(Crax Fasciolata) Life History Patterns in the Northern

UNIVERZA NA PRIMORSKEM 2020 FAKULTETA ZA MATEMATIKO, NARAVOSLOVJE IN

INFORMACIJSKE TEHNOLOGIJE

MASTER’S THESIS

(MAGISTRSKO DELO)

CAMERA TRAP BASED DATA ANALYSIS OF THE BARE-FACED CURASSOW (CRAX FASCIOLATA) LIFE HISTORY PATTERNS IN THE NORTHERN PANTANAL, BRAZIL R'S THESIS (ANALIZA VZORCEV ŽIVLJENJSKE ZGODOVINE MASTE GOLOLIČNE HOKOJKE (CRAX FASCIOLATA) NA OSNOVI PODATKOV PRIDOBLJENIH S FOTOPASTMI

NA OBMOČJU SEVERNEGA PANTANALA,

BRAZILIJA)

MARTIN SENIČ

MARTIN SENIČ UNIVERZA NA PRIMORSKEM FAKULTETA ZA MATEMATIKO, NARAVOSLOVJE IN INFORMACIJSKE TEHNOLOGIJE

Master's thesis (Magistrsko delo)

Camera trap based data analysis of the Bare-faced Curassow (Crax fasciolata) life history patterns in the northern Pantanal, Brazil

(Analiza vzorcev življenjske zgodovine gololične hokojke (Crax fasciolata) na osnovi podatkov pridobljenih s fotopastmi na območju severnega Pantanala, Brazilija)

Ime in priimek: Martin Senič Študijski program: Varstvo narave, 2. stopnja Mentor: prof. dr. Karl-Ludwig Schuchmann Somentor: doc. dr. Andrej Sovinc Delovna somentorica: mag. Kathrin Burs

Koper, maj 2020 Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 II

Ključna dokumentacijska informacija

Ime in PRIIMEK: Martin SENIČ

Naslov magistrskega dela: Analiza vzorcev življenjske zgodovine gololične hokojke (Crax fasciolata) na osnovi podatkov pridobljenih s fotopastmi na območju severnega Pantanala, Brazilija

Kraj: Koper Leto: 2020 Število listov: 94 Število slik: 22 Število tabel: 8 Število prilog: 15 Število strani prilog: 15 Število referenc: 74 Mentor: prof. dr. Karl-Ludwig Schuchmann Somentor: doc. dr. Andrej Sovinc Delovni somentorica: mag. Kathrin Burs UDK: 598.261.4(043.2) Ključne besede: Cracidae, Crax fasciolata, gololična hokojka, fotopasti, uspešnost zajema, vzorci dnevnih aktivnosti, socialna organizacija, starševska skrb, razmerje spolov, starost potomcev, paritvena sezona

Izvleček: Preučevali smo vzorce življenjske zgodovine gololične hokojke (Crax fasciolata, Aves, Cracidae) na območju parka SESC Baía das Pedras (pribl. 4200 ha), Poconé, severni del območja matogrossenskega Pantanala, Brazilija. Med julijem 2015 in decembrom 2017 smo člani raziskovalne enote CO.BRA (http://cobra.ic.ufmt.br/), s fotopastmi spremljali 37 lokacij. S fotopastmi smo zajeli 357 neodvisnih zabeležb vrste (554 ptic) na 26 lokacijah. Opazili smo vzorce aktivnosti, ki nakazujejo, da je bila uspešnost zajema posnetkov najvišja v sezoni umikajoče se vode. Dnevna aktivnost vrste je sledila bimodalnemu vzorcu. Ritmi dnevne aktivnosti so bili podobni med različnimi sezonami, spoloma in odraslimi s potomci ali brez njih. Število posnetih živali glede na neodvisen zajem vrste se je gibalo od 1 do 4 in v povprečju znašalo 1,55 ± 0,81 SE. Večjih enospolnih skupin odraslih ptic nismo zaznanih. Potomce smo vsaj enkrat zabeležili na osmih lokacijah v marcu, aprilu in maju 2016 ter v juniju, juliju, oktobru in novembru 2017. Večinoma smo jih zabeležili z obema staršema, kar nakazuje starševsko skrb s strani obeh staršev. Razmerje spolov odraslih osebkov smo z 1,05:1,00 obravnavali kot enakovredno, medtem ko je razmerje spolov pri potomcih z 0,51:1,00 izrazito v prid samicam. Izdelači in uporabili smo „Identifikacijski ključ starostnih razredov potomcev C. fasciolata“. Določitev starostnih razredov je omogočila zaključek, da se odrasle ptice parijo skozi vse leto. Rezultati študije prinašajo pomembna spoznanja o biologiji vrste, saj smo jih pridobili na območju z nizkimi antropogenimi vplivi. Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 III

Key words documentation

Name and SURNAME: Martin SENIČ

Title of the thesis: Camera trap based data analysis of the Bare-faced Curassow (Crax fasciolata) life history patterns in the northern Pantanal, Brazil

Place: Koper Year: 2020 Number of pages: 94 Number of figures: 22 Number of tables: 8 Number of appendices: 15 Number of appendices pages: 15 Number of references: 74 Mentor: Prof. Karl-Ludwig Schuchmann, PhD Co-Mentor: Assist. Prof. Andrej Sovinc, PhD Working Co-Mentor: Kathrin Burs, MSc UDC: 598.261.4(043.2) Keywords: Cracidae, Crax fasciolata, Bare-faced Curassow, camera trap, capture success, activity patterns, sex ratio, social organization, parental care, offspring age, breeding season

Abstract: Life history patterns of a population of the Bare-faced Curassow (Crax fasciolata, Aves, Cracidae) were studied at the SESC Baía das Pedras Park (c. 4200 ha), Poconé, Northern Pantanal of Mato Grosso, Brazil. Between July 2015 and December 2017, 37 locations were monitored by the Computational Bioacoustics Research Unit (CO.BRA; http://cobra.ic.ufmt.br/), using camera traps. The Bare-faced Curassow was detected on 357 independent occasions (554 independent individual captures) at 26 locations. Observed activity patterns suggest that the highest capture success rates occured during the receding water season. The species’ daily activity followed a bimodal pattern. Daily activity rhythms were similar between different seasons, gender, and adults with or without offspring. The number of captured individuals per single survey occasion ranged from 1 and 4, and averaged 1.55 ± 0.81 SE. Larger single sex aggregations of adult birds were not observed. Offspring were detected at least once at 8 locations, namely in March, April, and May 2016 and in June, July, October, and November 2017. They were mostly accompanied with both parents, suggesting parental care by both male and female. Offspring sex ratio was significantly female-skewed with 0.51:1.00, whereas adult sex ratio of 1.05:1.00 was considered equal. An “Age-class Identification Key for C. fasciolata Offspring” was elaborated and applied. Age assessment of offspring captures suggests that breeding occurs throughout the year. The study results are biologically meaningful since they were obtained in an area with low anthropogenic pressures. Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 IV

LIST OF CONTENTS

1 INTRODUCTION ...... 1 1.1 CRACIDS (CRACIDAE) ...... 1 1.1.1 General morphological characteristics ...... 1 1.1.2 General behavioral characteristics ...... 1 1.1.3 Geographical distribution of cracids ...... 2 1.1.4 Conservation status, main threats, and conservation objectives ...... 3 1.2 BARE-FACED CURASSOW (CRAX FASCIOLATA SPIX, 1825) ...... 4 1.2.1 Scientific classification ...... 4 1.2.2 Description of morphological characteristics ...... 5 1.2.3 Geographical range of distribution ...... 5 1.2.4 Habitat preferences ...... 7 1.2.5 Conservation status and main threats ...... 7 1.2.6 Historical and contemporary distribution of Bare-faced Curassow in the Brazilian Pantanal ...... 7 1.2.7 Existing measures and conservation actions ...... 8 1.2.8 Feeding habits ...... 8 1.2.9 Activity patterns and social organization ...... 9 1.2.10 Breeding habits ...... 9 1.3 PANTANAL ...... 11 1.3.1 Pantanal as an important bird sanctuary ...... 12 1.4 CAMERA TRAP BASED SURVEY OF CRAX SPECIES ...... 12 1.5 OBJECTIVES AND APPROACHES ...... 13 2 MATERIALS AND METHODS ...... 14 2.1 GENERAL DESCRIPTION OF A STUDY AREA ...... 14 2.2 PROJECT HISTORY AND CAMERA TRAPPING ...... 16 2.3 ACTIVITY PATTERNS, SOCIAL ORGANIZATION AND SEX RATIO ...... 16 2.3.1 Data Analysis ...... 16 2.3.2 Data comparison...... 17 2.4 OFFSPRING AGE IDENTIFICATION AND ASSESSMENT OF BREEDING SEASON ...... 18 2.4.1 Development and usefulness of »Age-class identification key for Crax fasciolata offspring« ...... 18 3 RESULTS ...... 20 3.1 ACTIVITY PATTERNS ...... 20 3.1.1 Daily activity patterns ...... 24 3.2 SOCIAL ORGANIZATION ...... 27 3.3 SEX RATIO ...... 30 Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 V 3.4 AGE-CLASS IDENTIFICATION ...... 31 3.4.1 Usage of Age-class identification key for Crax fasciolata offsprings ...... 31 3.4.2 Age-class identification key for Crax fasciolata offspring ...... 33 3.4.3 Results of practical use of Age-class identification key for C. fasciolata offsprings ...... 38 3.5 BREEDING SEASON ...... 40 3.5.1 Evaluation of breeding season phases considering estimated age-classes ..... 40 4 DISCUSSION ...... 45 4.1 CAMERA TRAPPING ...... 45 4.2 ACTIVITY PATTERNS ...... 45 4.2.1 Daily activity patterns ...... 46 4.3 SOCIAL ORGANIZATION ...... 47 4.3.1 Parental care ...... 48 4.4 SEX RATIO ...... 49 4.5 AGE-CLASS IDENTIFICATION KEY FOR C. FASCIOLATA OFFSPRING AND BREEDING SEASON EVALUATION ...... 51 5 CONCLUSIONS ...... 54 6 POVZETEK V SLOVENSKEM JEZIKU ...... 56 7 REFERENCES ...... 62

LIST OF TABLES

Table 1. Conservation status of different groups of Cracidae species (curassows, guans, and chachalacas) summarized after The IUCN Red List of Threatened Species ...... 3 Table 2. Number of independent camera trap captures / 30 min, capture success of the species, capture success of individuals, CSind / CSsp ratio of Bare-faced Curassows for different Years, Periods and Seasons...... 21 Table 3. Cumulative numbers of Bare-faced Curassows groups of at least two adult individuals of same sex or more than two adults of both sexes were recorded (from July 2015 to December 2017) ...... 28 Table 4. Available sex ratio data for different Crax species regarding summarized and adjusted data available from different surveys ...... 31 Table 5. List of estimated age-classes for all independent captures of Crax fasciolata offspring (from July 2015 to December 2017)...... 38 Table 6. Presentation of estimated egg laying and hatching range intervals for Crax fasciolata on monthly basis relative to the period when the captures were taken for different survey occasions where offsprings were detected during the survey at Baía das Pedras in the Pantanal of Mato Grosso, Brazil in year 2016 ...... 41 Table 7. Presentation of estimated egg laying and hatching range intervals for Crax fasciolata on monthly basis relative to the period when the captures were taken for different survey occasions where offsprings were detected during the survey at Baía das Pedras in the Pantanal of Mato Grosso, Brazil in year 2017 ...... 43 Table 8. Final presentation of overall egg laying and hatching range intervals estimation for Crax fasciolata on a monthly basis ...... 44

LIST OF FIGURES

Figure 1. Combined distribution range map for all species of Cracidae – guans, chachalacas, and curassows ...... 2 Figure 2. Adult female and male of Bare-faced Curassow (Crax fasciolata) ...... 5 Figure 3. Geographical range of distribution of Bare-faced Curassow (Crax fasciolata) .....6 Figure 4. Geographical position of Pantanal in relation to different countries and states ... 11 Figure 5. Geographical position of SESC Baía dasa Pedras Park (study area) in relation to different countries and states ...... 14 Figure 6. Orthophoto of SESC Baía dasa Pedras Park (study area) and related sample locations (numbers; Appendix A) ...... 15 Figure 7. Comparison of capture success of the species of Bare-faced Curassows between dry and wet period for 2016 and 2017 ...... 22 Figure 8. Comparison of capture success of individuals of Bare-faced Curassows between dry and wet period for 2016 and 2017 ...... 23 Figure 9. Number of independent species captures of Bare-faced Curassows (from July 2015 to December 2017) based on 1 h periods ...... 24 Figure 10. Number of captures of adult individuals of Bare-faced Curassows (from July 2015 to December 2017) based on 1 h periods ...... 24 Figure 11. Periodically based proportions (%) of independent captures of adult individuals of Bare-faced Curassows (from July 2015 to December 2017) based on 1 h periods ...... 25 Figure 12. Sexually based proportions (%) of independent capture of adult individuals of Bare-faced Curassows (from July 2015 to December 2017) based on 1 h periods ...... 25 Figure 13. Periodically based proportions (%) of independent species captures of Bare-faced Curassows (from July 2015 to December 2017) based on 1 h periods ...... 26 Figure 14. Proportions (%) of independent species captures of Bare-faced Curassows with and without offspring obtained during the survey (from July 2015 to December 2017) based on 2 h periods ...... 26 Figure 15. Proportions (%) of independent species captures divided on single male, single female, adult pair (male-female), adult with offspring and other forms of grouping of Bare- faced Curassow (from July 2015 to December 2017) for different years of study ...... 27 Figure 16. Proportions (%) of independent species captures of Bare-faced Curassow for overall study, different periods (dry, wet) and seasons (receding water, low water, rising water, high water) (from July 2015 to December 2017) divided on single female, single male, adult pair (male-female), adult with offspring and other forms of grouping ...... 28 Figure 17. Overall ratio of independent individual records of adults and offspring of the Bare-faced Curassow presented for different periods and seasons (based on cumulative data gathered from July 2015 to December 2017) ...... 29 Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 VIII Figure 18. Overall distribution of offspring grouping records of Bare-faced Curassow (based on cumulative data gathered from July 2015 to December 2017) ...... 30 Figure 19. Overall independent individual captures of different sexes of Bare-faced Curassow (from July 2015 to December 2017) presented for adults and offspring ...... 30 Figure 20. Offspring age-class identification key for Crax fasciolata specification spreadsheet...... 32 Figure 21. Plates and photographs of different age stages of different curassow species. .. 37 Figure 22. Female offspring, adult female and male offspring of Bare-faced Curassow (Crax fasciolata). Estimated age-class of the offspring is between 3 and 4 months of age...... 37 Figure 23. Sampling effort (y axis) expressed in days per each month (x axis) for years 2015, 2016 and 2017 and offspring presence (shapes on the x axis) in 2016 and 2017 ...... 40

LIST OF APPENDICES

APPENDIX A Geographic coordinates of sample locations (= sites)

APPENDIX B Sampling effort per sample location (= site) and per month

APPENDIX C Specification sheet for G2_17_2_1 placement

APPENDIX D Specification sheet for G6_16_1_1A placement

APPENDIX E Specification sheet for G6_16_1_2A placement

APPENDIX F Specification sheet for G6_16_1_3B placement

APPENDIX G Specification sheet for G7_16_1_1 placement

APPENDIX H Specification sheet for G11_16_1_1 placement

APPENDIX I Specification sheet for G23_17_1_3 placement

APPENDIX J Specification sheet for G29_17_2_3 placement

APPENDIX K Specification sheet for G29_17_2_4 placement

APPENDIX L Specification sheet for G33_17_2_1 placement

APPENDIX M Specification sheet for G36_16_1_1 placement

APPENDIX N Specification sheet for G36_17_1_2 placement

APPENDIX O Specification sheet for G36_17_1_3 placement

LIST OF ABBREVIATIONS

APP – Permanent Preservation Areas

ASR – Adult Sex Ratio

BAPP – Advanced Research Base of the Pantanal

CS – Capture Success

CO.BRA – Computational Bioacoustics Research Unit

HBW – Handbook of the Birds of the World

IBA – Important Bird and Biodiversity Areas

ICBP – International Council for Bird Preservation

INAU – Brazilian National Institute for Science and Technology in Wetlands

IUCN – International Union for Conservation of Nature

UFMT – Federal University of Mato Grosso

RAI – Relative Abundance Index

SESC – Social Service of Commerce

ACKNOWLEDGEMENTS

I would like to thank Prof. Dr. Karl-L. Schuchmann and Prof. Dr. Marinez Isaac Marques for having included me in the Sounds of the Pantanal – The Pantanal Automated Acoustic Biodiversity Monitoring Program (INAU / CO.BRA / UFMT) and for giving me the opportunity to become a part of their story. Thank you for enabling me to work on your data, which was essential for this thesis and thank you for enabling me to stay, study and train with you during the international internship and beyond.

A special thank goes to all of my supervisors, Prof. Dr. Schuchmann, Assist. Prof. Dr. Sovinc, and MSc. Kathrin Burs for all the guidance and advice. And especially for all the patience. Your effort to point me in the right direction when needed eventually paid off. Especially for me, as I profited greatly from the work we did together with many new lessons learned and I am glad to carry those into my future career. I hope that our paths of cooperation will cross again for many times in the future.

In addition, special thanks to Kathrin Burs for all the field experiences, guidance, conversations. Thank you for giving me the proper insight about field work in a harsh and dynamic environment. It was truly one of the greatest experiences of my life. And thank you for pushing me to finish my work too many times.

I would like to thank all of my colleagues who strongly supported my (not always the greatest) efforts to finish. You motivated me with kind reminders, work flexibility and I am especially grateful for all the patience. I hope I will be able to repay you for this when and if your motivation hits the hard ground.

Thank you to my family and friends for moral and other support during all my struggles. There are too many people to name. Many of you were the reason that I managed until the end. And I would especially like to thank my brother, who took care of all the things at home alone while I was gone for a couple of months. I am aware of your sacrifice and I am proud that you are my brother. I wish you all the best on your path which is already in the making.

I would like to dedicate this work to a special place. A place which grown very close to me. To the Pantanal. I hope you will flourish in all of your greatness for centuries to come, despite all the threats you are facing. Hopefully, this work will additionally contribute to your recognition as a refuge for species such as the Bare-faced Curassow. Wish you all the best and see you soon. Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 1

1 INTRODUCTION 1.1 CRACIDS (CRACIDAE)

Cracids are considered the most ancient family within the Order Galliformes comprising three subfamilies: curassows, guans, and chachalacas. Their distribution ranges throughout Neotropics and the extreme southern Nearctic Region. They are considered to be one of the most vulnerable taxa of birds worldwide. This is especially true of the curassows, due to a large knowledge gap in their biology to provide adequate measures for their conservation (del Hoyo 2019, BirdLife International 2019b, IUCN 2019a).

1.1.1 General morphological characteristics

In general, cracids (Cracidae) are medium to large-sized gallinaceous birds. They measure from 42 to 92 cm in lenght. The appearance of cracids is much like that of other gallinaceous birds indicated by a heavy body, a long neck, and a small head, often with a crest. In general, the beak is similar to that of a domestic chicken and normally medium-sized. The legs are long and strong, an adaptation to their ground-dwelling habits. Cracids have an elongated hind toe, ideal for perching in trees, their prefered nocturnal shelter. All taxa of this subfamily have characteristic, strongly rounded long tails, and concave wings with well- developed pectoral musculature. Sustained flights are rare as their flight capacity is moderate. Currasows are the poorest fliers among the Cracidae, besides they vary in consistency of plumage and a coloration which is usually discreet, although often attractive. There are various degrees of sexual dimorphism in the plumage of some curassows, whereas sexes in guans and chachalacas are alike. In general, the offspring’s plumage is very similar to that of adults, even when still noticeably smaller (del Hoyo 2019).

1.1.2 General behavioral characteristics

Not much known about cracid behavior and life histories, as most of the species live in tropical forests and wetlands, areas difficult to access. The subfamily of chachalacas are generally the best known group as they inhabit more open areas, frequently near settlements, and thus are much easier to observe (del Hoyo 2019).

