Biologging data enable animal ecologists to directly measure species’ fine-scale spatiotemporal responses come environments, together as activity – an important for our knowledge of biodiversity declines in the Anthropocene. Pet movement between source patches is a behavioral expression the multiple ecological processes that influence individual fitness. Defended area (PA) networks are a tool used to conserve biodiversity by maintain habitat patches across huge heterogeneous landscapes. However, our capability to design PA networks that conserve biodiversity relies on ours accurate knowledge of animal movement and also functional connectivity; this understanding is hardly ever tested in real-world situations due come the big geographic expanse of most PA networks. Using a tractable PA network mesocosm, we employ cutting-edge biologging modern technology to analyze animal movement decision in solution to a extremely heterogeneous landscape. We analysis these data to test, in a novel way, three common hypotheses around functional connectivity – structure corridors, least cost paths, and stepping stones. Consistently, pets moved along structurally self-similar corridors. In recommendation to the Aichi 2020 Biodiversity Targets, relying on varieties to “stepping stone” across habitat remnants might not accomplish protected area network conservation objectives.

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Researching pet ecology has always been clouded by obstacles observing free-roaming individuals. Brand-new biologging devices such together high-fix rate an international Positioning system (GPS) units quantify one animal’s space-use by providing refined monitorings of individual movement patterns. Activity is a behavioral expression the multiple environmental processes, the main of i m sorry is source selection within heterogeneous environments1: ecologists because of this need to fully understand this behavioral expression to implement reliable biodiversity conservation tools as landscapes proceed to quickly change2,3. Amplified spatial and also temporal resolution in dimensions of animal movement family member to see heterogeneity – such as how pets select, and also move between, partly or wholly disjunct resource patches to meet their energetic and life-history demands – greatly speeds up our knowledge of ecological processes and animal responses to changing environments.

Increasing landscape heterogeneity with land use readjust – reduction and spatial fragmentation of currently discontinuously dispersed resources – is a worldwide problem that affects individual animal movements, populations, and ultimately biodiversity persistence4,5,6,7. Pets must be able to move between disjunct source patches come sufficiently accomplish life-history requirements; hence landscape-scale practical connectivity is an important for species persistence8. For example, with detailed and also collaborative global monitoring of pet movements (i.e. Movebank.org, ICARUSinitiative.org), decreased movement in areas of high fragmentation has actually been observed across multiple taxa9. This might be in part due to enhancing proportion the landscape matrix – areas of greater risk and fewer sources that pets may or may not decide to overcome to accessibility the next resource patch10 – in ~ highly broke up anthropogenic landscapes. The worldwide area occupied by “working landscapes”– locations of high fragmentation and also interspersed natural and anthropogenic features11,12 – much exceeds that of undeveloped safeguarded areas13 so society currently relies heavily on functioning landscapes that contain protected locations to support biodiversity. Protected locations are often expected come anchor pet populations that usage the neighboring working landscapes, and the difficulties of this landscape technique to conservation is fine researched14,15.

The IUCN’s Convention on biological Diversity Aichi Target 11, inquiry 17% of the globe’s terrestrial area to it is in designated as protected, is swiftly approaching (2020). To support populations and metapopulations of big vagile varieties (such together mammals and also birds), protected areas too tiny for metapopulations need to be functionally associated into defended area networks (PANs)16. Useful connectivity is “the degree to i beg your pardon the see facilitates or impedes activity among resource patches”17. The bulk of sensible connectivity research examines the result of connectivity – such together gene flow, movement, or species distribution – from which practical connectivity is then inferred18. Without high-precision motion data, biologists struggled to reliably measure the process, quite than the outcome, of useful connectivity. Biologging helps to leg this gap.

Here, we focus on three of the most common, non-mututally exclusive theories of connectivity; despite these principles are defined under different terminologies througout the literature, we define our consumption of each below.

