Quaternary fossil vertebrates from continental Portugal: … Lopez... · 2019. 7. 10. · para a detecção de espécies vivas, resultando em que 38% dos tetrápodes terrestres vivos

Darío Estraviz López

Graduado em Biología

Quaternary fossil vertebrates from continental

Portugal: Paleobiodiversity, revision of

specimens and new localities

Dissertação para obtenção do Grau de Mestre em

Paleontologia

Orientador: Prof. Associado Octávio Mateus, Faculdade de Ciências e

Tecnologia da Universidade Nova de Lisboa

Júri:

Presidente: Prof. Auxiliar Paulo Alexandre Rodrigues Roque Legoinha (FCT-

UNL)

Arguente: Prof. Catedrático João Luís Cunha Cardoso (Universidade Aberta)

Vogal: Prof. Associado Octávio João Madeira Mateus (FCT-UNL)

Júri:

Junho 2019

i

Darío Estraviz López Graduado em Biología

Quaternary fossil vertebrates

from continental Portugal:

Paleobiodiversity, revision of

specimens and new localities

Dissertação para obtenção do Grau de Mestre em

Paleontologia

Orientador: Prof. Associado Octávio Mateus, Faculdade de

Ciências e Tecnologia da Universidade Nova de Lisboa

© Darío Estraviz López, FCT/UNL e UNL

A Faculdade de Ciências e Tecnologia e a Universidade Nova de Lisboa tem o direito, perpétuo e sem

limites geográficos, de arquivar e publicar esta dissertação através de exemplares impressos

reproduzidos em papel ou de forma digital, ou por qualquer outro meio conhecido ou que venha a ser

inventado, e de a divulgar através de repositórios científicos e de admitir a sua cópia e distribuição

com objetivos educacionais ou de investigação, não comerciais, desde que seja dado crédito ao autor e

editor.

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“In vertebrate paleontology, increasing knowledge leads to triumphant loss of clarity.”

Alfred Romer

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ACKNOWLEDGEMENTS

It is a strange feeling to finally write this. I have been working on this master thesis for one and a half

years and at the same time that I am happy about finishing it I feel like this is the end of an era: the end

of the straight road that took me from the preschool to this master, always with paleontology in mind. I

could have never reached this stage without the people that have accompanied me and for them are

these words:

First I have to say thanks to my family. This dream of mine is only possible because of their tireless

support. Thanks to my grandparents (Mariví, Josefa and Nicolás). Infinite thanks to my father Antonio

for understanding my passion for paleontology and caring for me while I am in other country, if the

computer where I am writing this is still working is thanks to you. Infinite thanks to my mother Silvia

and my stepfather Suso, I do not have enough words to tell how much I appreciate that you come

every few months to help me at my home in Portugal. All of you are the best family of the world and I

am so lucky of having you by my side. You all make me happy in every possible way.

I want to thank my advisor professor Octávio Mateus. He has given me an immense freedom to follow

the research path that I felt that was the most appropriate for each topic. The freedom to do my own

failures (and his help to get out of them) has made me a better scientist.

I would like to say thanks to Miguel Moreno Azanza, that have really acted a lot of time as a de facto

co-advisor, even when he had to take care of his real students. I won´t forget it. Also I would like to

thank Eduardo Puértolas Pascual for his comments on the thesis and his help to overcome concrete

problems during it.

Thanks to my fellow master students at the Lourinhã PaleoTeam for their companionship and support,

Hugo Campos (And his family that opened me their house in Algarve), João Pratas, André Saleiro,

Cátia Ribeiro, Francisco Costa, Vincent Cheng and Alexandra Fernandes; especially to my classmates

of the fifth master edition Filippo Maria Rotatoria and Alexandre Guillaume. I won´t forget the

dinners on Thursday nights in Almada!

Big thanks to the personnel of the Museum da Lourinhã Alexandre Audigane, Bruno Pereira, Carla

Alexandra Tomás, Carla Abreu, Jorge Beja and Raquel Furtado. Thanks for letting me use the

museum facilities and also for the paraffin heater to prevented me to freeze during winter while

working.

Thanks to all the other people that stood by my side in Lourinhã: Isabel Mateus, Marta Mateus, Simão

Mateus, Silvia Batista, Ana Luz, Micael Martinho, João Marinheiro, João Russo, Marco Marzola, João

Muchagata, Marta, Isabel and all the English class, my training partners at “Miga fight team” and

everyone else; you all complete my “second family” in Portugal and you make me feel at home in a

foreign country.

But this thesis couldn´t have been done without many people that work outside Lourinhã.

I would like to thank the entire GPS (Grupo Proteção Sicó) especially to Sérgio Medeiros and all the

other speleologists that have helped me several times to descend to the Algar do Vale da Pena, for

their disinterested help. I owe you my life and that is not an exaggeration!

Thanks to my former advisor at University of A Coruña, Aurora Grandal d´Anglade, that has helped

me immensely during the process of doing the chemical analysis of my samples. I promise I will get

something interesting for ancient DNA soon.

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Many thanks to the paleontologist that helped me with the identification of the fauna of Santa

Margarida: Pedro Callapez, Raquel Moyá Costa, Pavlos Piskoulis and Juan Manuel López García. I

really hope to be able to keep working with you in further studies about this locality in the future.

Thanks to Miguel Ramalho, Jorge Sequeira and all the personnel of the Geological Museum in Lisbon,

for letting me to visit their collections on short notice twice. Thanks to Jorge Graça, the Algarvian

photographer that contacted Octávio Mateus and discovered the Santa Margarida locality.

Also thanks to Alexandra Elbakyan, without her work I could have never done this thesis.

Finally thanks to the students of other editions, personnel of the Department of Earth Sciences of the

FCT-UNL and everyone that has helped me one way or the other and could not see his/her name on

this thesis, I thank you a lot.

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ABSTRACT

The Quaternary fossil record of Portugal is important for our understanding of the paleobiodiversity in

Iberia. In the present master thesis a series of studies augment our knowledge about this topic.

A census of Quaternary paleobiodiversity is carried out in order to test how reliable the fossil record is

for detecting living species, resulting in that ~38% of living terrestrial tetrapods are recognized in the

fossil record for Portugal, although the number of species recognized varies between groups.

The body mass of a Portuguese proboscidean (Palaeoloxodon antiquus) is calculated via numerical

methods for the first time (11metric tons) and morphometric comparisons of this species with

Mammuthus primigenius are presented using an extensive Proboscidean sample.

A new fossil brown bear (Ursus arctos) locality, Algar do Vale da Pena, with numerous claw mark in

the walls of the cave (the first of this type of marks described in Portugal) is presented and the fossil

bear remains identified and compared to a sample from NW Spain. The bears from Algar do Vale da

Pena contrast with other previously known Portuguese brown bear specimens by relative small size.

A new microvertebrate locality from Algarve, Santa Margarida, is presented. It is an extraordinary rich

site with one fossil for every two grams of sediment selected and processed. The locality provided the

first record of two arvicoline taxa in Portugal (Iberomys huescarensis and Victoriamys chalinei),

which allows giving a minimal age of around 800.000 YBP for at least part of it. This makes Santa

Margarida one of the oldest three localities in the Pleistocene of Portugal.

KEYWORDS

Holocene, Pleistocene, Mammalia, mammals, cave deposits, paleontology

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RESUMO

O registro fóssil Quaternário de Portugal é importante para nossa compreensão da paleobiodiversidade

na Ibéria. Na presente tese de mestrado, uma série de estudos aumenta o nosso conhecimento sobre a

matéria.

Um censo de paleobiodiversidade do Quaternário é realizado para testar a fiabilidade do registro fóssil

para a detecção de espécies vivas, resultando em que 38% dos tetrápodes terrestres vivos são

reconhecidos no registro fóssil de Portugal, ainda que este número varie entre grupos.

A massa corporal de um proboscídeo português (Palaeoloxodon antiquus) é calculada através de

métodos numéricos pela primeira vez (11 toneladas) e comparações morfométricas desta espécie com

Mammuthus primigenius são apresentadas, utilizando uma extensive amostra de proboscídeos.

É apresentada uma nova localidade de urso pardo (Ursus arctos); o Algar do Vale da Pena, com

numerosas marcas de garras nas paredes da gruta (as primeiras deste tipo de marcas descritas em

Portugal) e os restos fósseis de ursos são identificados e comparados com uma amostra procedente do

Noroeste de Espanha. Os ursos do Algar do Vale da Pena contrastam com outros exemplares

portugueses fósseis de urso pardo pelo seu tamanho relativamente pequeno.

Uma nova localidade de microvertebrados do Algarve, Santa Margarida, é apresentada. É um local

extraordinariamente rico, com um fóssil por cada dois gramas de sedimentos selecionados e

processados. A localidade forneceu o primeiro registro de dois taxa de Arvicolinae em Portugal

(Iberomys huescarensis e Victoriamys chalinei), o que permite determinar uma idade mínima de cerca

de 800.000 YBP para, pelo menos, parte da jazida. Isto faz de Santa Margarida uma das três

localidades mais antigas do Plistocénico de Portugal.

PALAVRAS-CHAVE

Holocénico, Plistocénico, Mammalia, mamíferos, depósitos de grutas, paleontologia.