Cracids tend to be fairly social. They can be found alone, but are more commonly observed in pairs or in small family groups. Large flocks are normally gathered where food is abundant, typically in fruiting trees that also attract other bird species. After feeding, cracids leave the tree and the flock splits up again into smaller ground dwelling units. They can occur in similar congregations at lakes or rivers when consuming water. The most typical Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 2 social units in cracids are pairs or females accompanied by a few offspring. There are also records with single sex flocks, observed outside the breeding season. Cracids are mostly diurnal, meaning that their daily activity is particularly pronounced with two peaks of their appearance. They are highly vocal, with various vocal characteristics among different individual groups. In general, cracids are vegetarian, but occasionally prey on insects, spiders, centipedes, molluscs, small rodents, and amphibians. Confirmed data on the breeding season of cracids is scarce and varies within their large distribution ranges. Some species, e.g., Razor-billed Curassow (Mitu tuberosum) and Wattled Curassow (Crax globulosa), most likely to be opportunistic breeders, i.e., reproduction is rather triggered by the availability of local food sources than by seasonality. They almost always build nests in trees, shrubs or vines. The nest can be built up to 20 m above the forest floor, but usually lower. Female and male both participate in nest building. Eggs are laid on alternate days. Incubation is carried out by female alone and it lasts c. 24 days in chachalacas, c. 28 in guans, and c. 29-32 days in curassows. The females leave their nest a few times per day in order to feed. There is virtually nothing known about breeding success of the cracids. In general, all taxa are non-migratory, but some local dispersal movements occur. Most regular movements within different elevations occur in mountain-dwelling species, e.g., Sira Curassow (Pauxi koepckeae) or Crested Guan (Penelope purpurascens). Movements can also be related to seasonal changes in temperature, precipitation, and wetlands water levels (del Hoyo 2019, del Hoyo and Motis 2004, Delacour and Amadon 2004).

1.1.3 Geographical distribution of cracids

Figure 1. Combined distribution range map for all species of Cracidae – guans, chachalacas, and curassows (source: HBW Alive, https://www.hbw.com/family/guans-chachalacas-curassows-cracidae; access: 21. 07. 2019). Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 3 The geographical distribution range of cracids reaches from the northern border of Mexico to Uruguay and Central Argentina (Figure 1). They are occupying a very wide altitudinal range of distribution from sea level up to 3900 m and from the tropical lowlands, with highest diversity, to the mountains in subtropical and temperate zones (del Hoyo and Motis 2004, del Hoyo 2019).

1.1.4 Conservation status, main threats, and conservation objectives

Del Hoyo (2019) outlined that even though there are still some large gaps of basic aspects of cracids biology, their conservation status is fairly well known. What we know about the status of cracids today is basically the result of: (1) Coordinated information exchange between field researchers working on cracid species under the ICBP/IUCN International Cracidae Specialist Group. As a result of this cooperation, a detailed Action Plan was prepared (Strahl 1990, cited in del Hoyo 2019), listing each species' status and conservation priorities, country by country. (2) Publication of the ICBP/IUCN Red Data Book for the Americas (Collar 1992, cited in del Hoyo 2019). In addition, the IUCN/SSC prepared a status survey and conservation action plan for Cracids in 2000 (Brooks et al.).

Table 1. Conservation status of different groups of Cracidae species (curassows, guans, and chachalacas) summarized after The IUCN Red List of Threatened Species (Birdlife International 2019b, IUCN 2019a). CRACIDS Conservation status TOTAL Curassows Guans Chachalacas

Extinct in the wild 1 0 0 1

Critically Endangered 4 1 0 5

Endangered 3 5 0 8

Threatened Vulnerable 3 4 1 8 Near Threatened 2 3 0 5 Least Concerned 3 12 14 29

Number of species 16 25 15 56

So far 21 out of the 56 known Cracidae species (curassows, guans, and chachalacas) are listed among one of the globally “Threatened” categories on the IUCN Red List of Threatened Species (Table 1; summarized by Birdlife International 2019b and IUCN 2019a). It is clear that the status of chachalacas is different from that of the guans and curassows (del Hoyo 2019). Only one species of chachalacas, Rufous-headed Chachalaca (Ortalis eryhroptera) is currently considered vulnerable. Out of 25 guan species there are 10 Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 4 considered threatened and three near threatened. For the curassow genera, the situation is even more alarming. Out of 15 species still existing in the wild, there are 10 species considered threatened, and two near threatened. It is important to mention that 4 out of 5 critically endangered cracid species belong to the curassow genera, which makes them the most threatened group from the Cracidae family. It is also important to mention that Bare- faced Curassow (Crax fasciolata), the focus species of my study, is listed as vulnerable (summarized by Birdlife International 2019b and IUCN 2019a).

According to del Hoyo (2019), the difference in conservation status between chachalacas and other cracids can be explained by habitat requirements. Guans and curassows are basically forest-dwelling species, often dependent on primary forest (del Hoyo 2019). However, there is alarming evidence of strong decrease of suitable habitat for all cracids, especially in Amazonia, due to human large-scale land exploitation (logging, soya and cotton monocultures; Delacour and Amadon 2004). Deforestation is seriously affecting cracid populations throughout the whole Neotropical Region and nearly all species of guans and currasows are facing severe declines in numbers and local extinctions as a result of habitat destruction and hunting pressure (Brooks and Fuller 2006, del Hoyo 2019). In a recent study (Rios et al. 2020) suggested that hunting pressure is the most important factor to consider in future conservation strategies for the Red-billed Curassow (Crax blumenbachii).

However, differences in overall status are also due to species' required range sizes and whether or not these are sufficiently large and within protected habitats. Therefore, both research and conservation measures must be jointly coordinated for taxon-specific requirements of cracid populations. Since cracids are somewhat easy to survey and because of their sensitivity to habitat loss and hunting, they are among the most important bioindicators for habitat quality. Only legal protection implementation agreements, stating the cracids are key species with important biological and ecological environmental roles, can stop the current alarming trends of decreasing population numbers (del Hoyo 2019).

1.2 BARE-FACED CURASSOW (CRAX FASCIOLATA SPIX, 1825) 1.2.1 Scientific classification

There are still some debates regarding the classification of different subspecies of the Bare- faced Curassow (Crax fasciolata Spix, 1825). Until recently there were three recognized subspecies: C. f. fasciolata, C. f. grayi and C. f. pinima (Delacour and Amadon 2004, Clay and Oren 2006). C. f. xavieri was suggested as the fourth subspecies (Nardelli 1993, cited in Clay and Oren 2006), but is now considered a possible aberration from captive individuals (Clay and Oren 2006, del Hoyo et al. 2019). However, according to del Hoyo et al. (2019) and BirdLife International (2019a, 2020), there is support to regard C. f. pinima as a separate Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 5 species, Crax pinima (Belem Curassow) and therefore only C. f. grayi and C. f. fasciolata are considered as the real subspecies of Crax fasciolata.

1.2.2 Description of morphological characteristics

There is a large degree of sexual dimorphism in the plumage of Bare-faced Curassow. Sexes can clearly be recognized even in fledglings.

Adult males are larger than females with 77 – 85 cm in length and have a body mass between 2700 and 2800 g. Females measure approximately 75 cm in length and have a body mass of 2200 to 2700 g. Both sexes have bare, blackish skin around the eye and long recurved crown feathers forming an expressive curled crest (Delacour and Amadon 2004; Ridgely 2010; del Hoyo et. al 2019; Figure 2)

Figure 2. Adult female and male of Bare-faced Curassow (Crax fasciolata). The photo was taken during the survey at Baía das Pedras in the Pantanal of Mato Grosso, Brazil in November 2017. Couple of birds moved close to each other while foraging near the Advanced Research Base of the Pantanal (BAPP) of the Federal University of Mato Grosso.

Males of C. fasciolata (Figure 2) are rather similar in appearance to C. pinina. They are largely black, but slightly glossed with greenish blue. Their curled crest is also black. Their belly, thigh tufts and tip of the tail are white, and they show a few white bars on the thigh feathers. Their slightly swollen cere is bright yellow in color. There is no knob or wattles. The horny beak is dusky with horn-colored blackish tip. Their eyes are dark brown, and their legs are rownish gray in color. Immature males look similar but show some yellow on bare skin below the eye (Delacour and Amadon 2004; Ridgely 2010; del Hoyo et. al 2019). Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 6 Females (Figure 2) are black or grayish-black above narrowly barred white on upper breast, back, wings and tail. The neck is just blackish. The tail has a dozen narrow white bars and the tip of the tail is white. The lower underparts except the breast are rich buff or fulvous. Also, the barring is on the breast and, in some birds, on the shoulders. The back is also buffy, rather than white. Young females are buffier than adults. Bicolored curled crest is well- developed, the feathers are mostly white, with curled black at the base. Their cere is dusky like the face, without the bright yellow of the male. The eyes are paler brown than of the male and their legs are fleshy pale pink in color (Delacour and Amadon 2004; Ridgely 2010; del Hoyo et. al 2019).

1.2.3 Geographical range of distribution

The Bare-faced Curassow (C. fasciolata) was extirpated from many parts of its original range (del Hoyo et al. 2019) but it is still distributed throughout an extensive area in central South America (Figure 3). The nominate subspecies Crax f. fasciolata is distributed in central and southwest Brazil, Paraguay and northern Argentina, while C. f. grayi occurs in eastern Bolivia (Clay and Oren 2006).

In Brazil, the Bare-faced Curassow is widely distributed and has been reported for Pará, Minas Gerais, Goiás, Mato Grosso, and north of Mato Grosso do Sul (BirdLife International 2019a). It is considered extinct in São Paulo state or extremely rare in Paraná, but still common in the Pantanal of Mato Grosso, Emas, Brasillia, and Araguaia National Parks (del Hoyo et al. 2019).

Figure 3. Geographical range of distribution of Bare-faced Curassow (Crax fasciolata; source: HBW Alive, https://www.hbw.com/species/bare-faced-curassow-crax-fasciolata; access: 21. 07. 2019). Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 7

1.2.4 Habitat preferences

Bare-faced Curassows usually occur in forest habitats, particularly in tropical lowland evergreen forest, semi-deciduous forest, and gallery forest, up to at least 900 meters of altitude (Stotz et al. 1996, del Hoyo and Motis 2004, del Hoyo et. al 2019). The species is frequently recorded at woodland edges and is not strictly tied to forest habitats than most other conspecifics. Occasionally they are noticed wandering along river or lake sandbanks, usually in early morning or late evening. They can also be observed walking onto roads, or appearing at small clearings, savannas, marsh edges, and often show a clear preference for river vicinities (del Hoyo and Motis 2004, Fernández-Duque et al. 2013, del Hoyo et. al 2019).

1.2.5 Conservation status and main threats

According to Stotz et al. (1996) the Bare-faced Curassow is considered as “fairly common” but its global population size has not been quantified up to date (Birlife International 2019). Although the Bare-faced Curassow still occupies a relatively large range (del Hoyo et al. 2019), with approximately 4.720.000 km2, it has disappeared from many parts of its former range of distribution. It is classified as “Vulnerable” on the IUCN Red List of threatened species with its current population trend in decrease (IUCN 2019a). The classification is based on a future deforestation model, which suggests that the species will lose further 24 - 36% of suitable habitats in Amazonia over the next 35 years, a percentile supported by other estimated habitat loss predictions of at least 17.8% (del Hoyo et al. 2019, Birdlife International 2019a). The species’ sensitivity to fragmentation, edge effects, and hunting adds additional impact on decline of its population. The suspected rate of population decline over the next three generations in Amazonia is therefore calculated to be 28%. Overall decline is assumed to be between 30 and 49%, considering the whole range of its current distribution. In the IUCN Cracid Action Plan, the Bare-faced Curassow is listed as High Conservation Priority. It is also suggested that more information is needed on its population size, trends, and habitat loss (BirdLife International 2019a).

1.2.6 Historical and contemporary distribution of Bare-faced Curassow in the Brazilian Pantanal

The Bare-faced Curassow is still common in some parts of its range of distribution, most notably in the Pantanal of Mato Grosso and Pantanal of Mato Grosso do Sul (Brazil; Cintra and Yamashita 1990). The level of its abundance is the highest in the Pantanal (del Hoyo and Motis 2004). The most recent survey on the Bare-faced Curassows density (Desbiez and Bernardo 2011) in the Pantanal of Mato Grosso has shown interesting results. The Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 8 highest density was found in forest landscapes (4.66 individuals / km2), the lowest density in floodplain landscapes (0.43 individuals / km2). Therefore the Pantanal is an ideal area to study C. fasciolata since the anthropogenic impacts, such as habitat destruction and hunting pressure, are low (Desbiez and Bernardo 2011). This allows us to study the species´ behavior under optimal natural conditions to plan and conduct suitable conservation actions.

1.2.7 Existing measures and conservation actions

BirdLife International (2019a) outlines that no targeted conservation and research actions for the Bare-faced Currasow are known, but that the species presumably occurs in a number of different, already established protected areas. It also outlines that there are only a few proposed conservation and research actions in general which include:

“The expansion of protected area network to effectively protect Important Bird and Biodiversity Areas (IBAs). Effectively resource and manage existing and new protected areas, utilizing emerging opportunities to finance protected area management with the joint aims of reducing carbon emissions and maximizing biodiversity conservation. As essential, conservation on private lands is suggested, as a result of expanded market pressures for sound land management and prevention of forest clearance on lands unsuitable for agriculture (Soares-Filho et al. 2006, cited in BirdLife International 2019a). Campaign against proposed changes to the Brazilian Forest Code which if implemented will lead to a decrease in the width of the areas of riverine forest protected as Permanent Preservation Areas (APPs), which function is to provide a vital corridors in fragmented landscapes.”

1.2.8 Feeding habits

As many other cradics, the Bare-faced Curassow is primarily a frugivorous species, but was also observed to feed on seeds, flowers, shoots, and invertebrates. Fruits are usually taken from the ground after falling from trees (Delacour and Amadon 2004). It is noted that the species usually forage individually or in pairs and that they visits salt-licks (del Hoyo et. al 2019). An observation of captive individuals confirmed that they ate 23 out of 38 items offered; 20 kinds of fruits, one flower, one shoot and one gastropod (Cândido Júnior 1996). The role of the Bare-faced Curassow as important seed disperser is still debated. It has been reported that seeds are non-viable when defecated but since little study has been conducted on this topic, they are still considered as a potential seed disperser and important for forest regeneration (Silva and Strahl 1991, Guix and Ruiz 2000, del Hoyo and Motis, Golcalves et al. 2010, Langanaro 2013). Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 9

1.2.9 Activity patterns and social organization

Bare-faced Curassows are mostly terrestrial in habits (Stotz et al. 1996). Usually they are sighted alone or in male-female pairs (Clay and Oren 2006, Desbiez and Bernardo 2011, Fernándes-Duque et al. 2013, Laino et al. 2018), although groups of males can be seen together sometimes (Delacour and Amadon 2004, del Hoyo and Motis 2004, Desbiez and Bernardo 2011). The peaks of their daily activity are around dusk and dawn (del Hoyo and Motis 2004). Desbiez and Bernardo (2011) reported observations of single sex groups of males suggesting the appearance of classical lekking behavior at four different occasions between January 2003 and October 2004. Lekking behavior is defined by an aggregation of males during reproductive period to establish courtship areas and engage in competitive displays to attract females (Fiske et al. 1998). The grandest formation was observed at the forest edge grassland near a small stream on January 22nd 2003. On that occasion eleven male Bare-faced Curassows were evenly distributed, about 15 to 20 meters from and all visible to each other, while only one female was observed near a male at the formations edge. However, the behavior was not fully registered as birds were disturbed during the observation. Summarized, C. fasciolata was observed single or in groups up to 12 birds Observations were made while conducting line transects (Desbiez and Bernardo 2011). Zalazar et al. (2017) reported observations of groupings with up to 6 individuals. With previous camera trap investigations, Bare-faced Curassows were captured single or in groups up to 5 birds (Fernándes-Duque et al. 2013, Laino et al. 2018). The birds were exclusively abundant during the day and more frequent during low or mild temperatures. The evaluation of daily activity patterns (hour of capture) showed a bimodal distribution of daily activity patterns (Fernándes-Duque et al. 2013, Laino et al. 2018). A similar bimodal activity pattern is also indicated in a survey on the Red-billed Curassow (Crax blumenbachii) in the Brazilian Atlantic forest (Srbec-Araujo et al. 2012) and in a survey on the Great Curassow (Crax rubra) in Southestern Mexico (Pérez-Irineo and Santos-Moreno 2017).

1.2.10 Breeding habits

Little information is available on the breeding habits of the Bare-faced Curassow, especially from their natural habitats. However, there is some data available for other curassow species, which will be used in my study to compare it with C. fasciolata. Cracids are considered monogamous, however males show a tendency to polygamy with two or three mates (Delacour and Amadon 2004). Notes from captivity for the Bare-faced Curassow suggest that usually both sexes are involved in nest-building, in a few cases males were found to build the nest alone. Usually only the female incubates. Both sexes contribute to parental care. Offspring curassows stay with their parents for some months (Coupe 1966, Campbell Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 10 and Lack 1985, Delacour and Amadon 2004). Leite et al. (2017) reported that a male offspring of Crax globulosa stayed with its mother for at least 10 months before separation.

1.2.10.1 Observations of Bare-faced Curassow in natural habitats

The reproductive period of the Bare-faced Curassow varies among the broad range of distribution. In Paraguay, offspring were observed in December (Krieg and Schumacher 1936). In Argentina (Pirané, Formosa), a nest with eggs was discovered end of November (de la Peña 1992). For Brazil only few records exist, e.g., In Minas Gerais (Casca D’Anta waterfall, 20º15’S 46º40’W), a pair with a single well-grown male offspring was observed between July and August 2005. Assumingly, the same pair was observed to breed again in late 2005, with a single (apparently male) offspring (Bruno et al. 2006). Another pair with a single small female offspring was observed in the Pantanal of Mato Grosso (north bank of the Rio Pixaim, without further specific Locality information) in the beginning of November 2006. At the end of December, an adult pair accompanied by a male and a female offspring was observed in Goiás (Emas National Park; Kirwan 2009).

1.2.10.2 Comparative observations of the Bare-faced Curassow in captivity

Observations from a breeding pair in captivity show that the female incubated alone and left the nest twice per day to feed. The male stayed in the vicinity of the nest. The incubation took about 30 days. Both parents took care of the offspring. Soon after hatching the offspring were already active. They followed either parent closely, staying below the adult’s elongated tail. Their diet during the first days after hatching consisted mostly of insects, but they began to take more fruits shortly thereafter. Since hatching, they were fed by their parents. However, from the start they were capable to drink water on their own from a provided bowl. At eight weeks of age the offspring were becoming darker in color and the curly crest feathers appeared (Faust and Faust 1963 cited in Delacour and Amadon 2004, Coupe 1966).

Unfortunately, most of the descriptions for the Bare-faced Curassow in this section came from single anecdotal case observations on the morphogenetic development of offspring. Therefore, detailed data on growth patterns under natural patterns are needed to better understand their life stages and parental care investments of this cracid species.

1.3 PANTANAL

The area of the Pantanal (18°00'00"S 56°30'00"W) contains seasonally flooded wetland which extends over the area coverage of 140.000 km2 (Coutinho 1994). The bigger portion of it is located in Western Brazil (states Mato Grosso and Mato Grosso do Sul) and it extends beyond the borders of Bolivia and Paraguay (Figure 4). This large alluvial plain is influenced by rivers which drain the Upper Paraguay basin. The Pantanal is surrounded and affected by four larger biomes: Amazon, Cerrado, Atlantic Forest, and Gran Chaco. This wetland ecosystem is one of the smallest Brazilian landscape units but one of the richest in terms of biodiversity due to influence of the adjacent biomes, together with it’s geological, climate, and relief characteristics and the seasonal hydrological cycles (Coutinho 1994, Gwynne et al. 2010, Boin et al. 2019, IUCN 2019b).

Figure 4. Geographical position of Pantanal (18°00'00"S 56°30'00"W) in relation to different countries and states. Source: Google Earth Pro (the main layer and map modification), ArcGIS (modification of layer for Pantanal from shape file to kmz file in program; shape file source: Bioscience, An Ecoregions-Based Approach to Protecting Half the Terrestrial Realm DOI: https://doi.org/10.1093/biosci/bix014).

1.3.1 Pantanal as an important bird sanctuary

The Pantanal supports one of the largest inland concentrations of water birds (Gwynne et al. 2010) and with 463 bird species in the floodplain habitat alone, this wetland harbors the highest bird species richness among wetlands worldwide (Tobelis and Tomas 2003, Wetlands International 2006). Tobelis and Tomas (2003) concluded that none of these 463 avian species are endemic to the Pantanal. It is an important refuge for many endangered species, such as the Hyacinth Macaw (Anodorhynchus hyacinthinus) and the Jabiru (Jabiru mycteria; Gwynee et al. 2010). The area also hosts species which were recently considered endemic for the adjacent Cerrado biome (Tubelis and Tomas 2003). Thus, the Pantanal can act as an important bird sanctuary, because Cerrado biome has become more threatened and fragmented over the last decades (Gwynne et al. 2010). There is also a strong potential that the Pantanal harbors even more species than we know, as many parts of this ecosystem are still poorly known due to limited accessibility (Tubelis and Tomas 2003).

