The evolution of wolves presence in Tuscany and the occurrence of wolf/dog hybrids

Marco Apollonio and Massimo Scandura CIRSeMAF e Dipartimento di Scienze della Natura e delle sue Risorse Università di Sassari TUSCANY

Area 22987 km2

Human density 163 in/km2

Over 90% hilly or mountain

Over 50% forested TUSCANY IS A KEY AREA FOR WOLF DISTRIBUTION IN ITALY

Fabbri et al. 2007 Molecular Ecology Zimen e Boitani 1975 Cagnolaro et al . 1974 Francisci e Guberti 1993 Wolf presence in municipalities 1996

Boitani e Ciucci 1996 Wolf presence in municipalities 2005

2009

2013

NonPresence segnalata not reported PresenzaOccasional sporadica presence PresenzaStable presence stabile WOLF DISTRIBUTION IN TUSCANY 2013

NonPresence segnalata not reported PresenzaOccasional sporadica presence PresenzaStable presence stabile WOLVES NUMBER

25 wolves Ciani 1989 ?

30-50 wolves Boitani 1989 WOLF MONITORING IN TUSCANY – PROMOTED BY THE TUSCAN REGIONAL GOVERNMENT FROM 2013

RESPONSIBLES OF MONITORING FOR PROVINCES IN 2013

AREZZO LUCA MATTIOLI E MARCO APOLLONIO

FIRENZE, PRATO, PISTOIA DUCCIO BERZI

LUCCA SIRIANIO LUCCARINI

MASSA PAOLO BONGI

PISA, LIVORNO VIVIANA VIVIANI (Polizia Prov Pisa e Livorno)

GROSSETO ANDREA SFORZI, LUCA MANGHI * (EXTERNAL CONTRIBUTION)

SIENA FEDERICO MORIMANDO

Wolf packs in Florence province 2013

Cod Località Province 21 Fiorenzuola 1 Firenze 22 Fiorenzuola 2 Firenze 23 Fiorenzuola 3 Firenze 24 Incisa Firenze 25 Marradi Firenze 26 Palazzolo sul Senio Firenze 27 Pontassieve Firenze 28 Reggello Firenze 29 Vicchio Firenze 30 Londa Firenze 31 Calenzano Firenze-Prato 31 Gambassi Firenze-Siena Wolf Packs in Tuscany 2013

71 packs

About 300-320 wolves

Underestimate UNGULATES IN TUSCANY

Species Roe deer Red deer Wild boar

Status 2011 163336 4181 149965

Absolut increase 2000- 71464 1581 71078 2011

Percent increase 2000- 44% 38% 47% 2011

Fallow Species Mouflon deer

Status 2011 8425 2270

Absolut change 2000- -39 -664 2011 ABOUT 330.000 UNGULATES IN TUSCANY Percent variation 0% -29% ROE DEER AND WILD BOAR REPRESENTS 95% 2000-2011 UNGULATES DENSITY IN EUROPEAN COUNTRIES

Austria Germany Tuscany Slovenia Switzerland Hungary Czech Republic Denmark United Kingdom Spain France Italy Slowakia Latvia Poland Belgium Sweden Lituania Serbia Estonia Netherlands Croatia Romania Norway Ireland Finland Portugal Greece 0 2 4 6 8 10 12 14 16 18 20 Density (heads/100ha) LONG TERM STUDY ON WOLF DISTRIBUTION AND BIOLOGY PROMOTED BY THE PROVINCIAL GOVERMENT OF AREZZO AND THE REGIONAL GOVERNMENT OF TUSCANY WOLF PACKS INCREASE: THE CASE OF AREZZO PROVINCE

Study area: mountain area of the central Apennines, province of Arezzo

Population monitored since 1993 Small packs (2-6 individuals; stable territories 100-150 km2)

4 PACKS 9 PACKS from Gazzola et al., 2007

Habitat variables in the municipalities of wolf-pack area (WPA) and no- wolf-pack area (no-WPA) in Arezzo Province, Italy, in 1998-2001 WPA

NO -WPA Study area: mountain area of the central Apennines, province of Arezzo

Population monitored since 1993 Small packs (2-6 individuals; stable territories 100-150 km2)

12 PACKS 21 PACKS Nowadays

21 PACKS WOLVES KILL RECORDS IN THE BERGAMO PROVINCE IN XVII CENTURY

Central Alps

Po valley ORIANI ET AL. 2014 WOLVES KILLS RECORDS IN THE BERGAMO PROVINCE IN XVIII CENTURY

Central Alps

Po valley

ORIANI ET AL. 2014 WOLVES KILLS RECORDS IN THE BERGAMO PROVINCE IN XIX CENTURY

Central Alps

Po valley

ORIANI ET AL. 2014 URBAN WOLVES URBANURBAN WOLVES WOLVES

Badia Agnano URBANURBAN WOLVES WOLVES

Anghiari URBAN WOLVES

Arezzo HYBRID WOLVES Paolo Savi – 1798-1871 Zoologist – Pisa University

Two wolves – one with white patches on flanks - attack a guarding dog and a lamb Courtesy of the Natural History Museum of Calci Composition by Paolo Savi XIX century

Hybrid carcasses distribution in Tuscany from 1978 to 2014

1978

Hybrid from 1978 to 2004 Hybrid from 2005 to 2014 Summary of data referred to the Tuscan population from several sources: CIRSeMAF -University of Sassari – Prov. Arezzo Braschi & Boitani 2013 (Life Ibriwolf) Greco 2009 (PhD thesis) Caniglia et al. 2013 EJWR

