Ecography ECOG-04347 Speed, J

Home , Alaska marmot, Alticola, Collared lemming, Singing vole

Ecography ECOG-04347 Speed, J. D. M., Skjelbred, I. Å., Barrio, I. C., Martin, M. D., Berteaux, D., Bueno, C. G., Christie, K. S., Forbes, B. C., Forbey, J., Fortin, D., Grytnes, J.-A., Hoset, K. S., Lecomte, N., Marteinsdóttir, B., Mosbacher, J. B., Pedersen, Å. Ø., Ravolainen, V., Rees, E. C., Skarin, A., Sokolova, N., Thornhill, A. H., Tombre, I. and Soininen, E. M. 2019. Trophic interactions and abiotic factors drive functional and phylogenetic structure of vertebrate herbivore communities across the Arctic tundra biome. – Ecography doi: 10.1111/ecog.04347

Supplementary material Appendix 1 Supplementary methods and results.

Table A1. Full list of 76 Arctic vertebrate herbivore species indicating those used in this study and those used by Barrio et al. (2016). GenBank accession numbers are shown for the four genetic markers for all included species. Species names according to IUCN and Birdlife International.

Species Used by Genetic Used in Notes cytB COI 12S ND4 Barrio et data current al. 2016 available study Dicrostonyx groenlandicus 1 1 1 KJ556713.1 JF456464.1 AF128937.1 Dicrostonyx hudsonius 1 1 1 AJ238436.1 JF456491.1 Dicrostonyx nelsoni 1 0 0 Dicrostonyx nunatakensis 0 0 0 Dicrostonyx richardsoni 1 1 1 AJ238435.1 JF443818.1 Dicrostonyx torquatus 1 1 1 KT867537.1 Dicrostonyx unalascensis 1 0 0 Dicrostonyx vinogradovi 1 0 0 Lemmus amurensis 0 1 1 FJ025979.1 Lemmus lemmus 1 1 1 JX483908.1 Lemmus portenkoi 1 0 0 Lemmus sibiricus 1 1 1 AY219144.1 Lemmus trimucronatus 1 1 1 AF119276.1 JF456714.1 AF128943.1 Synaptomys borealis 1 1 1 AF119259.1 JF457125.1 AF128932.1 Arvicola amphibius 1 1 1 KM005047.1 AY332681.1 AF128938.1 Alticola lemminus 1 1 1 KJ556633.1 Microtus miurus 1 1 1 EF608581.1 Microtus abbreviatus 1 1 1 AF163890.1

1 Species Used by Genetic Used in Notes cytB COI 12S ND4 Barrio et data current al. 2016 available study Microtus oeconomus 1 1 1 KP190237.1 KP190307.1 AJ616853.1 Microtus middendorffii 1 1 1 AF163898.1 HM137740.1 Microtus gregalis 1 1 1 KP190244.1 KP190315.1 Microtus levis 1 1 1 FJ641161.1 EF608582.1 FJ619947.1 EF608583.1 Microtus pennsylvanicus 1 1 1 KF948531.1 KM189812.1 JN393216.1 U83806.1 Microtus longicaudus 1 1 1 KF964344.1 JF456725.1 AF128936.1 Microtus chrotorrhinus 1 1 1 AF163893.1 Microtus xanthognathus 1 1 1 AF163907.1 Microtus hyperboreus 1 1 1 HM137736.1 Myodes gapperi 1 1 1 DQ323950.1 JQ350489.1 U83808.1 Myodes rutilus 1 1 1 JX477342.1 HM165297.1 Myodes rufocanus 1 1 1 KR059903.1 JF693313.1 Sicista betulina 1 1 1 KP715861.1 Apodemus sylvaticus 1 1 1 KM582049.1 KP869163.1 AJ311131.1 Urocitellus parryii 1 1 1 Synonym: AF157931.1 KM537933.1 Spermophilus parryii Marmota broweri 1 1 1 JN024621.1 Marmota camtschatica 1 1 1 AF100715.1 JF313276.1 Marmota monax 1 1 1 AF157953.1 JF456717.1 AY227529.1 JF313286.1 Marmota caligata 1 1 1 KJ458055.1 JF313275.1 Castor canadensis 1 1 1 KY321562.1 LC144616.1 AY012111.1 JQ663965.1 Ondatra zibethicus 1 1 1 KT376465.1 JF456977.1 JN315625.1 U83809.1 Erethizon dorsatum 1 1 1 KC463889.1 JF456596.1 AY012118.1 Ochotona hyperborea 1 1 1 KR076823.1 DQ347441.1 AY012127.1 EU549756.1 Ochotona turuchanensis 1 1 1 EF567056.1 DQ347468.1