1.4 CAMERA TRAP BASED SURVEY OF CRAX SPECIES

''In the study of rare and elusive wildlife, photographs not only provide confirmation of presence and identity, but photographic evidence may also provide insights into distribution, abundance, population dynamics and behavior as well.''… ''The simplest use of camera trapping is documentation of a species’ occurrence at a site. Species occurrence data is an important component of biodiversity surveys, as well as a fundamental aspect of range determination and IUCN status.'' (O’Brien and Kinnaird 2008). Biodiversity assessment studies based on camera trap data have become very common in the last decades (Trolliet et al. 2014, Caravaggi et al. 2017). Such surveys have many advantages. One of the most essential ones is the possibility of 24-hour monitoring without human disturbance leading to a high cost efficiency. Previous camera trap studies focusing on the Bare-faced Curassow (Fernández-Duque et al. 2013, Gomes et al. 2018, Laino et al. 2018) and a few other Crax species (Srbec-Araujo et al. 2012, Lafleur et al. 2014, Alves et al. 2017, Pardo et al. 2017, Pérez-Irineo and Santos-Moreno 2017, Whitworth et al. 2018), have documented the effectiveness of this method and led to a great amount of new data on aspects of life history, behavior, occurrence, and habit preferences.

The Pantanal is an ideal place for detailed research of Crax fasciolata since the anthropogenic impacts are low in this region (Desbiez and Bernardo 2011). Also, the survey area is protected, human activities are limited, and therefore the human impact on activity patterns is considered low. This provides an exceptional opportunity to observe a species´ ecological and behavioral characteristics in its natural environment. Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 13

1.5 OBJECTIVES AND APPROACHES

This study is part of a long-term international biodiversity assessment program on vertebrates and invertebrates called Sounds of the Pantanal – The Pantanal Automated Acoustic Biodiversity Monitoring Program in the northern Pantanal, Brazil, run by the Brazilian National Institute for Science and Technology in Wetlands (INAU), Federal University of Mato Grosso (UFMT), Computational Bioacoustics Research Unit (CO.BRA) and coordinated by Karl-L. Schuchmann and Marinez Isaac Marques (SISBIO permit no. 39095 KLS). For my study all camera trap data and metadata was provided by CO.BRA. My MSc. project on Crax fasciolata was suggested and supervised by Prof. Karl-L. Schuchmann and is based on the taxon-specific data with special focus on selected eco-ethological life history patterns. To achieve this goal I addressed the following topics (1, 2) and the listed qualitative and quantitative assumptions (H1-H6):

(1) Evaluation of activity patterns, sex ratio and social organization. (2) Evaluation of offspring’s age and assessment of breeding season.

H1: Capture success varies between different seasons, periods, and years. H2: Daily activity patterns are bimodal. H3: Adult birds form single sex aggregations. H4: Both parents invest in parental care. H5: No significant difference exists in the numbers of male and female individuals in camera trap captures. H6: Breeding occurs throughout the year.

2 MATERIALS AND METHODS 2.1 GENERAL DESCRIPTION OF A STUDY AREA

The study was conducted at SESC Baía das Pedras, one of the units of the SESC Pantanal Ecological Resort, located in the municipality of Poconé in the Northern Pantanal of Mato Grosso, Brazil (Figure 5, Figure 6, 16°29'55"S 56°24'46"W). It is a privately protected area of approximately 4200 ha, and location of the Advanced Research Base of the Pantanal (BAPP) of the Federal University of Mato Grosso. The field station, with a laboratory and lodging infrastructure (SESC Pantanal 2019), served as a base camp for this study.

Figure 5. Geographical position of Pantanal and SESC Baía dasa Pedras Park (study area; 16°29'55"S 56°24'46"W) in relation to different countries and states. Source: Google Earth Pro (the main layer and map modification), ArcGIS (modification of layer for Pantanal from shape file to kmz file in program; shape file source: Bioscience, An Ecoregions-Based Approach to Protecting Half the Terrestrial Realm DOI: https://doi.org/10.1093/biosci/bix014).

Figure 6. Orthophoto of SESC Baía dasa Pedras Park (study area, 16°29'55"S 56°24'46"W) and related sample locations (numbers; Appendix A). Source: Google Earth Pro (the main layer and map modification) as the borders and sample location were adjusted after Kathrin Burs (personal contact).

2.2 PROJECT HISTORY AND CAMERA TRAPPING

My work is embedded in INAU / CO.BRA program. I am using data from a long-term monitoring project (Pantanal Automated Acoustic Biodiversity Monitoring), which I joined from October to December 2017.

Between July 2015 and December 2017, 37 locations were monitored by the Mammalian study group of CO.BRA using camera traps. Cameras were placed following a computer generated, regular GRID (Hawth's Analysis Tools for ArcGIS) with a 1 km distance between different sample locations (= sample sites; Appendix A, Figure 6). The sample sites were fixed over different periods of sampling. The sampling effort (number of days - 24 hour cycles, when camera trap was active) varied between different sample sites (Appendix B), periods (dry = April to September, wet = October to March), seasons (receding water = April to June, low water = July to September, rising water = October to December, high water = January to March), and years (2015, 2016, 2017). In each location, sampling was continuous for a minimum of 5 days. Each sampling location was equipped with a single camera trap on a diel regime. Five different models of camera traps were used for the purposes of this study (RECONYX PC800, RECONYX HC600, UWAY VH400, BUSHNELL TROPHY CAM AGGRESSOR, BUSHNELL TROPHY CAM HD 2012). All models were infrared triggered, responding to temperature changes or movement of animals within the optical camera range, and producing images and videos. With every image or video taken, the cameras also recorded moon phase, temperature, time and date. Fixed camera trap height was 60 cm.

2.3 ACTIVITY PATTERNS, SOCIAL ORGANIZATION AND SEX RATIO 2.3.1 Data Analysis

Crax fasciolata individuals were properly identified (Delacour and Amadon 2004, Van Perlo 2009, Gwynne et al. 2011) from image and video sets from our 37 sample sites, and joined with metadata such as temperature, moon phases, and seasonal periodical activity patterns.

Time and date of data records (= captures) were gathered and pre-analysed according to independent captures. An independent capture was determined by the following approaches:

(1) Following the methodology of O'Brien et al. (2003), originally developed for the analyses of Relative Abundance Indices of mammal species, only images or videos of C. fasciolata Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 17 taken 30 minutes apart were counted as an independent capture, except when different individuals on subsequent records could clearly be recognized.

(2) According to suggested applications by Srbec-Araujo et al. (2012), for the Red Billed Curassow (Crax blumenbachii), I modified and adapted these for the purpose of my study as follows: Instead of a 60-min interval, the images or videos of Crax fasciolata were taken 30 min apart and were counted as an independent capture. The decision to take the 30-minute option into consideration was made for practical reasons to allow the comparison of the data with a parallel run mammal survey in the same grid area.

To achieve the goals of my study, two different capture success variables were defined as two different approaches were used. Capture success of individuals (CSind; synonymous to Relative Abundance Index (RAI) defined by O'Brien et al. 2003), which represents the number of independent records of recognized individuals per 100 days and capture success of the species (=CSsp; synonymous to Capture success defined by Srbec-Araujo et al. 2012), which represents the number of independent records of species per 100 days. The calculation of relationship between CSind and CSsp expressed with the CSind / CSsp ratio was possible, as the same time interval for independence was used for both approaches. The ratio represents the average group size of independent individual captures recorded per independent survey occasion.

2.3.2 Data comparison

The nonparametric Chi2 (= x2) test was applied for analyses of differences between seasonal and daily activity patterns. The considered level of significance was p > 0.05.

Capture success was compared between (three) different years, periods, and seasons. Furthermore, the same period and season was compared between different years. Capture success comparisons were carried out to identify the differences in activity patterns between different time periods. In addition, records were classified into different categories: “Single” - one adult male or one adult female, “Pair” - one adult male and one adult female or “Group” - any other composition of adult and offspring individuals. Furthermore, “Group” was divided into two subcategories: “Adult with offspring” and “Other form of grouping”. Then, based on the combined data from all three years, comparisons of capture frequency between different categories were carried out to identify different seasonal activity patterns.

The time data of independent captures for the whole data set was modified into one-hour intervals (5:00 to 05:59, 6:00 to 06:59, etc.) with the purpose to determine daily activity patterns. When data sets were smaller, time data were combined and analyzed by two-hour intervals (04:00 to 05:59, 06:00 to 7:59, etc.). Daily activity patterns were compared between Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 18 different periods (dry, wet) and sexes (female, male). As the offspring’s occurrence might depend on the occurrence of adult individuals, the daily activity pattern was analyzed (1) excluding and (2) including the offspring. In addition, daily activity patterns were compared between independent species captures where offspring were present and those where they were absent.

Based on data for independent individual captures from all three years, male and female capture frequencies were compared separately for adults and offspring. Comparisons were carried out to determine the adult and offspring sex ratio (number of males per one female) and identify if those were skewed (Hill et al. 2008, Fournier and Janik 2008, Martínez- Morales et al. 2009, Srbek-Araujo et al. 2012, Fernández-Duque et al. 2013, Alves et al. 2017, Pardo et al. 2017, Whitworth et al. 2018, Laino et al. 2018). Data from referenced studies were summarized and adjusted for comparisons between different studies and Crax species.

2.4 OFFSPRING AGE IDENTIFICATION AND ASSESSMENT OF BREEDING SEASON

For a more precise conclusion about the reproductive period of C. fasciolata based on camera trap data, an “Age-class identification key for C. fasciolata offspring” was prepared.

2.4.1 Development and usefulness of »Age-class identification key for Crax fasciolata offspring«

A possible way to reconstruct the onset of the breeding season is to identify »Age-classes for C. fasciolata offspring« and to consider reciprocally the developmental stages of offspring, e.g., hatching, incubation, and laying for each capture on a monthly basis.

The age identification key is based on the literature available for different curassow species (Guimaraes et al. 1935, cited in Vaurie 1968, Krieg and Schumacher 1936, cited in Delacour and Amadon 2004, Taibel 1940, Bronzini 1940, cited in Vaurie 1968, Bronzini 1943, cited in Vaurie 1968, Taibel 1953, cited in Vaurie 1968, Vaurie 1968, del Hoyo and Motis 2004, Delacour and Amadon 2004, Roer, cited in Delacour and Amadon 2004). However, studies considering age dependent characteristics are derived from offsprings in captivity and anecdotal observations from field studies, specifically on the Bare-faced Curassow, but they rarely examined their age (Krieg and Schumacher 1936, cited in Delacour and Amadon 2004, de la Peña 1992, cited in Delacour and Amadon 2004). Observations on the potential differences between natural and captive rearing of Cracidae species are missing. Since offspring in captivity are raised under optimal food conditions and under little environmental stress from predation, the existing literature can only be applied with caution when compared Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 19 to wild populations. The most important age assessment parameters I used are based on morphological characteristics which are clearly distinctive and comparable between different ages of offspring and adults.

I developed an age-class key to provide an approximate age assessment with a mean of variation of one months. For an estimate of the breeding season of C. fasciolata in the Northern Pantanal, Brazil, camera trap offspring captures were used for which the development stages since hatching were considered as an estimate. After such an estimation of age-class for the individual capture, the subtraction of the obtained age-class interval from the capture date followed. This resulted in an estimation of the hatching range interval. From the latter, the subtraction of additional 30 days (incubation period for C. fasciolata in captivity referred to Faust and Faust, cited in Delacour and Amadon 2004) was conducted to estimate egg laying range interval. The time period between egg laying and hatching range intervals represents the estimation of the incubation range. For an easier and uniform use of the identification key, I prepared a specification spreadsheet (Figure 20). In addition, the graphical presentation on a half-monthly basis was added for visualization of those estimations. The graphical (Figure 20) contains the information in which month the data (independent capture) was recorded and in which months laying and hatching occurred according to my estimation.

3 RESULTS

Camera traps were active for 4768 days and nights. 554 independent captures of Bare-faced Curassow individuals were obtained within 357 independent 30-minute capture periods. 26 (70.27%) out of 37 fixed sample sites captured the species during the study period. Offspring were captured at 8 (21.62%) sample sites. Due to logistics, availability of field researchers, accessibility of sample site, and / or camera trap malfunctioning the sampling effort per sample site and sample occasion differed (Appendix A).

3.1 ACTIVITY PATTERNS

The overall capture success of the species (= CSsp) was 7.49 (Table 2) of camera trap sampling effort. There is little evidence to conclude that there was a difference in capture success of the species between dry and wet periods (x2 = 2.91, df = 1, p = 0.088). As there were enough independent records of the species, the capture success in the four different seasons was compared. Capture success was highest in receding water season with 9.39, low and rising water season followed with 7.29 and 6.75 respectively. With 5.92, high water season deviates with the lowest frequency of captures. Overall, there is little evidence to conclude that there was a difference in capture success of the species between the different seasons (x2 = 7.14, df = 3, P = 0.067). The comparison of different seasons within the same period didn't show a significant difference neither between receding and low water season (x2 = 3.26, df = 1, P = 0.07078) nor between rising and high water season (x2 = 0.22, df = 1, P = 0.637).

The overall capture success of individuals (= CSind) was 11.62 of camera trap sampling (Table 2). There is evidence for a significant difference in capture success of individuals between dry and wet period (x2 = 10.44, df = 1, p = 0.001). The number of independent records of individuals was sufficient to analyze capture success variances between the different seasons. Capture success was highest in the receding water season with 17.10, low and rising water season followed with 10.61 and 10.24 respectively. With 6.98, the high water season deviates with the lowest frequency of captures. Overall there is significant evidence that there was a difference in capture success of individuals between different seasons (x2 = 35.36, df = 3, P < 0.00001). The comparison of the different seasons within the same period showed a significant evidence that there was a difference between receding and low water season (x2 = 19.41, df = 1, P = 0.00001), while there was no significant difference between rising and high water season (x2 = 3.27, df = 1, P = 0.070).

CSind / CSsp ratio (Table 2) for overall data sets was 1.55. Moreover, the average of detected individuals per survey occasion was higher during the dry period than during the wet period with 1.61 against 1.43, respectively. Divided into seasons, the highest overall average was detected during receding water season with 1.82, followed by rising and low water season with an average of 1.52 and 1.46 respectively. With 1.18, the high water season resulted with the lowest average of individuals per independent survey occasion.

Table 2. Number of independent camera trap captures / 30 min, capture success of the species (number of independent records of species per 100 sampling days), capture success of individuals (number of independent records of individuals per 100 sampling days), CSind / CSsp ratio (= average of detected individuals per independent survey occasion) of Bare-faced Curassows in Baia das Pedras in Pantanal of Mato Grosso, Brazil expressed for different Years, Periods and Seasons. Number of Number of Capture Capture independent independent Sampling CSind/CSsp Variables success of success of captures of captures of effort ratio the species individuals the species individuals Total 64 97 864 7.41 11.23 1.52

Dry 64 97 864 7.41 11.23 1.52 Period Wet ------

2015 Receding water ------

Low water 64 97 864 7.41 11.23 1.52 Season Rising water ------

High water ------Total 116 198 1180 9.83 16.78 1.71

Dry 81 147 794 10.20 18.51 1.81 Period Wet 35 51 386 9.07 13.21 1.46

2016 Receding water 80 146 766 10.44 19.06 1.83

Low water 1 1 28 3.57 3.57 1.00 Season Rising water 30 44 340 8.82 12.94 1.47

High water 5 7 46 10.87 15.22 1.40 Total 177 259 2724 6.50 9.51 1.46

Dry 101 151 1407 7.18 10.73 1.49 Period Wet 76 108 1317 5.77 8.20 1.42

2017 Receding water 21 38 310 6.77 12.26 1.81

Low water 80 113 1097 7.29 10.30 1.41 Season Rising water 53 82 890 5.96 9.21 1.55

High water 23 26 427 5.39 6.09 1.13 Total 357 554 4768 7.49 11.62 1.55

Dry 246 395 3065 8.03 12.89 1.61 Period Wet 111 159 1703 6.52 9.34 1.43

Overall Receding water 101 184 1076 9.39 17.10 1.82

Low water 145 211 1989 7.29 10.61 1.46 Season Rising water 83 126 1230 6.75 10.24 1.52

High water 28 33 473 5.92 6.98 1.18

Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 22 During 2015 camera trapping was carried out only in the dry period but it was considered in the overall evaluation between seasons.

The capture success of the species (= CSsp) was higher in 2016 than in 2017, with 9.83 against 6.50 respectively and a significant difference (x2 = 10.76, df = 1, P = 0.001). The proportion of capture success in the dry and wet period was similar in 2016 and 2017 despite the high difference between capture success of the species when compared on a yearly basis (Figure 7). However, the results show that there was no significant difference between dry and wet periods in 2016 (x2 = 0.21, df = 1, P = 0.650) and 2017 (x2 = 1.74, df = 1, P = 0.187).

The capture success of individuals (= CSind) in 2016 was higher than in 2017, with 16.78 against 9.51 respectively, and with a significance level of P < 0.00001 (x2 = 31.88, df = 1), but the proportion of capture success of individuals between dry and wet period was similar in 2016 and 2017 despite the high difference between capture success frequency of individuals on a yearly basis (Figure 8). However, the results also show that there is a similar level of significance of results between dry and wet periods in 2016 (x2 = 3.47, df = 1, P = 0.062) and 2017 (x2 = 3.93, df = 1, P = 0.047).

10.20 (52.94%) 9.07 (47.06%) 7.18 (55.44%) 5.77 (44.56%) Dry period

Wet period

the species the

of Capture success success Capture

2016 2017 Year Figure 7. Comparison of capture success of the species (number of independent records of the species per 100 sampling days) of Bare-faced Curassows between dry and wet period for 2016 and 2017. Data was obtained during the survey at Baía das Pedras in the Pantanal of Mato Grosso, Brazil. Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 23

18.51 (58.35%)

13.21 (41.65%) 10.73 (56.69%) 8.20 Dry period

(43.31%) Wet period

of individualsof Capture success success Capture

2016 2017 Year Figure 8. Comparison of capture success of individuals (number of independent records of individuals per 100 sampling days) of Bare-faced Curassows between dry and wet period for 2016 and 2017. Data was obtained during the survey at Baía das Pedras in the Pantanal of Mato Grosso, Brazil.

The data comparison between different seasons is based on independent records of the species and is explained in the context of sample size which was insufficient for receding and low water seasons in 2016 and thus was excluded from further analysis. Data from 2017 showed no significant difference between those two seasons (x2= 0.03, df = 1, P = 0.867). Results of the comparison between rising and high water season suggest that there is no difference in capture success neither for 2016 (x2 = 0.02, df = 1, P = 0.888), nor for 2017 (x2 = 0.07, df = 1, P = 0.790). Capture success between 2016 and 2017 did not vary between receding (x2 = 2.55, df = 1, P = 0.110), low (x2 = 0.53, df = 2, P = 0.767), rising (x2 = 2.37, df = 1, P = 0.123) and high water season (x2 = 1.13, df = 1, P = 0.288).

The data comparison based on independent records of individuals for the receding and low water season in 2016 was also excluded from further evaluation (explanation see above). Data from 2017 showed no significant difference between those two seasons (x2 = 0.60, df = 1, P = 0.438). Results of the comparison between rising and high water season suggest that there is no difference in capture success neither for 2016 (x2 = 0.02, df = 1, P = 0.885), nor for 2017 (x2 = 2.83, df = 1, P = 0.092). There is significant evidence of a difference in capture success between 2016 and 2017 for receding water season (x2 = 4.83, df = 1, P = 0.028). But comparison of results for low (x2 = 1.63, df = 2, P = 0.442), rising (x2 = 2.63, df = 1, P = 0.104) and high (x2 = 3.20, df = 1, P = 0.073) water season suggests that there is no difference in capture success between 2016 and 2017.

3.1.1 Daily activity patterns

The results show similar patterns for the independent records of the species (Figure 9) and the independent records of adult individuals (Figure 10). Independent captures of Bare-faced Curassows were recorded over a 13-h interval. The earliest capture was recorded at 05:04 h and the latest one at 17:54 h. A prominent peak occurs between 06:00 - 07:00 and the second, lower than the first, between 16:00 - 17:00 h. The lowest values of daily activity were noted between 13:00 – 14:00 h (Figure 9, Figure 10).

species captures species 10

Number of independentof Number 0

Time of the capture Figure 9. Number of independent species captures of Bare-faced Curassows obtained during the survey (from July 2015 to December 2017) at Baía das Pedras in the Pantanal of Mato Grosso, Brazil, based on 1 h periods (based on independent capture rates of 30 min interval).

90 80 70 60 50 40 30

of adult individuals adult of 20 Number of captures captures of Number 10 0

Time of the capture

Figure 10. Number of captures of adult individuals of Bare-faced Curassows obtained during the survey (from July 2015 to December 2017) at Baía das Pedras in the Pantanal of Mato Grosso, Brazil, based on 1 h periods (based on independent capture rates of 30 min interval). Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 25 The proportional data for different periods and sexes shows that the bimodal distribution of all independent records of adult individuals was also very similar between dry and wet period (Figure 11; x2 = 10.35; df = 12, P = 0.585) and between adult males and females (Figure 12; x2 = 5.13; df = 12, P = 0.953).