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RESUMO GRÁFICO | GRAPHICAL ABSTRACT

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INDEX OF MATERIALS

1. INTRODUCTION ............................................................................................................................... 1

1.1 History of the study of Quaternary fossil vertebrates from continental Portugal .............................. 1

1.2 Vertebrates in the Quaternary of Portugal ......................................................................................... 7

1.3 Aim, context and objectives of the present thesis............................................................................ 22

2. PALEOBIODIVERSITY OF THE QUATERNARY FOSSIL TETRAPODS IN CONTINENTAL

PORTUGAL .......................................................................................................................................... 23

2.1 Introduction and methods ................................................................................................................ 23

2.2 Lists of localities and fossil tetrapods of the Quaternary of Portugal .............................................. 23

2.3 Results ............................................................................................................................................. 36

2.4 Discussion ....................................................................................................................................... 41

2.5 Conclusions ..................................................................................................................................... 42

3.THE PALAEOLOXODON ANTIQUUS OF SANTO ANTÃO DO TOJAL ................................... 43

3.1 Introduction and methods ................................................................................................................ 43

3.2 Geographical and geological settings .............................................................................................. 43

3.3 The material ..................................................................................................................................... 44

3.4 Measurements of femur and tibia .................................................................................................... 48

3.5 Body mass estimation ...................................................................................................................... 52

3.6 Analysis of metrical data ................................................................................................................. 54

3.7 Conclusions ..................................................................................................................................... 66

4. ALGAR DO VALE DA PENA: A NEW FOSSIL BEAR SITE FROM PORTUGAL .................... 67

4.1 Overview ......................................................................................................................................... 67

4.2 Introduction ..................................................................................................................................... 67

4.3 Methods, expeditions to the cave and preparation of the material .................................................. 73

4.4 The bone assemblages ..................................................................................................................... 77

4.5 Results: The recovered bone remains .............................................................................................. 80

4.6 The claw marks.............................................................................................................................. 118

4.7 Discussion ..................................................................................................................................... 119

4.8 Conclusions ................................................................................................................................... 121

4.9 Further work .................................................................................................................................. 122

5. SANTA MARGARIDA: A NEW MICROVERTEBRATE LOCALITY FROM ALGARVE ...... 123

5.1 Overview ....................................................................................................................................... 123

5.2 Introduction: Location, discovery, geological settings and nature of the site ............................... 123

5.3 Preparation methods ...................................................................................................................... 126

5. 4 Results .......................................................................................................................................... 129

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5.5 Discussion ..................................................................................................................................... 143

5.6 Conclusions ................................................................................................................................... 146

6. CONCLUSIONS ............................................................................................................................. 147

Chapter 2: Paleobiodiversity of the Quaternary fossil tetrapods in continental Portugal .................... 147

Chapter 3: The Palaeoloxodon antiquus of Santo Antão do Tojal ...................................................... 157

Chapter 4: Algar do Vale Da Pena: A new fossil bear site from Portugal .......................................... 148

Chapter 5: Santa Margarida: A new microvertebrate locality from Algarve. ..................................... 148

7. REFERENCES ................................................................................................................................ 149

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INDEX OF FIGURES

Figure 1.1: Bust of Carlos Ribeiro at the Geological Museum, Lisbon (Photo by Octávio Mateus). ..... 1

Figure 1.2: Picture of Nery Delgado at exposition in the Geological Museum, Lisbon. ........................ 2

Figure 1.3: Hyena coprolite from Furninha cave recovered by Nery Delgado (MG). Note the glued

identification.Geological Museum (Lisbon) ............................................................................................ 2

Figure 1.4: Picture of Paul Choffat at exposition in the Geological Museum, Lisbon. ........................... 3

Figure 1.5: Georges Zbyszewski (Picture by unknown author). .............................................................. 4

Figure 1.6: Miguel Telles Antunes in 2009 (Photo by Natacha Cardoso). .............................................. 5

Figure 1.7: João Luis Cardoso in 2017 (Photo by Octávio Mateus). ...................................................... 6

Figure 1.8: “Contribuição para o conhecimento dos grandes mamíferos do Plistocénico Superior de

Portugal” ................................................................................................................................................. 6

Figure 1.9: Vertebra of Salamandra salamandra from Figueira Brava cave in dorsal view; scale bar

1mm. Catalog number of the specimen not provided in the original (Crespo et al., 2000). ................... 8

Figure 1.10: Mandible from an indeterminate lacertid recovered at the site of Santa Margarida

(Portugal). FCT-UNL. ............................................................................................................................. 8

Figure 1.11: Right dentary of Blanus cinereus from Figueira Brava in lingual view, scale bar 1 mm.

Catalog number of the specimen not provided in the original (Crespo et al., 2000). ............................. 9

Figure 1.12: Hyoplastron of Testudo hermanni from Figueira Brava in ventral view; scale bar 1 cm.

Catalog number of the specimen not provided in the original (Lapparent de Broin & Antunes, 2000). 9

Figure 1.13: Right femur of Bubo bubo (Linnaeus, 1758) from Furninha cave in anterior view.

Geological Museum, Lisbon. ................................................................................................................. 10

Figure 1.14: Right humerus of Cygnus olor from Furninha cave in anterior view. Geological Museum,

Lisbon. ................................................................................................................................................... 10

Figure 1.15: Pinguinus impennis proximal humerus from Furninha in lateral view. Geological

Museum, Lisbon. .................................................................................................................................... 12

Figure 1.16: Right hemimandible of Erinaceus europaeus from Furninha in lingual view. Geological

Museum, Lisbon. .................................................................................................................................... 13

Figure 1.17: Rodenia incisor recovered at the site of Santa Margarida (Portugal). FCT-UNL. .......... 13

Figure 1.18: Molar of Palaeoloxodon antiquus (MG 3561) recovered by Paul Choffat in Condeixa

during the XIX century in lateral view. ................................................................................................. 15

Figure 1.19: Partial molar of Stephanorhinus hemitoechus from Gruta da Serra de Molianos.


Figure 1.20: Type specimen of Equus ferus antunesi in lateral view (MG). Geological Museum, Lisbon.

............................................................................................................................................................... 16

Figure 1.21: Anterior mandibular fragment of Hippopotamus incognitus from Condeixa in dorsal

view. Geological Museum, Lisbon. ........................................................................................................ 17

Figure 1.22: Skull of female Cervus elaphus from Gruta das Fontainhas in lateral view. Geological

Museum, Lisbon. .................................................................................................................................... 18

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Figure 1.23: Distal part of a humerus of Capra pyrenaica in anterior view from Caldeirão cave.

Catalog number and scale of the specimen not provided in the original (Davis, 2002). ...................... 19

Figure 1.24: Fragment of left hemimandible of Cuon alpinus with P4 and fragment of M1 from

Escoural cave in oclusal (left) and labial (right) views. Catalog number of the specimen or not

provided in the original, scale do not appear in the image (Cardoso, 1993a). .................................... 20

Figure 1.25: Left Ursus arctos mandible from Furninha cave in lingual view. Geological Museum,

Lisbon. ................................................................................................................................................... 20

Figure 1.26: Skull of Hyaena hyaena prisca from Furninha cave in dorsal view. Geological Museum,

Lisbon. ................................................................................................................................................... 21

Figure 1.27: The leopard (Panthera pardus) skull from Algar da Manga Larga in anterolateral view.


Figure 2.2: Percentages of different parameters about the paleodiversity of the Quaternary in

Portugal. ................................................................................................................................................ 37

Figure 2.3: Status of the different big mammals that appear in the fossil record in Portugal. ............. 38

Figure 2.4: Percentage of species from each group in extant, fossil and fossil species (without cave

deposits). ................................................................................................................................................ 41

Figure 3.1: The location of the Santo Antão do Tojal Palaeoloxodon antiquus is marked by the

pushpin, in the Lisboa Peninsula (left) and at local level (right). Credit Google Earth. ...................... 43

Figure 3.2: General view of the Palaeoloxodon antiquus femur of Santo Antão do Tojal (MG 5788) at

display in the geological museum in posterior view. Scale bar 7 cm. The chalk mark signals a cut in

the bone where a stone tool was found, being it regarded by Zbyszewski as a proof that the animal was

butchered by Neanderthals. ................................................................................................................... 45

Figure 3.3: Right tibia of the Palaeoloxodon antiquus of Santo Antão do Tojal (MG 5793) at display in

the Geological Museum in anterior view. Scale bar 7 cm. ................................................................... 46

Figure 3.4: Proboscidean tibia of Santo Antão do Tojal (MG 5793) in left, anterior and right proximal

views. The principal anatomical features are marked, in white the tibial depression and in red the

medial articular surface. Scale bar 7 cm. ............................................................................................. 46

Figure 3.5: The spinal process of the proboscidean of Santo Antão do Tojal (MG 8081) in A) anterior

and B) posterior views. .......................................................................................................................... 47

Figure 3.6: The phalanx from the proboscidean of Santo Antão do Tojal in A) dorsal; B) lateral; C)

ventral and D) lateral views. ................................................................................................................. 48

Figure 3.7: Measurements of the femur of mammals by Von den Driesch, 1976. ................................. 49

Figure 3.8: Measurements of the tibia of mammals by Von den Driesch, 1976. ................................... 51

Figure 3.9: Plot of means for the previously selected measurements of the femur of Mammuthus

primigenius (left) and Palaeoloxodon antiquus (right) with an interval of confidence of 95%. All

measurements are in millimeters. .......................................................................................................... 57

Figure 3.10: Plot of means for the previously selected measurements of the tibia of Mammuthus

primigenius (left) and Palaeoloxodon antiquus (right) with an interval of confidence of 95%. All

measurements are in millimeters. .......................................................................................................... 58

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Figure 3.11: Measurements of Santo Antão do Tojal specimen (in red) compared to the means for them

in Mammuthus primigenius (left) and Palaeoloxodon antiquus (right) with a 95% confidence interval.

