Ecological Archives E090-184-D1
Kate E. Jones, Jon Bielby, Marcel Cardillo, Susanne A. Fritz, Justin O'Dell, C. David L. Orme, Kamran Safi, Wes Sechrest, Elizabeth H. Boakes, Chris Carbone, Christina Connolly, Michael J. Cutts, Janine K. Foster, Richard Grenyer, Michael Habib, Christopher A. Plaster, Samantha A. Price, Elizabeth A. Rigby, Janna Rist, Amber Teacher, Olaf R. P. Bininda-Emonds, John L. Gittleman, Georgina M. Mace, and Andy Purvis. 2009. PanTHERIA: a species-level database of life history, ecology, and geography of extant and recently extinct mammals. Ecology 90:2648.
INTRODUCTION
Macroecology and macroevolution are growing research  areas (Brown 1995, Gaston and Blackburn 2000), describing and testing hypotheses for many  large-scale patterns in biodiversity, such as species richness (Orme et al. 2005, Grenyer et al. 2006); extinction  risk (Purvis et al. 2000, Jones et al. 2003, Fisher and Owens 2004, Jetz et  al. 2004, Cardillo et al. 2005, Price and Gittleman 2007); evolutionary history (Sechrest et al. 2002, Bininda-Emonds et al. 2007,  Davies et al. 2008); geographic  range size (Gaston 2003, Orme et al. 2006), and interspecific allometric relationships (Carbone et al. 2007a, Carbone et al. 2007b). Such comparative studies require large amounts of  data, often covering a wide range of species or taxonomic groups in a number of  geographic regions. Researchers gathering these data face two general problems. First, the amount of time and effort required  to collect enough comparable information can be considerable; and second, the coverage  of the available data, which determines the feasibility of studying  different geographic regions or taxonomic groups, may be very inconsistent.
Here  we describe a global species-level data set of key life-history, ecological and  geographical traits of all known extant and recently extinct mammals  (PanTHERIA) developed for a number of macroecological and macroevolutionary  research projects. Additionally, we describe the tools, techniques and  protocols developed to minimize errors and biases given the disparate data  sources required to generate this data set and the large team of data-gatherers  used to collect these data (referred to here as data-miners). We also describe  our statistical methods developed to detect and, where possible, eliminate  outlying values, and to then derive a single value for each trait for each  species.
METADATA
CLASS I. DATA SET  DESCRIPTORS
A. Data set identity: PanTHERIA1.0: a species-level data set of life history,  ecology, and geography of extant and recently extinct mammals.
B. Data set identification code: 
  (1) PanTHERIA_1-0_WR93_Aug2008.txt
  (2) PanTHERIA_1-0_WR05_Aug2008.txt
C. Data set description:
Principal Investigator(s): Kate E. Jones, Institute  of Zoology, Zoological Society of London, Regents Park, London,   NW1 4RY, United    Kingdom. Olaf R.P. Bininda-Emonds, AG Systematik und Evolutionsbiologie, Carl von Ossietzky Universität   Oldenburg, 26111 Oldenburg, Germany. John  L. Gittleman, Odum School of Ecology, University of Georgia, Athens, Georgia  22902, USA. Georgina M. Mace, Centre for  Population Biology, Imperial College London, Silwood Park  Campus, Ascot, SL5 7PY, United    Kingdom. Andy Purvis, Division of Biology,  Imperial College London, Silwood Park Campus, Ascot, SL5 7PY, United Kingdom;  and Centre for Population Biology, Imperial College London, Silwood Park  Campus, Ascot, SL5 7PY, United Kingdom. 
Abstract: Analyses of life history,
ecological, and geographic trait differences among species, their causes,
correlates, and likely consequences are increasingly important for understanding
and conserving biodiversity in the face of rapid global change. Assembling multispecies
trait data from diverse literature sources into a single comprehensive data set
requires detailed consideration of methods to reliably compile data for
particular species, and to derive single estimates from multiple sources based
on different techniques and definitions. Here we describe PanTHERIA — a species-level
data set compiled for analysis of life history, ecology, and geography of all
known extant and recently extinct mammals. PanTHERIA is derived from a database
capable of holding multiple geo-referenced values for variables within a
species containing 100740 lines of biological data for extant and recently
extinct mammalian species, collected over a period of three years by 20
individuals. PanTHERIA also includes spatial databases of mammalian geographic
ranges and global climatic and anthropogenic variables. Here we detail how the
data fields are extracted and defined for PanTHERIA using a customized data
input format MammalForm; how data were collected from the literature, species
names and sources tracked, error-checking and validation procedures applied, and
how data were consolidated into species-level values for each variable. Tables
of the consolidated species-level values are made available for each of two recent
species-level taxonomic classifications of mammals, as well as associated
taxonomic synonymy conversion and data-input files. This study provides a
useful guide to prospective researchers on how to structure and codify life history,
ecological, geographic, and taxonomic data and methods to extract meaningful
species-level traits. It also provides comprehensive information on traits like
size, diet, environmental conditions, and ecology to permit macroecological and
macroevolutionary analyses of this important clade.
D. Key words:  behavior; biogeography; body size; climatic variables; comparative analyses; conservation; ecology; geographic range; life history; mammals; PanTHERIA; population density.
CLASS II. RESEARCH ORIGIN DESCRIPTORS
A. Overall project description
Identity: Understanding past, present, and future mammalian biodiversity.
Originators: Kate E. Jones, Institute of Zoology, Zoological Society of London,  Regents Park,  London, NW1 4RY, United    Kingdom. Olaf R.P. Bininda-Emonds, AG Systematik und Evolutionsbiologie, Carl von Ossietzky Universität   Oldenburg, 26111 Oldenburg, Germany. John  L. Gittleman, Odum School of Ecology, University of Georgia, Athens, Georgia  22902, USA. Georgina M. Mace, Centre for  Population Biology, Imperial College London, Silwood Park  Campus, Ascot, SL5 7PY, United    Kingdom. Andy Purvis, Division of Biology,  Imperial College London, Silwood Park Campus, Ascot, SL5 7PY, United Kingdom;  and Centre for Population Biology, Imperial College London, Silwood Park  Campus, Ascot, SL5 7PY, United Kingdom. 
Period of study: 2001 – continuing.
Objectives: 1.  To understand the patterns of mammalian biodiversity and extinction risk, and  the processes behind them. 2. As  necessary steps towards objective 1, to assemble comprehensive information on  the comparative ecology and life history, the geographic distribution and the  phylogeny of extant mammalian species.
Abstract: Species-richness and extinction risk are  distributed unevenly among mammalian taxa and among geographic regions. This project aims to document these patterns  across the entire class Mammalia and to use phylogenetic comparative analyses  to test a wide range of hypotheses for them.  Two research grants in particular supported the work, one entitled  “Threatening processes and the conservation status of contemporary mammals” (awarded by NERC to G.M. and A.P.) and one entitled “The macroecology and  biodiversity of geographic ranges in mammals”  (awarded by NSF to J.L.G.). The combination of the different facets of  mammalian biodiversity information has permitted large-scale analyses of (among  many others) diversification rates through time and correlates of extinction  risk, and identification of species particularly vulnerable to rapid decline if  threatening processes intensify.
Sources of funding: NERC grant (NER/A/S/2001/00581) to GM and AP, NSF grant (DEB/0129009) to JLG and funding from Conservation International. KEJ was additionally funded  through an Earth Institute Fellowship, Columbia University and the University of Virginia. SAF through the European Commission (FP6 Early-Stage Training Network HOTSPOTS)  and KS was funded through a Swiss National Science Foundation fellowship.
B. Specific subproject description
Site  description: Data collection from all extant and recently extinct mammalian  species.
Experimental  design: Data were collected from the literature or from the author’s  previously collated databases.
Research  methods:
Field/Laboratory: n/a
Data sources: Multiple values of  biological trait data were gathered with geo-references where available for  extant and recently extinct mammalian species from primary and secondary  sources. Data gathering was completed until June 2004. Data were initially  gathered systematically from the most relevant journals
 and secondary sources (e.g., Mammalian Species,  Journal of Mammalogy, Mammalia, Journal of Zoology, Hayssen et al. 1993) and the electronic search  engine Web of Science (WOS) and Zoological Record were used to locate further  sources for particular variables, clades or individual species. We reduced data  duplication by developing a dynamic online source reference library that was  consulted prior to searching for new sources or new data entry sessions. Additionally,  we incorporated previous data compilations held by the authors, i.e., Small  Mammal Life History Database (Mace 1979); Bat Life History Database (Jones 1998); Carnivore Body Size and Life History Database (Gittleman 1982, Bininda-Emonds 1998); unpublished Primate Life History Database from (Purvis et al. 2000); unpublished Population Density Database from (Carbone and Gittleman  2002).  In total we used data from 3143 sources  for the PanTHERIA data set (see Class V, Section F for a full reference list). We  also used spatial databases of mammalian geographic ranges and global climatic  and anthropogenic variables (see Class V, Section C for a full description).
Data  collection: Biological trait data were collected by 20 different data  miners over a period of 24 months from July 2002 to June 2004, using a standardized  data input protocol with a Visual Basic interface programmed within Excel  (MammalFormv2.xls see Class V, Section D).
Variables: Data were gathered from the literature for 25  types of ecological and life history information for any extant or recently  extinct species within class Mammalia (100740 data lines): 1. Activity Cycle; 2. Age  at Eye Opening; 3. Age at First  Birth; 4. Average Lifespan; 5. Body Mass; 6. Diet; 7. Dispersal  Age; 8. Adult Limb Length; 9. Gestation Length; 10. Group Composition & Size; 11. Growth Data; 12. Habitat Layer; 13. Head-Body  Length; 14. Interbirth Interval; 15. Litter size; 16. Litters Per Year; 17. Maximum Longevity; 18. Metabolic  Rate; 19. Migratory Behaviour; 20. Mortality Data;  21. Population  Density; 22. Ranging Behaviour; 23. Sexual Maturity Age; 24. Teat Number; and 25. Weaning Age. 30 specific variables (see  Class IV, Table 1) were extracted from the above data types for PanTHERIA  from a total of 94729 data lines (before error checking). Additionally, 4 variables were derived from extracted variables  within PanTHERIA and 19 variables were  calculated from other spatial data sources (see Class  V, Section C).
Taxonomy and systematics: Two species-level data output files were generated each  using a different taxonomy: (1) (Wilson and Reeder 1993) (PanTHERIA_1-0_WR93_Aug2008.txt) and (2) (Wilson and Reeder 2005) (PanTHERIA_1-0_WR05_Aug2008.txt). SppSynonymID1.0.txt  documents how species names or synonyms given in the original data sources were  tracked onto these current taxonomies (see Class V, Section B).
Project personnel: Kate E. Jones, Jon Bielby, Marcel Cardillo, Susanne A. Fritz, Justin  O’Dell, C. David L. Orme, Kamran Safi, Wes Sechrest, Elizabeth H. Boakes, Chris  Carbone, Christina Connolly, Michael J. Cutts, Janine K. Foster, Richard  Grenyer, Michael Habib, Christopher A. Plaster, Samantha A. Price, Elizabeth A.  Rigby, Janna Rist, Amber Teacher,  Olaf R.P. Bininda-Emonds, John L. Gittleman, Georgina M. Mace, and Andy Purvis.
CLASS III. DATA SET STATUS AND  ACCESSIBILITY
A. Status
Latest update: 14th June 2004.
Latest archive date: n/a
Metadata status: 8th Aug 2008, metadata are current.
Data verification: Data were carefully checked for quality and consistency (described in Class  V, Section B below).
B. Accessibility
Storage location and medium: The species-level data sheets are included here and  the raw data are held in an Access database at The Zoological Society of London.
Contact person: Kate E. Jones, Institute of Zoology, Zoological Society of London, Regents Park,  London, NW1 4RY, United    Kingdom. E-mail: [email protected].
Copyright restrictions: None.
Proprietary restrictions: None.
Costs:  None, the authors believe that scientific data collated using public funds  should be free for scientific use. We would appreciate researchers citing this  paper if using these data.
CLASS IV. DATA STRUCTURAL DESCRIPTORS
A.  Data Set File
Identity: (1)  PanTHERIA1.0_WR93_Aug2008.txt and (2) PanTHERIA_WR05_Aug2008.txt.
Size:  (1) 4629 records, not including header  row; (2) 5416 records,  not including header row.
Format  and storage mode: ASCII text, tab delimited, not compressed.
Header  information:  Headers describe the content of each column. We use the following abbreviations: MSW93 – Wilson and Reeder (1993)  Mammalian Species of the World 2nd Edition; MSW05 – Wilson and Reeder (2005) Mammalian Species of the World 3rd  Edition;  AET – Actual Evapotransipration Rate; C – centigrade; d –  days; dd – decimal degrees; deg – degrees; EXT – extrapolated; g –  grams; GR – geographic range; Grp – Group; HuPopDen – Human Population Density; Indiv – Individual; Isl - Island; Lat – Latitude; Len – Length; Long – Longitude; Max –  Maximum; Met – Metabolic; Min – Minimum; mLO2hr – milliliters of O2 per hr; mm – millimeters;n/km2 –  Number per km2; PET –  Potential Evapotranspiration Rate; Precip – Precipitation; Temp – Temperature; and 5p – 5th percentile. Words and abbreviations are  separated by underscores. Numeric values at the start of the entries in the  header refer to the data type from which the variable was extracted. For  example, “5-1_AdultBodyMass_g”, “5”  refers to the data type ‘Body Mass’ and “1” that it is the first variable to be  extracted from this data type.
Headers included are as follows: MSW93_Order or MSW05_Order;  MSW93_Family or MSW05_Family; MSW93_Genus or MSW05_Genus; MSW93_Species or  MSW05_Species; MSW93_Binomial or MSW05_Binomial; 1-1_ActivityCycle; 5-1_AdultBodyMass_g; 8-1_AdultForearmLen_mm; 13-1_AdultHeadBodyLen_mm; 2-1_AgeatEyeOpening_d; 3-1_AgeatFirstBirth_d; 18-1_BasalMetRate_mLO2hr; 5-2_BasalMetRateMass_g; 6-1_DietBreadth; 7-1_DispersalAge_d; 9-1_GestationLen_d; 12-1_HabitatBreadth; 22-1_HomeRange_km2; 22-2_HomeRange_Indiv_km2; 14-1_InterBirthInterval_d; 15-1_LitterSize; 16-1_LittersPerYear; 17-1_MaxLongevity_m; 5-3_NeonateBodyMass_g; 13-2_NeonateHeadBodyLen_mm; 21-1_PopulationDensity_n/km2; 10-1_PopulationGrpSize; 23-1_SexualMaturityAge_d; 10-2_SocialGrpSize; 24-1_TeatNumber; 12-2_Terrestriality; 6-2_TrophicLevel; 25-1_WeaningAge_d; 5-4_WeaningBodyMass_g; 13-3_WeaningHeadBodyLen_mm; References; 5-5_AdultBodyMass_g_EXT;  16-2_LittersPerYear_EXT; 5-6_NeonateBodyMass_g_EXT; 5-7_WeaningBodyMass_g_EXT; 26-1_GR_Area_km2; 26-2_GR_MaxLat_dd; 26-3_GR_MinLat_dd; 26-4_GR_MidRangeLat_dd; 26-5_GR_MaxLong_dd; 26-6_GR_MinLong_dd; 26- 7_GR_MidRangeLong_dd; 27-1_HuPopDen_Min_n/km2; 27-2_HuPopDen_Mean_n/km2; 27-3_HuPopDen_5p_n/km2; 27-4_HuPopDen_Change; 28-1_Precip_Mean_mm; 28-2_Temp_Mean_01degC; 29-1_AET_Mean_mm; and 29-2_PET_Mean_mm.
Alphanumeric attributes: Mixed.
Special characters/fields: If no information is available for a given record, this  is indicated by -999.
Authentication  procedures:
 Sums of the numeric columns for
 