Dry Wet All 18 16

14 (%) 12

captures captures 10

individuals 8 6

ndependent 4

I of adult of 2 0

Time of the capture

Figure 11. Periodically based proportions (%) of independent captures of adult individuals of Bare-faced Curassows obtained during the survey (from July 2015 to December 2017) at Baía das Pedras in the Pantanal of Mato Grosso, Brazil, based on 1 h periods (based on independent capture rates of 30 min interval).

Female Male All 18 16

14 (%) 12

captures 10 8 6

ndependent ndependent

I of adult individualsadult of 2 0

Time of the capture

Figure 12. Sexually based proportions (%) of independent captures of adult individuals of Bare-faced Curassows obtained during the survey (from July 2015 to December 2017) at Baía das Pedras in the Pantanal of Mato Grosso, Brazil, based on 1 h periods (based on independent capture rates of 30 min interval). Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 26 The cumulative results of all independent captures of the species are similar to the capture rates of adult individual Bare-faced Curassows. There is no significant difference in daily activity patterns between dry and wet periods (Figure 13; x2 = 10.10; df = 12, P = 0.607). There is also no significant difference between males and females when only captures with single adult animals were considered (x2 = 10.39; df = 12, P = 0.581). The comparison between all independent species captures without the presence of offspring with those with the presence of offspring resulted in insufficient evidence to conclude the difference in capture between those (Figure 14; x2 = 12.27; df = 12, P = 0.424).

Dry Wet All 12

(%) captures captures 6

species species 4 of

ndependent ndependent 2 I

Time of the capture

Figure 13. Periodically based proportions (%) of independent species captures of Bare-faced Curassows obtained during the survey (from July 2015 to December 2017) at Baía das Pedras in the Pantanal of Mato Grosso, Brazil, based on 1 h periods (based on independent capture rates of 30 min interval).

15 With offspring 10 of species (%) species of Without offspring

Independent captures captures Independent 5 All

Time of the capture

Figure 14. Proportions (%) of independent species captures of Bare-faced Curassows with and without offspring obtained during the survey (from July 2015 to December 2017) at Baía das Pedras in the Pantanal of Mato Grosso, Brazil, based on 2 h periods (based on independent capture rates of 30 min interval). Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 27

3.2 SOCIAL ORGANIZATION

The number of individuals of Bare-faced Curassows per independent capture ranged between 1 – 4 with an average of 1.55 ± 0.81 SE. As shown in Figure 15, overall captures with single individuals were higher (n = 217; 60.78%) than captures of any other formation (n=140; 39.22%). Single male captures (n = 118; 33.05%) were the most common, followed by single female captures (n = 99; 27.73%). There wasn’t any offspring identified outside of group formations. Pair formations (male-female) were the most common (n = 76; 21.29%), followed by adults with offspring (n=44; 12.32%), and other forms of grouping (n =20; 5.60%).

38.98 Single male

35.94 33.05

29.69 Single female

28.45

27.73

27.12

25.86

25.00 24.14

21.29 Adult pair

18.97 (male-female)

15.25

12.43 of species (%) species of 12.32 Adult with

9.38 offspring

Independent captures captures Independent 6.21

2.60 Other form of 2.59

0.00 grouping

2015 2016 2017 Ove ral l

Figure 15. Proportions (%) of independent species captures divided on single male, single female, adult pair (male-female), adult with offspring and other forms of grouping of Bare-faced Curassow obtained during the survey (from July 2015 to December 2017) at Baía das Pedras in the Pantanal of Mato Grosso, Brazil for different years of study (based on independent capture rates of 30 min interval).

Looking closer into the latter groups (Table 3), three to four individuals were recorded during eleven occasions (3.08% of all captures), while records of adult individual groups (single sex pairs) were identified during nine occasions (2.52% of all captures). Aggregations with offspring(s) were evaluated separately as their behavior is dependent on adult individuals, they were always recorded with a single adult parent or with a parent pair (male-female) but never with larger groups or same sex pairs. Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 28 Table 3. Cumulative numbers of Bare-faced Curassows groups of at least two adult individuals of same sex or more than two adults of both sexes were recorded during the survey (from July 2015 to December 2017) at Baía das Pedras in Pantanal of Mato Grosso, Brazil. Number of survey Form of grouping occasions

2 males 5 2 females 4 1 male and 2 females 5 2 males and 1 female 4 2 males and 2 females 2

Figure 16. Proportions (%) of independent species captures of Bare-faced Curassow for overall study, different periods (dry, wet) and seasons (receding water, low water, rising water, high water) obtained during the survey (from July 2015 to December 2017) at Baía das Pedras in the Pantanal of Mato Grosso, Brazil divided on single female, single male, adult pair (male-female), adult with offspring and other forms of grouping.

The proportions of the total data sets of individuals and group captures during different periods and seasons are shown in Figure 16. There was little difference in capture proportions of different single and group categories between the dry and wet periods (x2 = 2.31, df = 4, P = 0.677). However, there is significant evidence that there was difference between different single and group categories between all seasons (receding water, low water, rising water and high water; x2 = 50.01, df = 12, P < 0.00001) and between receding and low water season (x2 = 38.98, df = 4, P < 0.00001). The biggest difference occurred for groupings with offspring with an overall maximum of 28.71% during the receding and minimum of 2.07% during the low water season. These results also suggest that there is little Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 29 evidence to conclude that capture success differed between rising and high water season (x2 = 6.17, df = 3, P = 0.104; with the exclusion of other forms of grouping under the assumption that none of the expected values should be less than one). However, the presentation (Figure 16) shows some differences in almost all categories between rising and high water seasons. Focusing on captures of adults with offspring, the results indicate another (smaller than at receding water season) peak during the rising water season with 13.25% compared to high water season with 3.57%.

The ratio of independent offspring versus adult captures, resulted with two peaks of offspring captures when compared between different seasons (Figure 17). The most obvious one was in the receding water season with 24.46% and the second in the rising water season with 11.11% of offspring captures, as the low and high water seasons resulted with 2.37% and 3.03% of offspring captures, respectively.

97.63 96.97 88.27 87.34 90.57 88.89 75.54

24.46 11.73 12.66 9.43 11.11 2.37 3.03

Total Dry Wet Receding Low Rising High water water water water Period Season

Adult records (%) Offspring records (%)

Figure 17. Overall ratio of independent individual records of adults and offspring of the Bare-faced Curassow obtained during the survey at Baía das Pedras in the Pantanal of Mato Grosso, Brazil, presented for different periods and seasons (based on cumulative data gathered from July 2015 to December 2017).

Offspring were mostly observed with adult pairs (male-female; n=24; 54.55%), followed by observations with single females (n=13; 29.55%), and single males (n=7; 15.91%; Figure 18). The comparison of captures where offspring were present with captures where they were absent suggests that there was difference between different appearances (x2 = 20.71, df = 3, P = 0.0001). This result also indicates that there is little evidence to conclude that the appearance of offspring differed between groupings with male and female (x2 = 2.04, df = 1, P = 0.153). Furthermore, results also show that there was a difference in the appearances of offspring with single or paired adult individuals (x2 = 13.66, df = 1, P = 0.0002). Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 30

Offspring captured in aggregation with

Other 0% Male 16%

Pair 55% Female 29%

Figure 18. Overall distribution of offspring grouping records of Bare-faced Curassow obtained during the survey at Baía das Pedras in the Pantanal of Mato Grosso, Brazil (based on cumulative data gathered from July 2015 to December 2017).

3.3 SEX RATIO

Combining adult and offspring records, there were 272 (49.10%) independent captures of males, 279 (50.36%) of females and 3 (0.54%) captures where the sex could not be determined, resulting in a sex ratio of 0.97:1.00. Looking at adults and offspring separately, the sex of offspring was determined for 62 (95.38%) out of 65 independent captures, with 21 (32.31%) records for males, 41 (63.08%) for females, and a sex ratio of 0.51:1.00. Adult males were identified on 251 (51.33%), females on 238 (48.67%) captures, representing a sex ratio of 1.05:1.00 (Figure 19).

Total Adults Offsprings Female 279 238 41 Male 272 251 21 Undetermined 3 0 3

Figure 19. Overall independent individual captures of different sexes of Bare-faced Curassow obtained during the survey (from July 2015 to December 2017) at Baía das Pedras in the Pantanal of Mato Grosso, Brazil, presented for adults and offspring. Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 31 With the number of males per female, the published data indicates a male-skewed adult sex ratio (ASR) for Crax genera (Table 4) (Hill et al. 2008, Fournier and Janik 2008, Srbek- Araujo et al. 2012, Fernández-Duque et al. 2013, Alves et al. 2017, Whitworth et al. 2018, Laino et al. 2018, Martínez-Morales et al. 2009, Pardo et al. 2017). Based on a published data for Crax rubra, the results suggest a female-skewed offspring sex ratio with 0.59:1.00 (Fournier and Janik 2008).

Table 4. Available sex ratio data for different Crax species regarding summarized and adjusted data available from different surveys. Sex ratio is expressed with number of males per one female respectively. Legend of different approaches on which sex ratio was based: *visual detection; transect-based survey, **camera trap- based survey, ***data from territory mapping, ****distance sampling models, *****observed in captivity. Studied Adult Offspring Survey species sex ratio sex ratio Fournier and Janik Crax ***** - - - - 2008 rubra 0.59:1.00 Hill et al. Crax * *** - - - 2008 globulosa 1.20:1.00 2.40:1.00 Martínez-Morales Crax * - - - - et al. 2009 rubra griscomi 0.56:1.00 Srbek-Araujo Crax ** - - - - et al. 2012 blumenbachii 1.56:1.00 Fernández-Duque Crax * ** - - - et al. 2013 fasciolata 1.60:1.00 1.67:1.00 Alves et al. Crax ** ****3.10- - - - 2017 blumenbachii 1.60:1.00 3.60:1.00 Pardo et al. Crax ** - - - - 2017 rubra 0.71:1.00 Whitworth et al. Crax ** - - - - 2018 rubra 1.50:1.00 Laino et al. Crax ** - - - - 2018 fasciolata 1.26:1.00 This survey Crax ** ** - - - 2020 fasciolata 1.05:1.00 0.51:1.00

3.4 AGE-CLASS IDENTIFICATION 3.4.1 Usage of Age-class identification key for Crax fasciolata offspring

I developed an age-class identification key for C. fasciolata offspring with specific annotations and applications for other curassow taxa based on shared morphological characteristics and similar body mass. The identification key is composed of information required for an easy and user-friendly application.

The separation of all offspring captures considered for C. fasciolata includes each location and survey occasion. For this part the focus was on offspring captures. If an image or video was not evident in visual details, it was not considered further. Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 32 I prepared a specification spreadsheet (Figure 20) and used one for each separate survey occasion. After filling in all basic data information obtained from the captures, age evaluation expressed in days was concluded based on age-dependent external morphological characteristics categorized with the age-class identification key. Thereafter, additional information on the length of the incubation period was added. Both datasets combined allowed the estimation of egg laying and hatching time. The latter was estimated by the subtraction of age estimation from the capture timing while the onset of egg laying was estimated by subtracting the incubation period from the hatching range.

SPECIES BREEDING DETAILS LOCATION A simple subtraction of the days estimated in age estimation from the date of the capture taken is sufficient to approximate the hatching. Then we subtract the number of the days of PLACEMENT incubation period from the hatching estimation and we will get the approximation of egg FROM TO laying. The time in between egg laying and hatching is the incubation period. We are not DATE going to determine exact dates, but just the monthly periods. TIME Incubation period of the surveyed species: OFFSPRING AGE ESTIMATED CAPTURE TIMING ESTIMATED ESTIMATED EGG CAPTURES ESTIMATION INCUBATION PERIOD HATCHING RANGE LAYING RANGE - IN DAYS RANGE INDEX - SEX INDEX- PLACE DATE TIME STUDY MENT MIN MAX FROM TO Always in between FROM TO

NOTES ON PLACEMENT NOTES ON RESULTS

GRAPHICAL Location: _____ JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC Captures taken Egg laying range Hatching range Figure 20. Offspring age-class identification key for Crax fasciolata specification spreadsheet. The spreadsheet was prepared in a way that it can be used for other curassow species as well. It is designed for the data collection of easy-to-depict external morphological information from offspring of single survey occasion and age estimation data input in combination with identification key based on Heinroth (1931, cited in Delacour and Amadon 2004), Guimaraes et al. (1935, cited in Vaurie 1968), Krieg and Schumacher (1936, cited in Delacour and Amadon 2004), Taibel 1940, Bronzini (1940, 1943, cited in Vaurie 1968), Taibel (1953, cited in Vaurie 1968), Vaurie 1968, Coupe 1966, Nardeli (1981, 1993, cited in del Hoyo and Motis 2004), del Hoyo and Motis 2004, Delacour and Amadon 2004, and Roer (cited in Delacour and Amadon 2004). Estimation of the hatching range interval is based on a simple subtraction of the age-class estimation from the capture date obtained from the captures as the estimation of egg laying range interval based on a simple subtraction of incubation period of a species from the hatching range interval. Estimated incubation period range is considered as the time between the incubation and hatching range, respectively.

3.4.2 Age-class identification key for Crax fasciolata offspring

(a) “At little over eight weeks at captive breeding the curassow offspring were becoming darker in color and beginning to develop the curly crest” (originally noted for Bare-faced Curassow (Crax fasciolata) offspring reared in captivity; Coupe 1966).

- Curly crest is absent...... ….……………….... (b) Explanation: We can assume that the offspring is less than 60 days old. - Curly crest is present....……………...……………...... …………. (g) Explanation: No assumptions should be made here, as there is no clear implication about curly crest before that statement. - Curly crest is not well visible or there are some doubts ...... ………………………………………………Check other characteristics first

(b) “At twenty days the wings are well developed” (originally noted for Razor-billed Curassow (Crax mitu) in captivity; Heinroth 1931 cited in Delacour and Amadon 2004). Check picture number 3 in Figure 21 to get visual data. The original photograph is from Great Curassow (Crax rubra) and represents three weeks old offspring in captivity.

- The wings are well developed.…………………………………………...... (c) Explanation: Assuming the offspring is 20 or more days old. - The wings are not well developed.…………………………….…………....(d) Explanation: Assuming the offspring is less than 20 days old.

(c) “The tail quills grow more slowly but are well developed at forty days. At that age wings, tail, and breast are well feathered, head is still in full down” (originally noted for Razor-billed Curassow (Crax mitu) in captivity; Heinroth 1931 cited in Delacour and Amadon 2004). Check plates and photographs 3, 4, 8, 9, Figure 21 (see photograph 9, the length of tail for 45 days old Great Curassow (Crax rubra) in captivity as a comparative calibration parameter).

- The tail quills are well-developed, wings, tail, and breast are well-feathered; colors are getting stronger.…………………………………………………….(f) Explanation: Assuming the offspring is at least 40 days old. - The tail quills are not well-developed, wings, tail, and breast less feathered...………………………………...……………………………..……(e) Explanation: Assuming offspring is less than 40 days old. Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 34 (d) Assuming that the juvenile is less than 20 days old. Check plates and photographs in identification key (Figure 21). Based on data from different species, incl. Bare-faced Curassow. Important feather features considered here: resemblances and degree of camouflaging pattern serving as an approximation of the offspring’s age. For offspring younger than 20 days, the age evaluation can be achieved with high accuracy supported by literature data (Taibel 1940, Delacour and Amadon 2004, Nardeli 1981, 1993 cited in del Hoyo and Motis 2004).

(i.) Identification key: See plates and photographs 1, 2, 3, 5, 6, 7; Figure 21. (ii.) Sexes in Bare-faced Curassow offsprings are easily identifiable shortly after hatching. Delacour and Amadon (2004) indicated sex-specific markings of males being noticeably darker than in females, see plates 1, 2; Figure 21. Also, the colored pattern is clearly seeable and distinguishable between male and female when they are just a few weeks old (personal observations). (iii.) It is also important to note down the information for Alagous Curassow (Mitu mitu) (Nardeli 1981, 1993 cited in del Hoyo and Motis 2004): …"The down feathers begin to be lost after a week… …Juvenile feathers begin to emerge on the upper back, belly, breasts, flanks, and upper tail coverts after two weeks; flight feathers and rectrices emerge after four weeks…"

(e) Assuming that the offspring’s age is in an advanced immature stage, between 20 and 40 days. Check plates and photographs 3, 4, 8, 9; Figure 21 (photograph 9, length of tail for 45 days old Great Curassow in captivity) in identification key for more accurate assessment and “flight feathers and rectrices emerge after four weeks” (data from Alagoas Curassow immatures reared in captivity; del Hoyo and Motis 2004).

(f) Assuming that immature age is between 40 and 60 days.

(g) When the size of an immature is one third of the size of an adult and the plumage is much like that of an adult, than offspring is about 2 months old.

- The size of the offspring is more than a third of the size of an adult.…...... (h) - The size of the offspring is less than a third of the size of an adult...... (i)

Explanation: Before concluding age identification for older offspring, it is important to mention a few facts from literature, quoting Vaurie (1968): “Guimaraes, Bergamin, and Crvalho (1935), Bronzini (1940, 1943), and Taibel (1940, 1953) ... …noted the change in the plumage and stated that the downy- Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 35 plumage had been replaced by a plumage similar to that of the adult in about two months, or in less than three. The change took 50 days … … in the young of Crax fasciolata raised by Guimaraes, Bergamin, and Carvalho, but the change extended into the third month in one young of C. fasciolata raised by Bronzini (1940).”

Delacour and Amadon (2004) cited Roer (pers. observation) who observed a Wattled Curassow (Crax globulosa) offspring about two months old. Its plumage already looked very much like that of an adult, with the trace of some brownish mottling on its wings. It was only a third of the size of an adult. For the Razor-billed Curassow (Crax mitu), it is reported that the plumage was entirely black (like the adults’) after 65 days from hatching (Heinroth 1931 cited in Delacour and Amadon 2004), similar to information for the Great Curassow (Crax rubra) that immatures look like adults after 3 months (Taibel 1940, Delacour and Amadon 2004). Notes for Alagoas Curassow (Mitu mitu) suggest that at 90 days of age, the offspring curassow is almost indistinguishable from its parents (Nardeli 1981, 1993 cited in del Hoyo and Motis 2004). Drawing the conclusion and simplifying that the appearance of immatures after 2 or 3 months is similar to that of an adult (Vaurie 1968). Since Bare-faced Curassows are similar in body mass and size to both the Wattled Curassow and the Razor-billed Curassow, but less so when compared to the Great Curassow. Drawing the conclusion that the former two should be considered suitable for ontogenetical comparisons within the cracids.

(h) When size of the offspring is over one third of the size of an adult, then offspring is most likely more than 2 months old.

- The size of the offspring is more than one third but less than a half of the size of an adult………………………………...…………………………………...(j) - The size of the offspring is about a half of the size of an adult.……………. (k) - The size of the offspring is more than a half of the size of an adult……..……(l)

Explanation: There are few but strong supports that there are a male and a female offspring which were about 3 to 4 months (90 - 120 days) old when captured (Camera trap location 36; Figure 22).

Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 36 (i.) On the captures from July 2017, few-weeks old offspring are clearly visible. One is recognized as male (uniformly blackish wing feathers) and the other clearly shows a female (blackish wings feathers with narrow white stripes). In October 2017, when they are a half of the size of an adult, one male and one female occur at the same camera trap location. (ii.) The size of a 2-month-old offspring is about one third of the size of an adult one, so clearly those offspring are more than 2 months old. (iii.) Plumage is already adult-like, without any seeable juvenile patterns. (iv.) There is a report for the Razor-billed Curassow (Crax mitu) (Heinroth 1931 cited in Delacour and Amadon 2004) that the crest is less developed at eighty days. The presented image, however, shows a male with a well-developed crest, which provides support to the claim that it is older than 80 days. (v.) It cannot be assumed with confidence that those are the same individuals, but there is strong support that this might be the case due to the same trapping locality and also due to sex-specific features and the size.

Under the assumption of equal sex ratio of offspring at hatching, the possibility that one male and one female will hatch from two eggs is 50%. The same possibility in two separated cases that another pair of offspring with different sexes will occur at the same position is 25%. Under the assumption of unequal sex ratio of offspring at hatching, the possibility would be even lower. Additionally, the possibility would be lower under the assumption that Crax species usually, but not always lay two eggs. However, those are open systems and a conclusion regarding only information about the possibility of occurrence cannot be made, as migration cannot be excluded. But this gives additional support for the conclusion my conclusion (v.).

(i) When the size of immatures is less than a third of the size of an adult, then the offspring is less than two months old...Return and check other characteristics.

(j) When the size of the offspring is more than one third, but less than a half of the size of an adult, then the offspring is between two and three months of age.

(k) When the size of the offspring is half the size of an adult, then offspring is between three and four months of age.

(l) When the size of the offspring is more than a half of the size of an adult, then the offspring is over four months of age.

Figure 21. Plates and photographs of different age stages of different curassow species. Information from Taibel (1940), Delacour and Amadon (2004).