49 out of 49 (100%) putative hybrids had a wolf (W14) mtDNA haplotype Asymmetrical 20 Out of 25 (80%) male putative hybrids hybridization in Tuscany had a wolf Y chromosome haplotype

15 Out of 21 (71%) putative hybrids with dark /black coat had the KB allele (melanistic deletion) at the b-defensin gene Hybrid packs in Tuscany 2014

25 “hybrid packs”

Type 5 condition sensu Allendorf

Photo-trapping N= 11

Photo-trapping + genetics N = 7

Genetics N= 2

Direct observation N = 4

Direct observation + genetics N= 1 LONG TERM STUDY ON WOLF DISTRIBUTION AND BIOLOGY PROMOTED BY THE PROVINCIAL GOVERMENT OF AREZZO AND THE REGIONAL GOVERNMENT OF TUSCANY Spatial distribution of samples 1998-2014 – Province of Arezzo

NON-INVASIVE CARCASSES

480 samples 211 different individuals 185 wolves Distribution of introgressed individuals in the province of Arezzo

Darker dots correspond to more introgressed individuals Y chromosome haplotypes in wolves and hybrids

(samples)

(genotypes)

HAPLOTYPE FREQUENCIES name in Randi et WOLVES + HAPLOTYPE al 2014 HYBRIDS (n=79) DOGS (n=37) H1 YH17 0.570 0.027 H2 YH26 0.316 0.081 H3 YH05 0.101 0.865 H4 YH01 0.013 H5 - 0.027 Mitochondrial DNA haplotypes (samples, wolves+hybrids)

(genotypes)

HAPLOTYPE FREQUENCIES WOLVES + HAPLOTYPE HYBRIDS (n=80) DOGS (n=22) w14 0.975 w16 0.013 new 0.013 dog haps (13) 1.000 Province Municipality Date Sex Genotype Putative mtDNA STR -Y Phenotype status Arezzo Anghiari 24/10/2006 F AP03 Hybrid w14 - Mantello scuro, assenza di mascherina facciale Arezzo Arezzo 26/11/2009 M AP07 Hybrid w14 H2 (wolf) Colorazione tipica, ma assenza di barre nere nelle zampe anteriori. Arezzo Bibbiena 08/01/2011 M AP09 Wolf w14 H2 (wolf) Mantello anomalo, estesa mascherina facciale, unghie depigmentate Arezzo Sansepolcro 05/06/2012 F LU26 Wolf w14 - Sperone Arezzo Anghiari 17/08/2012 M AP24 Hybrid w14 H2 (wolf) Macchia bianca sullo sterno

Arezzo Anghiari 17/08/2012 F AP25 Hybrid w14 - Mantello scuro Arezzo Anghiari 07/09/2012 F AP26 Wolf w14 - Mantello anomalo, assenza di mascherina facciale

Arezzo Arezzo 22/10/2012 F AP22 Wolf w14 - Bandeggi anomali fianco e zampe; maschera grigia, unghie depigmentate

AP03 AP26 Spread of introgressed genes within the province of Arezzo

Genetic monitoring Alpe di 1 7 microsatellites 0,9 Catenaia 0,8 259 samples 0,7 0,6

wolf 0,5 1998-2005 Q 0,4 0,3 0,2 0,1 0 Individual 1 0,9 0,8 0,7 0,6

2006-2013 wolf 0,5 Q 0,4 0,3 0,2 0,1 0 Individual

Alpe di Poti 1 0,9 0,8 0,7 0,6

Since 2005 wolf 0,5 Q 0,4 0,3 0,2 0,1 0 Individual WOLVES AND HYBRIDS COMPARISON Preliminary insights •Ecology •Behaviour •Physiology

♂

Pack A

♀ ♂

PACK B

♀ WOLVES AND HYBRIDS : DIET Diet analyses LEVIN INDEX Method Alpe di Poti Alpe di Catenaia AV% 2,22 2,14 Waver biomass 2,05 2,04 Ciucci biomass 2,06 2,04

PIANKA INDEX Method O AV% 0,953 Waver biomass 0,955 Ciucci biomass 0,955 WOLVES AND HYBRIDS : ACOUSTIC COMUNICATION

Wolf howl versus hybrid howl

Italian wolf howls Hybrid wolf howls ( 7 packs, 135 howls) (2 packs, 20 howls)

Fundamental frequency (F0) Fundamental frequency (F0)  190-1350 Hz  290-990 Hz Duration Duration  0.75-9.55 sec  0.70-5.85 sec

Preliminary analyses show no differences in the acoustic structure between wolf and hybrid howls From Passilongo et al. 2010 WOLVES AND HYBRIDS : REPRODUCTIVE BIOLOGY ? REPRODUCTION Wild x Domestic Hybridization Evolutionary implications of introgressive hybridization  on wild populations  on domestic stocks

Genetic changes

Modifications in: o Anatomy (body mass, body growth, fatness) o Physiology (reproductive cycle, fertility, hormone production) o Morphology (coat colour) o Behaviour (aggressiveness, shyness, inclination towards humans)

WOLF/DOGS HYBRIDS REPRODUCTION

MAY BE •Earlier breeding •Larger litter size THE MONITORING ON WOLF DISTRIBUTION AND STATUS IN TUSCANY WAS FOUNDED BY THE REGIONAL GOVERNEMENT OF TUSCANY

THE LONG TERM STUDY ON WOLF IN THE PROVINCE OF AREZZO WAS FOUNDED BY THE PROVINCIAL GOVERNMENT OF AREZZO AND BY THE REGIONAL GOVERNMENT OF TUSCANY

Luca Mattioli, Antonello Canu, Daniela Passilongo, Elena Bassi,. Ilaria Firmio and the group of monitors greatly contributed to this presentation