  PanTHERIA1.0_WR93_Aug2008: (1) 1-1_ActivityCycle = -2977053; (2) 5-1_AdultBodyMass_g = 622610217.539998; (3) 8-1_AdultForearmLen_mm = -3679100.18999997; (4) 13-1_AdultHeadBodyLen_mm = -1646782.58; (5) 2-1_AgeatEyeOpening_d = -4147029.48; (6) 3-1_AgeatFirstBirth_d = -3824836.7; (7) 18-1_BasalMetRate_mLO2hr = -3028141.87; (8) 5-2_BasalMetRateMass_g = 375079.45; (9) 6-1_DietBreadth = -2467939; (10) 7-1_DispersalAge_d = -4421311.55999999; (11) 9-1_GestationLen_d = -3121092.90999998; (12) 12-1_HabitatBreadth = -1864542; (13) 22-1_HomeRange_km2 = -3759464.11216601; (14) 22-2_HomeRange_Indiv_km2 = -3844306.9899835; (15) 14-1_InterBirthInterval_d = -3706000.22; (16) 15-1_LitterSize -2147613.93000001; (17) 16-1_LittersPerYear = -3727577.23; (18) 17-1_MaxLongevity_m = -3419427.87; (19) 5-3_NeonateBodyMass_g = 7897056.36899; (20) 13-2_NeonateHeadBodyLen_mm = -4323358.91; (21) 21-1_PopulationDensity_n/km2 = -2939604.31021; (22) 10-1_PopulationGrpSize = -2882537.57; (23) 23-1_SexualMaturityAge_d = -2988989.75; (24) 10-2_SocialGrpSize = -3933491.88; (25) 24-1_TeatNumber = -3982491.86; (26) 12-2_Terrestriality = -1951847; (27) 6-2_TrophicLevel = -2469332; (28) 25-1_WeaningAge_d = -3343767.53999999; (29) 5-4_WeaningBodyMass_g = 22498611.15; (30) 13-3_WeaningHeadBodyLen_mm = -4533545.76; (31) 5-5_AdultBodyMass_g_EXT = -3175645.3; (32) 16-2_LittersPerYear_EXT -4236199.01000001; (33) 5-6_NeonateBodyMass_g_EXT =  -59305.9999999998;  (34) 5-7_WeaningBodyMass_g_EXT = 3973958.5; (35) 26-1_GR_Area_km2 = 7866119721; (36) 26-2_GR_MaxLat_dd = -299314.0548; (37) 26-3_GR_MinLat_dd = -359359.625670002; (38) 26-4_GR_MidRangeLat_dd -329337.231889999; (39) 26-5_GR_MaxLong_dd = -213097.97; (40) 26-6_GR_MinLong_dd = -307813.81389; (41) 26-7_GR_MidRangeLong_dd = -260457.328693106; (42) 27-1_HuPopDen_Min_n/km2 = -294370; (43) 27-2_HuPopDen_Mean_n/km2 = -22370.5697; (44) 27-3_HuPopDen_5p_n/km2 = -282940.5; (45) 27-4_HuPopDen_Change = -394196.188091501; (46) 28-1_Precip_Mean_mm = 25792.15; (47) 28-2_Temp_Mean_01degC = 300029.089999999; (48) 29-1_AET_Mean_mm = 3216946.29000001; and (49) 29-2_PET_Mean_mm = 4755394.55. .
Sums of the numeric columns for
 