2 Species Used by Genetic Used in Notes cytB COI 12S ND4 Barrio et data current al. 2016 available study Ochotona collaris 1 1 1 KP411020.1 Lepus americanus 1 1 1 KM261475.1 U58923.1 Lepus arcticus 1 1 1 HQ596461.1 JF443819.1 Lepus othus 1 1 1 HQ596479.1 Lepus timidus 1 1 1 DQ882959.1 HM232960.1 HM232960.1 Rangifer tarandus 1 1 1 Both wild and semi- KJ138217.1 JF443494.1 AY184438.1 domestic populations used in current study, while Barrio et al. 2016 used only wild populations Ovibos moschatus 1 1 1 U17862.1 JF443354.1 AY670662.1 Alces alces 1 1 1 KC337273.1 KX859263.1 Alces americanus 1 1 1 M98484.1 JF443170.1 Ovis dalli 1 1 1 AF034728.1 JF443359.1 AY670664.1 Ovis nivicola 1 1 1 AJ867265.1 Y09259.1 Bison bison 1 1 1 AF036273.1 JF443195.1 Ovis aries 1 1 0 Excluded as non-native domestic livestock within study region Cygnus columbianus 1 1 1 Including Cygnus EU585642.1 DQ433560.1 columbianus ssp. bewickii Cygnus cygnus 1 1 1 EU585643.1 GU571360.1 AY164523.1 Cygnus buccinator 1 1 1 AY509690.1 AY666404.1 U59667.1 Anser brachyrhynchus 1 1 1 EU585614.1 GU571244.1 Anser albifrons 1 1 1 EU585612.1 DQ433314.1 AY164531.1

3 Species Used by Genetic Used in Notes cytB COI 12S ND4 Barrio et data current al. 2016 available study Anser anser 1 1 1 AY427814.1 GU571242.1 AY164530.1 DQ468124.1 Anser fabalis 1 1 1 EU585618.1 FJ808625.1 AY164514.1 Anser erythropus 1 1 1 EU585617.1 GU571729.1 Chen caerulescens 1 1 1 Synonym: Anser DQ434537.1 caerulescens Chen rossii 1 1 1 EU914156.1 DQ434538.1 U83734.1 Chen canagica 1 1 1 EU585615.1 AF173714.1 Branta canadensis 1 1 1 EU585629.1 DQ434443.1 AF173715.1 Branta hutchinsii 1 1 1 AY072593.1 DQ434479.1 Branta bernicla 1 1 1 HM063580.1 GU571279.1 HM063557.1 Branta leucopsis 1 1 1 EU585630.1 GU571283.1 Branta ruficollis 1 1 1 EU585631.1 Anas penelope 1 1 1 AF059107.1 GU571239.1 AY164518.1 Anas americana 1 1 1 AF059103.1 DQ433309.1 Lagopus lagopus 1 1 1 EF571187.1 GU571438.1 AF222583.1 Lagopus muta 1 1 1 AY156346.1 DQ433738.1 KC785614.1 Lagopus leucura 0 1 1 AF230171.1 DQ433716.1 AF222584.1 Total 73 74 70

4 Figure A1 The phylogenetic tree of 70 Arctic vertebrate herbivore species. The five main clades are coloured. From top: Anseriformes (light blue), Galliformes (dark blue), Lagomorpha (yellow), Rodentia (red), Artiodactyla (green).

5 Table A2. Description of the functional traits used to develop the functional classification of Arctic vertebrate herbivores. Abbreviations for trait categories in the column Trait quantification refer to abbreviations used in Appendix S4. Numeric values for factors refer to the order for ordered factors.

Trait Trait function Trait quantification Variable type Importance of Forbs If the herbivore species feeds mainly on Scored 0-3 depending on the frequency in diet. Discrete, ordered from main arctic forbs, it will likely have a large impact 0; non-existing low to high importance functional on this plant functional group 1; low importance, (0-10% of an average diet or in diet groups of Graminoids If the herbivore species feeds mainly on known usage from only some populations; diet plants in the graminoids, it will likely have a large proportions above 0.5 should be infrequent) herbivore’s diet impact on this plant functional group 2; medium importance (10-50% of an average diet Shrubs If the herbivore species feeds mainly on or used by most populations during most seasons; shrubs, it will likely have a large impact proportions can vary from low to high) on this plant functional group 3; high importance (>50% of an average diet or Mosses If the herbivore species feeds mainly on known usage of medium to high proportions from mosses, it will likely have a large impact all populations and seasons) on this plant functional group Lichens If the herbivore species feeds mainly on lichens, it will likely have a large impact on this plant functional group Diet type How selective or generalist a herbivore According to Shipley et al 2009: Factor, ordered from species is will determine if its impacts Obligatory generalist (OG, 1) generalist to specialist affect only certain plant species, or the Facultative generalist (FG, 2) effects spread across several plant Facultative specialist (FS, 3) species Obligatory specialist (OS, 4)

Gut type Gut morphology will determine what Type of gut: Factor, ordered from plants or plant parts can be eaten by undifferentiated (U, 1) inefficient to efficient the herbivore hindgut fermenter (HF, 2) ruminant (R, 3)

