-Sep)

Uncertainty Low à Med à High

Morley et al. (2018) PlosOne Use of traits to model species’ vulnerability to climate change

wide

thermotolerance Abundance

narrow

Temperature (˚C) 0˚ +2˚ +4˚ +6˚ warming environment But what about species effects on each other? Predator-prey interactions from the coast to the open ocean

Brent Hughes, OSU John Healey, Vancouver Aquarium Albacore tuna as indicators for climatic & ecological change q Albacore tuna as ecosystem samplers q Move beyond q Identify trait-based parameters for diet shifts vs. environmental gradients q Apply those parameters to project for future ecological & environmental change Albacore tuna as indicators for climatic & ecological change

ü Broad & variable diet composition across environmental gradients

ü HMS – tracking change in thermal habitats

Hazen et al. 2013 ü Commercially valuable – NE Pacific with cross-jurisdictional CAN-US Treaty Trait-based predictions for future change

Conserved

Switching traits/prey with climate No relationship of prey traits in diets in traits of prey Predicted probability Predicted Cold Warm Average → Warming ocean Climate regimes Towards trait-based predictions for future change

trait * environment relationship

Predictive traits Taxonomic diet information

Environment Temperature + + Geography Not so fast…

o What is a typical albacore prey? o Which strong predictors for diet shifts? What is a typical albacore prey?

Global, historical review of albacore diets from late 1880’s to early 2000’s Table S5. Metrics showing prey species found in 86 albacore (Thunnus alalunga) stomachs in 2008, 2009, and 2010. Metrics are as follows: N is abundance, %N is relative abundance, FO is frequency of occurrence by stomach, %FO is % of stomachs containing that food item. Total mass is mass of prey items estimated from length, otolith size (fish), or beak metrics (), and total kJ is estimated from mass using energy density values from Glaser et al.(7). IRI is shown for the 4 most prevalent prey items. A hyphen (–) indicates that metric(s) could not be calculated because species were unidentified; ‘nd’ (‘no data’) indicates that conversion algorithms were not available for mass and kJ for that species.

Total mass Total Order/Family Prey Category N %N FO %FO (g) kJ IRI FISHES Carangidae symmetricus 179 11 35 41 452 2894 23.44 Clupeidae Sardinops sagax 51 3 25 29 122 887 4.97 Engraulidae Engraulis mordax 6 <1 2 2 16 105 Kyphosidae Medialuna californiensis 1 <1 1 1 nd nd Myctophidae Triphoturus mexicanus 6 <1 6 7 nd nd Scomberesocidae Cololabis saira 7 <1 5 6 <1 3 Scombridae Scomber japonicus 4 <1 2 2 0.1 1 Sebastidae Sebastes spp. 29 2 9 10 21 90 Fishes unid. 111 7 30 35 – –

CEPHALOPODS Argonautidae Argonauta argo 13 1 8 9 nd nd affinis 1 <1 1 1 0.1 0.5 Enoploteuthidae Abraliopsis felis 19 1 11 13 35 155 Gonatus spp. 176 11 44 51 154 678 13.50 Loliginidae Doryteuthis opalescens 79 5 3 3 372 1639 Octopodidae Octopus bimaculatus 24 2 10 12 3 11 Octopoteuthis sicula 19 1 7 8 10 46 Octopoteuthidae Octopoteuthis spp. 2 <1 1 1 1 6 Ocythoidae Ocythoe tuberculata 1 <1 1 1 nd nd Ommastrephidae Dosidicus gigas 1 <1 1 1 55 240 Onychoteuthidae Onychoteuthis borealijaponica 17 1 4 5 13 59 Ceph. unid. 143 9 31 36 – –

Diet table CRUSTACEA unid 20 1 6 7 nd nd Pleuroncodes planipes 2 <1 1 1 3 10 Euphausiacea Euphausiid 540 33 45 52 384 1190 35.88 Crust. unid. 35 2 5 6 – –

French diet

table from 14 the 1930’s Trait-variation in albacore diets across geography

Conserved

Switching traits/prey with climate No relationship of prey traits in diets in traits of prey Predicted probability Predicted

Geography Global historical review of albacore tuna diets

NE Pacific

• 31 published 200 papers from

1880’s to 2000’s ichness e r

NE Atlantic • 50 unique ulati v observations of m albacore diet 100 Mediterranean

composition Species cu NW Atlantic

SW Pacific

NE Indian SW Atlantic 0 1880 1920 1960 2000 Years sampled High prey diversityAncistrochei in albacore diets