    PanTHERIA1.0_WR05_Aug2008.txt:  (1) 1-1_ActivityCycle = -3750219; (2) 5-1_AdultBodyMass_g = 627931538.920001; (3) 8-1_AdultForearmLen_mm = -4458863.50999997; (4) 13-1_AdultHeadBodyLen_mm = -2390461.5599999900;  (5) 2-1_AgeatEyeOpening_d = -4928216.74;  (6) 3-1_AgeatFirstBirth_d = -4597710.77999999;  (7) 18-1_BasalMetRate_mLO2hr = -3761625.05;  (8) 5-2_ BasalMetRateMass_g = -229229.44;  (9) 6-1_DietBreadth = -3246141; (10) 7-1_DispersalAge_d = -5208296.92; (11) 9-1_GestationLen_d = -3898706.93999997;  (12) 12-1_HabitatBreadth = -2685754;  (13) 22-1_HomeRange_km2 = -4538723.81603595;  (14) 22-2_HomeRange_Indiv_km2 = -4580748.63033297;  (15) 14-1_InterBirthInterval_d = -4481414.57;  (16) 15-1_LitterSize = -2905774.13999999;  (17) 16-1_LittersPerYear = -4515784.58;  (18) 17-1_MaxLongevity_m = -4199598.31;  (19) 5-3_NeonateBodyMass_g = 7124135.98899836;  (20) 13-2_NeonateHeadBodyLen_mm = -5109030.42;  (21) 21-1_PopulationDensity_n/km2 = -3694898.15021332;  (22) 10-1_PopulationGrpSize = -3667882.43;  (23) 23-1_SexualMaturityAge_d = -3668760.78999999;  (24) 10-2_SocialGrpSize = -4699371.08;  (25) 24-1_TeatNumber = -4766713;  (26) 12-2_Terrestriality = -2773092;  (27) 6-2_TrophicLevel = -3247567;  (28) 25-1_WeaningAge_d = -4109794.78999999;  (29) 5-4_WeaningBodyMass_g = 21720528.5299999;  (30) 13-3_WeaningHeadBodyLen_mm = -5319075.47;  (31) 5-5_AdultBodyMass_g_EXT = -3579884.12;  (32) 16-2_LittersPerYear_EXT = -5016415.79; (33) 5-6_NeonateBodyMass_g_EXT = -948179.94; (34) 5-7_WeaningBodyMass_g_EXT = 3344775.98; (35) 26-1_GR_Area_km2 = 7755657748.19744; (36) 26-2_GR_MaxLat_dd = -687919.002494141; (37) 26-3_GR_MinLat_dd = -748817.5456675; (38) 26-4_GR_MidRangeLat_dd = -718368.751886152; (39) 26-5_GR_MaxLong_dd = -581739.880000002;  (40) 26-6_GR_MinLong_dd = -676884.413893151;  (41) 26-7_GR_MidRangeLong_dd = -629313.481315651;  (42) 27-1_HuPopDen_Min_n/km2 = -675228;  (43) 27-2_HuPopDen_Mean_n/km2 = -378781.01794477;  (44) 27-3_HuPopDen_5p_n/km2 = -662777;  (45) 27-4_HuPopDen_Change = -789755.285668012; (46) 28-1_Precip_Mean_mm = -334931.200000001; (47) 28-2_Temp_Mean_01degC = -35609.3699999999; (48) 30-1_AET_Mean_mm = 3142189.33;  and (49) 30-2_PET_Mean_mm = 4795170.44.
B. Variable definitions
30  specific variables were extracted for PanTHERIA (130). See Table 1 for  definitions, total number of data lines (before error checking), and species coverage (after error checking) according to the classifications of Wilson and Reeder (1993) (NSpp MSW93) and  Wilson and Reeder (2005) (NSpp MSW05). 23 variables (3153) were derived from these  variables or other data sources and are described  in Class V, Section C.
TABLE  1. Variable definitions.
  
    | Variable: Variable    Definition
 | Units | Data lines | NSpp MSW93 | NSpp MSW05 | 
  
    | (1) 1-1_ActivityCycle:    Activity cycle of each species measured for non-captive populations; adult or    age unspecified individuals, male, female, or sex unspecified individuals;    primary, secondary, or extrapolated sources; all measures of central tendency;    in all localities. Species were defined as (1) nocturnal only, (2)    nocturnal/crepuscular, cathemeral, crepuscular or diurnal/crepuscular and (3)    diurnal only. | NA | 2724 | 1646 | 1659 | 
  
    | (2) 5-1_AdultBodyMass_g: Mass of adult (or age unspecified) live or    freshly-killed specimens (excluding pregnant females) using captive, wild, provisioned, or unspecified populations; male, female, or sex unspecified individuals; primary, secondary, or extrapolated sources; all measures of    central tendency; in all localities. | grams | 17528 | 3473 | 3542 | 
  
    | (3) 8-1_AdultForearmLen_mm: Total    length from elbow to wrist of adult (or age unspecified) live, freshly-killed, or museum specimens using captive, wild, provisioned, or unspecified populations; male, female, or sex unspecified individuals; primary, secondary, or extrapolated sources; all measures of central tendency; in all localities. | mm | 1952 | 897 | 903 | 
  
    | (4) 13-1_AdultHeadBodyLen_mm: Total length    from tip of nose to anus or base of tail of adult (or age unspecified) live, freshly-killed, or museum specimens using captive, wild, provisioned, or unspecified populations; male, female, or sex unspecified individuals;    primary, secondary, or extrapolated sources; all measures of central tendency;    in all localities. | mm | 4941 | 1907 | 1941 | 
  
    | (5) 2-1_AgeatEyeOpening_d: Age at which    both eyes are fully open after birth using captive, wild, provisioned, or unspecified populations; male, female, or sex unspecified individuals;    primary, secondary, or extrapolated sources; all measures of central tendency;    in all localities | days | 710 | 471 | 476 | 
  
    | (6) 3-1_AgeatFirstBirth_d: Age at which    females give birth to their first litter (eutherians), or their young attach    to teats (metatherians) or hatch out (monotremes), using non-captive, wild, provisioned, or unspecified populations; primary, secondary, or extrapolated sources; all measures of central tendency; in all localities. | days | 780 | 440 | 445 | 
  
    | (7) 18-1_BasalMetRate_mLO2hr: Basal    metabolic rate of adult (or age unspecified) individual(s) using captive,    wild, provisioned, or unspecified populations; male, female, or sex unspecified individuals; primary, secondary, or extrapolated sources; all measures of    central tendency; in all localities. Metabolic rate was measured when    individual(s) were experiencing neither heat nor cold stress (i.e. are in    their thermoneutral zone); are resting and calm; and are post–absorptive (are    not digesting or absorbing a meal) and data were only accepted where there    was also a measure of body mass for the same individual(s). | mL.O2/hr | 971 | 559 | 573 | 
  