6 Trait Trait function Trait quantification Variable type Belowground feeding Belowground feeding by herbivores can Type of belowground feeding: Unordered factor have distinct impacts on vegetation Burrowing (B) Grubbing (G) None (N) Body mass Key variable in trophic ecology, Body mass (grams) Continuous reflecting both feeding and predation ecology. Mobility Wide ranging herbivores will have an Ability to move between the 100x100 km pixels Binary impact on vegetation over larger spatial used in analyses: scales than herbivores with reduced Yes (1) mobility No (0) Group size summer Larger groups of herbivores foraging Group size categories: Factor, ordered from together will have a more intense effect solitary (S, 1) solitary to large groups on vegetation and behaviour in relation small group (SG, 2) to predation family group (FG, 3) winter Larger groups of herbivores foraging large group (LG, 4) together will ha e a more intense effect on vegetation and behaviour in relation to predation Population dynamics (cyclicity) The effects of herbivores can also vary Prevalence of cyclic populations: Factor, ordered from low over time, from variable (cyclic) impacts non-cyclic (NC, 1) to high degree of when herbivores have peaks in cyclic/non-cyclic (C/NC, 2); when some populations temporal variation in abundance, to uniform (noncyclic). of a species cycle and others do not. population size These affect trophic dynamics in Cyclic (C, 3) relation to vegetation and plants. Litter/clutch size Larger litter/clutch sizes underlie Litter size, number of offspring per year. Continuous population dynamics, notably growth

7 Trait Trait function Trait quantification Variable type rates, with impacts on trophic dynamics. Wintering strategy Winter strategy controls trophic Wintering strategy: Factor, ordered from not interactions in the Arctic during winter. Not present in the Arctic during winter (NP, 1) present to active above Several species migrate out of the Arctic Active below ground (BS, 2) snow during winter. Active above ground (AAS, 3) Habitat Main habitat affects the type of trophic Habitat class: Factor interactions Terrestrial (T) Limnic (L)

8 Table A3. Functional traits of Arctic vertebrate herbivores related to what they eat: their diet and the contribution of different plant groups to their diet (forbs, graminoids, shrubs, mosses and lichens, scored 0-3 depending on the frequency in diet, with 3 being highest) , gut type, occurrence of belowground feeding (BG) ; and traits related to how much plant biomass they eat (body mass, g) , their population dynamics, reproductive output (litter size, nr of offspring per year) , group size in summer and winter, wintering strategy, habitat and their ability to move between 100x100 km pixels (mobility) . Diet type: OG obligatory generalist, FG facultative generalist, OS obligatory specialist, FS facultative specialist. Gut type: HF hindgut fermenter, R ruminant, U undifferentiated. Belowground feeding: B burrowing, G grubbing, N none. Population dynamics: C cyclic, NC non-cyclic, C/NC cyclic/non-cyclic when some populations of a species cycle and others do not. Group size: LG large group, FG family group, SG small group, S solitary (for analyses FG and SG were pooled together) . Winter: AAS active above ground, ABS active below ground, NP not present. Habitat: T terrestrial, L limnic. Data were compiled by different experts (see initials corresponding to author names, DE – Dorothee Ehrich, Arctic University of Norway, SR – Sonia Rozenfeld, Russian Academy of Sciences) ; relevant references are indicated by superscripts where needed. E Data extrapolated from species in the same genus. Species in rows shaded in grey were not included in the functional classification due to lack of phylogenetic data or status as livestock. However, they are included in this table for completeness. For all species, values provided represent averages across Arctic populations and subspecies.

oss raminoid hrub orb ichen winter F G S M L Winter Winter strategy Habitat Mobility References Order/Family Binomial Initials Diet type Gut Belowground mass Body Population dynamics size Litter size Group summer size Group 1–6 Rodentia/Cricetidae Dicrostonyx groenlandicus EMS OG 3 2 3 1 1 HF B 54.40 C 3.78 S S ABS T no 3,6–9 Rodentia/Cricetidae Dicrostonyx hudsonius NL OGE 2 2 2 1 0 HFE B 57.00 C 3.49 SE SE ABSE T no 3,10 Rodentia/Cricetidae Dicrostonyx nelsoni NL OGE 2 2 3 1 0 HFE BE 60.85 CE 3.5 SE SE ABSE TE no 3,11,12 Rodentia/Cricetidae Dicrostonyx nunatakensis NL OGE 2 2 3 1 0 HF B 60.82 CE 3E S S ABS T no 3,10,11,13 Rodentia/Cricetidae Dicrostonyx richardsoni NL OG 3 3 3 0 0 HF B 55.00 C 3.5 SG S ABS T no 3,6,14–18 Rodentia/Cricetidae Dicrostonyx torquatus NS OG 2 2 3 0 0 HF B 85.00 C 3.85 S S ABS T no 3,6,11,19 Rodentia/Cricetidae Dicrostonyx unalascensis EMS OG 3 2 3 1 0 HFE BE 60.84 C 5.6 SE SE ABSE TE no 3,20,21 Rodentia/Cricetidae Dicrostonyx vinogradovi DE OG 2 2 3 0 0 HF B 60.78 C 5.6 S S ABS T no 15,22 Rodentia/Cricetidae Lemmus amurensis NS OS 1 1 0 3 0 HF B 43.70 C 5.5 S S ABS T no 1,3,6,23,24 Rodentia/Cricetidae Lemmus lemmus EMS FG 1 3 1 3 0 HF B 47.50 C 5.1 S S ABS T no 3,21 Rodentia/Cricetidae Lemmus portenkoi DE FS 2 2 0 3 0 HF B 55.76 C 5.75 S S ABS T no 3,6,16,17,20,25,26 Rodentia/Cricetidae Lemmus sibiricus DE, NS FG 2 3 1 0 0 HF B 52.27 C 6.39 S S ABS T no 1,3–6,27,28 Rodentia/Cricetidae Lemmus trimucronatus EMS FG 1 3 1 3 0 HF B 69.82 C 6.86 S S ABS T no 3,6,11 Rodentia/Cricetidae Synaptomys borealis EMS FG 2 3 1 0 0 HFE B 21.30 C/NCE 4.27 SE SE ABS T noE 3,6,29–35 Rodentia/Cricetidae Arvicola amphibius KH OG 3 3 1 0 0 HF B 120.00 C/NC 4.76 S S ABS T no

