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Determinants of Distribu/on 1) The Niche Fundamental Niche: total range of abio/c environmental condi/ons in which a taxon can survive and reproduce From Hutchinson (1957) the n-‐dimensional hypervolume (or mul/dimensional space) that describes the range of abio/c environmental condi/ons in which a taxon can survive and reproduce (each abio/c factor is a single dimension) 1 Determinants of Distribu/on 1) The Niche 0 Growth Rate Temperature Temp (C) 0 Growth Rate 10 -‐10 20 Salinity (ppt) 10 -‐10 Growth Rate Niche Space Growth Ra te Growth Rate 10 Salinity 2 Determinants of Distribu/on 1) The Niche When do range boundaries coincide with the fundamental niche? Eastern Phoebe Northern wintering range limit hOp://upload.wikimedia.org/ wikipedia/commons/c/c7/ Eastern_Phoebe1.jpg -‐4oC January minimum isotherm (from Root 1988) Northern range limit is correlated with metabolic costs of thermoregula/on for at least 14 species (in mul/ples of basal metabolic rate: MR = ~2.5 x BMR) 3 Determinants of Distribu/on 1) The Niche When do range boundaries coincide with the fundamental niche? Northern range limit is correlated with metabolic costs of thermoregula/on for at least 14 species (MR = ~2.5 x BMR) Eastern Phoebe Northern wintering range limit (from Root 1988) hOp://upload.wikimedia.org/ wikipedia/commons/c/c7/ Eastern_Phoebe1.jpg http://ebird.org/plone/ebird/news/patterns-from-ebird-eastern-phoebe 4 Determinants of Distribu/on 1) The Niche Realized Niche: A subset of the fundamental niche comprising the actual environmental condi/ons in which a taxon survives and reproduces in nature, including bio/c factors (compe//on, preda/on, mutualism, etc). Balanus: realized niche ~ fundamental niche Chthamalus: constrained by compe//ve interac/ons to narrower realized niche How could we test this experimentally? Determinants of Distribu/on 1) The Niche Realized Niche: A subset of the fundamental niche comprising the actual environmental condi/ons in which a taxon survives and reproduces in nature, including bio/c factors (compe//on, preda/on, mutualism, etc). A natural experiment... Species A and B are closely related ~ similar niche Cordillera Vilcabamba Main Andes In the main Andes: -‐ both species present -‐ ranges do not overlap Terborgh & Weske 1975, Ecology Determinants of Distribu/on 1) The Niche Realized Niche: A subset of the fundamental niche comprising the actual environmental condi/ons in which a taxon survives and reproduces in nature, including bio/c factors (compe//on, preda/on, mutualism, etc). In a range isolated from the Andes: -‐ high eleva/on species absent -‐ low eleva/on species expands range upward Cordillera Vilcabamba Main Andes Cerros del Sira 100 km from Andes Ecological release: expansion of the realized niche of a species where few compe/tors exist but an undiminished range of resources and habitats is present Terborgh & Weske 1975, Ecology Determinants of Distribu/on 1) The Niche Important caveats to the niche as the main determinant of a species’ range: a) Species may occupy unfavourable areas b) Species may be absent in favourable areas 8 Determinants of Distribu/on 1) The Niche Important caveats to the niche as the main determinant of a species’ range: a) Species may occupy unfavourable areas • r = b + i – d – e • popula/ons can be a source (birth rate exceeds death rate; b > d) or sink (death rate exceeds birth rate; d > b) • sink popula/ons depend on immigra/on (i) from source popula/ons • peripheral popula/ons of a species range are ogen sink popula/ons b) Species may be absent in favourable areas • due to geographic barriers or isola/on c) Metapopula/on structure when suitable niche space is patchy, some patches may be occupied intermiOently 9 Determinants of Distribu/on 2) Metapopula?on structure Metapopula?on: a popula/on consis/ng of a set of subpopula/ons linked by a cycle of alterna/ng coloniza/on and ex/nc/on (Levins 1970) metapopula/on subpopula/ons 10 Determinants of Distribu/on 2) Metapopula?on structure Habitat 1 (source