Dor

Heteroteuthis dispar Liocr

Alloteuthis sub S T Helicocr yteuthis opalescens

s eutho e

i

is acirrhus

p

n niana

s yi i

i anchia reinhardti i f

l e nalis f or Cr

e t r Galiteuthis ar a

f Galiteuthis ph t er

us lesueu r

T w a anchia scabr

s Sthenoteuthis oualaniensisEucleoteuthis luminosa s Hy odaropsis eb anchia pf i enia megalops i

o

s

s

w

aloteuthis pelagica p Ommastrephes bar p e

o

o

i

i

l n

l

a i

a ulata

a Ille r r

n

b

Dosidicus gigas b apetella heathi apetella diaphana ii eff a ia lamarckii Or J J Eledone cirrhosa Octopus vulgar Or x coindetii A Scaeurgus unicirrhus A Pteroctopus tetr yroteuthis inf Octopus bimaculatus Argonauta nour mata Argonauta argo usta nithoteuthisnithoteuthis antillar v yllur er Argonauta nodosus a Ocythoe tuberculata lanae Haliphronremoctopus atlanticus violaceus inar T Ille i amp a pelagica T Opisthoteuthis calif odarodes sagittatus V yr x argentin a Car Pter T Atlanta peronii odarodes pacificus Eusergestes similis Eusergestes arcticus itanicus ygioteuthis microlampas Pter tr Sergia rob wii Pyroteuthis margar amii Gennadas elegans ygioteuthisAbr giardi olatilis Systellaspisapasiphae debilis ussulcatifrons arctus Acantheph us ar alis alia redfieldi P us maur Alpheus glaberur HistioteuthisL heteropsis um ynchus ycoteuthis lor Scyllaalir n viatumvegica P olybius henslo bre Histioteuthis bonnellii P Pleuroncodes planipes Histioteuthis re Axius stirh Octopoteuthis sicula itif Nyctiphanes austr Octopoteuthis nielseni iger er Nematoscelisyctiphanes megalops vnooidesr a Stylocheiron ab matus Mastigoteuthis dentata a Megan atulus v Euphausia pacifica Chiroteuthis v ersa Platyscelus o usculumis Br Platyscelus ar Ancistroteuthisachioteuthis lichtensteinii r yscelus macrocephalusuirostr i Platyscelusach serr Br yscelus cr On ach On ykia rob er Br ychoteuthisOn borealijaponica anii ustulumvillei ychoteuthis banksiiiisei Leptocotis ten losse usta Streetsia challengea b r arapronoe cr borealis P ylomer Berr Anch yteuthis anon Phrosina semilunata Gonatus calif imno macropa Pr ia galba Gonatus steenstychusr Hyper orniensis Themisto gaudichaudiibosa ana Gonatus berr Vibilia gib y upi Gonatus p Lanceola sa yi Neognathophausia gigas Gonatus oyrosn Engr Neognathophausia ingens Engr aulis mordax inii aulis encr yx Idotea metallica Engr asicolus Echinosquilla guer aulis japonicus Odontodactylusus darwiniihansenii Clupea pallasii Cosmocalan Sardinella aur Calanus finmarchicus Sardinops sagaxita Clausocalanus furcatus Sardina pilchardus Farranula gibbula Bathylagoides w Pseudevadne tergestina esethi Bathylagus stilbius Thaleichth Sebastes zacentrus ys pacificus Sebastes wilsoni Vinciguerria lucetia Sebastes proriger Bathophilus flemingi Sebastes bre vispinis Photonectes margarita Sebastes aleutian aldiviae Sebastes miniatusus Melanostomias v a Sebastes helv noptyx obscur Ster Sebastes maligeromaculatus diaphanaersii Sebastes diploproa Helicolen Argyropelecus olatof rius i Anarrhichtush dactylopter Maurolicus imper ueller Bath ys ocellatus us Maurolicus m Anoplopomayagon fimbr us agassiziisso us pentacanthus us r P Chlorophthalm en eristedion gr Arctoz yrenoides Eutr ingens Medialunaigla gu calir f ia Lestidiops sph acile Lestidiops r Diplodus sargusnardus Boops boops alepis coregonoidesyalina Spondyliosoma canthaor r Par alepis speciosa niensis Par Sudis h Capros aper Antigonia capros wii Lactor alepis affinis ao Magnisudis atlantica Halichoeres nicholsi ini Microstomia diaphana MacropaOmosudisr us pha lo r us Glyptocephalus zachir us nikpar eni Pleuronichth Anotopter Cithar Diaphus theta is us pacificus Anotopter Ar ular T noglossus imper r ichth Diaphus luetk T achur r ys decurrens i Seleneachu setapinnisr ys sordidus issoer T us symmetr ii r us Notoscopelus elongatus Cor achinotus o us tr us T r letonbeania cren Scomberesachuroyphaena hippur Tar achur ialis Myctophum punctatumElectrona r xican Cololabis sair Benthosema glaciale Belone belone icus wnsendi us Strongylur us japonicus us mingii i Ather v yctophum crocus mke Mugil cephalus atus ar Blennius ocellar itter japonicus Lobianchia gemellar ius Scomber scombr inopsis calif x saur Protom Centrolophus niger iphotur Icichth r is us T Br a e a icar Br us T Pter ama br atoscopelus to ar us us Diplospin ama orcini xilis achium r P is i Psenes pellucidus yctus crocodilus Cubiceps gr xim Katsuw epr actes asper Cer G achius leucopsar i ys lockingtoni atoscopelusatoscopelus maderensis w achium regale ycomb yctus alatus aiti e ylor ypter m or Cer us vulgar ilus simillim Cer yctus intr ama ipes