1. The corridor framework, arisen in working-landscape systems19,20,21 posits that long thin habitat patches, typically unsuitable for sustaining a varieties on your own, will facilitate sensible connectivity in between populations once physically connecting structurally comparable patches and also habitats (i.e. Source patches)21,22. The standard example is wooded hedgerows connecting remnant forest patches in agricultural landscapes wherein small mammals pick structurally self-similar landscape functions in which to travel20. 2. The least-cost paths (LCP) structure is acquired from ideas about light refraction23, and also is connected to electric circuit theory24; specifies connectivity no by physical similarity to undeveloped patches but solely through the cost a piece of landscape imparts on pet movement. Herein the see is a continuum of costs that correlate to the type, and density, the habitat features25; that does not think about habitat and also matrix as binary, separate, entities. 3. The stepping rock theory is derived from Island Biogeography theory26. Island Biogeography, and also the species-area relationship, demonstrates that types richness and turnover space outcomes that connectivity in between isolated and also distinct resource patches. Although controversial27 this theory says disjunct but physically proximate patches of comparable resources (i.e. Islands) facilitate connectivity; herein landscape procession (i.e. Oceans) and habitat (i.e. Islands) room acutely distinguished, together the matrix is unusable and discrete island source patches – represented by either polygonal see features, or raster cells, in a geographic Information system – facilitate movement. In terrestrial landscapes the hostility of the matrix is relaxed however still assumed to be strongly avoided, or “stepped across”.

The validity of this theories is of critical importance in landscape planning, particularly in planning new protected areas and also protected area networks. Despite the raising spread of functioning landscapes, and also the worldwide inititaive to rise protected locations to support biodiversity, we understand very tiny about just how well, or if, networks actually facilitate species’ movement within them (with part exceptions)28. Us ask even if it is a model PA network – the newly created UNESCO Beaverhills Biosphere (Fig. 1) – facilitates useful connectivity within a working landscape. We usage high-frequency biologging paired with a new movement and source selection statistical version in a PA network landscape mesocosm29 to examine whether the corridor, least cost path, or stepping stone framework of connectivity ideal explain practical connectivity for a model varieties subject to substantial connectivity and also biologging research30,31,32,33, the fisher (Pekania pennanti).


Figure 1

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We posed three non-mutually exclusive hypotheses and weighed proof for each across multiple candidate to adjust in one information-theoretic approach. We connected hypotheses to connectivity frameworks by measure up the distance in between movement steps and landscape attributes (measured as ‘distance to’ in meters; the proximity of animal movement come the sheet of a landscape feature), and also the movement price of polygonal landscape features (measured together ‘density’ in pixels/m2; the cost of pet movement v landscape attributes of different source densities; Fig. 3). If fishers room using protected areas as stepping stones throughout the working landscape matrix, us predict fisher to display highly tortuous movements and also short step lengths (high use) in ~ PAs and long linear movements between them; we mean a hopeful correlation between the presence of protected areas and tortuosity, and also a negative correlation between the existence of safeguarded areas and the step size (Fig. 3; defended Area Stepping Stones). If corridors facilitate connectivity, then we predict fishers should relocate along structurally self-similar see features; we expect a confident correlation between consecutive measures so the the thickness of each polygonal feature, and the distance to each linear feature, is similar between actions (Fig. 3; Corridors). Finally, if fisher relocate along least price pathways across the landscape, fisher should display tortuous and short step lengths within features clearly distinguished by high movement expense (high use), and linear however long activities within features with low movement prices (low use, for travel only); we mean the thickness of landscape attributes to be negatively correlated with tortuosity and positively associated with the length of movement steps (Fig. 3; Least cost Paths). We identify that a structure corridor may without doubt be a least expense pathway if it gives the course of the very least resistance.


Figure 3

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Parameters within every clogit version describing hypothesized frameworks for landscape connectivity throughout the Beaver Hills Biosphere. Street to (Dist) represents fisher activity along straight features, where as density (Dens) to represent the movement cost of a polygonal habitat function (high density = high cost). Every models affiliated a set of core model(*) variables that we hypothesized would be generally essential to fisher movement: CosTurnAngle + lnStepLength - Dist(DECID) + Dens(DECID) - Dist(CONIF) + Dens(CONIF) - Dist(MIXED) + Dens(MIXED) - Dist(WATER). Interactions are denoted by “:”.