............................................................................................................................................................... 59

Figure 3.12: Estimation of density for maximum length and minimum diaphysis width of the femur of

Mammuthus primigenius and Palaeoloxodon antiquus. ........................................................................ 60

Figure 3.13: Estimation of density for maximum length and minimum diaphysis width of the tibia of

Mammuthus primigenius and Palaeoloxodon antiquus ......................................................................... 61

Figure 3.14: From upper to bottom: Percentage of variation accounted by the first three Principal

Component of the femur; PC1, PC2 and PC3. In PC1 and PC2 the highest contribution comes from

the maximum length and the minimum diaphysis circumference respectively. ..................................... 63

Figure 3.15: PC1 vs PC2 of the femur of selected proboscideans. The Santo Antão do Tojal specimen

is marked with the red square, Mammuthus primigenius in blue and Palaeoloxodon antiquus in black.

............................................................................................................................................................... 63

Figure 3.16: From upper to bottom: Percentage of variation accounted by the first three Principal

Component of the tibia; PC1, PC2 and PC3. In PC1 and PC2 the highest contribution comes from the

maximum length and the minimum diaphysis circumference respectively. ........................................... 65

Figure 3.17: PC1 vs PC2 of the tibia of selected proboscideans. The Santo Antão do Tojal specimen is

marked with the red square, Mammuthus primigenius in blue and Palaeoloxodon antiquus in black. 65

Figure 4.1: Location of the Algar do Vale da Pena. Upper left, at peninsular level, it is situated in the

Portuguese Estremadura. Bottom, view at regional level, the closest city is Alcobaça. Upper right,

view at local level. Credit Google Earth. .............................................................................................. 67

Figure 4.2: Litostratigraphic log of the “Maciço Calcário Estremenho”, with the Candeeiros

Formation marked (Carvalho, 1996). .................................................................................................. 68

Figure 4.3: Location of the cave (red and orange dot) in the corresponding sheet (26 D, Caldas da

Rainha) of the geologic map (1:50000) of Portugal, including legend (Zbyszewski & de Matos, 1959).

............................................................................................................................................................... 68

Figure 4.4: Topography of the Algar do Vale da Pena. In black, some landmarks of the cave are

marked and in red, the bone findings (Modified from Amendoeira et al., 2015). ................................. 70

Figure 4.5: A) Entrance of the cave showing the first ramp and the top of two meters step. B) Picture

taken from the base of the two meters step, showing the second ramp. C) Final vertical well of the

entrance from the bottom. D) Picture from the bottom of the cave showing the final well (white line),

the second ramp (green line), the two meter step (red line) and part of the firs ramp (blue line).

Pictures by Carlos Ferreira. ................................................................................................................. 71

Figure 4.6: Claw marks (left) and dog skeleton (right) in the small conduct at the right of the cave. .. 72

Figure 4.7: Thick deposit of sands below the calcareous crust (left) and claw marks (right). .............. 72

Figure 4.8: Equipment needed to descend (in black) and climb out of the cave again (in red). Left; A)

Shunt, B) Descensor, C) Climbing fist. Right; D) Ventral croll and security knot to stop descending,

note how the fingers secure the shunt. ................................................................................................... 73

Figure 4.9: First excavation around the skull of “bear number one” in October 2016. ....................... 74

Figure 4.10: The main block as in October 2017, during the second expedition, no new diggings where

carried out this time. The skull is near the scale of seven centimeters, ................................................. 74

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Figure 4.11: A) Digging of the main block of “bear one” (note the camera of Rui Guerra). B) First

layer of plaster after the block was cut. C) Putting the block in the net. D) Moving the block with the

tree trunk. E) Final plastering of the block. F) Current state of the block. Pictures by Carlos Ferreira.

............................................................................................................................................................... 75

Figure 4.12: Before ( left) and after( right) for some ursid remains recovered in Algar do Vale da

Pena: a) distal radius, b) mandibular ramus and c) basicranium. ...................................................... 76

Figure 4.13: The main block of “bear” one during the second expedition. Scapula and protruding long

bones are marked with black arrows. In red, line where the block was cut, main skull with the scale of

7 cm and muddy area with numerous smaller bone remains. ............................................................... 77

Figure 4.14: Remains of the “bear number two” before (left) and after (right) removal of the block

that was hiding them. ............................................................................................................................. 78

Figure 4.15: Muddy sediment below the calcareous crust, during the preparation of the hemimandible

(ADVP 2.1). Scale 7 cm. ........................................................................................................................ 78

Figure 4.16: Material of bear number four encased in the calcareous crust. Left, humerus; right, femur

and pelvis. .............................................................................................................................................. 79

Figure 4.17: Distal humerus of the bear number five. ........................................................................... 79

Figure 4.18: Damaged scapula (ADVP 1.1) from “bear number one” (Ursus arctos) of Algar do Vale

da Pena. ................................................................................................................................................. 81

Figure 4.19: Right scapholunate bone (ADVP 1.3) from “bear number one” (Ursus arctos) of Algar do

Vale da Pena in A) proximal; B) distal; C) anterior and D) exterior views. ........................................ 81

Figure 4.20: Measurements for the scapholunate bone in ursids according to the system of Tsoukala &

Grandal-d’Anglade (2002). All of them could be recorded for the specimen. ...................................... 82

Figure 4.21: Transverse diameter vs anteroposterior diameter of the scapholunate of brown bears

(García Vázquez, 2015) as brown dots and cave bears (unpublished own data from Cova da Ceza

population) as green squares; plus the scapholunate ADVP 1.3, marked as a yellow star. ................ 83

Figure 4.22: Transverse diameter vs anteroposterior diameter of the scapholunate of brown bears

(data from García Vázquez, (2015)) as brown dots; plus the scapholunate ADVP 1.3, marked as a

yellow star. ............................................................................................................................................ 83

Figure 4.23: Right hamate bone (ADVP 1.4) from “bear number one” (Ursus arctos) of Algar do Vale

da Pena. ................................................................................................................................................. 84

Figure 4.24: Measurements for the scapholunate ursid bone according to the system of Tsoukala &

Grandal-d’Anglade (2002).The recorded measurement is marked. ...................................................... 84

Figure 4.25: Length of the right hamate bone (ADVP 1.4) from “bear number one” of Algar do Vale

da Pena compared to the length of the same bone of other brown bears (García Vázquez, 2015) .

Measurements are in mm according to the system of Tsoukala & Grandal-d’Anglade (2002). Two

groups (males and females) are classified according to this measurement. ......................................... 85

Figure 4.26: Second phalanx (ADVP 1.6) from “bear number one” (Ursus arctos) of Algar do Vale da

Pena in A) dorsal; B) ventral; C) proximal; D) distal views. ............................................................... 86

Figure 4.27: Measurements for the second phalanx of ursids according to the system of Tsoukala &

Grandal-d’Anglade (2002). All of them could be recorded. ................................................................. 86

Figure 4.28: Length vs distal transverse diameter of the second phalanx (ADVP 1.6, marked as blue

square) from “bear number one” (Ursus arctos) of Algar do Vale da Pena compared to the same

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ration of a recent brown bear, data from García Vázquez, (2015) . The 95% ellipses are drawn in the

graphs. Measurements are in mm according to the system of Tsoukala & Grandal-d’Anglade (2002).