oss raminoid hrub orb ichen F G S M L winter Order/Family Binomial Initials Diet type Gut Belowground mass Body Population dynamics size Litter size Group summer size Group Winter strategy Habitat Mobility References 3,36 Rodentia/Cricetidae Alticola lemminus NS FS 1 1 1 2 2 HF BE 33.56 NC 6.17E S S ABS T no 3,6,37,38 Rodentia/Cricetidae Microtus miurus EMS OG 2 2 2 1 1 HF B 41.00 C/NC 3.89 S FG ABS T no 3,6,39 Rodentia/Cricetidae Microtus abbreviatus EMS OGE 2 2 2 1 1 HFE BE 62.00 C 3 SE FGE ABSE TE no 3,6,23,40,41 Rodentia/Cricetidae Microtus oeconomus EMS OG 1 3 2 1 1 HF B 34.38 C 5.62 S S ABS T no 3,6,16,18,42–44 Rodentia/Cricetidae Microtus middendorffii NS FG 2 3 2 0 0 HF B 36.49 C 5.49 S S ABS T no 3,16,18,25,26,42,45–49 Rodentia/Cricetidae Microtus gregalis NS OG 3 3 2 0 0 HF B 47.50 C 9 FG S ABS T no 3,50–55 Rodentia/Cricetidae Microtus levis KH OG 3 3 0 0 0 HF B 35.49 C 5.2 S S ABS T no 3,6,56,57 Rodentia/Cricetidae Microtus pennsylvanicus EMS OG 2 2 1 0 0 HFE B 36.75 C 5.16 S S ABS T no 3,6,58,59 Rodentia/Cricetidae Microtus longicaudus EMS OG 2 1 1 0 0 HFE B 46.71 C 4.73 S S ABS T no 3,6,60,61 Rodentia/Cricetidae Microtus chrotorrhinus EMS OG 2 1 2 1 0 HFE B 39.00 NC 3.58 S S ABS T no 3,6,62 Rodentia/Cricetidae Microtus xanthognathus EMS OG 2 3 1 0 1 HFE B 125.75 NC 8.1 S S ABS T no 6,14,16,18,42,43,63 Rodentia/Muridae Microtus hyperboreus NL FG 2 3 2 0 0 HF B 36.00E C 5.49 S S ABS T no 3,6,64 Rodentia/Cricetidae Myodes gapperi EMS FG 1 1 2 1 1 HF B 19.83 NC 5.37 S S ABS T no 3,6,65–67 Rodentia/Cricetidae Myodes rutilus KH FG 1 1 2 1 2 HF N 19.94 C/NC 5.6 S FG ABS T no 3,6,23,68 Rodentia/Cricetidae Myodes rufocanus EMS OG 3 3 3 1 0 HF B 36.43 C 5.01 S S ABS T no 3,69 Rodentia/Dipodidae Sicista betulina NS OG 3 3 3 0 0 HF N 8.00 NC 5.15 S S NP T no 3,6,70–75 Rodentia/Muridae Apodemus sylvaticus NL OG 2 3 3 0 0 HF B 30.45 NC 5.16 S S ABS T no 3,6,76 Rodentia/Sciuridae Urocitellus parryii ICB FG 3 3 2 1 1 HF B 759.99 NC 6.5 LG FG NP T no 3,77,78 Rodentia/Sciuridae Marmota broweri ICB FG 3 3 1 1 1 HF B 3405.00 NC 4.5 LG FG NP T no 3,6,79 Rodentia/Sciuridae Marmota camtschatica ICB FG 3 3 1 1 1 HF B 3500.00 NC 4.99 LG FG NP T no 3,6,80,81 Rodentia/Sciuridae Marmota monax ICB FG 3 3 2 1 1 HF B 3801.72 NC 4.1 S FG NP T no 82,83 Rodentia/Sciuridae Marmota caligata ICB FG 3 3 1 1 1 HF B 4900.00 NC 3.9 LG FG NP T no 3,6,84,85 Rodentia/Castoridae Castor canadensis NL OG 1 1 3 0 0 HF N 21820.00 NC 3.6 SG SG AAS L no 3,6,86,87 Rodentia/Cricetidae Ondatra zibethicus ICB FG 2 3 0 0 0 HF B 1065.75 C 6.55 LG FG ABS L no 3,6,88–90 Rodentia/Erethizontidae Erethizon dorsatum DB FS 1 1 3 0 0 HF N 7085.33 C/NC 1 S S AAS T no 3,6,91,92 Lagomorpha/Ochotonidae Ochotona hyperborea EMS, ICB FG 2 1 2 0 1 HF N 120.00 NC 3.02 S S ABS T no 3,91,93 Lagomorpha/Ochotonidae Ochotona turuchanensis EMS, ICB FGE 2 1 2 0 1 HF N 142.15 NC 3.4 S S ABS T no 3,6,94,95 Lagomorpha/Ochotonidae Ochotona collaris EMS, ICB FG 1 3 3 1 1 HF N 129.00 NC 3.35 S S ABS T no 3,6,96–98 Lagomorpha/Leporidae Lepus americanus JF FG 2 1 3 0 0 HF N 1710.02 C/NC 3.54 S S AAS T no