p is niensis LampanNannobr Stenobr ach a ta y

on l us m niensis luccius productus u us petersii Nannobr us parf us

Lampan s us cuvier or us pelamis us tr Lampan s Mer ur acilis Chilar e ultistr

us atlanticus Gaidropsar r

Microgadus pro p us aequoreus us athron ur

e Nealotus t r

Mullus barbatus agon n

iatus achipter Melamphaes lugubr Chiasmodon niger s r Bar Micromesistius poutassou etr T agon Desmodema polystictum T

Rhinogobiops nicholsii Entelur Scopelogadus bispinosus etr

T

Syngnathus calif ~240 unique species and ~460 unique taxa identified Max frequency of occurrence observed for prey species Typical focal prey species There is a need to model change in complex ecological communities using generalisable traits Traits for mediating predator-prey interactions

+ + + +

Habitat Refuge/ Physical Body position & diel migration avoidance defense shape behaviour

Using published, grey literature & FishBase + SeaLifeBase (Froese & Pauly 2000) 100 Habitat attributes

75

benthic 50

Number of Species demersal 25 benthopelagic

0 reef−associated continental shelf oceanic coastal continental slope midwater Horizontal Habitat 0 m epipelagic tical Habitat

Horizontal habitat e r 200 m V mesopelagic 1000 m bathypelagic 2000 m 0 25 50 75 Number of Species Habitat attributes

benthic

demersal

100 benthopelagic

midwater 0 m 50 epipelagic tical Habitat

Number of Species ?

e r 200 m V mesopelagic 1000 m 0 bathypelagic UN 0 1 2000 m Diel Migration 0 25 50 75 Number of Species Physical defenses

100 100

50 50 Number of Species Number of Species

0 0 0 1 0 1 Refuge use Presence of physical defenses

Predator avoidance/Refugia use Body shape

75

50 Number of Species 25

0 compressiform globiform fusiform elongated eel−like Body Shape Traits for mediating predator-prey interactions

✓ ✓ ✗ ✗ ✓

+ + + +

Habitat Refuge/ Physical Body position & diel migration Avoidance defense shape behaviour Multivariate trait-based analyses of tuna diets

How much variation is explained by traits?

Species Species predictive traits

Y X2 • Frequency occurrence (FO%) Morphological • Abundance (%N) Habitat position • Biomass (%W) Predator avoidance Physical defenses Aggregation behaviour Explanatory variables Morphometrics

X1 Species traits

• Geography Explanatory variables • Time series Y~X1*X2 • Temperate metrics (SST, temp anomalies) Fourth-corner results • Primary productivity (Chl-a) (correlations) • PDO, ENSO indices