popula/on) Representa/on of source and sink habitats and rela/ve magnitude of the four processes that b1 > d1 determine growth and persistence of popula/ons “Excess” individuals in Habitat 1 disperse, resul/ng e1 > i1 in higher emigra/on than immigra/on Habitat 2 is able to persist due to higher i2 > e2 immigra/on than emigra/on, despite lower births than deaths b2 < d2 Metapopula/on source/sink dynamics more likely to occur toward the periphery of a species’ range Habitat 2 (sink popula/on) (based on the abundance-‐center hypothesis) Fig from ch. 4, Lomolino et al. 2006 Determinants of Distribu/on 2) Metapopula?on structure Åland Islands, Finland Glanville fri/llary buOerfly (Melitaea cinxia) Photograph courtesy of Hannu Aarnio. ~ 4000 dry meadows in 2012 Red = occupied Blue = unoccupied Ojanen, S.P et al. 2013 Ecology and Evolu>on 12 Determinants of Distribu/on 2) Metapopula?on structure 1993-‐1994 1994-‐1995 Index of synchrony Glanville fri/llary on the Åland Islands became more synchronous over /me, possibly due to increasing frequency of extreme weather events: 2005-‐2006 Year 2006-‐2007 Red down-‐poin/ng triangles = decline; Green up-‐poin/ng triangles = increase Size of the triangle ~ magnitude of per capita change Ojanen, S.P et al. 2013 Ecology and Evolu>on 13 Determinants of Distribu/on 3) Disturbance Habitat disturbance can cause abrupt range limits. Countless examples, but one in our research is the high Andean treeline: Peruvian farmers in highland communi/es set fires to maintain fresh grass sprouts for caOle Anthropogenic treeline at ~ 3400 m, upper limits of forest species High-‐eleva/on species that move upslope with climate change hit “grass ceiling” Manu Na/onal Park, Peru (photos M. Chappell, Z. Peterson) 14 Determinants of Distribu/on 3) Disturbance Disturbance-‐adapted animals? Urban adapters? American Crow (Corvus brachyrhynchos) Common Starling (Sturnus vulgaris) 15 Determinants of Distribu/on 4) Bio?c Interac?ons: Direct Compe??on Direct compe//on – tes/ng species replacements Monteverde, Tilarán Mountains, Costa Rica 1100-‐1800m Wood-Wrens (Henicorhina) Wood-Wrens (Henicorhina) White-breasted Wood-Wren Gray-breasted Wood-Wren White-breasted Wood-Wren Gray-breasted Wood-Wren Nightingale-Thrushes (Catharus) Legend Orange-billed Nightingale-Thrush Orange-billed Nightingale-Thrush Black-headed Nightingale-Thrush Black-headed Nightingale-Thrush Slaty-backed Nightingale-Thrush Slaty-backed Nightingale-Thrush 16 Determinants of Distribu/on 4) Bio?c Interac?ons: Direct Compe??on Target species: Wood-‐Wrens Gray-‐breasted Wood-‐Wren White-‐breasted Wood-‐Wren Determinants of Distribu/on 4) Bio?c Interac?ons: Direct Compe??on Target species: Nigh/ngale-‐Thrushes Slaty-‐backed Nigh/ngale-‐Thrush Black-‐headed Nigh/ngale-‐Thrush Orange-‐billed Nigh/ngale-‐Thrush Determinants of Distribu/on 4) Bio?c Interac?ons: Direct Compe??on Our previous example of species replacements and compe//on was a “natural experiment” with observa/onal evidence of ecological release. For birds that defend territories using song... we can use territory defense as a behavioural metric of compe//ve interac/ons ...and design an experiment Does interspecific compe//on reinforce range boundaries of species along eleva/onal gradients? Terborgh & Weske 1975, Ecology Jankowski et al. 2010, Ecology Determinants of Distribu/on 4) Bio?c Interac?ons: Direct Compe??on Our previous example of species replacements and compe//on was a “natural experiment” with observa/onal evidence of ecological release. For birds that defend territories using song... we can use territory defense as a behavioural metric of compe//ve interac/ons ...and design an experiment Does interspecific aggression reinforce range boundaries of species along eleva/onal gradients? Terborgh & Weske 1975, Ecology Jankowski et al. 2010, Ecology Behavioral responses recorded: • Closest approach to speaker (meters) • Latency to approach speaker (seconds) • Average length of inter-‐song intervals #@*!? Behavioral responses recorded: • Closest approach to speaker (meters) • Latency