    | (8) 5-2_BasalMetRateMass_g: Mass of    individual(s) from which the basal metabolic rate was taken. | grams | 971 | 559 | 573 | 
  
    | (9) 6-1_DietBreadth: Number of dietary    categories eaten by each species measured using any qualitative or    quantitative dietary measure, over any period of time, using any assessment    method, for non-captive or non-provisioned populations; adult or age    unspecified individuals, male, female, or sex unspecified individuals;    primary, secondary, or extrapolated sources; all measures of central tendency;    in all localities. Categories were defined as vertebrate, invertebrate,    fruit, flowers/nectar/pollen, leaves/branches/bark, seeds, grass and    roots/tubers. | No. of diet categories | 4631 | 2153 | 2161 | 
  
    | (10) 7-1_DispersalAge_d: Age at which young    permanently leave the parent, parental group, social group or any associated    territories, using non-captive, wild, non-provisioned or unspecified    populations; male, female, or sex unspecified individuals; primary, secondary, or extrapolated sources; all measures of central tendency; in all localities. | days | 175 | 145 | 144 | 
  
    | (11) 9-1_GestationLen_d: Length of time of non-inactive fetal growth, using captive, wild, provisioned, or unspecified populations; male, female, or sex unspecified individuals; primary, secondary, or extrapolated sources; all measures of central tendency; in all localities. Gestation was measured between specified start and end points as follows: Start points – conception, fertilization, first observed copulation, fertilization, implantation, laying, palpably pregnant, removal of pouch young, capture (except marsupials) or unspecified. End points – birth, hatching or unspecified. | days | 6537 | 1354 | 1362 | 
  
    | (12) 12-1_HabitatBreadth: Number of habitat layers used by each species measured using any qualitative or quantitative time measure, for non-captive populations; adult or age unspecified individuals, male, female, or sex unspecified individuals; primary, secondary, or extrapolated sources; all measures of central tendency; in all localities. Categories were defined as above ground dwelling, aquatic, fossorial and ground dwelling. | No. of habitat layers | 3933 | 2759 | 2724 | 
  
    | (13) 22-1_HomeRange_km2: Size of the area within which everyday activities of individuals or groups (of any type) are typically restricted, estimated by either direct observation, radio telemetry, trapping or unspecified methods over any duration of time, using non-captive populations; male, female, or sex unspecified individuals; primary, secondary, or extrapolated sources; all measures of central tendency; in all localities. | km2 | 3269 | 699 | 707 | 
  
    | (14) 22-2_HomeRange_Indiv_km2: Size of the area within which everyday activities of individuals are typically restricted, estimated by either direct observation, radio telemetry, trapping or unspecified methods over any duration of time, using non-captive populations; male, female, or sex unspecified individuals; primary, secondary, or extrapolated sources; all measures of central tendency; in all localities. | km2 | 2675 | 624 | 626 | 
  
    | (15) 14-1_InterBirthInterval_d: The length of time between successive births of the same female(s) after a successful or unspecified litter using non-captive, wild, provisioned, or unspecified populations; primary, secondary, or extrapolated sources; all measures of central tendency; in all localities. | days | 1732 | 687 | 698 | 
  
    | (16) 15-1_LitterSize: Number of offspring born per litter per female, either counted before birth, at birth or after birth, using captive, wild, provisioned, or unspecified populations; male, female, or sex unspecified individuals; primary, secondary, or extrapolated sources; all measures of central tendency; in all localities. | no. of offspring | 15020 | 2473 | 2501 | 
  
    | (17) 16-1_LittersPerYear: Number of litters per female per year using non-captive, wild, provisioned, or unspecified populations; male, female, or sex unspecified individuals; primary, secondary, or extrapolated sources; all measures of central tendency; in all localities. | no. of litters | 2501 | 896 | 894 | 
  
    | (18) 17-1_MaxLongevity_m: Maximum adult age    measured either through direct observation, capture-recapture estimates, projected    from physical wear or unspecified, using captive, wild, provisioned, or unspecified populations; male, female, or sex unspecified individuals;    primary, secondary, or extrapolated sources; in all localities. | months | 3294 | 1009 | 1013 | 
  
    | (19) 5-3_NeonatalBodyMass_g: Mass of live    or freshly-killed specimens of infants at either a near term embryonic stage,    birth, immediately after birth or up to an age of seven days after birth,    using captive, wild, provisioned, or unspecified populations; male, female, or sex unspecified individuals; primary, secondary, or extrapolated sources; all    measures of central tendency; in all localities. | grams | 4308 | 1078 | 1085 | 
  
    | (20) 13-2_NeonateHeadBodyLen_mm : Total    length from tip of nose to anus or base of tail of live, freshly-killed, or museum    specimens of infants at either a near term embryonic stage, birth,    immediately after birth or up to an age of seven days after birth, using    captive, wild, provisioned, or unspecified populations; male, female, or sex unspecified individuals; primary, secondary, or extrapolated sources; all    measures of central tendency; in all localities. | mm | 346 | 226 | 226 | 
  
    | (21) 21-1_PopulationDensity: Number of    individuals per square kilometer, estimated with either direct, indirect or    unspecified counts, measured in any area size within a human, ecological or    unspecified boundary, over any duration of time, using non-captive,    non-provisioned populations; male, female, or sex unspecified individuals;    primary, secondary, or extrapolated sources; all measures of central tendency;    in all localities. | number /km2 | 5244 | 929 | 956 | 
  
    | (22) 10-1_PopulationGrpSize: Number of    individuals, adults or definition unspecified in a group that spends the    majority of their time in a 24 hour cycle together, measured over any    duration of time, using non-captive populations; male, female, or sex unspecified individuals; primary, secondary, or extrapolated sources; all    measures of central tendency; in all localities. | no. of individuals, adults or unspecified | 905 | 386 | 388 | 
  
    | (23) 23-1_SexualMaturityAge_d: Age when    individuals are first physically capable of reproducing, defined as either    physically sexually mature, age at first mating or unspecified (males and    females), age at first estrus or age at first pregnancy (females only), age    at spermatogenesis or age at testes descent (males only), using captive,    wild, provisioned, or unspecified populations; male, female, or sex unspecified individuals, primary, secondary, or extrapolated sources; all measures of    central tendency; in all localities.  | days | 4680 | 1042 | 1051 | 
  
    | (24) 10-2_SocialGrpSize: Number of    individuals, adults or definition unspecified in a group that spends the    majority of their time in a 24 hour cycle together where there is some    indication that these individuals form a social cohesive unit, measured over    any duration of time, using non-captive populations; male, female, or sex unspecified individuals; primary, secondary, or extrapolated sources; all    measures of central tendency; in all localities. | no. of individuals, adults or unspecified | 1598 | 687 | 707 | 
  
    | (25) 24-1_TeatNumber: Total number of teats    present, using captive, wild, provisioned, or unspecified populations; male, female, or sex unspecified individuals; primary, secondary, or extrapolated sources; all measures of central tendency; in all localities. | No. of teats | 841 | 639 | 641 | 
  
    | (26) 12-2_Terrestriality: Degree of    terrestriality of each species measured using any qualitative or quantitative    time measure, for non-captive populations; adult or age unspecified    individuals, male, female, or sex unspecified individuals; primary, secondary, or extrapolated sources; all measures of central tendency; in all localities.    Species were defined as (1) fossorial and/or ground dwelling only and (2)    above ground dwelling. | NA | 3933 | 2671 | 2636 | 
  
    | (27) 6-2_TrophicLevel: Trophic level of    each species measured using any qualitative or quantitative dietary measure,    over any period of time, using any assessment method, for non-captive or    non-provisioned populations; adult or age unspecified individuals, male, female, or sex unspecified individuals; primary, secondary, or extrapolated sources; all measures of central tendency; in all localities. Species were    defined as (1) herbivore (not vertebrate and/or invertebrate), (2) omnivore    (vertebrate and/or invertebrate plus any of the other categories) and (3)    carnivore (vertebrate and/or invertebrate only) | NA | 4644 | 2153 | 2161 | 
  
    | (28) 25-1_WeaningAge_d: Age when primary    nutritional dependency on the mother ends and independent foraging begins to    make a major contribution to the offspring’s energy requirements, measured as    either weaning/lactation length, nutritionally independent, first solid food,    last observed nursing, age at first flight (bats only), age at pouch exit or    length of teat Attachment (marsupials only) or unspecified definition, using    captive, wild, provisioned, or unspecified populations; male, female, or sex unspecified individuals; primary, secondary, or extrapolated sources; all    measures of central tendency; in all localities. | days | 4858 | 1147 | 1164 | 
  