oss raminoid hrub orb ichen F G S M L winter Order/Family Binomial Initials Diet type Gut Belowground mass Body Population dynamics size Litter size Group summer size Group Winter strategy Habitat Mobility References 3,6,99 Lagomorpha/Leporidae Lepus arcticus JF FG 1 1 3 1 1 HF N 4405.04 NC 5.62 S S AAS T no 3,6,100 Lagomorpha/Leporidae Lepus othus JF FG 1 2 3 0 0 HF N 4805.96 NC 6.15 S S AAS T no 3,6,101,102 Lagomorpha/Leporidae Lepus timidus JF FG 2 2 3 1 1 HF N 3048.00 NC 3.16 S S AAS T no ÅP, VTR, 3,103–112 Artiodactyla/Cervidae Rangifer tarandus SR, DF FG 2 3 2 1 2 R N 86033.98 NC 1 LG LG AAS T yes 3,6,104,111–116 Artiodactyla/Bovidae Ovibos moschatus JBM FG 2 3 2 1 1 R N 340501.06 NC 1.01 SG SG AAS T yes 3,6,117,118 Artiodactyla/Cervidae Alces alces JDMS FS 2 1 3 0 0 R N 356998.16 C/NC 1.25 S S AAS T yes 3,6,119 Artiodactyla/Cervidae Alces americanus JF FG 2 2 3 1 1 R N 541460.44 NC 1 S S AAS T yes 3,6,120–123 Artiodactyla/Bovidae Ovis dalli EMS, ICB OG 2 3 2 0 1 R N 55650.62 NC 1.22 FG LG AAS T yes 3,6,124,125 Artiodactyla/Bovidae Ovis nivicola BM FG 1 3 2 1 2 R N 90000.00 NC 1 LG LG AAS T yes 3,6,126–132 Artiodactyla/Bovidae Bison bison DF FG 1 3 1 0 1 R N 579255.28 NC 0.98 LG LG AAS T yes 3,6,133–137 Artiodactyla/Bovidae Ovis aries BM FG 2 3 2 1 1 R N 50000.00 NC 1.19 FG LG NP T no 138–144 Anseriformes/Anatidae Cygnus columbianus ER FG 3 3 2 1 0 U G 6298.81 NC 3.85 LG LG NP L yes 3,145–148 Anseriformes/Anatidae Cygnus cygnus ER FG 3 3 1 1 0 U G 9349.99 NC 4.47 LG LG NP L yes 3,149–151 Anseriformes/Anatidae Cygnus buccinator ER FG 3 3 2 2 0 U G 11071.13 NC 5 LG LG NP L yes 3,152–154 Anseriformes/Anatidae Anser brachyrhynchus JDMS FG 3 3 1 1 0 U G 2642.04 NC 4 LG LG NP L yes 3,155,156 Anseriformes/Anatidae Anser albifrons SR FG 2 3 2 1 0 U G 2506.39 NC 5 LG LG NP L yes 3,157 Anseriformes/Anatidae Anser anser IT, SR FG 1 3 1 0 0 U G 3302.41 NC 5 LG LG NP L yes 3,155,158,159 Anseriformes/Anatidae Anser fabalis IT, SR FG 3 3 2 1 0 U G 2754.73 NC 4 LG LG NP L yes 3,158–161 Anseriformes/Anatidae Anser erythropus SR FG 1 3 1 0 0 U N 1755.50 NC 5 LG LG NP L yes 3,155,162,163 Anseriformes/Anatidae Chen caerulescens EMS, JDMS FG 2 3 1 1 0 U G 2636.15 NC 4 LG LG NP L yes 3,155,163,164 Anseriformes/Anatidae Chen rossii EMS, JDMS FG 2 3 1 2 0 U N 1635.99 NC 4 LG LG NP L yes 3,155 Anseriformes/Anatidae Chen canagica SR FS 1 3 1 0 0 U N 2136.49 NC 4.9 LG LG NP L yes 3,165 Anseriformes/Anatidae Branta canadensis NL FG 2 3 1 0 0 U G 2811.68 NC 4 LG LG NP L yes 165,166 Anseriformes/Anatidae Branta hutchinsii NL FG 2 3 1 0 0 U G 2050.00 NC 4 LG LG NP L yes 3,154,158,159,167–169 Anseriformes/Anatidae Branta bernicla JDMS, SR FG 2 2 1 2 0 U N 1277.91 NC 4 LG LG NP L yes 3,153,154,158,159,170 Anseriformes/Anatidae Branta leucopsis JDMS, SR FG 2 3 1 2 0 U N 1683.97 NC 5 LG LG NP L yes –173 3,156,174 Anseriformes/Anatidae Branta ruficollis SR FS 3 2 2 0 0 U N 1226.47 NC 5 LG LG NP L yes 3,155,175 Anseriformes/Anatidae Anas penelope NL FG 3 3 0 0 0 U N 770.03 NC 8 SG LG NP L yes