............................................................................................................................................................... 87

Figure 4.29: Proximal epiphysis of the third right metatarsal (ADVP 1.7) from “bear number one”

(Ursus arctos) of Algar do Vale da Pena in A) exterior; B) dorsal; C) interior and D) ventral views. 88

Figure 4.30: Measurements for the third metatarsal of ursids according to the system of Tsoukala &

Grandal-d’Anglade (2002).The recorded measurements are marked. ................................................. 88

Figure 4.31: Proximal transverse diameter vs Proximal anteroposterior diameter of the third

metatarsal (ADVP 1.6, marked as blue square) from “bear number one” (Ursus arctos) of Algar do

Vale da Pena (As green triangle) compared to the same ration of fossil brown bears from NW Spain,

data from García Vázquez, (2015) . Measurements are in mm according to the system of Tsoukala &

Grandal-d’Anglade (2002). ................................................................................................................... 89

Figure 4.32: Distal epiphysis of the right radius (ADVP 1.8) from “bear number one” (Ursus arctos) of

Algar do Vale da Pena in A) anterior; B) exterior; C) Posterior; D) Interior and E) Distal views. ....... 90

Figure 4.33: Measurements for the radius of ursids according to the system of Tsoukala & Grandal-

d’Anglade (2002). Recorded measurements are marked. ..................................................................... 90

Figure 4.34: Transverse diameter of distal radius from ADVP 1.8 and fossil and recent brown bears

from NW Spain and Portugal (Cardoso, 1993; García Vázquez, 2015). Specimens with more than 60

mm are classified as male and less than 58 mm as females. ................................................................. 91

Figure 4.35: Transverse diameter vs anteroposterior diameter of distal radius from ADVP 1.8 (Green

in the graph) and fossil and recent brown bears from NW Spain (García Vázquez, 2015). Two clusters

of individuals are classified as females or males. ................................................................................. 92

Figure 4.36: Distal distal part of the ulnar diaphysis (ADVP 1.9) from “bear number one” (Ursus

arctos) of Algar do Vale da Pena in A) anterior; B) exterior; C) Posterior and D) Interior views. ..... 92

Figure 4.37: Measurements for the ulna of ursids according to the system of Tsoukala & Grandal-

d’Anglade (2002). Taken measurement is marked with a circle. .......................................................... 93

Figure 4.38: Minimum anteroposterior diameter of the ulnar distal diaphysis (ADVP 1.9) from “bear

number one” (Ursus arctos) of Algar do Vale da Pena compared graphically to the same measurement

of fossil and recent brown bears from NW Spain (García Vázquez, 2015). Suggested males and

females are marked................................................................................................................................ 94

Figure 4.39: Posterior part of right hemimandible (ADVP 2.1) from “bear number two” (Ursus

arctos) of Algar do Vale da Pena in A) dorsal; B) posterior; C) exterior; D) anterior; E) interior and

F) ventral views. .................................................................................................................................... 94

Figure 4.40: Measurements for the mandible of ursids according to the system of Tsoukala & Grandal-

d’Anglade (2002). Taken measurements are marked with a circle. ...................................................... 95

Figure 4.41: Total height of the mandible vs length from the mandibular condyle to the third molar

from ADVP 2.1 (blue square in the graph), fossil and recent brown bears from NW Spain as black dots

(García Vázquez, 2015) and cave bears as green triangles (unpublished data from Cova da Ceza).

Two clusters of individuals are classified as females or males. ............................................................ 97

Figure 4.42: Total height of the mandible vs height of the mandibular condyle from ADVP 2.1 (blue

square in the graph) and fossil and recent brown bears from NW Spain as black dots (García Vázquez,

2015) and cave bears as red inverted triangles (unpublished data from Cova da Ceza). .................... 97

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Figure 4.43: Diameter of the mandible between M2 and M3 vs height of the mandibular corpus

labially at M3 from ADVP 2.1 (blue square in the graph) and fossil and recent brown bears from NW

Spain as black dots (García Vázquez, 2015) and a cave bear as a green diamond (unpublished own

data from Cova da Ceza). ...................................................................................................................... 98

Figure 4.44: Partial occipital and parietal bones (ADVP 2.2) from the “bear two” (Ursus arctos) of

Algar do Vale da Pena in A) dorsal; B) posterior and C) anterior views. ............................................ 99

Figure 4.45: Temporal and partial parietal bones (ADVP 2.3) from the “bear two” (Ursus arctos) of

Algar do Vale da Pena in A) exterior and B) interior views. ................................................................ 99

Figure 4.46: Partial canine teeth (ADVP 2.5) from the “bear two” (Ursus arctos) of Algar do Vale da

Pena in lateral view. ............................................................................................................................ 100

Figure 4.47: Fourth upper premolar (ADVP 2.6) from the “bear two” (Ursus arctos) of Algar do Vale

da Pena in A) oclusal, B) lingual and C) labial views. ....................................................................... 100

Figure 4.48: Measurements for the fourth upper premolar of ursids according to the system of

Tsoukala & Grandal-d’Anglade (2002). ............................................................................................. 102

Figure 4.49: Breadth vs length of the fourth upper premolar ADVP 2.6 (blue square in the graph) and

the same measurements of recent and fossil brown bears from NW Spain as black dots (García

Vázquez, 2015). ................................................................................................................................... 102

Figure 4.50: First upper molar (ADVP 2.7) from the “bear two” (Ursus arctos) of Algar do Vale da

Pena in A) oclusal, B) lingual and C) labial views. ............................................................................ 103

Figure 4.51: Measurements for the first upper molar according to the system of Tsoukala & Grandal-

d’Anglade (2002). Taken measurements are marked with a circle. .................................................... 103

Figure 4.52: Breadth vs length of the first upper premolar ADVP 2.7 (blue square in the graph) and

the same measurements of recent and fossil brown bears from NW Spain as black dots (García

Vázquez, 2015) and cave bears as orange triangles from Liñares cave, Galicia, NW Spain (González

& Martelli, 2003). ................................................................................................................................ 105

Figure 4.53: Second lower molar (ADVP 2.8) from the “bear two” (Ursus arctos) of Algar do Vale da


Figure 4.54: Measurements for the second lower molar according to the system of Tsoukala &

Grandal-d’Anglade (2002). ................................................................................................................. 107

Figure 4.55: Breadth vs length in mm of the second lower molar ADVP 2.8 (blue square in the graph)

and the same measurements of recent and fossil brown bears from NW Spain as green

diamonds(García Vázquez, 2015) plus cave bears from A Ceza cave as black dots, Galicia, NW Spain

(Unpublished own data). ..................................................................................................................... 107

Figure 4.56: Third lower molar (ADVP 2.9) from the “bear two” (Ursus arctos) of Algar do Vale da


Figure 4.57: Measurements for the third lower molar according to the system of Tsoukala & Grandal-

d’Anglade (2002). ................................................................................................................................ 108

Figure 4.58: Breadth vs length in mm of the third lower molar ADVP 2.9 (black dot in the graph) and

the same measurements of recent and fossil brown bears from NW Spain as blue diamonds (García

Vázquez, 2015) plus cave bears from A Ceza, Liñares and Eirós caves, Galicia, NW Spain (Grandal-

d’Anglade, 1993; González & Martelli, 2003; unpublished own data) as purple triangles. .............. 110

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Figure 4.59: Third phalanx (ADVP 2.10) from the “bear two” (Ursus arctos) of Algar do Vale da

Pena in A) Distal; B) and D) Lateral; C) Proximal view. ................................................................... 110

Figure 4.60: Measurements for the third phalanx of ursids according to the system of Tsoukala &

Grandal-d’Anglade (2002). Taken measurements are marked with a circle. ..................................... 111

Figure 4.61: Height and diameter of the third phalanx (ADVP 2.10) (black dot in the graph) and the

same measurements of recent and fossil brown bears from NW Spain as blue squares (García

Vázquez, 2015) plus cave bears from A Ceza, Liñares and Eirós caves, Galicia, NW Spain (Grandal-

d’Anglade, 1993; González & Martelli, 2003; unpublished own data) as crimson diamonds. 95%

ellipses are drawn................................................................................................................................ 112

Figure 4.62: Distal epiphysis of femur (ADVP 3.1) from the “bear three” of Algar do Vale da Pena in

A) distal B) lateral views. .................................................................................................................... 112

Figure 4.63: Anteroposterior diameter of the distal epiphysis of the femur (ADVP 3.1) and the same

measurement of recent and fossil brown bears from NW Spain (García Vázquez, 2015). ................. 113

Figure 4.64: Vertebral centrum (ADVP 3.2) from the “bear three” of Algar do Vale da Pena in A)

dorsal and B) anterior views. .............................................................................................................. 114

Figure 4.65: Proximal epiphysis of second right metatarsal (ADVP 4.1) from the “bear four” of Algar

do Vale da Pena in A) Interior; B) Ventral; C) Exterior and D) Dorsal. ........................................... 114

Figure 4.66: Measurements for the second metatarsal according to the system of Tsoukala & Grandal-

d’Anglade (2002).Taken measurements are marked with a circle. ..................................................... 115

Figure 4.67: Anteroposterior vs transversal diameter of the proximal epiphysis (upper) and diaphysis

(bottom) of the second metatarsal, ADVP 4.1 (blue square) and the same measurements of recent and

fossil brown bears from NW Spain as black dots (García Vázquez, 2015). ........................................ 116

Figure 4.68: Proximal epiphysis of fifth right metatarsal (ADVP 4.2) from the “bear four” of Algar do

Vale da Pena in A) Interior; B) Ventral; C) Exterior and D) Dorsal views........................................ 116

Figure 4.69: Measurements for the fifth metatarsal according to the system of Tsoukala & Grandal-

d’Anglade (2002). Taken measurements are marked with a circle. .................................................... 117

Figure 4.70: Anteroposterior proximal diameter vs transversal diameter of diaphysis of the fifth

metatarsal, ADVP 4.2 (blue square) and the same measurements of recent and fossil brown bears

from NW Spain as black dots (García Vázquez, 2015). ...................................................................... 117

Figure 4.71: Claw marks in the walls of the Algar do Vale da Pena. Some of them are today partially

covered by calcareous crusts. Scale bar 7 cm. .................................................................................... 118

Figure 5.1: Location of the site of Santa Margarida. Figure by Cátia Ribeiro. Credit of the maps,

google maps. ........................................................................................................................................ 123