Of the 14 fisher captured and collared, we obtained GPS data from 10 individuals (5 males: 5 females) comprising 17% that the approximated population34. These 10 collars gathered 19,578 gps fixes, over an average of 32.97 days (minimum = 4.87 days, maximum = 90.79 days) of consistent movement data per individual. Fisher action lengths over a 5-min settle interval approximated a log normal distribution with many small steps (mean = 105.47 m, s.e. = 1.85 m, min = 0.06 m, max = 2972.0 m), and also turn angle were ~ above average little and positive, describe directional motion behaviour (mean = 0.08 rad, s.e. = 0.0001 rad).

Corridor models containing organic features finest explain movement throughout a heterogeneous defended area network

The corridor version of sensible connectivity, in which pets move amongst structurally comparable features across the landscape, finest explained fisher movement throughout this PA network. This version received the highest possible AIC load of evidence throughout 6 the the 10 fisher people (86–99%), and second highest because that the 2 the the remaining 4 individuals. 4 individuals verified support because that the least expense paths version as the 2nd highest AIC weight of evidence, and no individuals showed assistance for the stepping rock hypothesis (Fig. 4). All models generally had sufficient concordance (concordance of optimal AIC models ranged indigenous 0.594 come 0.647).


Figure 4

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High fix-rate general practitioners movement telemetry data from six of 10 fisher people showed the greatest relative support for a corridor structure of sensible connectivity when compared to one of two people least price paths or stepping stone framework hypotheses throughout the heterogeneous see mesocosm of Alberta’s Beaver Hills Biosphere. The counting of height models showing support because that each theory is demonstrated in black, and also the count of second-best models mirroring support for each theory is demonstrated in grey. Hypotheses encompass the corridor theory of movement, a global model, the least price paths theory of movement, a stepping rock hypothesis that movement, and also a core design representing species-specific habitat selection.


Among fisher individuals, polygonal natural and also anthropogenic features ideal explained the observed variance in fisher steps. Density of defended areas, distance to nearest safeguarded areas, and interactions in between PAs and both movement step size (lnStepLength) and turning angle (cosTurnAngle) to be rarely far-reaching in top models, saying fishers walk not distinguish the PA network native other features in the landscape (Fig. A1.1). Only two individuals’ activity was defined by PAs, yet neither stood for the top version for the individual. One fisher individual’s activity varied with PA thickness of the current (iSSA parameter slope and also significance; ß = 0.20 ± 0.04, p A1.1,A.1.2). This occurred despite the presence and proximity of protected areas. Together, these activity patterns suggest fishers predominantly offered natural attributes as corridors, rather than crossing multiple landscape functions with different movement costs, or utilizing stepping stones in ~ or in between protected habitat patches.

Individual pet responses to defended areas

If PAs conserve element habitat patches, we predicted separation, personal, instance fisher to display highly tortuous and short movement actions as an indication of high residency time within PAs. However, we discovered variable assistance for this principle within separation, personal, instance movements: 40% that fisher displayed linear and long steps, 20% of fisher displayed linear and also short steps, 20% that fisher presented tortuous and also short steps, and 20% that fisher had no variation in their actions as lock only emerged in protected areas. Therefore, 40% that the data assistance the prominence of PAs because that connectivity, and also 60% of the data carry out not. Across individual movements within PAs, step size (Pearson correlation; r = 0.003, df = 214150, p = 0.11) and turning angles (r 35, and (4) the prestige of considering matrix effects on predicted movement and behavioural variations within protected areas.


We unexpectedly found that protected locations alone had actually very little effect on the activity choices, and also therefore see connectivity, of a midsized Nearctic mammal. When detailed a see of high heterogeneity, fisher repetitively used organic polygonal functions as motion corridors, regardless of the visibility of a highly heterogeneous landscape spanned by a network of thousands of protected locations of widely varying sizes. Certainly, PAs added to the lot of remnant indigenous habitat ~ above this landscape which is otherwise introgressed by agriculture, rural residency, transportation, and also petroleum extraction. However, the moraine soils have made this see less perfect for agriculture, and also hence landowners have retained aboriginal vegetation, which has actually proven crucial for fisher movement. When planning and implementing defended area networks, the visibility of lot of protected areas alone may not facilitate see connectivity uneven structurally connected by natural, polygonal, see features. The ecological integrity that the rapidly arising protected locations under Aichi Target 11 may depend top top the visibility of herbal landscape features in between them; this might be of an important importance because that today’s abundant working landscapes.