    | (29) 5-4_WeaningBodyMass_g: Mass of live or    freshly-killed specimens of weanlings, using captive, wild, provisioned, or unspecified populations; male, female, or sex unspecified individuals;    primary, secondary, or extrapolated sources; all measures of central tendency;    in all localities. | grams | 1106 | 481 | 487 | 
  
    | (30) 13-3_WeaningHeadBodyLen_mm: Total    length from tip of nose to anus or base of tail of live, freshly-killed, or museum specimens of weanlings, using captive, wild, provisioned, or unspecified populations; male, female, or sex unspecified individuals;    primary, secondary, or extrapolated sources; all measures of central tendency;    in all localities. | mm | 60 | 46 | 47 | 
  
    | References: Index numbers referring to sources of data for    each species according to the list of references given in section F below. |  |  |  |  | 
Calculating  measures of central tendency
We often obtained more than one valid  datum for a particular variable within a species, e.g., from different subspecies,  populations, years or localities (mean per species per variable for species  with data = 3.3, n = 94729 data lines). For  each continuous variable, we calculated a single measure of central tendency  for each species from the different estimates (as described in Table 2). Categorical  variables and values for maximum longevity were treated differently and are as  described in Table 1. Two factors may influence the measures of central  tendency obtained. Firstly, outliers may bias the value. We ameliorated this  problem by calculating the median of the multiple values for each species,  which is less sensitive to outliers than the mean. For variables measured as  real numbers (e.g. body mass), we calculated medians of unique values, to guard  against duplicate entries, but for the variables that are more often measured  as integers, i.e. litters per year and teat number, we calculated medians of  all values (see Table 2). Secondly, different combinations of the delimiters  used for each variable (e.g., age, sex, captivity status, definition of  maturity stage) may influence the values of central tendency obtained in two ways:  different combinations of the delimiters may actually give unrealistic  information (e.g., using captive data to estimate interbirth intervals), or may  merely give systematically different values (e.g. male and female body sizes in  sexually dimorphic species), so that a species value will be shaped by the  relative frequencies of the different delimiters.
 To  minimize the first problem of including incorrect values, we removed a priori  from each variable the delimiter combinations that would generate unrealistic  data (see Table 1). To ameliorate the second problem while still maximizing  sample sizes, we estimated species values for those variables that contained  delimiters that were likely to produce significantly different values (15  variables, see Table 2) using a general linear model (GLM) setting species and  the appropriate delimiters as factors (performed separately for each order for (2) 5-1_AdultBodyMass_g and (16) 15-1_LitterSize, and across the whole class  for the other 13 variables). For example, the model fitted to sexual maturity  age was: ln(sexual maturity age) = species + sex + sexual maturity definition.  This is because males and females may reach maturity at different ages, and the  recorded maturity age may also depend on the way it is defined. For each  factor, we assigned the level with the largest number of values as the base  level, against which the model could calculate offsets for all of the other  levels (see Table 2). The offsets were then subtracted from each of the  original values to give an adjusted value. For cases where there were single  values for a species, we calculated adjusted values for these species using as  offsets the mean offset value for all other species. In effect, the GLM can be  thought of as converting all of the data into the same currency (that of the  base factor), by adding an offset that depends upon the delimiters and which is  calculated directly from the data themselves. Hence, for age at sexual maturity  the offset for values measured as “age at first pregnancy” is 0.04 (on a log  scale), relative to the baseline definition, so each original value measured in  this way is reduced by this amount.
For these 15 traits,  a central tendency measure was then calculated from the logged adjusted values  using unique medians (see Table 2). This method, therefore, takes into account  the fact that not all data values for some variables are in the same currency,  though they may all contain useful information about the same underlying  variable. At the same time, it allows a central tendency measure of a variable  to be calculated for all species for which any data have been recorded,  avoiding the need to discard valid data. We term this process consolidation,  and values generated in this way consolidated values.
In one case it was not  clear whether to remove the delimiters a priori or to include them in a GLM (home  range area measured for either groups or individuals – see Table 1 and 2). For this  case, we consolidated the data first including all values of range measure and  performing a GLM with group as a factor ((13) 22-1_HomeRange_km2) and second, excluding the data lines for group  home ranges ((14) 22-2_HomeRange_Indiv_km2).  Details of the GLM model specifications and the consolidated values for each  variable are given in Table 2. The consolidated values analysis was implemented  in the program R (R Development Core Team  2005).
TABLE 2. Methods to generate measures of central  tendency for species-level traits.
  
    | Variable | GLM model specification[method]
 | Base factor | Central tendency measure | 
  
    | (1) 1-1_ActivityCycle | None | None | Categorical | 
  
    | (2) 5-1_AdultBodyMass_g  | ln(body mass) = species + sex  [within orders] | Sex = both | Unique median  | 
  
    | (3) 8-1_AdultForearmLen_mm | ln(forearm length) = species + sex [across class] | Sex = unspecified | Unique median  | 
  
    | (4) 13-1_AdultHeadBodyLen_mm | ln(head-body length) = species + sex [across class] | Sex = unspecified | Unique median  | 
  
    | (5) 2-1_AgeatEyeOpening_d | None | None | Unique median | 
  
    | (6) 3-1_AgeatFirstBirth_d | None | None | Unique median | 
  
    | (7) 18-1_BasalMetRate_mLO2hr | None | None | Unique median | 
  
    | (8) 5-2_BasalMetRateMass_g | None | None | Unique median | 
  
    | (9) 6-1_DietBreadth | None | None | Sum of diet categories | 
  
    | (10) 7-1_DispersalAge | ln(dispersal age) = species + sex [across class] | Sex = unspecified | Unique median | 
  
    | (11) 9-1_GestationLen_d | ln(gestation length) = species + start point + end point    [across class] | Start point =  unspecified, End point = unspecified | Unique median | 
  
    | (12) 12-1_HabitatBreadth | None | None | Sum of habitat layers | 
  
    | (13) 22-1_HomeRange_km2 | ln(home range) = species + sex + estimation method + range  measure [across class] | Sex = unspecified, Estimation method = unspecified, Range  measure = individual | Unique median | 
  
    | (14) 22-2_HomeRange_Indiv_km2 | ln(home range) = species + sex + estimation method    [[across class] | Sex = unspecified, Estimation method = unspecified | Unique median | 
  
    | (15) 14-1_InterBirthInterval_d | None | None | Unique median | 
  
    | (16) 15-1_LitterSize | ln(litter size) = species + litter size definition [within    orders] | Litter size definition = unspecified | Unique median | 
  
    | (17) 16-1_LittersPerYear | None | None | Full median | 
  
    | (18) 17-1_MaxLongevity  | None | None | Maximum | 
  
    | (19) 5-3_NeonateBodyMass  | ln(body mass) = species + sex + age [across class] | Sex = unspecified, Age = neonate | Unique median | 
  
    | (20) 13-2_NeonateHeadBodyLen_mm | ln(head-body length) = species + sex + age [across class] | Sex = unspecified, Age = neonate | Unique median | 
  
    | (21) 21-1_PopulationDensity_n/km2 | ln(population density) = species + estimation method + study    area type [across class] | Estimation method = unspecified, Study area type =    unspecified | Unique median  | 
  
    | (22) 10-1_PopulationGrpSize | None | None | Full median | 
  
    | (23) 23-1_SexualMaturityAge_d | ln(sexual maturity age) = species + sex + sexual maturity    definition [across class] | Sex = female, Sexual maturity definition = unspecified | Unique median  | 
  
    | (24) 10-2_SocialGrpSize | None | None | Full median | 
  
    | (25) 24-1_Teat Number | None | None | Full median | 
  
    | (26) 12-2_Terrestriality | None | None | Categorical | 
  
    | (27) 6-2_TrophicLevel | None | None | Categorical | 
  
    | (28) 25-1_WeaningAge | ln(weaning age) = species + weaning age definition [across    class] | Weaning age definition = weaning/lactation length | Unique median  | 
  
    | (29) 5-4_WeaningBodyMass_g | ln(body mass) = species + sex [across class] | Sex = unspecified | Unique median | 
  
    | (30) 13-3_WeaningHeadBodyLen_mm | ln(head-body length) = species + sex [across class] | Sex = unspecified | Unique median | 
 