Figure 5.2: Maximum concentration point of fossil bearing blocks in the wall of the site as it was

discovered in 2017. Note the difference of color between the bone bearing breccified blocks of

cemented terra rossa and the grey tone of the calcareous rocks. Photo by Octávio Mateus. .............. 124

Figure 5.3: Location of Santa Margarida site. In blue the calcareous Picavessa Formation. Modified

by Cátia Ribeiro from the geologic map of Algarve by LNEG. ........................................................... 124

Figure 5.4: Left; comparison of freshly cut cemented terra rossa block and the underlying grayish

calcareous rock of the Picavessa formation. Right; calcareous crust covering the “terra rossa”..... 125

Figure 5.5: A) Abundant microfossils in rich block; B) Rabbit humerus in situ; C) and D) larger long

bones protruding from block. .............................................................................................................. 125

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Figure 5.6: Upper left, possible leporid phalanx in dorsolateral view; upper right, arvicoline mandible

in lingual view; bottom left, shrew mandible in labial view; bottom right, lizard mandible in labial

view. ..................................................................................................................................................... 126

Figure 5.7: Left, consolidated arvicoline mandible with PARALOID B72™; center, extraction of the

mandible using a air-scribe and right the extracted mandible. ........................................................... 126

Figure 5.8: Right, rodent teeth mounted in the plastic box and other fossils stored in similar settings

(Scale 7cm); left, the binocular lenses used to take the pictures (Photo by Alexadre Guillaume). ..... 127

Figure 5.9: Workflow followed in the preparation of the Santa Margarida material (Portugal) in the

Museum of Lourinhã............................................................................................................................ 128

Figure 5.10: Amount of processed sediment from Santa Margarida (Portugal) at Museum of Lourinhã.

............................................................................................................................................................. 129

Figure 5.11: Paralaoma servilis from Santa Margarida in A) lateral, B) ventral and C) dorsal views.

............................................................................................................................................................. 130

Figure 5.12: Bisected shell of Pomatia elegans from Santa Margarida. ............................................. 130

Figure 5.13: Lacertid mandible from Santa Margarida (Portugal) in lingual view............................ 131

Figure 5.14: Lacertid mandible from Santa Margarida (Portugal) in A) lingual and B) oclusal views.

............................................................................................................................................................. 131

Figure 5.15: Unicusp teeth ascribed to a bat from Santa Margarida (Portugal) in meso-lingual view

(Image by Cátia Ribeiro). .................................................................................................................... 132

Figure 5.16: Right upper second bat molar from Santa Margarida (Portugal) in oclusal view. The

rectangle marks the w shaped labial ridge and the circle the lingually recurved hypocone shelf and the

strong postprotocrista. Metastyle/paracone (1,3 mm) and metastyle/lingual (2 mm) lengths are

marked. Drawing by Pavlos Piskoulis. ................................................................................................ 132

Figure 5.17: Soricidae upper incisors recovered from Santa Margarida (Portugal) in labial view. A)

Sorex, B) and C) Crocidura. ................................................................................................................ 133

Figure 5.18: Broken second upper molar of Crocidura recovered from Santa Margarida (Portugal) in

oclusal view. Labial part to the right. ................................................................................................. 133

Figure 5.19: Fragment of Crocidura mandible (bearing P4, M1 and M2) recovered from Santa

Margarida (Portugal) in A) oclusal, B) lingual and C) labial views. ................................................. 134

Figure 5.20: Tip of a rodent incisor from Santa Margarida (Portugal). ............................................. 135

Figure 5.21: A) third lower molar and B) first upper molar of Apodemus cf. sylvaticus from Santa

Margarida (Portugal) in oclusal view. ................................................................................................ 135

Figure 5.22: Left upper first or second molar of Eliomys quercinus from Santa Margarida (Portugal)

in oclusal view. .................................................................................................................................... 136

Figure 5.23: Left upper first molar of Allocricetus bursae from Santa Margarida (Portugal) in oclusal

view. ..................................................................................................................................................... 136

Figure 5.24: Nomenclature and measurements for an arvicoline first lower tooth according to López-

García et al., (2015). ABBREVIATIONS: a: length of the anteroconid complex; ACC, anteroconid

complex; AC, anterior cap; BRA, buccal reentrant angle; BSA, buccal salient angle; c: width of the

opening of triangles T4-T5; L, length; La, mean width of T4; Li, mean width of T5; LRA, lingual re-

xxi

entrant angle; LSA, lingual salient angle; PL, posterior lobe; TTC, trigonidtalonid complex, T1-T7,

triangles 1-7; W, width. ....................................................................................................................... 137

Figure 5.25: Right lower first molar of Victoriamys chalinei from Santa Margarida (Portugal) in

oclusal view. ........................................................................................................................................ 137

Figure 5.26: Left lower first molar of Iberomys huescarensis from Santa Margarida (Portugal) in

oclusal view. ........................................................................................................................................ 138

Figure 5.27: Right and left (respectively) lower first molars of Iberomys brecciensis from Santa


Figure 5.28: Left lower first molar (without the posterior lobe) of Iberomys cabrerae from Santa


Figure 5.29: A/L x100 vs C/W x100 ratios for a collection of arvicoline teeth. Victoriamys chalinei

from El Chaparral locality appear as pink circles. Allophaiomys lavocati from El Chaparral appear

as blue pluses symbols. Iberomys huescarensis, from a collection of Iberian and Italian sites appear as

black dots. Data from López-García et al., (2012, 2015). The specimens from Santa Margarida

(Portugal) appear marked with a red circle. ....................................................................................... 140

Figure 5.30: A/L x100 vs La/Li x100 ratios for a collection of arvicoline teeth. Victoriamys chalinei

from El Chaparral locality appear as red inverted triangles. Allophaiomys lavocati from El Chaparral

appear as blue squares. Iberomys huescarensis, from a collection of Iberian and Italian sites appear

as black dots. Data from López-García et al., (2012, 2015). The specimens from Santa Margarida

(Portugal) appear marked with a red circle. ....................................................................................... 140

Figure 5.31: A/L x100 vs La/Li x100 ratios for Iberomys brecciensis (green squares) and I. chalinei

(gold inverted triangles) from a collection of Iberian and Italian sites. Data from López-García et al.,

(2015). The specimens from Santa Margarida (Portugal) appear marked with a red circle. ............. 141

Figure 5.32: Left distal Oryctolagus cuniculus humerus from Santa Margarida (Portugal) in A)

Anterior; B) Exterior; C) Posterior and D) Medial views. ................................................................. 142

Figure 5.33: Diverse postcranial elements of microvertebrates recovered from the sample of Santa

Margarida. A) Distal femur in posterolateral view, B) Partial distal humerus? in anterior view and C)

Distal radius in lateral view. ............................................................................................................... 142

Figure 5.34: Evolution of the first lower molar of the genus Iberomys with examples from the site of

Santa Margarida (Portugal). ............................................................................................................... 144

Figure 5.35: The two possible stratigraphy scenarios that explain the faunal assemblage recovered

from the ex situ blocks from Santa Margarida (Portugal). Upper: Fossils from different but similar

layers that became mixed during preparation. Lower: Reworked fossils situated in the same layer as

the younger ones. ................................................................................................................................. 145

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INDEX OF TABLES

Table 1.1: Presence of different species of Microtus/Iberomys in different Quaternary sites in

Portugal, data from Antunes et al., 1986b, Antunes et al., 1989; Póvoas et al., 1992; Jeannet,

2000; Moreno-García & Pimenta, 2002; Bicho et al., 2003; Valente & Haws, 2006; Pimenta,

2014, López-García et al., 2018). ........................................................................................ 14

Table 2.1: Localities of the Quaternary of Portugal. ............................................................. 23

Figure 2.1: Satellital picture showing the location of 30 sites with Quaternary fossil

vertebrates in Portugal, mentioned in the table 2.1. In black, the localities situated in the

Lusitanian basin and in red the ones situated in the Algarve basin. Credit Google Earth. .... 25

Table 2.2: Amphibians of the Quaternary of Portugal. ......................................................... 26

Table 2.3: Reptiles of the Quaternary of Portugal. ............................................................... 26

Table 2.4: Birds of the Quaternary of Portugal. .................................................................... 27

Table 2.5: Mammals of the Quaternary of Portugal.............................................................. 31

Table 2.6: Resume table showing the number, percentage and status of extant and fossil

species of the Quaternary of Portugal.................................................................................. 36

Table 2.7: Table showing the tetrapods and localities known outside the cave deposits in the

Quaternary of Portugal. ....................................................................................................... 38

Table 2.8: Resume table showing the number, percentage and status of extant and fossil

species in the Quaternary of Portugal, excluding the cave deposits. ................................... 40

Table 3.1: Measurements of the phalanx of the proboscidean of Santo Antão do Tojal.