Figure 22. Female offspring, adult female and male offspring of Bare-faced Curassow (Crax fasciolata). Estimated age-class of the offspring is between 3 and 4 months of age.

3.4.3 Results of practical use of Age-class identification key for C. fasciolata offsprings

Out of 65 independent captures of C. fasciolata offspring, 12 (18.46%) were excluded due to the low quality of records (Table 5). Additionally, there were 11 (16.92%) independent captures where the estimation of age was more than 120 days old.

Table 5. List of estimated age-classes for all independent captures of Crax fasciolata offspring obtained during the survey (from July 2015 to December 2017) conducted at Baía das Pedras in the Pantanal of Mato Grosso, Brazil. Sample Capture taken Age estimation Location Placement Sex index Date Time Min Max 1_65 G2 G2_17_2_1 12.06.2017 06:44 M 30 40 2_65 G2 G2_17_2_1 14.06.2017 16:01 M Excluded (evaluation not possible) 3_65 G2 G2_17_2_1 14.06.2017 16:01 M Excluded (evaluation not possible) 4_65 G2 G2_17_2_1 15.06.2017 14:32 M 30 40 5_65 G2 G2_17_2_1 16.06.2017 16:49 M 40 60 6_65 G2 G2_17_2_1 16.06.2017 16:49 M 40 60 7_65 G2 G2_17_2_1 19.06.2017 16:09 M Approximately 60 8_65 G2 G2_17_2_1 21.06.2017 15:44 M Excluded (evaluation not possible) 9_65 G2 G2_17_2_1 24.06.2017 06:58 M 40 60 10_65 G6 G6_16_1_1A 31.03.2016 16:13 M 40 60 11_65 G6 G6_16_1_1A 3.04.2016 06:58 F 40 60 12_65 G6 G6_16_1_1A 3.04.2016 06:58 M 40 60 13_65 G6 G6_16_1_1A 13.04.2016 07:34 M 120< 14_65 G6 G6_16_1_1A 13.04.2016 07:34 F 120< 15_65 G6 G6_16_1_1A 13.04.2016 10:48 F 90 120 16_65 G6 G6_16_1_1A 13.04.2016 13:24 F 90 120 17_65 G6 G6_16_1_1A 15.04.2016 06:49 M Excluded (evaluation not possible) 18_65 G6 G6_16_1_1A 15.04.2016 06:49 F Excluded (evaluation not possible) 19_65 G6 G6_16_1_1A 15.04.2016 11:35 F 60 90 20_65 G6 G6_16_1_1A 15.04.2016 15:03 F 40 60 21_65 G6 G6_16_1_1A 18.04.2016 10:57 F 120< 22_65 G6 G6_16_1_1A 18.04.2016 10:57 F 120< 23_65 G6 G6_16_1_2A 19.04.2016 07:41 M 40 60 24_65 G6 G6_16_1_2A 19.04.2016 07:41 F 40 60 25_65 G6 G6_16_1_2A 19.04.2016 07:41 F 40 60 26_65 G6 G6_16_1_2A 19.04.2016 12:23 F 90 120 27_65 G6 G6_16_1_2A 25.04.2016 07:51 F 120< 28_65 G6 G6_16_1_2A 26.04.2016 11:53 F 120< Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 39

29_65 G6 G6_16_1_2A 26.04.2016 11:53 F 120< 30_65 G6 G6_16_1_2A 27.04.2016 10:04 F 90 120 31_65 G6 G6_16_1_2A 27.04.2016 10:04 F 90 120 32_65 G6 G6_16_1_3B 14.05.2016 09:53 F 120< 33_65 G6 G6_16_1_3B 14.05.2016 09:53 F 120< 34_65 G7 G7_16_1_1 20.05.2016 16:19 F 40 60 35_65 G7 G7_16_1_1 20.05.2016 16:19 F 40 60 36_65 G11 G11_16_1_1 16.05.2016 10:23 F Excluded (evaluation not possible) 37_65 G11 G11_16_1_1 16.05.2016 17:07 F 90 120 38_65 G11 G11_16_1_1 16.05.2016 17:07 M 90 120 39_65 G11 G11_16_1_1 17.05.2016 06:08 F 90 120 40_65 G11 G11_16_1_1 20.05.2016 08:52 F Excluded (evaluation not possible) 41_65 G11 G11_16_1_1 20.05.2016 08:53 M Excluded (evaluation not possible) 42_65 G11 G11_16_1_1 24.05.2016 08:08 F 90 120 43_65 G11 G11_16_1_1 24.05.2016 08:08 M 90 120 44_65 G23 G23_17_1_3 16.10.2017 12:02 F Excluded (evaluation not possible) 45_65 G23 G23_17_1_3 29.10.2017 16.30 U 0 7 46_65 G23 G23_17_1_3 29.10.2017 16.30 U 0 7 47_65 G29 G29_17_2_3 29.10.2017 15:24 F Excluded (evaluation not possible) 48_65 G29 G29_17_2_3 2.11.2017 05:09 F Approximately 60 49_65 G29 G29_17_2_3 4.11.2017 06:27 F Excluded (evaluation not possible) 50_65 G29 G29_17_2_4 16.11.2017 06:02 F Excluded (evaluation not possible) 51_65 G29 G29_17_2_4 21.11.2017 16:37 F 90 120 52_65 G29 G29_17_2_4 22.11.2017 05:46 F Excluded (evaluation not possible) 53_65 G29 G29_17_2_4 22.11.2017 08:42 F 90 120 54_65 G29 G29_17_2_4 24.11.2017 05:04 F 120< 55_65 G29 G29_17_2_4 24.11.2017 05:04 M 120< 56_65 G33 G33_17_2_1 27.06.2017 10:27 F 40 60 57_65 G36 G36_16_1_1 27.05.2016 06:56 F 40 60 58_65 G36 G36_16_1_1 27.05.2016 06:56 F 40 60 59_65 G36 G36_17_1_2 23.07.2017 05:04 M 14 21 60_65 G36 G36_17_1_2 23.07.2017 05:04 F 14 21 61_65 G36 G36_17_1_2 23.07.2017 14:55 M 14 21 62_65 G36 G36_17_1_2 23.07.2017 14:55 F 14 21 63_65 G36 G36_17_1_2 25.07.2017 10:12 U 14 21 64_65 G36 G36_17_1_3 22.10.2017 08:01 M 90 120 65_65 G36 G36_17_1_3 22.10.2017 08:01 F 90 120

3.5 BREEDING SEASON

During different years of the study, field work was carried out in different time periods. The overall sampling effort varied from year to year and from month to month (Figure 23). The offspring were detected at different sampling locations between March and May 2016 but also in June, July, October and November 2017 (Table 5, Table 6, Table 7, Table 8, Figure 23). Although the highest overall sampling effort was conducted in August (936 sampling days), no offspring were detected. In March, the month with the lowest sample effort, offspring were successfully recorded.

1000

900

800

700

600 Sampling effort in 2017 500 Sampling effort in 2016 400 Sampling effort in 2015

300 Offspring presence in 2017 Sampling effort (days) effort Sampling 200 Offspring presence in 2016

100

Figure 23. Sampling effort (y axis) expressed in days per each month (x axis) for years 2015, 2016 and 2017 is presented with stacked bars (expressed with different shades of gray). Offspring presence is noted with the shapes on the x axis. The triangle represents their presence during the surveys in 2016 as the circle presents those in 2017.

3.5.1 Evaluation of breeding season phases considering estimated age-classes

For more detailed information about egg laying, incubation, and hatching intervals per independent capture (see the Appendix C to O). A summary of all estimations per independent capture inside the same survey occasion are graphically presented in Table 6 for the captures from 2016 and in Table 7 for the captures in 2017.

At location G6 the survey occasions were combined as they were consecutive. There were at least three (maybe even four) family groups captured with offspring estimated to be in different ages-classes respectively. They were clearly distinguishable as the offspring of the Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 41 different groups belonging to different age-classes. Age estimation ranged from 40 days to more than 120 days (Table 5). Therefore, the estimated overall egg laying and hatching range interval is large. The estimated egg laying period for the offspring evaluated at G6 is between November and February and the estimated hatching period is between December and March (Table 6 - G6_16_1_1A/2A).

At location G7 there was just one family group captured. Age estimation was between 40 and 56 days (Table 5). Therefore, the estimated egg laying period is between the second half of February and the first half of March as the estimated hatching period is between the second half of March and the first half of April (Table 6 - G7_16_1_1).

At location G11 there was at least one family group captured. Age estimation was between 90 and 120 days (Table 5). Therefore, the estimated egg laying period is between the second half of December and January as the estimated hatching period is between the second half of January and February (Table 6 - G11_16_1_1).

Table 6. Presentation of estimated egg laying and hatching range intervals for Crax fasciolata on a monthly basis relative to the period when the captures were taken for different survey occasions where offspring were detected during the survey at Baía das Pedras in the Pantanal of Mato Grosso, Brazil in year 2016. GRAPHICAL G6_16_1_1A/2A JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC Captures taken X X Egg laying range X X X X X X X Hatching range X X X X X X X G7_16_1_1 JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC Captures taken X Egg laying range X X Hatching range X X G11_16_1_1 JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC Captures taken X Egg laying range X X X Hatching range X X X G36_16_1_1 JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC Captures taken X Egg laying range X X X Hatching range X X X 2016 JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC Captures taken X X X Egg laying range X X X X X X X X X X Hatching range X X X X X X X X X X

Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 42 Location G36 was the most interesting location to evaluate more closely as captures of offspring were taken during three different time periods (May 2016, July 2017, and October 2017).

The age estimation for the captures taken in May 2016 was between 40 and 60 days (Table 5). Therefore, the eggs were laid in March and hatched in April (Table 6 - G36_16_1_1). Only one family group was captured.

The age estimation for the captures taken in July 2017 was between 2 to 3 weeks (Table 5). Therefore, the concluded estimation is that the eggs were laid in the beginning of June and hatched in the beginning of July (Table 6 - G36_17_1_2). The offspring were clearly recognizable between the sexes on most of the captures. One male and one female were recognized within a single family grouping, respectively. There is strong support that there is just one family group captured as all the captures occurred in the range of a few days.

The age estimation for the captures taken in October 2017 was between 90 to 120 days (Table 5). Therefore, the estimated egg laying period is between the second half of May and June as the estimated hatching period is between the second half of June and July (Table 7 - G36_17_1_3). Here it is important to note down that two offspring, one female and one male, were observed within a single family grouping. There is a good implication that this is the same family group as the one captured in July 2017, because the estimated hatching and egg laying intervals overlap among those two positions (Table 7; check and compare the graphical for G36_17_1_2 and G36_17_1_3). Under the assumption that the sex ratio is equal between the genders at hatching, the possibility of capturing another pair structure of one male and one female offspring in two different survey occasions is 25%.

Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 43 Table 7. Presentation of estimated egg laying and hatching range intervals for Crax fasciolata on a monthly basis relative to the period when the captures were taken for different survey occasions where offspring were detected during the survey at Baía das Pedras in the Pantanal of Mato Grosso, Brazil in year 2017. GRAPHICAL G2_17_2_1 JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC Captures taken X X Egg laying range X X X Hatching range X X X G23_17_1_3 JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC Captures taken X Egg laying range X Hatching range X G29_17_2_3 JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC Captures taken X Egg laying range X Hatching range X G29_17_2_4 JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC Captures taken X Egg laying range X X X Hatching range X X X G33_17_2_1 JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC Captures taken X Egg laying range X X X Hatching range X X X G36_17_1_2 JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC Captures taken X Egg laying range X Hatching range X G36_17_1_3 JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC Captures taken X Egg laying range X X X Hatching range X X X 2017 JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC Captures taken X X X X X X Egg laying range X X X X X X X X X X Hatching range X X X X X X X X X X

At location G2 there was at least one family group captured, with estimated offspring age between 30 and 60 days (Table 5). Therefore, the estimated egg laying period is between the second half of March and April and the estimated hatching period is between the second half of April and May (Table 7 – G2_17_2_1).

At location G23 there were two family groups captured but one of them was excluded from further examination because of the low quality of the captures. The age estimation for the offspring from non-excluded captures was between 0 and 7 days (Table 5). Therefore, the concluded estimation is that the eggs were laid at the end of September and hatched at the Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 44 end of October (Table 7 – G23_17_1_3). It is also important to note down that sex determination was not possible for those captures.

At location G29 there were two consecutive survey occasions. (Table 7 – G29_17_2_3 and G29_17_2_4).

The age estimation of offspring for the captures taken in the first survey (between 29th October and 4th November 2017) was approximately 60 days (Table 5). Therefore, I estimated that the eggs were laid at the beginning of August and hatched at the beginning of September. There were at least two different family groups captured, but some of the data was excluded because of the low quality of camera traps data.

In the second survey (captures taken between 16th and 24th November 2017), there were also at least two different family groups captured, but one of them was excluded because the estimated age for the offspring was over 120 days (Table 5). The age estimation of offspring was between 90 and 120 days. Therefore, the estimated egg laying period is between the second half of June and July and the estimated hatching period is between the second half of July and August.

At location G33 there was one family group captured, with estimated offspring age between 40 and 60 days (Table 5). Therefore, the egg laying period was estimated to be between the second half of March and April and the estimated hatching period between the second half of April and May (Table 7 – G33_17_2_1).

3.5.1.1 The results of the evaluation of the breeding season phases considering estimated age-classes

The results of the evaluation of different breeding phases suggesting that the breeding season for Crax fasciolata lasts throughout the year (Table 8).

Table 8. Final presentation of overall egg laying and hatching range intervals estimation for Crax fasciolata on a monthly basis. OVERALL JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC Captures taken x x x x x x x Egg laying range x x x x x x x x x x x Hatching range x x x x x x x x x x x

4 DISCUSSION 4.1 CAMERA TRAPPING

Camera trap studies on the life history of Bare-faced Curassow (Fernández-Duque et al. 2013, Gomes et al. 2018, Laino et al. 2018) and other Crax species (Srbec-Araujo et al. 2012, Lafleur et al. 2014, Alves et al. 2017, Pardo et al. 2017, Pérez-Irineo and Santos- Moreno 2017, Whitworth et al. 2018) have documented the importance of the scientific application of this methodology and presented a great amount of new data on several aspects of their behavior, occurrence, and habitat preferences. Unfortunately, there is no common protocol on the best practice of camera traps applications for cracids, which limits comparisons of the biological and statistical approaches of previous studies.

My grid-based camera trap study provides novel information on many different aspects of the Bare-faced Curassow´s life history, which extends those of Desbiez and Bernardo (2011), Fernández-Duque et al. (2013), Gomes et al. (2018), Zalazar et al. (2018), and Laino et al. (2018), demonstrating the usage of camera traps as a crucial tool for a better understanding of their biology. In addition, the results of my study are especially important since the field work was carried out in the remote and protected areas of the northern Pantanal of Mato Grosso, Brazil, with low human disturbance impacts. Most notably, this is the first extensive research on C. fasciolata to the best of my knowledge.

Despite the evident usefulness of camera trapping as a tool for detailed biological studies, there are some limitations worth noting in my study. The greatest limitation was that the sampling effort varied noticeably between different sampling periods and sampling sites, as a result of inaccessibility of study sites due to flooding. In future studies, sampling effort should be focused on forest and riparian habitats as those are recognized as important “social gathering areas” for C. fasciolata (personal observations, Desbiez and Bernardo 2011, Fernández-Duque et al. 2013), which could help elucidate intraspecific social behaviors.

4.2 ACTIVITY PATTERNS

This subchapter covers the discussion of hypothesis number 1: “Capture success varies between different seasons, periods, and years.”

Overall capture success in my study was higher than described in previous investigations on the Crax genera (Srbec-Araujo et al. 2012, Fernández-Duque et al. 2013, Perez- Irineo and Santos-Moreno 2017, Pardo et al. 2017). Only the investigation of Laino et al. (2018) had a higher capture success. Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 46 The observed activity patterns of Crax fasciolata in this study suggest that overall capture success of individuals (CSind) did vary significantly between dry and wet periods and different seasons. As for the capture success of the species (CSsp), there was no significant difference between periods and seasons. The observed activity patterns also suggest that CSind and CSsp varied significantly between 2016 and 2017.

The highest observed CSind and CSsp rates were during the receding water season. Furthermore, the average of detected individuals per survey occasion was also the highest during the latter season. Higher average values of detected individuals could be the result of higher offspring foraging activity as capture success of offspring individuals occurred mostly with their close-by parents during the receding water season.

Further research is needed to confirm the drivers for differences in activity patterns between different years, periods, and seasons. The difference in sampling effort per sampling site and sampling period needs to be considered. For February and March there is a lack of data, therefore the results presented for the high water season could be misleading. Kattan et al. (2015) already concluded that some cracids’ (Crax daubentoni, Mitu salvini, Penelope perspicax) movements occur as a response to rainfall seasonality or resource availability corresponding with gathering around water sources during the dry season or around seasonally abundant food sources. This is also the case with C. fasciolata, as observed by higher capture rates near rivers (Fernández-Duque et al. 2013). This could explain the changes in capture success and activity patterns as a response to limited food resources. Further studies with the focus on habitat phenology are needed to fully understand the difference between capture success and average number of individuals per sample occasion.

4.2.1 Daily activity patterns

This subchapter covers the discussion of hypothesis number 2: “Daily activity patterns are bimodal.“

The observed daily appearance of Crax fasciolata in my study supports previously observed bimodal distribution of daily activity patterns for Crax fasciolata (del Hoyo and Motis 2004, Fernándes-Duque et al. 2013, Laino et al. 2018) as well as for other Crax species (Crax blumenbachii; Srbek-Araujo et al. 2012, Crax rubra; Pérez-Irineo and Santos-Moreno 2017). The amount of obtained data allowed me to disentangle the data set on smaller units and make further comparisons between different periods, sexes, and most importantly, between independent species captures with offspring presence and all other captures. Daily activity patterns were similar between different periods and sexes. Furthermore, the most notable comparison is the one between daily activity patterns of adults with offspring and other independent species captures. Daily activity patterns did not vary where offspring Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 47 individuals were present or absent. This suggests that the breeding season does not have a significant impact on daily activity patterns.

It is important to outline that the results of daily activity patterns based on two different approaches were compared where possible. Both approaches were defined with a 30-minute interval for capture independence. However, they differed in definition of independent capture: (1) Based on independent records of clearly recognized adult individuals. Offspring were not considered as independent captures under the assumed codependency to parental daily activity patterns. (2) Based on independent species records. Both approaches resulted with similar daily activity patterns, suggesting that there is no difference between periods, sexes or offspring presence / absence.

4.3 SOCIAL ORGANIZATION

This subchapter covers the discussion of hypothesis number 3: “Adult birds form single sex aggregations.”

Studies on the social organization of C. fasciolata are rare. During a line-transect study, Desbiez and Bernardo (2011) recorded larger groups with more than 2 individuals (3 – 12) at 20 different occasions (10% of total records) and observed a potential classical lekking behavior of the species for the first time. No other report on observation of lekking behavior for C. fasciolata was made. Their study was conducted between July 2002 and October 2004 inside a 200 km2 area in the central Pantanal, Brazil (18°59' S, 56°39'). Before that, Kreig and Schumacher (1936, cited in Delacour and Amadon 2004) reported observations of similar size single sex groups (8 – 11) of males in the forests of eastern Paraguay. A line- transect and a camera trap study was carried between October 2010 and July 2012 within Estancia Guaycolec, Argentina (25°54'S, 58°13'W) in the Mirikiná Reserve (11 km2) by Fernándes-Duque et al. (2013). Groups with more than 2 C. fasciolata individuals (3 – 5) were observed at 6 different occasions (32% of total records) while conducting line transects. Camera trapping proofed more than 2 individuals (3 – 5) at 5 different occasions (4% of total records) while individual birds averaged with 1.23 ± 0.59 SE, and those observed while conducting transect observations averaged 2.05 ± 1.28 SE individuals during independent surveys. Zalazar et al. (2017) reported 1 – 6 individuals with an average occurrence of 2.5 ± 1.5 SE while conducting transect surveys in the eastern area of the province of Formosa, northern Argentina, from June to August 2016. In a recent study, Laino et al. (2018) reported larger groups of C. fasciolata with more than 2 individuals (3 – 5) observed at 10 different occasions (8.8% of total records) with an average of 1.45 ± 0.83 SE individuals per survey occasion. Their study was conducted between November 2016 and November 2017 inside a 40 km2 area in the south of Paraguayan Chaco (24°58'S, 57°22'W). Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 48 During my research there was no observation of any behavior which would indicate a potential lekking behavior of C. fasciolata. Moreover, there was no observation of larger single sex aggregations similar to those reported by Kreig and Schumacher (1936, cited in Delacour and Amadon 2004) and Desbiez and Bernardo (2011). The highest observed single sex adult grouping were 2 individuals, suggesting that single sex aggregations are present, but maybe limited to fewer individuals. Therefore, a firm conclusion about larger single sex aggregations of C. fasciolata cannot be made. This aspect of their biology could be underestimated as camera trap detection of single animals, male-female pairs or groups composed from different sexes is higher than single sex pairs or group detections.