CLASS V. SUPPLEMENTAL DESCRIPTORS
A.  Data acquisition.
Data  forms: We developed a standardized data input protocol using a Visual Basic  interface programmed within Excel (MammalFormv2.xls) and input protocols (MammalFormv2Protocols.pdf),  see Class V, Section D. Data lines were  outputted as text files and stored in an Access database.
Location of completed data forms: Raw data is held in an Access database at The  Zoological Society of London. These data and input protocols are tools we used  to develop the database which are no longer current. We are now in the process  of updating these protocols and adding additional data through the development  of an online database web portal (YouTHERIA) (contact KEJ for further details).
B.  Quality assurance/quality control procedures.
The accuracy of these data  was verified by a number of methods:
Taxonomic Accuracy: For each datum, its genus, species, subspecies and  authority (publication author) was recorded as given in the source. These  original taxon names were transformed  into valid Latin binomials using the most comprehensive mammalian taxonomy  available, that of Wilson and Reeder. We used the two most recent versions (Wilson and Reeder 1993, 2005) producing a separate data table for each (PanTHERIA1.0_WR93_Aug2008.txt  and PanTHERIA1.0_WR05_Aug2008.txt). In most cases, there was an exact match  between source names and currently valid names. Where there was a difference,  spelling discrepancies accounted for the majority of the mismatches. Further  differences were caused because the species was described since version  compilation. In these cases the names were valid but unrecognized. The  remaining differences were caused by changes to the name given in the source  (synonymisation). We used the synonym lists given in Wilson and Reeder (1993, 2005) to trace the original name to its current usage. By  keeping the taxonomic name given in each datum’s source, we are able to  accommodate many changes in taxonomy and classification without losing data. However,  in some cases it was not possible to unambiguously determine taxonomic affiliation  (2002 data lines for Wilson and Reeder 1993); (2147 data lines for Wilson and Reeder 2005) and these data were not used in any subsequent  analysis. This taxonomic conversion file (SppSynonymID1.0.txt) is available  from Class V, Section D. 
    Input Error:  At monthly intervals, all newly inputted data were transformed into a common  taxonomic and unit currency (e.g., data on population densities converted into  individuals per km2 from the different units given in the sources).  All new data were checked for obvious aberrant values, and returned to the  original data miner for checking and, where appropriate, correction.
Outlier Analysis:  We identified data values in the specific variables that were significant  outliers from general trends (standardized residual >+3). These  outliers were then checked and, where necessary, corrected or excluded from  subsequent analysis. Two types of outliers were identified; firstly, an ANOVA  was performed with taxonomic level as the grouping variable to identify data  lines unusual with respect to their order, family or genus. Secondly, an ANCOVA  was performed to identify outliers that were unusual with respect to their  allometric relationship with adult body mass. We generated a mass variable for  checking the other variables using an earlier version of (2) 5-1_ AdultBodyMass_g that had been  checked for taxonomic outliers and summarized into species value medians (but  not adjusted). This file was used to check (4) 13-1_AdultHeadBodyLen_mm and (3) 8-1_AdultForearmLen_mm files to create checked, species value  medians (but not adjusted) versions of these traits. Then we generated  additional mass values from head-body lengths or forearm lengths for species  that were missing mass data using the following equations. For MSW93 species: non-bats  ln Adult Body Mass = -9.570 + 2.842 * (ln Adult Head-Body Length) and bats ln Adult  Body Mass = -8.255 + 2.852 * (ln Adult Forearm Length). For MSW05 species: non-bats  ln Adult Body Mass = -9.580 + 2.844 * (ln Adult Head Body Length) and bats ln Adult  Body Mass = -8.298 + 2.864 * (ln Adult Forearm Length). Using  this combined mass file we performed allometric checks for all the remaining  variables except those variables that take qualitative values and could not be  checked this way, i.e.; (1) 1-1_ActivityCycle,  (9) 6-1_DietBreadth, (12) 12-1_HabitatBreadth,  (26) 12-2_Terrestriality and (27) 6-2_TrophicLevel. The outlier analysis  was implemented in R (R Development Core Team 2005).
    Data  Duplication: As our data were collected from both primary and secondary  literature sources, data from primary sources may be duplicated within  secondary sources. This is easy to identify when the secondary source lists all  primary data separately as the duplicate values will be identical. However,  when the secondary source summarizes the data value for a number of primary  sources then the value will be different and duplication harder to identify. To  ameliorate these problems we firstly removed from subsequent analysis  non-obvious duplications that we knew of (e.g., bat body mass data in (Smith et al. 2003) is all from the primary data  in (Jones 1998) which is already incorporated  into the database). Secondly, we used techniques to calculate a value of  central tendency that minimized the effects of identical duplicate values (see Calculating  measures of central tendency in Class IV, Section B).
C. Related material
We used our existing  variables and other spatial databases to calculate 23 additional variables.
Derived  variables from existing data. Many  of our variables are highly intercorrelated (i.e., one explains a high degree of  variation in the other). This is particularly true of size variables in general  (Calder 1996). For example, head-body length is an excellent  predictor of body mass. We used this fact to estimate fitted values for some  variables by fitting across-species GLMs using the species medians of the consolidated values (see Class IV, Section B) for the following 4 variables:  (31) 5-5_AdultBodyMass_g_EXT (368 MSW93  species, 394 MSW05 species); (32) 16-2_LittersPerYear_EXT (388, 394 species); (33) 5-6_NeonateBodyMass_g_EXT (22, 24  species); (34) 5-7_WeaningBodyMass_g_EXT  (24, 25 species) (see Table 3 for details  of the models). The analysis was implemented in R (R Development Core Team  2005).
Table 3. Models for Extrapolated Variables. Which of the three bat models was used depended on whether a species had a value for adult head-body length, adult forearm length, or both.