“Thickness” refers to anteroposterior diameter of the bone and “width” to the mediolateral

diameter of it. All the measurements are in millimeters. ....................................................... 48

Table 3.2: Measurements of 48 individual femurs of Palaeoloxodon antiquus and

Mammuthus primigenius taken from the literature. The specimen of Santo Antão do Tojal is

marked. All measurements are in millimeters. ..................................................................... 49

Table 3.3: Measurements of 30 individual tibias of Palaeoloxodon antiquus and Mammuthus

primigenius taken from the literature. The specimen of Santo Antão do Tojal is marked. In

order to fit the table, abbreviations have been used: ML, maximum length; AL, articular

length; PW, proximal width; PT, proximal thickness; PAW, proximal articular width; PAT,

proximal articular thickness; MDW, minimum diaphysis width; MDT, minimum diaphysis

thickness; MDC, minimum diaphysis circumference; DW, distal width; DT, distal thickness;

DAW, distal articular width; DAT, Distal articular thickness. All measurements are in

millimeters. .......................................................................................................................... 51

Table 3.4: Maximum length vs minimum diaphysis circumference ratios of femur and tibia, for

selected Palaeoloxodon antiquus specimens. All measurements in mm.”E” stands for

“estimated”. ......................................................................................................................... 53

Table 3.5: Femur vs tibia lengths for selected Palaeoloxodon antiquus specimens. All

measurements in mm.”E” stands for “estimated”. ................................................................ 53

Table 3.6: Shoulder height and body masses of different Palaeoloxodon antiquus specimens.

The specimens below to the Santo Antão do Tojal one had not been calculated using the

same methods as presented. Possible males are marked in blue. ....................................... 54

Table 3.7: Number of individual measurements taken for proboscidean femurs and tibias of

the considered sample. Percentage of specimens where the measurement has been taken

compared to the total is also presented. The measurements that have been taken in 50% or

more of the specimens are marked. .................................................................................... 55

xxiii

Table 3.8: Mean for selected measurements in the hind limb of Palaeoloxodon antiquus and

Mammuthus primigenius, with percentage of difference between the species for each

measurement and bone. All measurements are given in millimeters. ................................. 56

Table 4.1: Measurements from the right scapholunate bone (ADVP 1.3) from “bear number

one” (Ursus arctos) of Algar do Vale da Pena compared to a set of measurements of other

brown bears (García Vázquez, 2015) and cave bears (unpublished own data from Cova da

Ceza population). Measurements are in mm according to the system of Tsoukala & Grandal-

d’Anglade (2002). ................................................................................................................ 82

Table 4.2: Length of the right hamate bone (ADVP 1.4) from “bear number one” (Ursus

arctos) of Algar do Vale da Pena compared to the length of the same bone of other brown

bears (García Vázquez, 2015) . Measurements are in mm according to the system of

Tsoukala & Grandal-d’Anglade (2002). ................................................................................ 85

Table 4.3: Length vs distal transverse diameter of the second phalanx (ADVP 1.6) from “bear

number one” (Ursus arctos) of Algar do Vale da Pena compared to the length of the same

bone of a recent brown bear (García Vázquez, 2015) . Measurements are in mm according

to the system of Tsoukala & Grandal-d’Anglade (2002). ...................................................... 87

Table 4.4: Proximal transverse diameter and proximal anteroposterior diameter of the third

metatarsal (ADVP 1.7) from “bear number one” (Ursus arctos) of Algar do Vale da Pena

compared to same measurements of fossil brown bears from NW Spain (García Vázquez,

2015) . Measurements are in mm according to the system of Tsoukala & Grandal-d’Anglade

(2002). ................................................................................................................................. 89

Table 4.5: Measurements from the radio (ADVP 1.8) from “bear number one” of Algar do

Vale da Pena compared to same ratio of fossil brown bears from NW Spain (García

Vázquez, 2015) as well as some of Portuguese caves. Measurements are in mm according

to the system of Tsoukala & Grandal-d’Anglade (2002). ...................................................... 91

Table 4.6: Minimum anteroposterior diameter of the ulnar distal diaphysis (ADVP 1.9) from

“bear number one” (Ursus arctos) of Algar do Vale da Pena compared to the same

measurement of fossil and recent brown bears from NW Spain (García Vázquez, 2015).

Measurements are in mm according to the system of Tsoukala & Grandal-d’Anglade (2002).

............................................................................................................................................ 93

Table 4.7: Measurements from the partial hemimandible (ADVP 2.1) from “bear number two”

(Ursus arctos) of Algar do Vale da Pena compared to same measurements of fossil brown

bears from NW Spain (García Vázquez, 2015) as well as some cave bears from the same

region (Unpublished own data from Cova de A Ceza). Measurements are in mm according to

the system of Tsoukala & Grandal-d’Anglade (2002). .......................................................... 96

Table 4.8: Measurements from the fourth upper premolar (ADVP 2.6) from “bear number two”

of Algar do Vale da Pena compared to broad an length of the same tooth of fossil and recent

brown bears from NW Spain (García Vázquez, 2015). Measurements are in mm according to

the system of Tsoukala & Grandal-d’Anglade (2002). .........................................................101

Table 4.9: Length and breadth from the first upper molar (ADVP 2.7) from “bear number two”

of Algar do Vale da Pena compared to breadth an length of the same tooth of fossil and

recent brown bears from NW Spain (García Vázquez, 2015) and cave bears from Liñares

cave, Galicia, NW Spain (González & Martelli, 2003). Measurements are in mm according to

the system of Tsoukala & Grandal-d’Anglade (2002). .........................................................104

Table 4.10: Breadth and length of the second lower molar (ADVP 2.8) from the “bear two” of

Algar do Vale da Pena compared to breadth an length of the same tooth of fossil and recent

brown bears from NW Spain (García Vázquez, 2015) and cave bears from A Ceza cave,

xxiv

Galicia, NW Spain (Unpublished own data). Measurements are in mm according to the

system of Tsoukala & Grandal-d’Anglade (2002). ...............................................................106

Table 4.11: Breadth and length of the third lower molar (ADVP 2.9) from the “bear two” of

Algar do Vale da Pena compared to breadth an length of the same tooth of fossil and recent

brown bears from NW Spain (García Vázquez, 2015) and cave bears from A Ceza, Liñares

and Eirós caves, Galicia, NW Spain (Grandal-d’Anglade, 1993; González & Martelli, 2003;

unpublished own data). Measurements are in mm according to the system of Tsoukala &

Grandal-d’Anglade (2002). .................................................................................................109

Table 4.12: Height and diameter of the third phalanx (ADVP 2.10) from the “bear two” of

Algar do Vale da Pena compared to the height and diameter of the same phalanx of fossil

and recent brown bears from NW Spain (García Vázquez, 2015) and cave bears from A

Ceza, Liñares and Eirós caves, Galicia, NW Spain (Grandal-d’Anglade, 1993; González &

Martelli, 2003; unpublished own data). Measurements are in mm according to the system of

Tsoukala & Grandal-d’Anglade (2002). ...............................................................................111

Table 4.13: Anteroposterior diameter of the distal epiphysis of the femur (ADVP 3.1) from the

“bear three” of Algar do Vale da Pena compared to the same measurement of fossil and

recent brown bears from NW Spain (García Vázquez, 2015). Measurements are in mm

according to the system of Tsoukala & Grandal-d’Anglade (2002). .....................................113

Table 4.14: Measurements of the second metatarsal (ADVP 4.1) from the “bear four” of Algar

do Vale da Pena compared to the same measurement of fossil and recent brown bears from

NW Spain (García Vázquez, 2015). Measurements are in mm according to the system of


Table 4.15: Measurements of the fifth metatarsal (ADVP 4.2) from the “bear four” of Algar do

Vale da Pena compared to the same measurement of fossil and recent brown bears from

NW Spain (García Vázquez, 2015). Measurements are in mm according to the system of


Table 4.16: Nitrogen and Carbon percentages in the samples of ADVP 1.10 and ADVP 3.4

from Algar do Vale da Pena (Portugal). ..............................................................................122

Table 5.1: Results of the tests for determining the product to be use in the chemical

preparation of Santa Margarida samples. ...........................................................................127

Table 5.2: Number of specimens from broad groups recovered from the processed sediment

of Santa Margarida. ............................................................................................................129

Table 5.3: Measurements and ratios of the most complete lacertid mandible fragment picked

from the disaggregated sediments of Santa Margarida, Portugal, following the methods of

Blain et al., 2007: h, is height of the teeth; d, its diameter and a, the height of the teeth that

protrudes over the dental crest. All measurements are in mm. ...........................................131

Table 5.4: Classification of the different Soricidae remains recovered from Santa Margarida

(Portugal). ..........................................................................................................................133

Table 5.5: Classification of the different rodent dental remains recovered from Santa

Margarida (Portugal). ..........................................................................................................134

Table 5.6: Measurements and ratios of the arvicoline first lower molars from Santa

Margarida ...........................................................................................................................139

xxv

LIST OF ABBREVIATIONS

Institutions

EAVP: European Association of Vertebrate Paleontology.

MG: Geological Museum of Lisbon.

FCT-UNL: Facultade de Ciência e tecnología da Universidade Nova de Lisboa.

LNEG: Laboratório Nacional de Energía e Geología.

ML: Museu da Lourinhã.

GPS: Grupo de Proteção Sicó.

GEM: Grupo de espeleologia e Montanhismo.

Other

PC: Principal Components.