to approach speaker (seconds) • Average length of inter-‐song intervals #@*!? Determinants of Distribu/on 4) Bio?c Interac?ons: Direct Compe??on If species are interspecifically aggressive at contact zones, this supports the hypothesis of direct compe//on Playback protocol to test species aggressive responses: Trial 1 Trial 2 Observa/on (Control) Playback (Congener/ Conspecific) Observa/on (Control) Playback (Congener/ Conspecific) 8 min. obs 3 min. song, 5 min. obs 8 min. obs 3 min. song, 5 min. obs Determinants of Distribu/on 4) Bio?c Interac?ons: Direct Compe??on Closest Approach (m) Wood-‐Wrens 20 15 Gray-breasted W hite-breasted Song played to bird: a a Control Congener (other species) 10 b b 5 b b Conspecific (same species) 0 Gray-‐breasted (s/mulus) White-‐breasted (response) Graybreasted White-breasted Jankowski et al. 2010, Ecology Determinants of Distribu/on Closest Approach Closest Approach (m) to Speaker (m) 4) Bio?c Interac?ons: Direct Compe??on Nigh/ngale-‐Thrushes 20 Slaty-backed Black-headed a a 15 Song played to bird: a Control b 10 c Congener (other species) b 5 Conspecific (same species) 0 20 15 Black-headed Orange-billed a a b Slatybacked X 10 X Black-headed c b 5 0 CNTR CNGN CNSP b CNTR CNGN CNSP Orange-billed Determinants of Distribu/on 4) Bio?c Interac?ons: Diffuse Compe??on Diffuse compe??on: the combined effect of compe//on with many other species – one species is nega/vely affected by numerous other species that collec/vely cause significant deple/on of shared resources (MacArthur 1972). Add more nest boxes (i.e., cavi/es) more cavity nesters less open cup nesters Western bluebird (Sialia mexicana) American robin (Turdus migratorius) (Bock et al. 1992) 26 20 40 60 % Nest Depredated 80 100 Nest preda/on For tropical birds, nest preda/on is a major source of mortality In Manu, Peru, we are gathering data to understand: 1) how nest preda/on changes with eleva/on? 2) how does the nest predator community change with eleva/on? 0 5) Bio?c Interac?ons: Preda?on Daily Nest Preda/on Rate Determinants of Distribu/on 350$600& 750$1100& 1250$1600& Eleva/on (m) 2500$3000& 27 pearance did not decrease with elevation (linear regression, t = 0.8, P = 0.445, R2 = 0.100). The highest incidence of canary egg disappearance was at 1,650 m. I found no relationship between elevation and incidence of unknown 5) Bio?c Preda?on marksInterac?ons: (linear regression, t = ¡0.4, P = 0.681, R2 = 0.030). Burhans 2 by snakes factors mu lower-elev DiVeren predation Meta-‐analysis and ar/ficial nest study show linear decrease imposed b in nest preda/on rate with eleva/on (Central America) change alo that at low redate mo nest crypsi elevations dominate, ment in sit This stu standing o tropical fo studies, da 0 500 1000 1500 2000 2500 3000 (Faaborg 2 Skutch'1985,'Boyle'2008' Eleva/on (m) multiple s Fig. 3 Relationship between elevation and proportion of nests depreimpractica Propor/on of nests depredated Determinants of Distribu/on Determinants of Distribu/on 5) Bio?c Interac?ons: Preda?on 40 60 80 100 % Nest Daily Depredated Nest Preda/on Rate How do rates of nest preda/on change with eleva/on in Manu Na/onal Park, Peru? 0 20 Eleva/on (m) 350$600& 750$1100& 1250$1600& 2500$3000& 29 Determinants of Distribu/on 5) Bio?c Interac?ons: Preda?on 40 60 80 100 % Nest Daily Depredated Nest Preda/on Rate How do rates of nest preda/on change with eleva/on in Manu Na/onal Park, Peru? 0 20 Eleva/on (m) 350$600& 750$1100& 1250$1600& 2500$3000& 30 20 40 60 80 100 How does the nest predator community change with eleva/on in Manu Na/onal Park, Peru? 0 5) Bio?c Interac?ons: Preda?on % Nest Depredated Daily Nest Preda/on Rate Determinants of Distribu/on 350$600& 750$1100& 1250$1600& Eleva/on (m) 2500$3000& 31 20 40 60 % Nest Depredated 80 100 Catching the culprits... 