The highest observed number of C. fasciolata per single survey occasion, with 4 different individuals, was lower than in the study of Fernándes-Duque et al. (2013) and Laino et al. (2018). Also, very similar average values of all individual captures were obtained as this investigation averaged with 1.55 ± 0.81 SE per survey occasion. However, all those values were higher in the studies conducted by transect observation (Desbiez and Bernardo 2011, Fernándes- Duque et al. 2013, Zalazar et al. 2017) than in camera trap based records (Fernándes- Duque et al. 2013, Laino et al. 2018, this survey 2020). There are also some disadvantages to line transect surveys compared to camera trapping. Male curassows are supposed to be visually and aurally more detectable because of their sexual behavior and a display different than that of behaviorally more tamed females. The latter are more cryptic in coloration which make them even more difficult to depict (Sedaghatkish and Brooks 1999, Delacour and Amadon 2004, Desbiez and Bernardo 2011, Pardo et al. 2017). Therefore, line transect investigations have a higher tendency to deal with male-biased results.

In summary, I could not confirm lekking behavior in male groupings in my camera trap study. In future studies, single sex groupings, polygamy, and lekking behavior should be additionally investigated. The implementation of camera traps on previously or newly identified “meeting points” near streams and rivers should be considered in future studies.

4.3.1 Parental care

This subchapter covers the discussion of hypothesis number 4: “Both parents invest in parental care.”

There is a lack of information on breeding habits and parental care of the Bare-faced Curassow, especially from their natural habitats. However, there are a few anecdotal observations of offspring joining their parents. We know from the observations in captivity that both parents take care of the offspring (Coupe 1966), which is also the case in their natural habitats, where C. fasciolata offspring were observed with parent birds. But such reports are the exception (Bruno et al. 2006, Kirwan 2009). Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 49 The investigations where camera trapping was carried out were so far not as successful at capturing C. fasciolata offspring, despite the captures of adult (paired) individuals being common (Fernández-Duque et al. 2013, Gomes et al. 2018, Zalazar et al. 2018, Laino et al. 2018). This holds for other Curassow species as well. The females of C. rubra were captured together with their offspring on only two different occasions: (1) during the investigation at San Juan-La Selva Biological Corridor in Northeastern Costa Rica between July 2009 and July 2011 (Lafleur et al. 2014); (2) from March 2011 to June 2013 at Los Chimalapas region, southeastern Mexico (Perez- Irineo and Santos-Moreno 2017). On one occasion, two females of C. blumenbachii with two offspring females were captured at Vale Natural Reserve, Linhares, Espírito Santo State, Brazil between June 2005 and October 2008 (Srbec-Araujo et al. 2012). Beirne et al. (2017) reported one Sira curassow (Pauxi koepckeae) offspring with an adult while conducting a study at Sira Communal Reserve, Peruvian Andes between March and September 2015.

In general, the observations of offspring individuals of C. fasciolata or other Curassow species tend to be scarce, even when great effort is put into the data collection. However, in my study such captures were not so rare as reported from previous research. Offspring were captured at least once at 8 out of 37 locations.

Offspring captures associated with adult individuals confirmed the previously reported parental care of both parents (notes from captivity, Coupe 1966). However, capturing offspring with only one parent by a camera trap is more likely to happen than capturing offspring with both parents, since one parent is often further away from the family group and thus undetected by the wildlife camera (personal observations). Following that, there is a great possibility that records collected by camera trapping are biased towards single adults with offspring as such a detectability is higher. Therefore, a higher number of captures where offspring appear with an adult pair represents an even more solid suggestion that parental care from both parents is rather the norm than an exception.

My camera trap based study confirms the parental care for C. fasciolata indicated by several records of offspring led by parent birds.

4.4 SEX RATIO

This subchapter covers the discussion of hypothesis number 5: “No significant difference exists in the numbers of male and female individuals in camera trap captures.”

During my study the observed proportion in sex ratio regarding all independent records of adult individuals (thereafter: Adult Sex Ratio or ASR) showed similar results in capture frequencies between males and females. Therefore, ASR with only a small tendency towards Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 50 male captures was considered as equal rather than male-skewed. The observed proportion in sex ratio regarding all independent records of offspring individuals was significantly biased towards females. This is similar to the observed sex ratio in the study of Crax rubra (Table 4; Fournier and Janik 2008). However, in the case of offspring sex ratio, only offsprings from different age classes were considered, but not hatchlings. Under the assumption of gathering sufficiently large data set, this is still a good approximation of the reality within the population, as the probability for individual capture of either female or male offspring by a camera trap should be similar under the assumption that their sex ratio is equal. Therefore, one could assume that female-skewed sex ratio of offspring could be a norm within the populations of Crax species, if similar results are obtained in future investigations.

Donald (2007) suggests that a skewed sex ratio is common in birds and that it’s most often is male-skewed. He also suggests that a higher female mortality rather than a biased offspring sex ratio is the main driver of male-skewed ASR in birds and that there is no quantitative evidence that ASR of one male to one female represents the norm for the bird species. The identification of possible causes for skewed ASR within the surveyed population is important. It is also important to identify sex ratio within the stable population and set the baseline for future studies. Studies on different Curassow species with a skewed sex ratio were either biased towards females (Martínez-Morales et al. 2009, Pardo et al. 2017) or males (Hill et al. 2008, Srbek-Araujo et al. 2012, Fernández-Duque et al. 2013, Alves et al. 2017, Whitworth et al. 2018, Laino et al. 2018).

Studies on Cozumel Curassows (Crax rubra griscomi) (Martínez-Morales et al. 2009) concluded that increased feral dog population on Cozumel Island, potentially combined with hunting pressure, may represent the main reason for the switch from sex ratio similar to those in our study towards strongly female-skewed one in a single decade (from 1995 to 2005). The female-skewed sex ratio of hatchlings was suggested as cause for biased ASR (Martínez-Morales et al. 2009). Whitworth et al. (2018) concluded that the newly observed male-skewed capture ratio of Great Curassows marks the success of hunting elimitation programs. In addition, these authors suggested that the absence of hunting presents a possibility for mesopredator release which can pray on nesting females.

Nest predation may be one of the causes for higher male-skewed values in ASR as females are more vulnerable to predation when protecting their nests (Alves et al. 2017). During the incubation period, females are leaving the nest only a few times per day, while males stay close to the nest site (Coupe 1966). This observation of female behavior could result in lower female detection rates by camera traps.

Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 51 This investigation was carried out inside a protected area. Anthropogenic impacts were low, people movement was restricted or logistically limited. Hunting and deforestation are not allowed and were not observed during the study period. Also, feral dogs were not detected. Desbiez and Bernardo (2011) concluded that hunting is not considered a major threat for Crax fasciolata in the Pantanal region. The female-male ASR was significantly less inclined towards male-skewed (it resulted almost equal) than in any other study to the best of our knowledge where female-male ASR was considered for Crax species and yielded male- skewed results (Srbek-Araujo et al. 2012, Fernández-Duque et al. 2013, Alves et al. 2017, Whitworth et al. 2018, Laino et al. 2018).

Camera traps are a valuable tool for sex ratio evaluation. Alves et al. (2017) suggested camera trap data as more accurate than transect-derived data while evaluating the sex composition of the population. Existing studies on C. fasciolata (Fernández-Duque et al. 2013, Laino et al. 2018) where the sex ratio was possible to conclude were male-skewed in ASR. Causes were not specified as this was not the focus of those studies. It is not necessarily true that male-skewed ASR represents a norm for C. fasciolata or any other Crax species. Also, if so, it has not yet been determined to what degree the ASR is normally skewed. The investigation of Laino et al. (2018) also resulted with the lowest deviation in sex ratio in comparison with different studies and this investigation (Table 4). A male-skewed ASR is perhaps more of a norm for C. fasciolata, but more detailed field studies are required to confirm this bias.

4.5 AGE-CLASS IDENTIFICATION KEY FOR C. FASCIOLATA OFFSPRING AND BREEDING SEASON EVALUATION

This subchapter covers the discussion of hypothesis number 6: “Breeding occurs throughout the year.”

The main goal of this investigation was to identify the reproductive period of C. fasciolata based on camera trap data. To achieve this goal, an “Age-class identification key for C. fasciolata offspring” was developed (and used for the first time). The main challenge addressed with age-class identification was to go “back in time” and estimate in a temporal context the offspring’s developmental stages (e.g., hatching, incubation, and laying) for each capture on a monthly basis. Summarizing those stages resulted with estimation of the breeding season.

The age-class identification key for C. fasciolata offspring turned out to be a valuable tool to estimate the onset of the reproduction period. My age key was the most accurate for younger offspring up to 60 days old as age-dependent morphological characteristics are more pronounced during this period. Moreover, the age key can be used with limitations for Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 52 offspring whose estimated age is 120 days; thereafter, the advancements of external morphological characteristics toward adolescence do not allow age-class classifications with confidence.

Overall camera trap sampling effort did vary between different locations and months (during the same and between different years of study). There was a particular lack of data on camera trap records for February and March because of low sampling effort. This could bias the results of this investigation as it significantly lowers the possibility of offspring detection. However, the age key is playing a crucial role here, as the offspring’s age estimation enables the identification if the reproductive period is already present during such gap-based captures from the following months. Regardless of the overall lowest sampling effort in March, one detection was made at the end of the month. Moreover, during the highest overall sampling effort in August and the third highest overall sampling effort in September, offspring were not detected. The lack of detection does not necessarily mean that there was lower presence of offspring during this period because there occurred behavioral, environmental and/or possibly human disturbances which caused biases during the investigation that could impact the detectability of the species and its offspring. There is additional support for this assumption as two out of three locations where offspring were detected after September resulted with the estimated age suggesting that they were hatched before or as late as in August. Therefore, they could already be detected by camera traps. But even if the sampling was carried out strongly during these months, the locations where offspring had been previously detected were not sampled during that period. As Crax are considered to be territorial (Zalazar et al. 2018), this could have impacted the presence of aggregations with offspring at other locations. However, it could also explain their later detection when the sampling was carried out at previously not sampled locations, with them already reaching a certain age. In summary, camera trapping is not sufficient enough to determine the breeding season just through observing the offspring's presence without considering their age. In the presented example, the age-class evaluation did take us “back in time” with the estimation that the breeding season also occurs during August and September even if the offspring were not detected during this period.

The reproductive period of Crax fasciolata was successfully identified during this investigation. Offspring of different ages were captured at least once during 7 out of 12 months. The estimation of their age suggests that breeding occurs throughout the year. My study represents an update of available data on the reproduction biology of the elusive C. fasciolata by applying modern methodological tools such as camera traps.

Another possible use of the identification key is to assess the minimal number of different family groups captured during the same survey occasion. This is possible when different captures observed inside the same time period result with different age-class evaluations. In Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 53 future studies, further evaluation of possible environmental aspects which could also affect the activity patterns, offspring detection, and aspects of the breeding season should be taken into consideration. Furthermore, age identification should be improved with special focus on potential habitats selected for nesting and parental care. With some modifications, my age key of offspring ages might be usefully applied and tested for suitability for researching other Crax species. Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 54

5 CONCLUSIONS

Cracids are sensitive to habitat loss and therefore considered a useful bioindicator of habitat quality. Curassows are one of the most threatened taxa within Cracidae. One of them, the Bare-faced Curassow, the focus species of this study, usually occurs in Neotropical lowland forests with a clear preference for riverine areas. In general, the Bare-faced Curassow is terrestrial in habits, especially when foraging, and most active around dawn and dusk (with a bimodal daily activity pattern). Mostly, these birds are sighted either alone or in pairs. Single sex groupings of males are considered to form leks (not confirmed in this study). The breeding season appears to vary among its distribution range, but it is not firmly concluded. The Bare-faced Curassow´s conservation status is “Vulnerable” despite its large range of distribution. In the IUCN Cracid Action Plan, it is listed as High conservation priority. It is still relatively common in the Pantanal of Mato Grosso since anthropogenic impacts, such as habitat destruction and hunting pressures, are low in this region.

This survey was carried out at Parque SESC Baía das Pedras, located in the municipality of Poconé in the Northern Pantanal of Mato Grosso, Brazil (16°29'55"S 56°24'46"W). The area is protected and human activities are limited, thus creating a low impact of disturbance. Such a situation allowed for life history studies of an intact population under natural conditions. I used camera traps in my study on the Bare-faced Curassow, which turned out to be a useful tool for gathering novel information on the species´ activity patterns, sex ratio, social organization, offspring’s age, and the breeding season.

Hypothesis 1: “Capture success varies between different seasons, periods, and years.” Confirmed.

Two different approaches were used for evaluating the data set: capture success of individuals (CSind) and capture success of the species (=CSsp). Both were based on independent captures, modified for the purposes of this investigation. Overall capture success of this study was higher than described in previous investigations on other curassows. There were differences in capture success depending on the year, season and period. However, further research on habitat type preferences is needed to fully understand the local activity movement patterns of the Bare-faced Curassow.

Hypothesis 2: “Daily activity patterns are bimodal.” Confirmed.

A clear bimodal distribution of daily activity patterns is characteristic for Crax fasciolata. My data results suggest that periods, gender, and presence or absence of offspring with their parents’ don´t have an impact on the species’ daily activity patterns. Future investigations Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 55 should address daily activity patterns regarding environment-dependent aspects (temperature, precipitation, and moon phase) and different habitat types.

Hypothesis 3: “Adult birds form single sex aggregations.” Neither confirmed nor disproved.

My study results didn´t provide evidence of larger single sex aggregation or lekking behavior for the Bare-faced Curassow previously suggested in another study. Camera trapping appears to be a less suitable approach for observation of social behavior when considering larger aggregations. However, it should be seriously considered as a great complement to line transect studies while implemented on known “meeting points” near streams and rivers.

Hypothesis 4: “Both parents invest in parental care.” Confirmed.

There is a lack of information about breeding habits and parental care of the Bare-faced Curassow, especially from their natural habitats. Offspring were mostly recorded with both parents, indicating that parental care in Crax fasciolata is the rule. Camera trapping was useful when addressing their parental behavior. Therefore, future investigations should additionally address their breeding behavior throughout camera trap observations and try to enrich scarce data sets available up to this point.

Hypothesis 5: “No significant difference exists in the numbers of male and female individuals in camera trap captures.” Partially confirmed.

Adult sex ratio (ASR) was concluded as equal while offspring sex ratio was distinctly female-skewed. Further research is needed to firmly confirm that this is the rule for Crax fasciolata. However, changes in ASR rates while conducting repeated camera trap observations should be more seriously considered as one of the first indicators that there is some significant change inside the surveyed population.

Hypothesis 6: “Breeding occurs throughout the year.” Confirmed.

The age-class identification key for C. fasciolata offspring was developed and used for the first time. It provides a good tool for a more accurate age assessment and estimation of the reproductive period. My results suggest that breeding for the Bare-faced Curassow in the Baía das Pedras, Pantanal of Mato Grosso, Brazil occurs throughout the whole year.

6 POVZETEK V SLOVENSKEM JEZIKU

Predstavniki kracidov (Cracidae) so zaradi občutljivosti na spremembe v habitatu prepoznani kot indikatorske vrste (del Hoyo 2019). V nalogi smo preučevali gololično hokojko (Crax fasciolata) na območju severnega Pantanala v nižinskih gozdovih, ki so njen značilni habitat (del Hoyo in sod. 2019). Gololična hokojka je prizemna vrsta (Stotz in sod. 1996), zlasti pri iskanju hrane, najbolj aktivna okoli zore in mraka (z bimodalnim vzorcem dnevnih aktivnosti; del Hoyo in Motis 2004, Fernándes-Duque in sod. 2013). Večinoma se ptice zadržujejo posamezno ali v parih (del Hoyo 2019, del Hoyo and Motis 2004, Delacour and Amadon 2004). Desbiez in Bernardo (2011) sta zabeležila pojav enospolnih sestojev samcev, ki nakazuje na potencialno vedenje dvorjenja na t. i. rastišču. Izgleda, da se reproduktivno obdobje gololične hokojke razlikuje glede na širok razpon njene razširjenosti, vendar pa le to ni dokončno določeno (Krieg in Schumacher 1936 – cit. po Delacour in Amadon 2004, Bruno in sod. 2006, Kirwan 2009). Gololična hokojka se po IUCN-ovem rdečem seznamu vrst, kljub njeni široki razširjenosti, uvršča med vrste s statusom ogroženosti "ranljiva" (angl. "Vulnerable"; IUCN 2019a). Na območju Pantanala, Mato Grosso je še vedno razmeroma pogosta, saj so antropogeni vplivi, kot sta uničenje gozdov in lovski pritisk, v tej regiji nizki (Desbiez in Bernardo 2011). To omogoča proučevanje vedenjskih karakteristik vrste v optimalnih naravnih pogojih, ključno za načrtovanje in izvedbo ustreznih ukrepov za ohranjanje.

Naša študija na gololični hokojki temelji na taksonomsko specifičnih podatkih s posebnim poudarkom na izbranih vzorcih eko-etološke življenjske zgodovine. Za dosego ciljev te naloge smo obravnavali naslednje teme (1, 2) in zastavili preverbo naslednjih hipotez (H1- H6):

(1) Ovrednotenje vzorcev dejavnosti, razmerja med spoloma in družbene organizacije. (2) Ovrednotenje starosti potomcev in ocena reproduktivnega obdobja.

H1: Uspešnost metode zajema podatkov s fotopastmi se razlikuje med različnimi sezonami, obdobji in leti. H2: Dnevni vzorci aktivnosti so bimodalni. H3: Odrasle ptice tvorijo enospolne sestoje. H4: Oba starša skrbita za svoje potomce. H5: Na posnetkih fotopasti ni bistvene razlike v številu zabeleženih moških in ženskih posameznikov. H6: Parjenje poteka skozi vse leto.

Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 57 V nadaljevanju so predstavljeni materiali in metode. Raziskava je bila izvedena v parku SESC Baía das Pedras, Poconé, severni Pantanal, Mato Grosso, Brazilija (16°29'55"S 56°24'46"W). Gre za zasebno zavarovano območje s približno 4200 ha, kjer so človeške dejavnosti omejene. Te razmere so omogočile raziskavo življenjske zgodovine nedotaknjene populacije v optimalnih pogojih. To delo je vpeto v program INAU / CO.BRA in obravnava podatke dolgoročnega spremljanja biotske raznolikosti (Pantanal Automated Acoustic Biodiversity Monitoring), ki sem se mu pridružil od oktobra do decembra 2017.

Med julijem 2015 in decembrom 2017 smo z uporabo kamer (tako imenovanih fotopasti) spremljali 37 različnih lokacij. Fotopasti so bile postavljene na podlagi računalniško generiranega območja (regular GRID; Hawth's Analysis Tools for ArcGIS) z razdaljo 1 km med različnimi vzorčnimi lokacijami. Vzorčenje (število dni oz. 24 urnih ciklov aktivne kamere pasti) se je razlikovalo med različnimi vzorčnimi lokacijami, obdobji (suho: april - september, deževno: oktober - marec), sezonami (umikajoča se voda: april - junij, nizka voda: julij - september, naraščajoča voda: oktober - december, visoka voda: januar - marec) in leti (2015, 2016, 2017). Na vsaki lokaciji je neprekinjeno vzorčenje potekalo najmanj 5 dni. Fotopasti so bile nameščene na višini 60 cm.

Zbrane podatke smo obravnavali skozi dva različna pristopa: (1) Uspešnost zajema (= zabeležbe) posnetkov različnih posameznikov (CSind; O'Brien in sod. 2003) in uspešnost zajema vrste (CSsp; Srbec-Araujo in sod. 2012) glede na vzorčenje (z enoto: število zajemov / 100 kamera dni). Oba temeljita na neodvisnih posnetkih s 30-minutnim intervalom in sta modificirana za namene te raziskave. Za analize razlik med sezonskimi in dnevnimi vzorci smo uporabili neparametričen Hi2-test. Za statistično značilne razlike smo privzeli p > 0,05.

Uspešnost zajema posnetkov (CSind in CSsp) smo primerjali med različnimi leti, obdobji in sezonami. Na podlagi združenih podatkov iz vseh treh let smo izvedli primerjave pogostnosti zajema med različnimi kategorijami (samec, samica, par, odrasli s potomstvom in druge oblike združevanj). S tem smo ugotavljali različne sezonske vzorce aktivnosti. Dnevne vzorce aktivnosti smo primerjali med različnimi obdobji (suho, deževno) in spoloma (ženska, moški). Ker je pojav potomcev lahko odvisen od pojavnosti odraslih posameznikov, smo analizirali vzorec dnevne aktivnosti (1) brez in (2) vključno s potomci. Na podlagi podatkov zajema posnetkov različnih posameznikov za obdobje vseh treh let, smo primerjali frekvence zajema med spoloma. In sicer ločeno za odrasle ptice in njihove potomce. S tem smo ugotavljali ali je razmerje med spoloma enakomerno oziroma v prid posameznemu spolu.