Fourth corner analyses using mvabund in R – Brown et al. (2014) Methods Ecol Evol Species traits Physical traits physical defense 0.4

elongate Body shape fusiform 0.3

globiform

compressiform 0.2 Habitat use traits refuge use

diel migrant 0.1 Horizontal reef-associated habitat oceanic 0.0 continent slope

continent shelf −0.1 coastal

Vertical habitat bathypelagic

mesopelagic −0.2

epipelagic

benthopelagic −0.3

demersal

benthic −0.4

acific acific anean

SW P NE P NE Atlantic Mediterr Ocean Basin Species traits Physical traits physical defense 0.4

elongate Body shape fusiform 0.3 Horizontal reef−associated coastal continental shelf continental slope globiform Habitat oceanic

compressiform 75 0.2

Habitat use traits ance(%) refuge use 50 diel migrant 0.1

Horizontal reef-associated habitat 25

oceanic

0.0 Frequency of Occu r 0 continent slope Mediterranean NE Atlantic NE Pacific SW Pacific Ocean Basin continent shelf −0.1 coastal benthic benthopelagic mesopelagic Vertical Habitat demersal epipelagic bathypelagic Vertical habitat bathypelagic −0.2 mesopelagic 75 ance(%)

epipelagic 50 benthopelagic −0.3

25 demersal

benthic −0.4 0 Frequency of Occu r Mediterranean NE Atlantic NE Pacific SW Pacific Ocean Basin acific acific anean

SW P NE P NE Atlantic Mediterr Ocean Basin Species traits Physical traits physical defense 0.4 Body Shape compressiform globiform fusiform elongated eel−like

elongate Body shape 75 fusiform 0.3 ance(%)

globiform 50 compressiform 0.2 Habitat use traits 25 refuge use

diel migrant 0.1 Frequency of Occu r 0 Horizontal reef-associated Mediterranean NE Atlantic NE Pacific SW Pacific habitat Ocean Basin oceanic 0.0 continent slope

continent shelf Diel Migration UN 0 1

−0.1 coastal

75 Vertical habitat bathypelagic ance(%)

−0.2 mesopelagic 50 epipelagic −0.3 benthopelagic 25

demersal

Frequency of Occu r 0 benthic −0.4 Mediterranean NE Atlantic NE Pacific SW Pacific Ocean Basin

acific acific anean

SW P NE P NE Atlantic Mediterr Ocean Basin Species traits Physical traits 0.4 physical defense Physical Defenses 0 1 elongate Body shape fusiform 0.3 75

globiform ance(%)

compressiform 0.2 50 Habitat use traits refuge use diel migrant 0.1 25 Horizontal reef-associated habitat

oceanic Frequency of Occu r 0 0.0 continent slope Mediterranean NE Atlantic NE Pacific SW Pacific Ocean Basin continent shelf −0.1 coastal Refuge 0 1

Vertical habitat bathypelagic −0.2 75 mesopelagic ance(%) epipelagic −0.3 50 benthopelagic demersal

benthic −0.4 25

acific acific anean Frequency of Occu r 0 SW P NE P NE Atlantic Mediterranean NE Atlantic NE Pacific SW Pacific Mediterr Ocean Basin Ocean Basin Trait-based predictions for future change

Conserved

Switching traits with climate

No relationship of prey traits in diets in traits of prey Predicted probability Predicted

Geography Trait-based predictions for future change

Conserved

Switching traits with climate

No relationship of prey traits in diets in traits of prey Predicted probability Predicted Cold Warm Average → Warming ocean Climate regimes Next steps: trait-mediated diet shifts in the NE Pacific

NE Pacific 100

75

50

um frequency of occurrence (%) 25 Maxi m

0 1949 1968 1969 1983 2002 2005−06 2008−10

PDO Cool Phase Warm Phase Neutral Take home points q Trait-based diet assessment can be done for tuna q Numerous complementary models to add to our analytical framework q Requires diet data at the level of inference q Ongoing effort for trait database on which to build analyses Stephanie Green & Cole Brookson @ U Alberta

Larry Crowder @ Hokpins Marine Station

Michael Jacox, Elliott Hazen & Steven Bograd @ NOAA Environmental Research Division Poster session tonight @ 6pm Cole Brookson à on trait-based mechanistic models for predator-prey interactions Piglet (Helicocranchia pfefferi) Black swallower (Chiasmodon niger) Questions? E/V Nautilus NOAA Okeanos Explorer Program Vampire squid (Vampyroteuthis infernalis) MBARI

Diaphanous hatchet fish (Sternopyx diaphana) Arturo Angulo, Research Gate