Measuring protected area network connectivity via outcomes – such as genetic diversity36 or varieties occurrence37, is that course vital to understanding exactly how landscape planning devices conserve biodiversity. However, when connectivity outcomes are shown to falter or fail, that is research study on connectivity procedure – motion – the reveals the instrument behind blockages. By assessing functional connectivity trends using high-frequency biologging within a tractable mesocosm PA network, we help to know the effect of the landscape procession on PA network efficacy38,39,40.

The theory of Island Biogeography26 first provided the theoretical framework that functional connectivity of discrete habitat spot (therein, the ability of archipelago to receive new species, generating varieties richness and turnover) relies on your size and also isolation from other perfect patches. In this framework, the room between habitat spot – the landscape matrix10 – is important, however purely inhospitable. This is true the oceanic islands, yet in terrestrial studies the matrix gives a continuous of suitability41 that creates a spectrum of connectedness42. The matrix can carry out varying levels of facilitation, or impediment, for useful connectivity38, types richness in remnant woodland patches41, or protected area efficacy39. This spectrum that connectedness in procession facilitation is types specific17, yet our results here also highlight that it may change for separation, personal, instance animals.

Animal-defined corridors are crucial consideration for connectivity, as animal responses to ecological heterogeneity demonstrate individual variation35 and also potential because that plasticity43,44. In a comparable study come ours, whereby the landscape matrix had a better proportion of urban landscape, LaPoint et al.31 show fisher movement data at regional scales best supports a corridor version of useful connectivity, but an ext importantly, the fisher-defined corridors are composed of a selection of land cover types. Ours novel finding is in reflecting individual variation between functional connectivity frameworks, together 4 the 10 fisher iSSA models ideal supported a version with every possible, non-correlated, predictors (i.e. Worldwide model) quite than a model representing any certain hypothesis. This result may highlight plasticity in separation, personal, instance responses to ecological heterogeneity, one individual’s capability to use multiple connectivity frameworks to piece together sources (i.e. Non-mutually exclusive hypotheses), or distinctions in PA quality amongst areas these fishers occupied. We demonstrate that PA network useful connectivity deserve to be enhanced by combine individual behavioural data, rather than suspect a uniform an answer by individuals to structure connectivity. This ‘animal-defined’ corridors quantified in heterogeneous landscapes45 will help to parameterize the functional components of connectivity throughout seasons, and both natural (e.g. Forest fires) and anthropogenic (e.g. Crop rotation, development) disturbances31,46,47.

It is currently clear that the matrix is just as beneficial as that remaining herbal (non-anthropogenic) habitat patches, quite than the extent and also density that PAs alone, in ~ a landscape. This is partly due to the fact that land in ~ PAs space no much longer 100% natural: if suitable habitat loss, whether in ~ or in between PAs, is above 80%, matrix high quality no longer buffers extinction thresholds40. Andren48 demonstrated a comparable finding throughout birds and also mammals; he qualities this finding to the fact that over 70% the habitat loss, percolation theory predicts the the effects of habitat loss and habitat fragmentation will compound. Corridors are meant to get rid of this “percolation effect” by forcing connections between habitat patches and also thus staying clear of the compounding effects of habitat loss and habitat fragmentation. Here, in ours empirical test of these concepts, prioritizing herbal features between PAs best facilitates useful connectivity within PA networks by providing corridors for fisher movement. We complete this an outcome is really likely similar for other forest-dependent species that consist of the bulk of biodiversity on this49,50, and also other, Nearctic forested landscapes.


There is considerable monetary and political resources investment in PA protection13. However, the investment frequently ends at the PA border, and mechanics of biodiversity conservation in the matrix is left purely to hope. Here, us test eco-friendly theory to display that conservation tools cannot rely on one of two people proximity or expect – we require to far better understand animal responses to ecological heterogeneity, and we need a planned and protected procession designed from correct theoretical underpinnings to provide effective biodiversity conservation throughout PA networks.