Za natančnejšo obravnavo paritvenega obdobja gololične hokojke na podlagi podatkov fotopasti, smo pripravili „Identifikacijski ključ starostnih razredov potomcev C. fasciolata“. Identifikacija starostnega razreda s pomočjo ključa omogoča določitev razvojnih faz Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 58 potomcev, kot so izvalitev, inkubacija in polaganje za vsak posnetek z natančnostjo do enega meseca. Ključ temelji na literaturi, ki je na voljo za različne vrste hokojk (Guimaraes in sod. 1935 - cit. po Vaurie 1968, Krieg in Schumacher 1936 - cit. po Delacour in Amadon 2004, Taibel 1940, Bronzini 1940 - cit. po Vaurie 1968, Bronzini 1943 - cit. po Vaurie 1968, Taibel 1953 - cit. po Vaurie 1968, Vaurie 1968, del Hoyo in Motis 2004, Delacour in Amadon 2004, Roer - cit. po Delacour in Amadon 2004).

V nadeljevanju je za vsako izmed hipotez predstavljen izlušček rezultatov s pripadajočo diskusijo.

Kamere so bile aktivne 4768 dni in noči (= kamera dni). Znotraj neodvisnih 30-minutnih intervalov zajema smo s fotopastmi zajeli 357 opažanj vrste (554 ptic). Gololično hokojko smo pri tem bila zabeleželi na 26 (70,27%) od skupno 37 vzorčnih lokacijah. Število kamera dni se je razlikovalo glede na posamezno lokacijo in na posamezno priložnost vzorčenja. In sicer, kot posledica logistike, razpoložljivosti teresnkih raziskovalcev, dostopnost do vzorčne lokacije in / ali okvaro fotopasti (Priloga A).

Opaženi vzorci aktivnosti gololične hokojke nakazujejo, da se je skupna uspešnost zajema

(= zabeležb) posnetkov posameznikov (CSind) močno razlikovala med suhim in deževnim obdobjem (x2 = 10,44, df = 1, p = 0,001) in med različnimi sezonami (x2 = 35,36, df = 3, P

<0,00001). Kar zadeva uspešnost zajema vrste (CSsp), nismo zaznali statistično značilne razlike med različnimi obdobji (x2 = 2,91, df = 1, p = 0,088) in sezonami (x2 = 7,14, df = 3, P = 0,067). Najvišji stopnji CSind in CSsp smo opazili med sezono umikajoče se vode z 11,62 in 9,39. Poleg tega smo tekom slednje, z 1,82 zabeležili tudi najvišje povprečje zabeleženih posameznikov na neodvisen zajem (30 minutni interval).

Gololične hokojke smo zabeležili znotraj 13 urnega intervala. Z najzgodnejšo zabeležbo ob 05:04 in najkasnejšo ob 17:54 uri. Izrazit vrh dnevnih aktivnosti smo zabeležili med 06:00 in 07:00 ter drugi, nižji od prvega, med 16:00 in 17:00 uro. Najnižje vrednosti dnevnih aktivnosti smo zabeležili med 13:00 in 14:00 uro. Opažena dnevna pojavnost gololične hokojke podpira predhodno opaženo bimodalno razporeditev dnevnih vzorcev aktivnosti za to vrsto (del Hoyo in Motis 2004, Fernándes-Duque et al. 2013, Laino in sod. 2018). Dnevni vzorci aktivnosti so bili med različnimi obdobji in spoli podobni. Prav tako se niso razlikovali med prisotnostjo in odsotnostjo potomcev. Slednje nakazuje, da reproduktivna sezona nima pomembnega vpliva na vzorce dnevnih aktivnosti.

Raziskave socialne organizacije gololične hokojke so redke. Med našo raziskavo nismo zaznali nobenega vedenja, ki bi pri gololični hokojki nakazovalo potencialno vedenje dvorjenja na t.i. rastišču. Poleg tega nismo zaznali večjih enospolnih sestojev (agregatov), podobnih tistim, ki sta jih poročala Kreig in Schumacher (1936 - cit po. Delacour in Amadon Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 59 2004) ter Desbiez in Bernardo (2011). Število posameznikov gololične hokojke glede na neodvisen zajem posnetkov (30-minutni interval) se je gibalo med 1 in 4, s povprečno vrednostjo 1,55 ± 0,81 SE. Glede na neodvisen zajem posnetkov so bili najpogosteje zabeleženi posamezni samci (n = 118; 33,05%), sledila je zabeležba samic (n = 99; 27,73%), nato pari (samec-samica; n = 76; 21.29%), odrasli s potomstvom (n=44; 12.32%) in druge oblike združevanj (n =20; 5.60%).

Evidentirana opazovanja potomcev gololične hokojke in drugih vrst hokojk so redka. Vendar tekom naše študije zabeležbe potomcev niso redkost. Te smo zabeležili vsaj enkrat na osmih (21,62%) od 37 lokacij. Potomce smo večinoma opazili z odraslimi pari (samec-samica; n = 24; 54,55%), sledila so opažanja s posameznimi samicami (n = 13; 29,55%) in samci (n = 7; 15,91%; Slika 18). Primerjave zabeležb, kjer smo zabeležili prisotnost potomcev, s tistimi, kjer potomcev nismo zabeležili, kažejo, da je obstajala razlika med različnimi pojavi (x2 = 20,71, df = 3, P = 0,0001). Z zabeležbami potomcev skupaj z odraslimi pticami smo potrdili starševsko skrb s strani obeh staršev (opombe iz ujetništva, Coupe 1966).

Spol potomcev gololične hokojke smo določili za 62 (95,38%) od 65 neodvisnih zabeležb. Od tega 21 zabeležb (32,31%) samcev in 41 (63,08%) samic. Razmerje med spoloma potomcev je z 0,51: 1,00 izrazito nagnjeno proti samicam. Med odraslimi pticami smo zabeležili 251 (51,33%) samcev in 238 (48,67%) samic, z razmerjem med spoloma odraslih ptic 1,05:1,00. Naši rezultati nakazujejo enakovredno razmerje med spoloma odraslih osebkov. Obstoječe študije gololične hokojke (Fernández-Duque in sod. 2013, Laino in sod. 2018), iz katerih je bilo mogoče zaključiti razmerje med spoloma odraslih ptic na zajetih posnetkih, nakazujejo, da je slednje v prid samcem. Fotopasti so se izkazale kot dragoceno orodje za ocenjevanje razmerja med spoloma. Ocenjevanje spolne sestave s fotopastmi je natančnejše od uporabe transektnih metod (Alves in sod. 2017).

Izmed 65 neodvisnih zabeležb potomcev gololične hokojke smo jih 12 (18,46%) izključili iz nadaljnje obravnave, saj je bila zaradi nizke kakovosti posnetkov identifikacija njihove starosti onemogočena. Dodatno smo izključili 11 (16,92%) zabeležb, pri katerih je bila ocena starosti več kot 120 dni. Napor vzorčenja se je razlikoval med različnimi časovnimi obdobji. Potomci smo zabeležili na različnih lokacijah vzorčenja med marcem in majem 2016, ter v juniju, juliju, oktobru in novembru 2017. „Identifikacijski ključ starostnih razredov potomcev C. fasciolata“ se je izkazal za dragoceno orodje pri oceni začetka obdobja razmnoževanja. Ocena njihove starosti nakazuje, da se reproduktivno obdobje pojavlja skozi vse leto. Naša študija predstavlja posodobitev razpoložljivih podatkov o reprodukcijski biologiji te “skrivnostne” vrste, z uporabo sodobnih metodoloških orodij, kot so fotopasti. Z nekaj spremembami je ključ za določanje starosti mogoče uporabiti in preizkusiti, ali je primeren za aplikacijo na drugih Crax vrstah.

Hipoteza 1: “Uspešnost metode zajema podatkov s fotopastmi se razlikuje med različnimi sezonami, obdobji in leti. ” Potrjena.

Skupni uspeh zajema posnetkov je bil višji, kot je navedeno v raziskavah, opravljenih na drugih vrstah hokojk. Uspeh zajema se razlikuje med različnimi sezonami, obdobji in leti. Vendar pa so potrebne nadaljnje raziskave preferenc habitatnih tipov, preden bomo v celoti razumeli lokalne vzorce gibanja gololične hokojke.

Hipoteza 2: “Dnevni vzorci aktivnosti so bimodalni. ” Potrjena.

Za gololično hokojko je značilna jasna bimodalna porazdelitev vzorcev dnevnih aktivnosti. Rezultati nakazujejo, da različna obdobja, spoli in prisotnost ali odsotnost potomcev ob njihovih starših nimajo vpliva na vzorce dnevnih aktivnosti. Nadaljnje raziskave bi morale naslavljati vzorce dnevnih aktivnosti v odvisnosti od okolijskih dejavnikov (temperatura, padavine, lunine faze) in različnih habitatnih tipov.

Hipoteza 3: “Odrasle ptice tvorijo enospolne sestoje.” Niti potrjena, niti ovržena.

Rezultati niso zagotovili dokaza o tvorbi večjih enospolnih sestojev (agregacij) odraslih ptic, niti vedenja, ki bi pri gololičnih hokojkah nakazovalo na potencialno vedenje dvorjenja na t.i. rastišču, kot je bilo pred tem že predlagano (Desbiez in Bernardo 2011). Zdi se, da je opazovanje s fotopastmi manj primeren pristop za spremljanje socialnega vedenja, kadar opazujemo združevanja v večje skupine. Kljub temu pa bi morali resno razmisliti o tej metodologiji kot dopolnilu transektnim raziskavam med njihovim izvajanjem na znanih "stičiščih" ob rekah in jezerih.

Hipoteza 4: “Oba starša skrbita za svoje potomce.” Potrjena.

V obstoječi literaturi primanjkuje informacij o paritvenih navadah in starševski skrbi za potomce gololične hokojke, še posebej iz njenih naravnih habitatov. Potomci so bili večinoma zabeleženi z obema staršema, kar nakazuje, da je starševska skrb s strani obeh staršev za gololično hokojko pravilo. Uporaba fotopasti se je torej izkazala kot koristna pri obravnavi starševskega vedenja. Zato bi morale prihodnje raziskave dodatno obravnavati njihovo paritveno vedenje s pomočjo fotopasti in tako poskusiti obogatiti redke podatke, ki so nam na voljo do tega trenutka.

Senič M. Camera trap based data analysis of Crax fasciolata life history patterns in the northern Pantanal. Univerza na Primorskem, Fakulteta za matematiko, naravoslovje in informacijske tehnologije, 2020 61 Hipoteza 5: “Na posnetkih fotopasti ni bistvene razlike v številu zabeleženih moških in ženskih posameznikov. ” Delno potrjena.

Razmerje med spoloma odraslih ptic smo obravnavali kot podobno, medtem ko je razmerje med spoloma potomcev izrazito v prid samicam. Za dokončno potrditev, da je takšen vzorec za gololično hokojko pravilo, so potrebne nadaljnje raziskave. Ne glede na to bi morali spremembe vrednosti razmerja med spoloma odraslih ptic med ponavljajočim se opazovanjem s fotopastmi resneje obravnavati kot enega izmed prvih pokazateljev, da je prišlo do izrazitih sprememb znotraj opazovane populacije.

Hipoteza 6: “Parjenje poteka skozi vse leto. ” Potrjena.

Razvili smo „Identifikacijski ključ starostnih razredov potomcev C. fasciolata“ in ga prvič tudi uporabili. Ključ zagotavlja dobro orodje za natančnejšo oceno starosti in oceno paritvenega obdobja. Rezultati nakazujejo, da se na območju raziskave gololične hokojke parijo vse leto.

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Wetlands International 2006. Waterbird Population Estimates 4th edition. Wageningen (Netherlands): Wetlands International.

Whitworth A., Beirne C., Flatt E., Huarcaya R. P., Cruz Diaz C. J., Forsyth A., Molnár P. K., Vargas Soto J. S. 2018. Secundary forest is utilized by Great Curassows (Crax rubra) and Great Tinamous (Tinamus major) in the absence of hunting. The Condor, 120(4):852- 862. American Ornithological Society.

Zalazar S., Benitez A. L., Di Giacomo A. S. 2018. Determining the factors that influence the occurrence of Bare-faced Curassows (Crax fasciolata) in Humid Chaco, northern Argentina. Avian conservation and Ecology. Volume 13, Issue 2, Article 1.

APPENDICES

APPENDIX A Geographic coordinates of sample locations (= sites)

Geographic coodrinate

Location Latitude Longitude G1 16° 29' 58.99"S 56° 24' 35.28"W G2 16° 30' 31.36"S 56° 24' 35.39"W G3 16° 29' 26.05"S 56° 26' 16.12"W G4 16° 29' 27.20"S 56° 25' 09.41"W G5 16° 29' 26.30"S 56° 25' 42.56"W G6 16° 30' 31.36"S 56° 25' 08.09"W G7 16° 29' 58.13"S 56° 26' 16.55"W G8 16° 30' 32.08"S 56° 25' 42.10"W G9 16° 29' 58.45"S 56° 24' 01.48"W G10 16° 29' 59.21"S 56° 25' 08.80"W G11 16° 31' 04.26"S 56° 25' 41.63"W G12 16° 31' 04.30"S 56° 25' 09.01"W G13 16° 30' 31.82"S 56° 24' 01.22"W G14 16° 28' 54.12"S 56° 26' 16.73"W G15 16° 28' 54.80"S 56° 26' 49.74"W G16 16° 29' 59.32"S 56° 25' 42.78"W G17 16° 31' 03.79"S 56° 24' 34.88"W G18 16° 29' 26.56"S 56° 27' 23.87"W G19 16° 31' 37.42"S 56° 23' 59.42"W G20 16° 32' 07.80"S 56° 24' 35.50"W G21 16° 31' 35.83"S 56° 25' 09.05"W G22 16° 31' 03.72"S 56° 26' 16.76"W G23 16° 31' 36.95"S 56° 26' 49.81"W G24 16° 30' 32.08"S 56° 26' 49.38"W G25 16° 30' 31.64"S 56° 27' 24.37"W G26 16° 32' 41.24"S 56° 24' 34.85"W G28 16° 31' 03.76"S 56° 24' 02.52"W G29 16° 29' 56.47"S 56° 27' 27.58"W G30 16° 29' 26.05"S 56° 26' 51.25"W G31 16° 31' 36.55"S 56° 24' 34.60"W G32 16° 31' 36.37"S 56° 26' 16.19"W G33 16° 32' 08.95"S 56° 26' 16.26"W G34 16° 31' 36.01"S 56° 25' 43.10"W G35 16° 29' 59.14"S 56° 26' 49.06"W G36 16° 30' 35.06"S 56° 26' 13.20"W G37 16° 32' 08.05"S 56° 25' 09.05"W G38 16° 32' 09.74"S 56° 25' 42.85"W

APPENDIX B Sampling effort per sample location (= site) and per month

JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC TOTAL G1 17 0 6 15 31 45 31 18 0 0 15 22 200 G2 31 1 7 29 30 15 3 11 14 1 18 26 186 G3 24 0 0 0 32 15 5 45 52 7 28 0 208 G4 24 0 0 0 29 21 6 52 2 0 24 0 158 G5 17 0 0 19 17 18 5 36 0 3 28 0 143 G6 23 0 5 30 31 18 26 19 13 20 14 25 224 G7 18 0 0 0 11 16 9 13 0 0 15 0 82 G8 31 3 0 0 0 15 4 15 0 14 7 18 107 G9 17 0 0 33 30 14 13 42 20 14 30 25 238 G10 14 0 0 28 31 16 6 41 5 0 23 27 191 G11 19 3 0 0 17 15 13 18 16 17 27 14 159 G12 0 0 0 0 0 0 24 6 21 17 0 16 84 G13 29 0 0 21 23 21 23 11 14 12 4 21 179 G14 21 0 0 25 31 18 4 57 30 12 8 0 206 G15 21 0 0 0 19 18 4 31 34 6 25 0 158 G16 19 0 0 0 22 17 1 88 0 16 7 12 182 G17 25 0 0 18 20 6 18 14 26 14 16 21 178 G18 21 0 0 25 31 18 4 57 30 12 25 0 223 G19 0 0 0 0 0 9 21 19 2 14 14 0 79 G20 0 0 0 0 0 0 25 23 2 14 13 1 78 G21 0 0 0 0 0 2 26 27 9 17 0 14 95 G22 0 0 0 0 0 0 15 39 37 30 0 17 138 G23 0 0 0 0 14 0 27 24 0 18 0 0 83 G24 0 0 0 0 0 0 0 49 5 10 6 14 84 G25 0 0 0 0 0 0 0 22 7 10 15 0 54 G26 0 0 0 0 0 0 10 23 2 14 0 0 49 G28 0 0 0 0 0 9 21 19 2 14 0 0 65 G29 13 4 0 0 0 16 28 18 16 6 25 0 126 G30 19 4 0 0 0 16 28 18 11 6 25 0 127 G31 0 0 0 0 0 0 22 14 22 17 0 0 75 G32 0 0 0 0 14 4 20 23 10 18 0 22 111 G33 11 3 0 0 14 8 31 20 0 17 0 0 104 G34 0 0 0 0 0 0 21 6 24 18 0 16 85 G35 20 3 0 0 0 0 0 15 19 3 13 0 73 G36 0 0 0 0 14 0 27 3 17 10 27 17 115 G37 0 0 0 0 0 2 26 0 16 17 0 14 75 G38 0 0 0 0 0 0 14 0 14 18 0 0 46 TOTAL 434 21 18 243 461 372 561 936 492 436 452 342 4768

APPENDIX C Specification sheet for G2_17_2_1 placement

SPECIES Crax fasciolata BREEDING DETAILS LOCATION G2 A simple subtraction of the days estimated in age estimation from the date of the capture taken is sufficient to approximate the hatching. Then we subtract the number of the days of incubation PLACEMENT G2_17_2_1 period from the hatching estimation and we will get the approximation of egg laying. The time in FROM TO between egg laying and hatching is the incubation period. We are not going to determine exact DATE 11.6.2017 26.6.2017 dates, but just the monthly periods. TIME 14:42 08:57 Incubation period of the surveyed species: 30 OFFSPRING AGE ESTIMATED CAPTURE TIMING ESTIMATED ESTIMATED EGG CAPTURES ESTIMATION INCUBATION PERIOD HATCHING RANGE LAYING RANGE - IN DAYS RANGE INDEX - SEX INDEX- PLACE DATE TIME STUDY MENT MIN MAX FROM TO Always in between FROM TO 1_65 1_9 12.6.2017 06:44 M 30 40 3.5.2017 13.5.2017 Apr-May 3.4.2017 13.4.2017 2_65 2_9 14.6.2017 16:01 M 30 40 5.5.2017 15.5.2017 Apr-May 5.4.2017 15.4.2017 3_65 3_9 14.6.2017 16:01 M 30 40 5.5.2017 15.5.2017 Apr-May 5.4.2017 15.4.2017 4_65 4_9 15.6.2017 14:32 M 30 40 6.5.2017 16.5.2017 Apr-May 6.4.2017 16.4.2017 5_65 5_9 16.6.2017 16:49 M 40 60 17.4.2017 7.5.2017 Apr-May 18.3.2017 7.4.2017 6_65 6_9 16.6.2017 16:49 M 40 60 17.4.2017 7.5.2017 Apr-May 18.3.2017 7.4.2017 7_65 7_9 19.6.2017 16:09 M 60 20.4.2017 Mar-Apr 21.3.2017 8_65 8_9 21.6.2017 15:44 M 40 60 22.4.2017 22.5.2017 Mar-Apr 23.3.2017 22.4.2017 9_65 9_9 24.6.2017 06:58 M 40 60 25.4.2017 15.5.2017 Mar-Apr 26.3.2017 15.4.2017 OVERALL 30 60 17.4.2017 17.5.2017 Mar-May 18.3.2017 17.4.2017 NOTES ON PLACEMENT NOTES ON RESULTS In study cases with indexes 1_65 and 4_65, we also made the estimation with the help of picture comparisons in the determination key and not just based on description. Exclusion of study cases with indexes 2_65, 3_65 and 8_65 because offspring are not clearly visible on those captures. The eggs were laid between March and April, and hatched between April and May.