YBP: Years Before Present.

xxvi

1

1.INTRODUCTION

1.1 History of the study of Quaternary fossil vertebrates from continental

Portugal

This section has been partially published in an abstract at meeting of EAVP 2018 (Estraviz-López &

Mateus, 2018b). The study of the remains of Quaternary fossil faunal assemblages cannot be

understood (mostly in their early years) without its close relation with archeological works and the

more general geologic survey. The discovery of tools first and then human remains (nowadays we now

that they were in fact younger) in the terraces of the Somme River in France, related with the remains

of extinct fauna like rhinoceroses and elephants in the beginning of the second half of the XIX

century, sparked the interest of the scientific community about the antiquity of humanity and the

search of new bone material and artifacts in Pleistocene deposits. Also, other two factors in this period

made the interest in the study of the past rise in the scientific community as never before: The

publication of “The origin of species” by Charles Darwin and the outbreak of the nationalist sentiment

that made the cult peoples of Europe to search for clues and material proof in the past, even competing

with other nations and scientists (Cardoso, 2015a).

In Portugal, the scientific study of the Quaternary started thanks to the work of the members of the 2nd

Geological Commission of Portugal in 1857; mostly Carlos Ribeiro and Nery Delgado, whose work

proved that it was possible to know about the distant past (well before the written texts, oral traditions

and collective memory) just thanks to the study of material remains (being them bones or tools)

coming from alluvial and cave deposits (Cardoso, 2015a).

Carlos Ribeiro (1813-1884) (Figure1.1) was the father of the prehistoric archeology (and therefore the

Quaternary paleontology) in Portugal. From humble origins he developed a career in the army as

military engineer that he left in 1847, being the leader of the Geological Commission of Portugal with

Pereira da Costa (more focused to collection management than fieldwork) since 1857 until his death

(Daveau, 2000). He published a surprisingly accurate description of the Quaternary deposits, that in

that moment also included some of Neogene origin, in the basins of Tagus and Sado rivers that he

never finished (a second part of that work was never published) (Cardoso, 2015b) and several other

works including descriptions of silex and quartzite tools from the terrains of those rivers (Ribeiro,

1871). He raised the question about how old the humanity could be and pointed towards the possibility

that some tools that he collected could be from Tertiary origin (Cardoso, 2000) . He also carried

fieldwork, excavating the prehistoric settlement of Leceias in 1878 and the cave of Ponte da Laje in

1879 among other diggings (Cardoso, 2013), probably searching for material to show to other fellow

scientist in the important IX session of the International Congress of Anthropology and Prehistoric

Archaeology that was carried in Lisbon in September of 1880; the culmination of 25 years of his work.

Figure 1.1: Bust of Carlos Ribeiro at the Geological Museum, Lisbon (Photo by Octávio Mateus).

2

Joaquim Felipe Nery Delgado (1835-1908) (Figure 1.2) was also a major figure in the study of the

Quaternary in Portugal. He graduated as military engineer in 1855 just in time to incorporate as

adjunct to the Director of the Geological Comission of Portugal in 1857 that he later lead since the

death of Carlos Ribeiro in 1884 to his own death in 1908, with more than 51 years of study and

research in the areas of geologic cartography and prehistoric archeology on his back. As geologist his

major interest was the study of the Paleozoic rocks in Portugal, mostly in Alentejo as well as the

nature of the enigmatic marks of “bilobites” (Cruziana) (Cardoso, 2008a).

Figure 1.2: Picture of Nery Delgado at exposition in the Geological Museum, Lisbon.

In 1865 and 1866 he undergone the first scientific excavation of a cave occupied by prehistoric

humans, Casa da Moura, in the municipality of Óbidos and Cova da Moura, Lourinhã and Bombarral

municipalities. He later published his findings, along with other excavations in nearby caves in his

work “Da Existência do Homem no Nosso Solo em Tempos Mui Remotos Provada pelo Estudo das

Cavernas. Notícia ácerca das Grutas da Cesareda” published in 1867 that included a human skull,

which being published before the Cro Magnon man would be the first Paleolithic anatomically modern

human remains known (Delgado, 1867; Zilhão, 1993). The same year of 1865 he started some

preliminary diggings in the Furninha cave in Peniche, but it was not until 1879 and 1880 when he

undertook this pivotal excavation in the history of the Portuguese prehistoric archeology and

Quaternary paleontology (Delgado, 1884). The methodology of this excavation is even nowadays

exemplar, with each piece of bone or tool identified not only with the site but with the layer from

where it came and with its coordinates in a grid system (Figure 1.3). So Nery Delgado was a pioneer

of the 3D location of the archeological artifacts (Cardoso, 2015a).

Figure 1.3: Hyena coprolite from Furninha cave recovered by Nery Delgado (MG). Note the glued

identification.Geological Museum (Lisbon)

3

Both Nery Delgado and Carlos Ribeiro were respected scientists in his time, being his work known

across Europe, proof of that is the correspondence that they kept with other fellow foreign scientists

(Cardoso & Avila de Melo, 2001) and the visits that they did to other countries, like for example Nery

Delgado to Spain in 1878 (Carneiro, 2007). The quality of their work put Portugal in the frontline of

this type of studies in Europe at that time, and all of this crystallized in the celebration of the IX

session of the International Congress of Anthropology and Prehistoric Archaeology in Lisbon, 1880.

Some new diggings were made to provide with new information and pieces to show in the congress,

for example the interesting site of Mealhada where some proboscidean remains were uncovered

(Cardoso, 1993).

In the last years of the XIX century and the first of the XX century Nery Delgado focused more in the

study of the geology of the Paleozoic of Portugal until his death (Delgado, 1886) but the study of the

Quaternary animal remains continued with less intensity. Paul Choffat was another important geologist

and paleontologist of those years, that inherited the lead of Nery Delgado over the geological services

and overall the study of the geology in Portugal; he uncovered some hippopotamus and proboscidean

remains in the calcareous tuffs of Condeixa in 1895 and 1898. Years later, in 1909, he also promoted

the diggings in the Pleistocene sites of Algar de João Ramos and Molianos (Cardoso, 1993b).

Paul Choffat (figure 1.4) realized that the materials recovered by Nery Delgado and Carlos Ribeiro

deserved an attention that he (being focused on the investigation of the Paleozoic and Mesozoic of

Portugal) could not properly give, so thanks to his initiative the classic Portuguese material of sites

like Furninha and Fontainhas was studied by Edouard Harlé, from Toulouse (One of the top experts in

the Pleistocene of that time). He published a work that would be a major reference for the study of

Quaternary paleontology in Portugal during the next 80 years, cataloging the fauna found until that

moment and comparing it with other countries of Europe (Harlé, 1910; Cardoso, 1993b).

Figure 1.4: Picture of Paul Choffat at exposition in the Geological Museum, Lisbon.

After this outburst of the early XX century the study of the Quaternary animals in Portugal entered a

phase of stasis. Paul Choffat said: “By reasons that we should not comment, although that our faculties

are filled with professors of great erudition, there are few of them that are dedicated to the progress of

observation sciences”, that was his way of saying that there was not much interest in carrying

fieldwork in Portugal during those years (Daveau, 2000). We should wait until the middle of the

century and the works of Octávio da Veiga Ferreira and mostly Georges Zbyszewski to restart the

findings in the Quaternary of the Portuguese caves and alluvial deposits.

4

Octávio da Veiga Ferreira (1917-1997) was a mine engineer that incorporated to the Geological

Services in 1950. He already had carried some archeological excavations by that time and during his

many years of activity he was a major figure in the Iberian archeology. In the area of paleontology,

although it was not his main research field, he published 32 papers, four of them about Pleistocene

vertebrates and one about Holocene mollusks (Cardoso, 2008 b). One of his major contributions is his

study about the rhinoceroses of Portugal (Veiga Ferreira, 1975).

Georges Zbyszewski (1909-1999) (Figure 1.5) was the direct superior of Octávio da Veiga Ferreira;

born in Russia, but later nationalized French citizen. He visited Portugal for first time in 1935

searching for clues about the tectonic movements in the Portuguese littoral during the Quaternary for

his doctoral thesis. In 1940 he incorporated to the Geological Services and during the next 40 years he

covered an immense array of geological topics, writing around 300 scientific publications from the

geological cartography of Portugal, to the volcanism in Açores and Madeira, to the Mesozoic reptiles

and over all of that; more than 100 publications about Quaternary lithic industries and fauna (Soares

de Carvalho & Cardoso, 1999; Teixeira, 1979). During the years “Zby” (as he was called by friends

and family) reunited a lot of information about the recent fluvial terraces of the Tagus River and its

tributaries in the work “Le Quaternaire du Portugal” published in 1957. A lot of publications were

made from mammal material coming from those terraces, either Miocene or Pleistocene, for example

some remains like phalanges and tusks of Paleoloxodon antiquus (Falconer & Cautley, 1847) from

that area and the classic site of Mealhada. Is also remarkable the different excavations and publications

that he did with the material of a lot of different Pleistocene caves like: Almonda (1941 and 1947),

lapa da Bogalheira (1941), the caves of Maceira and Vimeiro (1949), the cave of Ponte da Laje

(excavated before by Carlos Ribeiro) (1957), Gruta das Salemas (1962), Gruta da Columbeira (1963),

Alcobertas (1971), and lapa do Bugio the same year. Beside all of this he also studied the Pleistocene

faunas coming from Algoz, in Algarve (Soares de Carvalho & Cardoso, 1999; Teixeira, 1979). All this

work would have been probably impossible without the hard work of dozens of unrecognized

collectors that were the eyes and ears of not only him but also great scientist before and after him.