oss raminoid hrub orb ichen F G S M L winter Order/Family Binomial Initials Diet type Gut Belowground mass Body Population dynamics size Litter size Group summer size Group Winter strategy Habitat Mobility References 3,176 Anseriformes/Anatidae Anas americana NL FG 3 3 0 0 0 U N 754.61 NC 8 SG LG NP L yes 3,177–179 Galliformes/Phasianidae Lagopus lagopus KC FG 1 1 3 1 0 HF N 566.86 C/NC 9 S LG AAS T yes 3,178,180–183 Galliformes/Phasianidae Lagopus muta KC FS 1 1 3 1 0 HF N 535.30 C/NC 9 S LG AAS T yes 3,184–186 Galliformes/Phasianidae Lagopus leucura KC FG 1 1 3 0 0 HF N 354.97 C/NC 5.5 S LG AAS T yes

12 Body mass (grams log) Gut type Group size S Group size W

40 30 30 25 30 20 20 20 15 10 10 10 5

0 5 10 15 0 0 0 2 4 6 8 10 12 14 solitary solitary ruminant large_group large_group small_group small_group undifferentiated hindgut_fermenter Popul dynamics Habitat type Wintering strategy Litter size 50 30 40 40 25 30 20 30 20 15 20 10 10 10 5

0 0 0 0 5 10 15 20 0 2 4 6 8 limnic cyclic terrestrial noncyclic not_present cyclic_noncyclic active_below_snow active_above_snow Belowground feeding Mobility Diet type Forbs in diet 30 40 25 20 30

15 20 10 10 5

0 0 10 20 30 40 0 0 5 10 15 20 25 30 no yes 1 2 3 none ob_gen ob_spe fac_gen fac_spe grubbing burrowing

Graminoids in diet Shrubs in diet Mosses in diet Lichens in diet 0 10 20 30 40 0 5 10 15 20 0 5 10 20 30 0 10 20 30 40

1 2 3 0 1 2 3 0 1 2 3 0 1 2

Figure A2 Distribution of trait values between species. See Table S2 for a description of traits and trait categories, and Table S3 for trait descriptions by species.

13 Table A4 Phylogenetic signal (K) of continuous and ordinal functional traits. Higher values (values >1) show phylogenetic conservatism of traits. Gut type was the most phylogenetically-conserved trait across herbivore species, followed by mobility, wintering strategy and habitat type. Diet type and composition were amongst the most phylogenetically- diversified traits.

Trait K Body mass 0.003 Litter clutch size 0 Habitat type 2.094 Belowground feeding 0.001 Mobility 8.33 Forbs in diet 0 Graminoids in diet 0 Shrub in diet 0.001 Mosses in diet 0 Lichens in diet 0.094 Group size 1.362 Group size summer 0.531 Group size winter 1.362 Population dynamics 0.295 Gut type 5.155 Diet type 0 Wintering strategy 2.556

14 Table A5 Summary table of factorial analysis of mixed data, showing eigenvalues and percent of variance explained by the first six components.

Component Eigenvalue Percentage of variance Cumulative percentage of variance

1 6.10 33.92 33.92

2 3.68 20.44 54.35

3 1.73 9.60 63.95

4 1.43 7.94 71.90

5 0.90 5.00 76.90

6 0.87 4.84 81.74

15 (a) Quantitative variables (b) Quaulitative variables

1.0

body_mass_log gut_type_order active_above_snow

0.5 diet_item_shrub diet_item_lichen

diet_type_order 2 no belowground feeding

group_size_winter_order 0.0 diet_item_moss group_size_summer_order mobile Dim 2 (20.44%) 2Dim (20.44%) 2Dim terrestrial population_dynamics_order 0

diet_item_graminoid immobile -0.5 diet_item_forb limnic Litter_clutch_size active_below_snow not_present grubbing burrowing

-2 -1.0

-1.0 -0.5 0.0 0.5 1.0 -2 -1 0 1 2 3 4

Dim 1 (33.92%) Dim 1 (33.92%)

Figure A3 (a) Quantitative variables plotted as vectors along the first two axes from the factorial analysis of mixed data and (b) qualitative variable levels plotted along the first two axis. Contributions of each of the first two axes are shown.