The present state that biologging science allows for data arsenal that solidifies the underpinnings the connectivity conservation. Combining these high-resolution data (e.g. General practitioners telemetry top top Movebank.org), with thorough GIS data in ~ an iSSA framework, allows for boosted precision and sophistication 보다 ever before for experimentation long-standing hypotheses with environmental data. Our PA network mesocosm analysis suggests that herbal habitat in ~ landscape matrices are just as essential as organic habitat within safeguarded areas: factor to consider of natural functions within the matrix have to receive greater management consideration. In addition to creating new protected locations under the Aichi Target 1116, concentrating on preserving or restoring natural landscape attributes within the procession of existing PA networks, or developing PA networks within existing landscapes of high natural features, will certainly greatly aid conservation objectives. Utilizing biologging, these objectives have the right to now be regularly monitored and adjusted as important within an adaptive monitoring framework51, elucidating vital patterns that an obstacle our conceptual understanding of pet ecology and also conservation science. We present that raising the extent of the global protected area network is not a stand-alone systems to connecting protected areas; the conservation of natural landscape features between PAs is the mortar that binds them together.


Data collection throughout the mesocosm

The Beaver Hills Biosphere (BHB) covers around 1,596 km2 of glacial moraine in east-central Alberta, Canada (53.381167°N, 113.062976°W; Fig. 1). This heterogeneous see is created of natural, anthropogenic, and protected area (PA) habitats (Table 1). Organic habitats can be either within or external of PAs and consists primarily of indigenous aspen parkland (Populus tremuloides and P. Balsamifera), and interspersed tiny waterbodies, meadows, and also patches of white spruce (Picea glauca). 7 hundred and sixty-three PAs the varying dimension (mean ± SE = 78.4 ± 29.0 km2), condition (from neighborhood conservation easements regulated by non-government institutions to provincial and also national parks), and also isolation (measured as the distance in between protected areas; mean ± SE = 0.95 ± 0.004 km) conserve these native features across the BHB – but these PAs likewise have a level of anthropogenic advance within them, including grazing areas, considerable recreation, and roads (Fig. 2). The remainder of the see is written of extensive resource extraction in the form of oil and gas, agriculture, forestry, and rural-residential development. The resulting matrix surrounding the BHB’s protected locations is a mosaic of private lands, roads, and farming that separate the BHB native tracts of contiguous forest in various other parts that the province.


Table 1 street to (Dist), and also density around (Dens), the end of both used and available fisher actions were quantified across 15 landscape attributes within the Beaver Hills Biosphere.
Full dimension table

From November 2015 through February 2016 we provided covered cage trap (Tomahawk 109, Tomahawk, WI) come live-capture 10 fisher50. We supplied a combination of ketamine (concentration = 100 mg/ml, dose = 12 mg/kg) and midazolam (concentration = 5 mg/ml, dose = 0.3 mg/kg) come sedate individuals; us monitored an essential rates and fitted individuals with general practitioners tracking collars (E-obs Collar 1 A; Grünwald, Germany). Collars included a gps microchip, ultra-high frequency transmitter for telemetry and also data download, and tri-axial accelerometer; the gps was programmed to take a GPS-fix every 5 minutes if the individual was moving higher than 10 cm/s. Spatial capture-recapture modeling of these data estimate the BHB fisher population to it is in at many 58 people (3.91 fishers/100 km2)34. We therefore obtained general practitioners telemetry data from at least 17% the the modern population. All research was approved by the InnoTech Alberta pet Care Committee (2070M-A02/048/15-P01), and also all research techniques were perform in accordance with Canadian council on animal Care.

Integrated step selection analysis

Functional connectivity deserve to be measure up as activity of pets in relation to see structure. The used motion path that individual pets can be acquired from GPS-fixes accumulated from GPS-telemetry collars. For each used GPS-fix in ~ this study, we generated 10 random available steps and also turn angles, and also compared in a used-available, or “case-control”, design52 (Fig. 1). These observed steps and turn angle were assigned a “1”, whereas obtainable (i.e. Created in GME) steps and turn angle a “0” and together stand for the binomial response variable in our conditional logistic regressions52.