APPENDIX D Specification sheet for G6_16_1_1A placement

SPECIES Crax fasciolata BREEDING DETAILS LOCATION G6 A simple subtraction of the days estimated in age estimation from the date of the capture taken is sufficient to approximate the hatching. Then we subtract the number of the days of incubation PLACEMENT G6_16_1_1A period from the hatching estimation and we will get the approximation of egg laying. The time in FROM TO between egg laying and hatching is the incubation period. We are not going to determine exact DATE 26.3.2016 18.4.2016 dates, but just the monthly periods. TIME 15:50 14:27 Incubation period of the surveyed species: 30 OFFSPRING AGE ESTIMATED CAPTURE TIMING ESTIMATED ESTIMATED EGG CAPTURES ESTIMATION INCUBATION PERIOD HATCHING RANGE LAYING RANGE - IN DAYS RANGE INDEX - SEX INDEX- PLACE DATE TIME STUDY MENT MIN MAX FROM TO Always in between FROM TO 10_65 1_13 31.3.2016 16:13 M 40 60 2.2.2016 22.2.2016 Jan-Feb 3.1.2016 23.1.2016 11_65 2_13 3.4.2016 06:58 F 40 60 3.2.2016 23.2.2016 Jan-Feb 4.1.2016 24.1.2016 12_65 3_13 3.4.2016 06:58 M 40 60 3.2.2016 23.2.2016 Jan-Feb 4.1.2016 24.1.2016 13_65 4_13 13.4.2016 07:34 M 120< 5.12.2015 ? Nov-? 15.11.2015 ? 14_65 5_13 13.4.2016 07:34 F 120< 5.12.2015 ? Nov-? 15.11.2015 ? 15_65 6_13 13.4.2016 10:48 F 90 120 15.12.2015 14.1.2016 Nov-Jan 15.11.2015 15.12.2015 16_65 7_13 13.4.2016 13:24 F 90 120 15.12.2015 14.1.2016 Nov-Jan 15.11.2015 15.12.2015 17_65 8_13 15.4.2016 06:49 M 30 40 6.3.2016 16.3.2016 Feb-Mar 5.2.2016 15.2.2016 18_65 9_13 15.4.2016 06:49 F 30 40 6.3.2016 16.3.2016 Feb-Mar 5.2.2016 15.2.2016 19_65 10_13 15.4.2016 11:35 F 60 90 16.1.2016 15.2.2016 Dec-Feb 17.12.2015 16.1.2016 20_65 11_13 15.4.2016 15:03 F 40 60 17.2.2017 6.3.2016 Jan-Mar 18.1.2016 7.2.2016 21_65 12_13 18.4.2016 10:57 F 120< 23.12.2015 ? Nov-? 23.11.2015 ? 22_65 13_13 18.4.2016 10:57 F 120< 23.12.2015 ? Nov-? 23.11.2015 ? OVERALL 40 120 15.12.2015 6.3.2016 Nov-Mar 15.11.2015 7.2.2016 NOTES ON PLACEMENT NOTES ON RESULTS Exclusion of study cases with indexes 17_65 and 18_65 because offspring are not clearly visible on those captures. Very low quality of some captures. Exclusion of study cases with indexes 13_65, 14_65, 21_65 and 22_65 because it was not possible to estimate age more accurately based on our determination key. Study case 12_65 is not well visible, but as it is in a family aggregation with the study case 11_65, the assumption about the same age therefore is solid. The eggs were laid between November and February and hatched between December and March. From the captures we can estimate that there were at least three family aggregations.

APPENDIX E Specification sheet for G6_16_1_2A placement

SPECIES Crax fasciolata BREEDING DETAILS LOCATION G6 A simple subtraction of the days estimated in age estimation from the date of the capture taken is sufficient to approximate the hatching. Then we subtract the number of the days of PLACEMENT G6_16_1_2A incubation period from the hatching estimation and we will get the approximation of egg FROM TO laying. The time in between egg laying and hatching is the incubation period. We are not going DATE 18.4.2016 4.5.2016 to determine exact dates, but just the monthly periods. TIME 16:49 12:01 Incubation period of the surveyed species: 30 OFFSPRING AGE ESTIMATED CAPTURE TIMING ESTIMATED ESTIMATED EGG CAPTURES ESTIMATION INCUBATION PERIOD HATCHING RANGE LAYING RANGE - IN DAYS RANGE INDEX - SEX INDEX- PLACE DATE TIME STUDY MENT MIN MAX FROM TO Always in between FROM TO 23_65 1_9 19.4.2016 07:41 M 40 60 19.2.2016 10.3.2016 Jan-Mar 20.1.2016 9.2.2016 24_65 2_9 19.4.2016 07:41 F 40 60 19.2.2016 10.3.2016 Jan-Mar 20.1.2016 9.2.2016 25_65 3_9 19.4.2016 07:41 F 40 60 19.2.2016 10.3.2016 Jan-Mar 20.1.2016 9.2.2016 26_65 4_9 19.4.2016 12:23 F 90 120 21.12.2015 20.1.2016 Nov-Jan 21.11.2015 21.12.2015 27_65 5_9 25.4.2016 07:51 F 120< 27.12.2015 ? Nov-? 27.11.2015 ? 28_65 6_9 26.4.2016 11:53 F 120< 28.12.2015 ? Nov-? 28.11.2015 ? 29_65 7_9 26.4.2016 11:53 F 120< 28.12.2015 ? Nov-? 28.11.2015 ? 30_65 8_9 27.4.2016 10:04 F 90 120 29.12.2015 28.1.2016 Nov-Jan 29.11.2015 29.12.2015 31_65 9_9 27.4.2016 10:04 F 90 120 29.12.2015 28.1.2016 Nov-Jan 29.11.2015 29.12.2015 OVERALL 40 120 21.12.2015 10.3.2016 Nov-Mar 21.11.2015 9.2.2016 NOTES ON PLACEMENT NOTES ON RESULTS Exclusion of study case with index 27_65 because it was not possible to estimate age more accurately. Exclusion of study cases with indexes 28_65 and 29_65 because offspring are not clearly visible on those captures. Study cases 24_65 and 25-65 are not well visible, but as they are in a family aggregation with study case 23_65 the assumption about the same age therefore is solid. The eggs were laid between November and February and hatched between December and March. From the captures we can estimate that there were at least three family aggregations.

APPENDIX F Specification sheet for G6_16_1_3B placement

32_65 1_2 14.5.2016 09:53 F 120< ? 16.1.2016 ? Dec-? 17.12.2015 ? 33_65 2_2 14.5.2016 09:53 F 120< ? 16.1.2016 ? Dec-? 17.12.2015 ? OVERALL 120< ? 16.1.2016 ? Dec-? 17.12.2015 ? NOTES ON PLACEMENT NOTES ON RESULTS

Exclusion of study cases with indexes 32_65 and 33_63 because it was not possible to estimate age more accurately. In this case we cannot conclude the age of captured animals.

APPENDIX G Specification sheet for G7_16_1_1 placement

SPECIES Crax fasciolata BREEDING DETAILS LOCATION G7 A simple subtraction of the days estimated in age estimation from the date of the capture taken is sufficient to approximate the hatching. Then we subtract the number of the days of incubation PLACEMENT G7_16_1_1 period from the hatching estimation and we will get the approximation of egg laying. The time in FROM TO between egg laying and hatching is the incubation period. We are not going to determine exact DATE 20.5.2016 3.6.2016 dates, but just the monthly periods. TIME 11:43 12:00 Incubation period of the surveyed species: 30 OFFSPRING AGE ESTIMATED CAPTURE TIMING ESTIMATED ESTIMATED EGG CAPTURES ESTIMATION INCUBATION PERIOD HATCHING RANGE LAYING RANGE - IN DAYS RANGE INDEX - SEX INDEX- PLACE DATE TIME STUDY MENT MIN MAX FROM TO Always in between FROM TO

34_65 1_2 20.5.2016 16:19 F 40 60 21.3.2016 10.4.2016 Feb-Apr 20.2.2016 11.3.2016 35_65 2_2 20.5.2016 16:19 F 40 60 21.3.2016 10.4.2016 Feb-Apr 20.2.2016 11.3.2016 OVERALL 40 60 21.3.2016 10.4.2016 Feb-Apr 20.2.2016 11.3.2016 NOTES ON PLACEMENT NOTES ON RESULTS

The eggs were laid between February and March and hatched between March and April.

APPENDIX H Specification sheet for G11_16_1_1 placement

SPECIES Crax fasciolata BREEDING DETAILS LOCATION G11 A simple subtraction of the days estimated in age estimation from the date of the capture taken is sufficient to approximate the hatching. Then we subtract the number of the days of incubation PLACEMENT G11_16_1_1 period from the hatching estimation and we will get the approximation of egg laying. The time in FROM TO between egg laying and hatching is the incubation period. We are not going to determine exact DATE 9.5.2016 26.5.2016 dates, but just the monthly periods. TIME 09:23 06:52 Incubation period of the surveyed species: 30 OFFSPRING AGE ESTIMATED CAPTURE TIMING ESTIMATED ESTIMATED EGG CAPTURES ESTIMATION INCUBATION PERIOD HATCHING RANGE LAYING RANGE - IN DAYS RANGE INDEX - SEX INDEX- PLACE DATE TIME STUDY MENT MIN MAX FROM TO Always in between FROM TO

36_65 1_8 16.5.2016 10:23 F 60 90 16.2.2016 17.3.2016 Jan-Mar 17.1.2016 16.2.2016 37_65 2_8 16.5.2016 17:07 F 90 120 17.1.2016 16.2.2016 Dec-Feb 18.12.2015 17.1.2016 38_65 3_8 16.5.2016 17:07 M 90 120 17.1.2016 16.2.2016 Dec-Feb 18.12.2015 17.1.2016 39_65 4_8 17.5.2016 06:08 F 90 120 18.1.2016 17.2.2016 Dec-Feb 19.12.2015 18.1.2016 40_65 5_8 20.5.2016 08:52 F 90 120 21.1.2016 21.3.2.016 Dec-Mar 22.12.2015 20.2.2016 41_65 6_8 20.5.2016 08:53 M 90 120 21.1.2016 21.3.2016 Dec-Mar 22.12.2015 20.2.2016 42_65 7_8 24.5.2016 08:08 F 90 120 25.1.2016 24.2.2016 Dec-Jan 26.12.2015 25.1.2016 43_65 8_8 24.5.2016 08:08 M 90 120 25.1.2016 24.2.2016 Dec-Jan 26.12.2015 25.1.2016 OVERALL 60 120 17.1.2016 17.3.2016 Dec-Mar 18.12.2015 16.2.2016 NOTES ON PLACEMENT NOTES ON RESULTS Exclusion of study cases with indexes 40_65 and 41_65 because it was not possible to estimate age more accurately. The eggs were laid between December and February and hatched between January and March.

APPENDIX I Specification sheet for G23_17_1_3 placement

SPECIES Crax fasciolata BREEDING DETAILS LOCATION G23 A simple subtraction of the days estimated in age estimation from the date of the capture taken is sufficient to approximate the hatching. Then we subtract the number of the days of incubation period PLACEMENT G23_17_1_3 from the hatching estimation and we will get the approximation of egg laying. The time in between egg FROM TO laying and hatching is the incubation period. We are not going to determine exact dates, but just the DATE 12.10.2017 30.10.2017 monthly periods. TIME 13:09 10:47 Incubation period of the surveyed species: 30 OFFSPRING AGE ESTIMATED CAPTURE TIMING ESTIMATED ESTIMATED EGG CAPTURES ESTIMATION INCUBATION PERIOD HATCHING RANGE LAYING RANGE - IN DAYS RANGE INDEX - SEX INDEX- PLACE DATE TIME STUDY MENT MIN MAX FROM TO Always in between FROM TO

44_65 1_3 16.10.2017 12:02 F 60 120 18.6.2017 17.8.2017 May-Aug 19.5.2017 18.7.2017 45_65 2_3 29.10.2017 16.30 U 0 7 22.10.2017 29.10.2017 Sep-Oct 23.9.2017 30.9.2017 46_65 3_3 29.10.2017 16.30 U 0 7 22.10.2017 29.10.2017 Sep-Oct 23.9.2017 30.9.2017 OVERALL 0 7 22.10.2017 29.10.2017 Sep-Oct 23.9.2017 30.9.2017 NOTES ON PLACEMENT NOTES ON RESULTS Exclusion of study case with index 44_65 because it was not possible to estimate age more accurately. The eggs were laid in September and hatched in October.

APPENDIX J Specification sheet for G29_17_2_3 placement

47_65 1_3 29.10.2017 15:24 F 90 120 31.7.2017 30.8.2017 Jul-Aug 1.7.2017 31.7.2017 48_65 2_3 2.11.2017 05:09 F 60 3.9.2017 Aug-Sep 4.8.2017 49_65 3_3 4.11.2017 06:27 F 90 120 6.8.2017 5.9.2017 Jul-Aug 7.7.2017 6.8.2017 OVERALL 60 3.9.2017 Sep-Oct 4.8.2017 NOTES ON PLACEMENT NOTES ON RESULTS

Exclusion of study cases with indexes 47_65 and 49_65 because offspring are not clearly visible on those captures. The eggs were laid in August and hatched in September.

APPENDIX K Specification sheet for G29_17_2_4 placement

SPECIES Crax fasciolata BREEDING DETAILS LOCATION G29 A simple subtraction of the days estimated in age estimation from the date of the capture taken is sufficient to approximate the hatching. Then we subtract the number of the days of incubation period PLACEMENT G29_17_2_4 from the hatching estimation and we will get the approximation of egg laying. The time in between FROM TO egg laying and hatching is the incubation period. We are not going to determine exact dates, but just DATE 8.11.2017 25.11.2017 the monthly periods. TIME 10:35 11:56 Incubation period of the surveyed species: 30 OFFSPRING AGE ESTIMATED CAPTURE TIMING ESTIMATED ESTIMATED EGG CAPTURES ESTIMATION INCUBATION PERIOD HATCHING RANGE LAYING RANGE - IN DAYS RANGE INDEX - SEX INDEX- PLACE DATE TIME STUDY MENT MIN MAX FROM TO Always in between FROM TO 50_65 1_6 16.11.2017 06:02 F 90< ? 18.8.2017 ? Jul-? 19.7.2017 ? 51_65 2_6 21.11.2017 16:37 F 90 120 24.7.2017 23.8.2017 Jun-Aug 24.6.2017 24.7.2017 52_65 3_6 22.11.2017 05:46 F 90 120 25.7.2017 24.8.2017 Jun-Aug 24.6.2017 24.7.2017 53_65 4_6 22.11.2017 08:42 F 90 120 25.7.2017 24.8.2017 Jun-Aug 25.6.2017 25.7.2017 54_65 5_6 24.11.2017 05:04 F 120< ? 27.7.2017 ? Jun-? 27.6.2017 ? 55_65 6_6 24.11.2017 05:04 M 120< ? 27.7.2017 ? Jun-? 27.6.2017 ? OVERALL 90 120 24.7.2017 24.8.2017 Jun-Aug 24.6.2017 25.7.2017 NOTES ON PLACEMENT NOTES ON RESULTS Exclusion of study cases with indexes 50_65 and 52_65 because offspring are not clearly visible on those captures to make a solid conclusions. Exclusion of study cases with indexes 54_65 and 55_65 because it was not possible to estimate age more accurately. The eggs were laid between June and July and hatched between July and August. From the captures we can estimate that there were at least two family aggregations.

APPENDIX L Specification sheet for G33_17_2_1 placement

SPECIES Crax fasciolata BREEDING DETAILS LOCATION G33 A simple subtraction of the days estimated in age estimation from the date of the capture taken is sufficient to approximate the hatching. Then we subtract the number of the days of incubation period PLACEMENT G33_17_2_1 from the hatching estimation and we will get the approximation of egg laying. The time in between FROM TO egg laying and hatching is the incubation period. We are not going to determine exact dates, but just DATE 26.6.2017 11.7.2017 the monthly periods. TIME 14:13 14:04 Incubation period of the surveyed species: 30 OFFSPRING AGE ESTIMATED CAPTURE TIMING ESTIMATED ESTIMATED EGG CAPTURES ESTIMATION INCUBATION PERIOD HATCHING RANGE LAYING RANGE - IN DAYS RANGE INDEX - SEX INDEX- PLACE DATE TIME STUDY MENT MIN MAX FROM TO Always in between FROM TO

56_65 1_1 27.6.2017 10:27 F 40 60 28.4.2017 18.5.2017 Mar-May 29.3.2017 18.4.2017 OVERALL 40 60 28.4.2017 18.5.2017 Mar-May 29.3.2017 18.4.2017 NOTES ON PLACEMENT NOTES ON RESULTS The eggs were laid between March and April and hatched between April and May.

APPENDIX M Specification sheet for G36_16_1_1 placement

SPECIES Crax fasciolata BREEDING DETAILS LOCATION G36 A simple subtraction of the days estimated in age estimation from the date of the capture taken is sufficient to approximate the hatching. Then we subtract the number of the days of incubation PLACEMENT G36_16_1_1 period from the hatching estimation and we will get the approximation of egg laying. The time in FROM TO between egg laying and hatching is the incubation period. We are not going to determine exact DATE 17.5.2016 31.5.2016 dates, but just the monthly periods. TIME 12:22 11:51 Incubation period of the surveyed species: 30 OFFSPRING AGE ESTIMATED CAPTURE TIMING ESTIMATED ESTIMATED EGG CAPTURES ESTIMATION INCUBATION PERIOD HATCHING RANGE LAYING RANGE - IN DAYS RANGE INDEX - SEX INDEX- PLACE DATE TIME STUDY MENT MIN MAX FROM TO Always in between FROM TO

57_65 1_2 27.5.2016 06:56 F 40 60 28.3.2016 18.4.2016 Mar-Apr 29.2.2016 18.3.2016 58_65 2_2 27.5.2016 06:56 F 40 60 28.3.2016 18.4.2016 Mar-Apr 29.2.2016 18.3.2016 OVERALL 40 60 28.3.2016 18.4.2016 Mar-Apr 29.2.2016 18.3.2016 NOTES ON PLACEMENT NOTES ON RESULTS The eggs were laid in March and hatched in April.

APPENDIX N Specification sheet for G36_17_1_2 placement

SPECIES Crax fasciolata BREEDING DETAILS LOCATION G36 A simple subtraction of the days estimated in age estimation from the date of the capture taken is sufficient to approximate the hatching. Then we subtract the number of the days of incubation period PLACEMENT G36_17_1_2 from the hatching estimation and we will get the approximation of egg laying. The time in between FROM TO egg laying and hatching is the incubation period. We are not going to determine exact dates, but just DATE 20.7.2017 3.8.2017 the monthly periods. TIME 10:54 08:00 Incubation period of the surveyed species: 30 OFFSPRING AGE ESTIMATED CAPTURE TIMING ESTIMATED ESTIMATED EGG CAPTURES ESTIMATION INCUBATION PERIOD HATCHING RANGE LAYING RANGE - IN DAYS RANGE INDEX - SEX INDEX- PLACE DATE TIME STUDY MENT MIN MAX FROM TO Always in between FROM TO

59_65 1_5 23.7.2017 05:04 M 14 21 2.7.2017 9.7.2017 Jun - Jul 2.6.2017 9.6.2017 60_65 2_5 23.7.2017 05:04 F 14 21 2.7.2017 9.7.2017 Jun - Jul 2.6.2017 9.6.2017 61_65 3_5 23.7.2017 14:55 M 14 21 2.7.2017 9.7.2017 Jun - Jul 2.6.2017 9.6.2017 62_65 4_5 23.7.2017 14:55 F 14 21 2.7.2017 9.7.2017 Jun - Jul 2.6.2017 9.6.2017 63_65 5_5 25.7.2017 10:12 U 14 21 4.7.2017 11.7.2017 Jun - Jul 4.6.2017 11.6.2017 OVERALL 14 21 4.7.2017 11.7.2017 Jun - Jul 4.6.2017 11.6.2017 NOTES ON PLACEMENT NOTES ON RESULTS The eggs were laid in June and hatched in July.

APPENDIX O Specification sheet for G36_17_1_3 placement

SPECIES Crax fasciolata BREEDING DETAILS LOCATION G36 A simple subtraction of the days estimated in age estimation from the date of the capture taken is sufficient to approximate the hatching. Then we subtract the number of the days of incubation period PLACEMENT G36_17_1_3 from the hatching estimation and we will get the approximation of egg laying. The time in between egg FROM TO laying and hatching is the incubation period. We are not going to determine exact dates, but just the DATE 21.10.2017 6.11.2017 monthly periods. TIME 13:58 12:14 Incubation period of the surveyed species: 30 OFFSPRING AGE ESTIMATED CAPTURE TIMING ESTIMATED ESTIMATED EGG CAPTURES ESTIMATION INCUBATION PERIOD HATCHING RANGE LAYING RANGE - IN DAYS RANGE INDEX - SEX INDEX- PLACE DATE TIME STUDY MENT MIN MAX FROM TO Always in between FROM TO

64_65 1_2 22.10.2017 08:01 M 90 120 24.6.2017 24.7.2017 May-Jul 25.5.2017 24.6.2017 65_65 2_2 22.10.2017 08:01 F 90 120 24.6.2017 24.7.2017 May-Jul 25.5.2017 24.6.2017 OVERALL 90 120 24.6.2017 24.7.2017 May-Jul 25.5.2017 24.6.2017 NOTES ON PLACEMENT NOTES ON RESULTS The eggs were laid between May and June and hatched between June and July.