Derived variables  from other data. We derived 19  variables using data from other sources. Firstly, we used digital geographic  range maps of all extant, non-marine mammals from Sechrest  (2003) and converted them to the two taxonomies used: Wilson and Reeder (1993, 4271 species) and Wilson  and Reeder (2005, 4668 species). These ranges were used to generate 7 measures of  geographic range extent and occupancy. Spatial analyses were performed using  ArcGIS (version 9.0) (ESRI 2005) for areas and R (R Development Core Team 2005) for geographic coordinates. The calculated variables  were as follows: (35) 26-1_GR_Area_km2  (4271, 4668 species), calculated using the total extent of a species range with  a global equal-area projection (Mollweide); (36) 26-2_GR_MaxLat_dd (4271, 4668 species), maximum latitudinal extent  of each species range calculated using a global geographic projection (decimal  degrees); (37) 26-3_GR_MinLat_dd (4271,  4668 species), minimum latitudinal extent of each species range calculated  using a global geographic projection (decimal degrees); (38) 26-4_GR_MidRangeLat_dd (4271, 4668 species),  median latitudinal extent of each species range calculated using a global  geographic projection (decimal degrees); (39) 26-5_GR_MaxLong_dd (4271, 4668 species), maximum longitudinal  extent of each species range calculated using a global geographic projection  (decimal degrees); (40) 26-6_GR_MinLong_dd  (4271, 4668 species), minimum longitudinal extent of each species range  calculated using a global geographic projection (decimal degrees); (41) 26-7_GR_MidRangeLong_dd (4271, 4668 species),  median longitudinal extent of each species range calculated using a global  geographic projection (decimal degrees). Secondly, we generated a number of  variables for each species describing the anthropogenic and environmental conditions  within each geographic range  based on the extent of the digital species range maps. The species ranges were  first gridded at 0.5 degrees using a global geographic projection and species  were scored as present in each cell if any of the range intersected that cell. We  then calculated summary statistics for each species by summarizing the value for  each anthropogenic and environmental variable (gridded to the same extent) for  all the cells the species was present. Except for the variables 27-1_HuPopDen_Min_n/km2 and 27-3_HuPopDen_5p_n/km2, we adjusted the  summary statistics for each variable for the fact that area varies with  latitude by multiplying each cell by the area and then dividing by the total  area of  cells for that variable for each  species. The anthropogenic and environmental variables  calculated across the geographic ranges of each species ranges were as follows:  (46) 27-1_HuPopDen_Min_n/km2 (4271, 4668  species), minimum human population density (persons per km2) using  the Gridded Population of the World (GPW) (CIESIN and CIAT 2005) for 1995; (47) 27-2_HuPopDen_Mean_n/km2  (4271, 4668 species), mean human population density (persons per km2)  using the Gridded Population of the World (GPW) (CIESIN and CIAT 2005) for 1995; (48) 27-3_HuPopDen_5p_n/km2  (4271, 4668 species), 5th percentile human population density  (persons per km2) using the Gridded Population of the World (GPW) (CIESIN and CIAT 2005) for 1995; (49) 27-4_HuPopDen_Change  (4234, 4625 species), mean rate of increase in human population density using  the Gridded Population of the World (GPW) (CIESIN and CIAT 2005) for 1990 and 1995 as: (19951990)/1990; (50) 28-1_Precip_Mean_mm (4160, 4533 species)  mean monthly precipitation (mm) calculated using data from
  ftp://ftp.ngdc.noaa.gov/Solid_Earth/Ecosystems/GEDII_a/datasets/a03/lc.htm;  (51) 28-2_Temp_Mean_01degC (4160, 4533  species), mean monthly temperature (0.1°C) calculated using data  from
 ftp://ftp.ngdc.noaa.gov/Solid_Earth/Ecosystems/GEDII_a/datasets/a03/lc.htm;  (52) 30-1_AET_Mean_mm  (3973, 4305 species), mean monthly AET (Actual  Evapotranspiration Rate) from 1920 to 1980 (mm) calculated using the Global  Resource Information Database of UNEP and is available from
 http://www.grid.unep.ch/data/grid/gnv183.php),  and (53) 30-2_PET_Mean_mm (3973, 4305  species), mean monthly PET (Potential Evapotranspiration Rate) from 1920 to  1980 (mm) calculated using the Global Resource Information Database of UNEP and  is available from
 http://www.grid.unep.ch/data/grid/gnv183.php).
D. Computer programs and data processing algorithms.
The following files were  developed to collate or generate our species-level data sheets:
(1) MammalFormv2.1.xls. A Visual Basic program within Excel that uses a standardized  data input protocol to generate text files which are imported into PanTHERIA.
(2) MammalFormv2.1.pdf. Protocols to inputting data into  MammalForm v2.
(3) SppSynonymID1.0.txt. Taxonomic conversion file used to convert  names in original sources into the taxonomy of Wilson and Reeder (1993) and Wilson  and Reeder (2005).
(4) MammalFormVB.zip. Provides VBA code for the data input forms, along with a binary file which contains the layout of each form and fields within it. MammalForm.bas contains the bulk of the code for the main form. 
E. Archiving: n/a
F. Publications and results
   Publications citing the data set
  These data and earlier versions of these  data sets have been cited in the following publications: Jones et al. (2005); Cardillo et al. (2005); Cardillo et al. (2006); Ezenwa et al. (2006); Grenyer et al. (2006); Jeschke and Strayer (2006); Bielby et al. (2007); Lindenfors et  al. (2007a) Lindenfors et al. (2007b); Pedersen et al. (2007); Price and Gittleman (2007); Cardillo et al. (2008); Davies et al. (2008); Shai et al. (2008); Welch et al. (2008); Jones et al. (in press); Cardillo and Purvis (in preparation). This version of PanTHERIA differs from earlier  versions because of minor corrections in the values for the related material. Earlier  versions can be obtained from the first author on request.
  Reference list for the data set: References are given as indexed values in  accordance with the following reference list.
  1.         Abbas, A. 1991. Feeding  Strategy of Coypu (Myocastor coypus)  in Central Western France. Journal of Zoology 224:385-401.
2.         Abe, Y. 1931. A Synopsis  of the Leporine Mammals of Japan.  Journal of Science of the Hiroshima   University, Series B,  Zoology 1:45-63.
3.         Abenspergtraun, M., C.  R. Dickman, and E. S. Deboer. 1991. Patch Use and Prey Defense in a Mammalian  Myrmecophage, the Echidna (Tachyglossus  aculeatus) (Monotremata, Tachyglossidae) - a Test of Foraging Efficiency in  Captive and Free-Ranging Animals. Journal of Zoology 225:481-493.
4.         Abenspergtraun, M., and  E. S. Deboer. 1992. The Foraging Ecology of a Termite-Eating and Ant-Eating  Specialist, the Echidna Tachyglossus  aculeatus (Monotremata, Tachyglossidae). Journal of Zoology 226:243-257.
5.         Acevedo, A., and S.  Burkhart. 1998. Seasonal distribution of bottlenose (Tursiops truncatus) and pan-tropical spotted (Stenella attenuata) dolphins (Cetacea: Delphinidae) in Golfo Dulce, Costa Rica. Revista de Biologia  Tropical 46.
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286.    Bernard, R. T. F. 1994.  Reproductive synchrony and annual variaiton in foetal growth rate in the  long-fingered bat Miniopterus schreibersii. Journal of Zoology 232:485-490.
287.    Bernard, R. F. T., D. C. D.  Happold, and M. Happold. 1995. Sperm storage in the Banana Bat (Pipistrellus  nanus) from a tropical latitude in Africa. Bat  Research News 36:48.
288.    Bernard, R. T. F., and J.  N. Tsita. 1995. Seasonally monoestrous reproduction in the molossid bat,  Tadarida aegyptiaca from low temperate latitudes (33 degrees South) in South Africa.  South African Journal of  Zoology 30:18-22. 
289.    Bernard, R. T. F., F. P. D.  Cotterill, and R. A. Fergusson. 1996. On the occurrence of a short period of  delayed implantation in Schreiber's long-fingered bat (Miniopterus schreibersii) from a tropical latitude in Zimbabwe.  Journal of Zoology 238:13.
290.    Berrow, S. D., B. Holmes,  and O. R. Kiely. 1996. Distribution and abundance of bottle-nosed Dolphins Tursiops truncatus (Montagu) in the  Shannon Estuary. Biology and Environment 96B:1-9 
291.    Berry, R. J. 1968. The Skokholm mouse:  chance and change. Skokholm Bird Observatory Reports 1967:30-35.
292.    Berry, A. J., T. J. C. Anderson, J. N. Amos,  and J. M. Cook. 1987. Spool-and-line tracking of giant rats in New Guinea.  Journal of Zoology 213:299-303.
293.    Berteaux, D. 2000.  Energetic costs of heating ingested food in mammalian herbivores. Journal of  Mammalogy 81:683-690.
294.    Berthoud, G. 1978. Note  preliminaire sur les deplacements du herrison europeen (Erinaceus europaeus). Terre Vie 32:73-82. 
295.    Bertolino, S., C. Viano,  and I. Currado. 2001. Population dynamics,  breeding patterns and spatial use of the garden dormouse (Eliomys quercinus) in an Alpine habitat. Journal of Zoology 253:513-521. 
296.    Best, T. L. 1973.  Ecological seperation of three genera of pocket gophers (Geomyidae). Ecology 54:1311-1319.
297.    Best, T. L., and J. A.  Lackey. 1985. Dipodomys gravipes.  Mammalian Species 236:1-4.
298.    Best, R. C., and A. Y.  Harada. 1985. Food-Habits of the Silky Anteater (Cyclopes didactylus) in the Central Amazon. Journal of Mammalogy 66:780-781. 