16 60 dim1 dim2 dim3 dim4 50 dim5 dim6

20 variable contribution to dimensions contribution variable

0 Mobility Gut type Diet type Diet Body mass Body Habitat type Habitat Forbs in diet Forbs in Shrubs in diet in Shrubs Lichens in diet in Lichens Mosses in diet in Mosses Litter/clutch size Litter/clutch Group size winter size Group Graminoids in diet in Graminoids Wintering strategy Wintering Group size summer size Group Population dynamics Population Belowground feeding Belowground

Figure A4 Contributions of each variable to the six dimensions of the factorial analysis of mixed data. Variable contributions sum to 100 for each dimension.

17 Figure A5 Functional classification of Arctic vertebrate herbivores based on hierarchical clustering of factorial analysis of mixed data. The three main functional groups are coloured. From top: (1) limnic-habitat associated herbivores, migrating outside the Arctic for winter, undifferentiated guts for which graminoids are an important diet component (waterfowl; paragon Anser anser, green), (2) immobile, burrowing species with hindgut fermenting digestive physiology (paragon Synaptomys borealis, black), and (3) large-bodied facultative-generalist species for which shrubs and lichens are an important diet component (paragon Lepus timidus, red). See Figure S7 for functional group characterisation by traits. 18 cluster 1 cluster 2 cluster 3

Alces americanus Ovis nivicola Alces alces Rangifer tarandus Bison bison 4 Ovibos moschatus

Ovis dalli Erethizon dorsatum

Lepus arcticus Lepus timidus LagopusCastor leucura canadensis Lepus americanusLepus othus Lagopus muta cluster 2 2 Lagopus lagopus

OOchotonachotonaOchotona turuchanensis hyperborea collaris Myodes rutilus

Dim 2 (20.44%) Dim2 Alticola lemminus

Myodes gapperi Chen canagica 0 Marmota monax Branta ruficollis AnserBranta erythropus bernicla

Microtus chrotorrhinus Marmota caligata Chen rossii Microtus miurus Marmota broweri Microtus abbreviatus Sicista betulina Marmota camtschatica DicrostonyxDicrostonyx groenlandicus torquatusLemmus amurensis Urocitellus parryii Brantacluster Anserleucopsis 3anser Apodemus sylvaticus AnserCygnus albifrons columbianus Microtus oeconomus Branta canadensis Dicrostonyx hudsonius BrantaChenCygnusAnser caerulescens hutchinsii fabalis buccinator Microtus longicaudus Microtuscluster xanthognathus1 Cygnus cygnus MMicrotus icrotusDicrostonyxLemmusS ynaptomysmiddendorffiihyperboreus richardsonilemmus borealis Anser brachyrhynchus AnasAnas americanapenelope MicrotusMyodes Lemmuspennsylvanicus rufocanus trimucronatus LemmusArvicola sibiricus amphibius

-2 Ondatra zibethicus

MicrotusMicrotus gregalis levis 0 5 -5

Dim 1 (33.92%)

Figure A6. Functional classification plotted along the first two dimensions of the hierarchical clustering on principle components. Species labels are coloured according to clusters to match Figure S5.

19 body_mass_log.x Litter_clutch_size.x diet_item_forb.x diet_item_graminoid.x 1.0 1.0

12 8 0.8 0.8 10 6 0.6 0.6 8 3 3 0.4 0.4 6 4 2 2 1 1 4 0.2 0.2 2

2 0.0 0.0 Burrowing Large-bodied Limnic Burrowing Large-bodied Limnic Burrowing Large-bodied Limnic Burrowing Large-bodied Limnic hindgut-fermenters facultative generalists migrators hindgut-fermenters facultative generalists migrators hindgut-fermenters facultative generalists migrators hindgut-fermenters facultative generalists migrators

diet_item_shrub.x diet_item_moss.x diet_item_lichen.x Diet_type 1.0 1.0 1.0 1.0

0.8 0.8 0.8 0.8

0.6 0.6 0.6 0.6

3 3 2 ob_spe 0.4 0.4 0.4 0.4 2 2 1 ob_gen 1 1 0 fac_spe 0.2 0 0.2 0 0.2 0.2 fac_gen

0.0 0.0 0.0 0.0 Burrowing Large-bodied Limnic Burrowing Large-bodied Limnic Burrowing Large-bodied Limnic Burrowing Large-bodied Limnic hindgut-fermenters facultative generalists migrators hindgut-fermenters facultative generalists migrators hindgut-fermenters facultative generalists migrators hindgut-fermenters facultative generalists migrators

gut_type Belowground_feeding.x Mobility.x group_size_summer

1.0 1.0 1.0 1.0

0.8 0.8 0.8 0.8

0.6 0.6 0.6 0.6

undifferentiated no belowground feeding mobile solitary 0.4 0.4 0.4 0.4 ruminant grubbing immobile large_group hindgut_fermenter burrowing small_group 0.2 0.2 0.2 0.2

group_size_winter Population_dynamics wintering_strategy Habitat_type.x

1.0 1.0 1.0 1.0

0.8 0.8 0.8 0.8

0.6 0.6 0.6 0.6

solitary noncyclic not_present terrestrial 0.4 0.4 0.4 0.4 large_group cyclic_noncyclic active_below_snow limnic small_group cyclic active_above_snow 0.2 0.2 0.2 0.2