Step lengths, i m sorry are characterized as straight-line distances between successive GPS-fixes, measure the rate of an animal (i.e. M/5 min) and can be supplied as an calculation of pet residency time within habitat features: much shorter steps indicate longer residency time53,54. Using the movement.ssf duty in GME (www.spatialeclogy.com/gme/), easily accessible fisher step lengths were sampled native a log in normal circulation parameterized on supplied step lengths for each separation, personal, instance (distribution form varied between 3.31–4.45, distribution scale varied between 1.99–1.40). Step lengths were ln-transformed (lnStepLength) sensu Avgar et al.52,55, and also are one estimator of the selection-free rate of animal movement. Available turn angle were sampled indigenous a uniform distribution between –π and also π radians52 and are identified as the angular deviation between two headings; these values were cosine-transformed (cosTurnAngle), which transitions a circular measure (radians) right into a direct measure between −1 and 155,56; values approaching 1 represent straight movement57. Therefore, procedures without a proceeding action (i.e. The very first step accumulated for every individual) were eliminated from the analysis. We carried out an analysis examining habitat choice and movement for every of 10 individuals to quantify assistance for each connectivity framework throughout the BHB.

Landscape functions as model covariates

To check the impact of landscape functions on step length, we used ArcGIS v10.3 (ESRI, Redlands, CA, USA) geographic Information device to quantify landscape heterogeneity. We offered the LandSat digital map inventory indigenous the Beaver Hills Biosphere (Land Management frame 2015) to quantify the distance of the end points the fisher steps (m) to every landscape feature, and also the density of landscape features, across 15 category representing natural, anthropogenic, and also protected locations (PAs); bare landscape, crops, deciduous forests, mixed forests, coniferous forests, wetlands, development, forage, grasslands, lakes, shrubs, protected areas, rail lines, roads, streams, and PAs (Table 1). Landscape features, whether polygonal or linear, were converted to a raster to calculate density; they to be measured together the thickness of a buffer approximately the end allude of every step, inside the buffer radius was determined by the average fisher step size (106 m) and measured together the number of raster pixels/m2. We scaled these procedures to permit comparison the coefficients within regression models together a measure of described variance; this scaled distances, and density, measures comprise the predictor variables in our conditional logistic regressions.

Our final data set comprised 214,148 used and accessible steps across 10 fisher individuals, including (1) used/available status, action length, revolve angle, fisher ID, UTM coordinates, and also strata of available steps; and also (2) the partnership of each step to landscape predictor variables – one of two people the distance to, and/or density, the 15 landscape attributes (Table 1). Distance to was truncated come 1000 m, together we did not suspect fisher come respond to landscape functions that were any kind of further way.

Statistical analyses

We created statistical models to check the three hypotheses of see connectivity acquired from current evaluation methods: corridors, least price paths, and also stepping stones (Fig. 3). We developed a ‘core model’ of assumed habitat features explaining variation of fisher movement in a non-human overcame landscape and included this design within every of ours connectivity hypotheses (Fig. 3). LnStepLength (m) represents the direct displacement between consecutive procedures – a proxy for pet speed together the time between steps is continuous (m/5-minutes) – vice versa, the motion directionality, or tortuosity, is explained by the cosine that the turning angle58. Including lnStepLength and also cosTurnAngle as model predictors in ~ a clogit regression extends the step selection function framework53 into an incorporated Step an option Analysis (iSSA), accounting for pet movement speed and also directionality within selected habitat features52.

From previous research study we mean tortuosity (i.e. Turn angle) and speed (i.e. Action length) to influence fisher movement and fisher to select locations of deciduous forest, coniferous forest, and mixed forest, while remaining proximate come water bodies31,32,59,60. These space what we define as ‘high’ movement expense habitats for fisher; landscape attributes that receive a high relationship of residency time. We because of this included the cosTurnAngle, lnStepLength, distance to wetlands, deciduous, coniferous, and mixed forests, and also density that deciduous, coniferous, and mixed woodlands within ours core design (Fig. 3).

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We competed five conditional logistic regression models (Fig. 3) in an info Theoretic technique using Akaike information Criterion values61 (AIC; Fig. 3) for each fisher individual. In a comparable approach to Prokopenko et al.55 we supplied the clogit role in the survival package62 in R (v3.2.2)63, to perform conditional logistic regression models; the solution variable was actions observed (0/1), and each strata was assigned to paired used:available actions (1/0)55. We completely explored ours data64 ensuring every clogit model assumptions were met. All statistical analyses were carried out in R63, and also results room presented as mean ± SE uneven otherwise specified.