0 5) Bio?c Interac?ons: Preda?on Daily Nest Preda/on Rate Determinants of Distribu/on 350$600& 750$1100& 1250$1600& Eleva/on (m) 2500$3000& 32 Determinants of Distribu/on 5) Bio?c Interac?ons: Preda?on Learning lots about the natural history of tropical birds...including who eats whom Cock-‐of-‐the-‐Rock nestling and adult (Rupicola peruviana) hOp://cdn-‐7.itsnature.org/wp-‐content/uploads/2010/04/Cock-‐of-‐the-‐Rock-‐antbird.jpg Determinants of Distribu/on -0.02 1000 1000 1500 2000 2500 1500 2000 2500 Eleva.on&(meters)& Eleva/on (m) elevation 3000 3000 Figure&2.&Rela.onship&between&bird&species&turnover&and& Daily&nest&preda.on&& Change in d aily nest preda/on rates 0.06 0.04 0.00 0.45 0.02 0.55 Species&Turnover& Daily&Preda.on& 0.35 turnover Species turnover is high = many species in the community have range boundaries in the same place 0.25 Species turnover Species&turnover&(Simpson&Index)& 5) Bio?c Interac?ons: Preda?on Determinants of Distribu/on -0.02 1000 1000 1500 2000 2500 1500 2000 2500 Eleva.on&(meters)& Eleva/on (m) elevation 3000 3000 Figure&2.&Rela.onship&between&bird&species&turnover&and& Daily&nest&preda.on&& Change in d aily nest preda/on rates 0.06 0.04 0.00 0.45 0.02 0.55 Species&Turnover& Daily&Preda.on& 0.35 turnover Highest change in species composi/on (species turnover) corresponds with the highest change in rates of daily nest preda/on 0.25 Species turnover Species&turnover&(Simpson&Index)& 5) Bio?c Interac?ons: Preda?on Determinants of Distribu/on 5) Bio?c Interac?ons: Preda?on Brown tree snake (Boiga irregularis) Spread of brown tree snake on Guam (arrows) and subsequent declines in numbers of na/ve birds across years Savidge 1987 Ecology 36 Determinants of Distribu/on 5) Bio?c Interac?ons: Preda?on Brown tree snake (Boiga irregularis) USGS Timeline of the Brown tree snake travel across Guam hOp://www.fort.usgs.gov/resources/educa/on/bts/invasion/colonize.asp#dispersal Determinants of Distribu/on 6) Bio?c Interac?ons: Mutualism Clark’s nutcracker (Nucifraga columbiana) and whitebark pine (Pinus albicaulis) Whitebark, limber, Colorado pinyon, single-‐leaf pinyon and southwestern white pines depend on nutcrackers for seed dispersal This interac/on has changed the trees’ seeds, cones, and even overall shape in comparison with other wind-‐dispersed pine species hOp://www.allaboutbirds.org/guide/clarks_nutcracker/lifehistory 38 Determinants of Distribu/on 6) Bio?c Interac?ons: Mutualism Indo-‐Pacific clownfish (Amphiprion clarkii) and two sea anemones (Heterac>s malu and Macrodactyla doreensis) Mutualisms are seldom perfect predictors of distribu/on. 39 Determinants of Distribu/on 6) Bio?c Interac?ons: Mutualism Timing of clownfish radia/on with interac/ng sea anemone species shown for each clownfish species 40 Determinants of Distribu/on 6) Bio?c Interac?ons: Mutualism LeOers A-‐F represent Indo-‐Pacific provinces 26 pairs of tropical clownfish occupy ~ 10 species of sea anemones Clownfish have a mucus layer to avoid s/ngs from anemones Anemones provide shelter and protec/on from predators to clownfish 41 Determinants of Distribu/on 6) Bio?c Interac?ons: Mutualism Radia/on of clownfish has two geographical replicates. Phylogeny of the east African clade (Genus Amphiprion) and pie charts coloured according to mutualis/c sea anemone species (with sea anemone species shown below) 42 Determinants of Distribu/on 7) Historical Factors Sapsuckers (Sphyrapicus spp.) ! Wood Warblers (Vermivora spp.) ! Vireos (Vireo spp.) ! ! Wood Warblers (Oporornis spp.) Several species of NA birds have geographical divides in the rockies Wood Warblers (Setophega spp.) ! Weir & Schluter 2004 ! Sparrows (Passerella spp.) million years ago ! ! ! ! Pacific! Interior! Boreal! Flycatchers (Empidonax spp.) Chickadees (Poecile spp.) Determinants of Distribu/on 7) Historical Factors Several areas with high concentra/on of species with secondary contact as ranges tracked glacial retreat Many of these “suture zones” cluster around the Rocky Mountains of BC ice cover! 1 - 4 ! refguia! secondary contact! Cannings & Cannings 2004 A-C! Determinants of Distribu/on mourning winter wren ! Bird species with contact zones in the BC rockies ! pacific wren townsend’s ! macgillivray’s ! ! myrtle ! ! audubon’s ! hermit ! black-‐throated green Toews et al. 2011 ! black-‐fronted goldman’s !