299.    Best, T. L., and B.  Hoditschek. 1986. Relationships between environmental variation and the  reproductive biology of Ord's kangaroo rat. Mammalia 50:173-183.
300.    Best, T. L. 1986. Dipodomys elephantinus. Mammalian  Species 255:1-4.
301.    Best, T. L. 1988. Dipodomys nelsoni. Mammalian Species 326:1-4.
302.    Best, T. L. 1988. Dipodomys spectabilis. Mammalian Species 311:1-10.
303.    Best, T. L., N. J.  Hildreth, and C. Jones. 1989. Dipodomys  deserti. Mammalian Species 339:1-8.
304.    Best, T. L., C. L. Lewis,  K. Caesar, and A. S. Titus. 1990. Ammospermophilus  interpres. Mammalian Species 365:1-6.
305.    Best, T. L., A. S. Titus,  K. Caesar, and C. L. Lewis. 1990. Ammospermophilus  harrisii. Mammalian Species 366:1-7.
306.    Best, T. L., A. S. Titus,  C. L. Lewis, and K. Caesar. 1990. Ammospermophilus  nelsoni. Mammalian Species 367:1-7.
307.    Best, T. L., K. Caesar, A.  M. Titus, and C. L. Lewis. 1990. Ammospermophilus  insularis. Mammalian Species 364:1-4.
308.    Best, T. L. 1991. Dipodomys nitratoides. Mammalian Species 381:1-7.
309.    Best, T. L., and H. H.  Thomas. 1991. Dipodomys insularis.  Mammalian Species 374:1-3.
310.    Best, T. L., and H. H.  Thomas. 1991. Spermophilus madrensis.  Mammalian Species 378:1-2.
311.    Best, T. L. 1992. Dipodomys venustus. Mammalian Species 403:1-4.
312.    Best, P. B., and H. Ruther.  1992. Aerial photogrammetry of southern right whales, Eubalaena australis. Journal of Zoology 228:595-614.
313.    Best, T. L. 1992. Dipodomys margaritae. Mammalian Species 400:1-3.
314.    Best, T. L., J. L. Bartig,  and S. L. Burt. 1992. Tamias canipes.  Mammalian Species 411:1-5. 
315.    Best, T. L., and J. A.  Lackey. 1992. Chaetodipus artus.  Mammalian Species 418:1-3.
316.    Best, T. L., and J. A.  Lackey. 1992. Chaetodipus pernix. Mammalian  Species 420:1-3.
317.    Best, T. L., and T. H.  Henry. 1993. Lepus alleni. Pages 1-8 in A. S. o. Mammalogists, editor.  Mammalian Species.
318.    Best, T. L. 1993. Tamias palmeri. Mammalian Species 443:1-6.
319.    Best, T. L. 1993. Tamias ruficaudus. Mammalian Species 452:1-7.
320.    Best, T. L. 1993. Tamias sonomae. Mammalian Species 444:1-5.
321.    Best, T. L., S. L. Burt,  and J. L. Bartig. 1993. Tamias durangae.  Mammalian Species 437:1-4.
322.    Best, T. L. 1993. Chaetodipus lineatus. Mammalian Species 451:1-3. 
323.    Best, T. L. 1993. Perognathus inornatus. Mammalian Species 450:1-5. 
324.    Best, T. L., and T. H.  Henry. 1993. Lepus callotis.  Mammalian Species:1-6. 
325.    Best, T. L., and T. H.  Henry. 1994. Lepus arcticus.  Mammalian Species:1-9.
326.    Best, T. L., and T. H.  Henry. 1994. Lepus othus. Mammalian  Species:1-5.
327.    Best, T. L. 1994. Chaetodipus nelsoni. Mammalian Species 484:1-6.
328.    Best, T. L., R. G. Clawson,  and J. A. Clawson. 1994. Tamias  panamintinus. Mammalian Species 468:1-7.
329.    Best, T. L., R. G. Clawson,  and J. A. Clawson. 1994. Tamias speciosus.  Mammalian Species 478:1-9.
330.    Best, T. L., and N. J.  Granai. 1994. Tamias merriami.  Mammalian Species 476:1-9.
331.    Best, T. L., and N. J.  Granai. 1994. Tamias obscurus.  Mammalian Species 472:1-6.
332.    Best, T. L., and M. P.  Skupski. 1994. Perognathus flavus.  Mammalian Species 471:1-10.
333.    Best, T. L., and M. P.  Skupski. 1994. Perognathus merriami.  Mammalian Species 473:1-7.
334.    Best, T. L. 1994. Perognathus alticolus. Mammalian Species 463:1-4. 
335.    Best, T. L., S. L. Burt,  and J. L. Bartig. 1994. Tamias  quadrivittatus. Mammalian Species 466:1-7. 
336.    Best, T. L. 1995. Sciurus alleni. Mammalian Species 501:1-4.
337.    Best, T. L. 1995. Sciurus deppei. Mammalian Species 505:1-5.
338.    Best, T. L. 1995. Sciurus variegatoides. Mammalian Species 500:1-6.
339.    Best, T. L. 1995. Spermophilus adocetus. Mammalian Species 504:1-4.
340.    Best, T. L. 1995. Spermophilus mohavensis. Mammalian  Species 509:1-7.
341.    Best, T. L., and G.  Ceballos. 1995. Spermophilus perotensis.  Mammalian Species 507:1-3.
342.    Best, T. L., and S. Riedel.  1995. Sciurus arizonensis. Mammalian  Species 496:1-5.
343.    Best, T. L. 1995. Sciurus colliaei. Mammalian Species 497:1-4. 
344.    Best, T. L. 1995. Sciurus nayaritensis. Mammalian Species 492:1-5. 
345.    Best, T. L. 1995. Sciurus oculatus. Mammalian Species 498:1-3. 
346.    Best, T. L. 1995. Spermophilus annulatus. Mammalian  Species 508:1-4. 
347.    Best, T. L., H. A. Ruiz  Puna, and L. S. Leon Paniagua. 1995. Sciurus  yucatanensis. Mammalian Species 506:1-4. 
348.    Best, T. L. 1996. Lepus californicus. Pages 1-10 in M. American Society of, editor.  Mammalian Species.
349.    Best, P. B. 1996. Evidence  of migration by Bryde's whales from the offshore population in the southeast Atlantic. Report of the International Whaling  Commission:315-322.
350.    Best, P. B., A. Brandao,  and D. S. Butterworth. 2001. Demographic parameters of southern right whales  off South Africa.  Journal of Cetacean Research and Management Special Issue 2:161-169.
351.    Best, P. B. 2001. Distribution  and population separation of Bryde's whale Balaenoptera edeni off southern Africa. Marine Ecology Progress Series 220:277-289.
352.    Best, P. B., V. M.  Peddemors, V. G. Cockcroft, and N. Rice. 2001. Mortalities of right whales and  related anthropogenic factors in South African waters, 1963-1998. Journal of  Cetacean Research and Management Special Issue Specia.
353.    Best, P. B., J. L.  Bannister, R. L. J. Brownell, and G. P. Donovan. 2001. Report of the Workshop  on Status and Trends of Western North Atlantic  Right Whales. The Journal of Cetacean Research and Management 2:61-87. 
354.    Bezerra, A. M. R., F. H. G.  Rodrigues, and A. P. Carmignotto. 2001. Predation of rodents by the yellow  armadillo (Euphractus sexcinctus) in  Cerrado of the Central Brazil. Mammalia 65:86-88. 
355.    Bhat, H. R. 1968. Dobson's  Long-tingued fruit bat, Eonycteris  spelaea (DOBSON) from Kumaon Hills, U.P. An addition to the Chiropteran  fauna of India.  Journal of the Bombay  Natural History Society 64:550-551.
356.    Bhat, H. R., M. A. Sreenivasan,  and P. G. Jacob. 1980. Breeding cycle of Eonycteris  spelaea (DOBSON 1871) (Chiroptera, Pteropidae, Macroglossinae) in India.  Mammalia 44:343-347. 
357.    Bhat, S. K., and D. N.  Mathew. 1984. Observations on the Breeding Biology of the Western   Ghats Squirrel, Funambulus-Tristriatus Waterhouse. Mammalia 48:573-584.
358.    Bhatnagar, K. P. 1978.  Breech presentation in the Hairy-legged vampire Diphylla ecaudata. Journal of Mammalogy 59:864-866. 
359.    Bhatnagar, K. P. 1978. Head  presentation in Artibeus jamaicensis with some notes on parturition. Mammalia 42:359-363. 
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G. History of data set usage
December 2003. Patrik  Lindenfors and Charlie Nunn requested the use of the carnivore trait data for  the project ‘Host  correlates of parasite characteristics in carnivores’ (analysis  outputs: Lindenfors et al. 2007b).
  July 2004. Sonia Altzier  requested use of hoofed mammal trait data for a project ‘Host correlates of the  diversity,  characteristics and prevalence of parasites in hoofed mammals’ (analysis  outputs: Ezenwa et al. 2006, Pedersen et al. 2007).
August 2004. Chris Carbone requested the use of the carnivore  population density and diet trait data for the project ‘Patterns in population  density and body mass in meat-eating carnivores’.
Jan 2005. Jonathan Jeschke and Dave Strayer requested the use of  European and North American mammal trait data for the project ‘Species introductions’  (analysis outputs: Jeschke and Strayer 2006).
February 2006. Shai Meiri requested the use of the mammal trait data  for the project ‘Body sizes of insular and mainland mammals’ (analysis outputs: Meiri et al. 2008).
May  2005. Patrik Lindenfors requested the use of the mammal trait data for the  project ‘Sexual Size Dimorphism in Mammals’ (analysis  outputs: Lindenfors et al. 2007a).
May  2005 - John Welch and Lindell Bromham requested the use of mammal trait data  for the project ‘Gene and lineage  effects in rates of molecular evolution’ {analysis  outputs: \Welch, 2008 #163}.
March  2006 – Simon Goodman requested the use mammal trait data for the project  ‘Comparative analysis of immunophysiological parameters with ecological and  life history correlates of disease exposure in ungulates’.
January  2007 - Tim Blackburn and Craig White  requested use of the mammalian trait data for the project ‘Population energy  use’.
January  2007 - Vincent Savolainen and Thomas Galewski requested the use of the mammalian  trait data for the project ‘Sexual selection and horn lengths in bovids’.
H. Data set update history
    Review history: n/a
    Questions and comments from secondary users: n/a
ACKNOWLEDGMENTS
This work was financially supported  through NSF (Grant No. DEB/0129009); NERC (Grant No. NER/A/S/2001/00581); Earth  Institute Fellowship, Columbia   University and University  of Virginia (KEJ); and the European Commission (FP6 Early-Stage Training  Network HOTSPOTS) (SAF). We also thank Louise Endler, Sophie   Gauss and Paul Racey for assistance and two anonymous  referees for comments on the manuscript.
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