20 Figure A7 Characterisation of three main functional groups shown in Figure S5. Boxplots are shown for the continuous trait variables (body mass and litter/clutch size), with median, interquartile range and 1.5 times interquartile range shown by lines, boxes and whiskers respectively. Spineplots are shown for the factorial trait variables, with the colour showing the trait order, the box width the number of species in each functional group and the box height the conditional relative frequency of the trait in the functional group.

21 Figure A8. Mantel statistics (Pearson correlation coefficients) between species distance matrices based on factorial anlaysis of mixed data. Correlation coefficients between the full 16-trait dataset and random selection of nine to 15 traits.

22 Table A6. List of Arctic species assumed to be predators of Arctic herbivores (excluding humans) and thus included in the variable predator diversity.

Class Order Family Species Aves Strigiformes Strigidae Aegolius funereus Aves Falconiformes Accipitridae Aquila chrysaetos Aves Strigiformes Strigidae Asio flammeus Aves Strigiformes Strigidae Bubo bubo Aves Strigiformes Strigidae Bubo scandiacus Aves Strigiformes Strigidae Bubo virginianus Aves Falconiformes Accipitridae Buteo lagopus Aves Falconiformes Accipitridae Circus cyaneus Aves Passeriformes Corvidae Corvus corax Aves Falconiformes Falconidae Falco columbarius Aves Falconiformes Falconidae Falco peregrinus Aves Falconiformes Falconidae Falco rusticolus Aves Falconiformes Accipitridae Haliaeetus albicilla Aves Falconiformes Accipitridae Haliaeetus leucocephalus Aves Falconiformes Accipitridae Haliaeetus pelagicus Aves Passeriformes Corvidae Perisoreus canadensis Aves Charadriiformes Stercorariidae Stercorarius longicaudus Aves Charadriiformes Stercorariidae Stercorarius parasiticus Aves Charadriiformes Stercorariidae Stercorarius pomarinus Aves Charadriiformes Stercorariidae Stercorarius skua Aves Strigiformes Strigidae Strix nebulosa Aves Strigiformes Strigidae Strix uralensis Aves Strigiformes Strigidae Sturnia ulula Mammalia Carnivora Canidae Canis latrans Mammalia Carnivora Canidae Canis lupus Mammalia Carnivora Mustelidae Gulo gulo Mammalia Carnivora Felidae Lynx canadensis Mammalia Carnivora Felidae Lynx lynx Mammalia Carnivora Mustelidae Martes zibellina Mammalia Carnivora Mustelidae Mustela erminea Mammalia Carnivora Mustelidae Mustela nivalis Mammalia Carnivora Mustelidae Neovison vison Mammalia Carnivora Ursidae Ursus arctos Mammalia Carnivora Ursidae Ursus maritimus Mammalia Carnivora Canidae Vulpes lagopus Mammalia Carnivora Canidae Vulpes vulpes

23 Figure A9 Pairwise plots between continuous environmental variables and response variables. Response variables are standardised as the residuals of relationships with species richness

24 Figure A10 Pairwise correlation plots between all continuous environmental variables. Lower panel show data and upper panels the absolute Pearson correlation coefficients, with text size proportional to the value.

25 Table A7 Generalised variance inflation factors (GVIF) estimated across environmental variables for each GLS model. Variables are standardised as the residuals of the relationships with species richness

Variable df Phylogenetic Functional Functional diversity diversity convergence

NDVI 1 2.04 1.81 1.96

Winter minimum 1 temperature 1.07 1.07 1.05

Habitat heterogeneity 1 1.01 1.02 1.02

Topographic 1 heterogeneity 1.07 1.03 1.05

Ice fee 1 1.01 1.02 1.01

Predator diversity 1 2.04 1.81 1.97

Zoogeographic region 2 1.01 1.06 1.02

26 Figure A11. Variograms showing spatial autocorrelation at increasing distance for each global GLS model fitted with exponential variance-covariance structures including coordinates of cell centroids as spatial variables.

27 Figure A12. Spatial patterns in diversity in terms of species richness, phylogenetic diversity, functional diversity and functional convergence of avian and mammalian Arctic vertebrate herbivore. Note species, phylogenetic and functional diversity are plotted on the same colour scale.

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Appendix 2 Multiple sequence alignment of markers COI, cytB, 12S, and ND4. The file is in

FASTA format. Provided here https://doi.org/10.6084/m9.figshare.6165923.v2

Appendix 3 Herbivore diversity maps as GIS layers. Provided here https://doi.org/10.6084/m9.figshare.6165923.v2