Figure 1.5: Georges Zbyszewski (Picture by unknown author).

The work of Veiga Ferreira and Zbyszewski in the Geological services of Portugal served well to kick

start the next phase of the study of the Quaternary faunas in Portugal, with the lead of Miguel Telles

Antunes first and João Luis Cardoso later. A good visualization of this is the work that both of them

published in 1990 to synthesize their many years of study in this matter with the new studies of the

next generation in the decade of 1980 (Zbyszewski & Veiga Ferreira, 1990).

5

Before continuing it should be noted the work of Torres Pérez-Hidalgo, a Spanish researcher that in

1979 published an extensive study about the bears of Portugal, including measurements of old material

recovered even in the XIX century and agreeing with the work of Harlé in that there is no other

recognized fossil bear in the country besides the brown bear, Ursus arctos Linnaeus, 1758 (Torres

Pérez-Hidalgo, 1979).

Miguel Telles Antunes (1937) (Figure 1.6) is probably until this day the most prolific figure in the

History of the Portuguese paleontology. He started his career in the University of Lisbon in 1960 after

his doctorate in geology and in 1974 he moved to the Universidade Nova de Lisboa. During his many

years in paleontology he covered practically all the topics in the area writing more than 450 scientific

articles and directing 9 doctoral theses, from the Miocene vertebrates in the area of Lisbon, to the

Eocene of Silveirinha (Baixo Mondego), or the dinosaur faunas in the Lusitanian Basin

(https://www.fct.unl.pt/noticias/2016/12/sessao-de-homenagem-ao-professor-doutor-miguel-telles-

antunes). His collaboration with Zbyszewski at the end of the decade of 1970 produced some new

insights into the Neogene of the Tagus Basin and during the decade of 1980 he carried several studies

in Pleistocene material: The faunas of Morgadinho and Goldra in Algarve, the review of the Algoz

fauna (also from Algarve) and the Mealhada material (that lead to the identification of Homotherium

latidens Owen, 1846 in Portugal) and also the studies carried out in the cave contexts of Figueira

Brava (where he impulse excavations), Caldeirão cave (where he recognized Castor fiber Linnaeus,

1758 for first time in Portugal) (Antunes, 1989a) or Pedreira das Salemas (Cardoso, 1993a). Over all

of this work, the impulse that Antunes gave to the study of Pleistocene faunas through the former

CEPUNL (Centro de Estratigrafía e Paleobiología da Universidade Nova de Lisboa) is everlasting

even today, with numerous radiometric datings and training of new personal.

Figure 1.6: Miguel Telles Antunes in 2009 (Photo by Natacha Cardoso).

More or less at the same time during the decade of 1980 and the beginning of the 1990, two

personalities started producing scientific investigation at high level. João Zilhão (1957) now professor

of the University of Barcelona is an archaeologist, therefore his core area is situated more in the

paleoanthropology, but his work in the Almonda cave system (Zilhão et al., 1991) and other caves in

the Estremadura like Caldeirão cave produced a wide array of Pleistocene faunal material (Zilhão,

1987). This material was subject of studies the following years.

6

The other personality is João Luis Cardoso (1956) (Figure 1.7) the most well known researcher in the

study of Quaternary faunas from Portugal. Professor of the Universidade Aberta de Lisboa has written

more than 550 publications, including books, chapters of books and scientific articles; a lot of them

about the paleontology and archeozoology, but also in archeology and geology (https://www.cm-

obidos.pt/jornadas-arqueologia). In the second half of the decade of 1980 and the first of the decade of

1990 he worked in the CEPUNL publishing together with Antunes or in solitary plenty of scientific

articles such the first reference of several species in the Pleistocene of Portugal, like Hippopotamus

incognitus Faure, 1984 in Portugal from the deposits of Mealhada (Antunes et al., 1988), Cuon alpinus

europaeus Bourguignat, 1868 (Cardoso, 1992), Panthera spelaea Goldfuss, 1810 (Antunes &

Cardoso, 1986), Crocuta crocuta intermedia (M. de Serres, 1828) (Cardoso, 1994), Equus hydruntinus

Regalia, 1905 (Cardoso, 1995), Dicerorhinus hemitoechus (Falconer, 1868) (Cardoso, 1990),

Mammuthus primigenius Blumebach, 1799 (Antunes & Cardoso, 1992) and Rupicarpa rupicapra

(Linnaeus, 1758) (Cardoso & Antunes, 1989). Also he has described a new subspecies of horse from

the Pleistocene of Portugal, Equus ferus antunesi (Cardoso & Eisenmann, 1989).

Figure 1.7: João Luis Cardoso in 2017 (Photo by Octávio Mateus).

But his main contribution on this period is his doctoral thesis defended in October, 1992 and published

in 1993 by the municipality of Oeiras titled “Contribuição para o conhecimento dos grandes

mamíferos do Plistocénico Superior de Portugal” (Figure 1.8); a book of more than 500 pages that is

the most comprehensive and at the same time detailed study ever made in Portugal about the

Quaternary big mammal fauna of the country, covering remains of 26 taxa from 28 localities across all

Portugal (Cardoso, 1993a).

Figure 1.8: “Contribuição para o conhecimento dos grandes mamíferos do Plistocénico Superior de Portugal”

7

During the next 20 years João Luis Cardoso has continued to develop some more work in that

investigation line, doing a catalog of Iberian equids (Morales-Muñiz et al., 1998), describing a

mammoth lamella near Lisbon (Cardoso & Regala, 2002) or a particularly strange leopard in the Algar

da Manga Larga (Cardoso & Regala, 2006) plus some important syntheses about the study of

Quaternary mammals and archeozoology (Cardoso, 1993 b; Cardoso, 2002) or the life and work of

important scientist that we have covered before (Cardoso & Avila de Melo, 2001; Cardoso, 2008a;

Cardoso, 2015b). Beside his labor, since the year 2000 other works have started to fill the gaps in our

knowledge of the fauna of that time period that Cardoso already noted: the study of Quaternary bird

fossils and microfauna like rodents, amphibians and reptiles (Cardoso, 2002).

About the Quaternary birds, two works have vastly augmented our knowledge during the last years. In

2010 a comprehensive thesis about the birds of this age in Portugal was presented, in which bird

remains of more than 13 sites were studied for first time or reviewed (Figueiredo, 2010) and also

recently another work was published about avian remains found in archeological contexts across the

country (Pimenta et al., 2015) making our understanding of the Quaternary avifauna of Portugal

improve considerably. Meanwhile, on amphibians and reptiles, only few publications about

Pleistocene material from Portugal have been made (Jiménez Fuentes et al., 1998; Crespo, 2002). The

micromammals (and even more the fishes) are still poorly studied compared to Spain but there are

some works published (Antunes et al., 1986a; Antunes et al., 1989), mostly related to archeological

contexts (Monteiro, 2016; Póvoas et al., 1992) although new works aim to a change this (López-

García et al., 2018).

1.2 Vertebrates in the Quaternary of Portugal

Quaternary fishes, amphibians, reptiles and micromammals are almost always treated in an

archaezoological way, not to study the small vertebrate’s ecology or paleodistribution per se (Dean &

Carvalho, 2011; Nabais & Rodrigues, 2017; Pereira et al., 2017).

Studied fish remains from the Pleistocene and Holocene of Portugal overwhelmingly come from

archeological contexts, mostly Holocene (Pereira et al., 2017) although some are older, Pleistocene,

like in Caldeirão cave (Zilhão, 1997). There are marine fishes recognized (although rare) in the after

mentioned cave and in Lapa do Suão or Lapedo. They appear in bigger numbers in Lapa do Picareiro

and Lapa dos Coelhos (Bicho & Haws, 2008). Fish remains recovered in geological context are the

pharyngeal teeth of cyprinids from the Pleistocene site of Morgadinho, Algarve (Antunes et al.,

1986a).

The Quaternary amphibians have been recognized in numerous recent archeological sites of Holocene

age; for example they have been found in the Mesolithic shell midden of Cabeço de Morros, although

not specific identification is given (Detry, 2008). According to Hockett & Haws (2009) amphibian

remains have been found in three out of the eight archeological sites treated in that work: Figueira

Brava, Picareiro and Lagar Velho. In the Lagar Velho site four amphibian species were recognized,

being two of them unidentified anurans and the other two cf. Triturus marmoratus (Latreille, 1800)

and cf. Salamandra salamandra (Linnaeus, 1758) (Moreno-García & Pimenta, 2002). In Picareiro

cave two species were recognized: Bufo bufo (Linnaeus, 1758) and Pelobates cultripes Cuvier, 1829

(Hockett & Haws, 2009). The cave of Figueira Brava has yielded remains of Salamandra salamandra

(Figure 1.9), Pelobates cultripes and one indeterminate anuran (Crespo et al., 2000). The largest

amphibian assemblage of this time period from Portugal comes from Guia, Algarve, a site in geologic

seasonal context with no archaeological remains known and five species of amphibians being

identified: Pelobates cultripes (Almost ¾ of all fossil remains), Pleurodeles waltl (Michahelles, 1830),

Bufo bufo, Epidalea calamita Laurenti, 1758 and Pelophylax perezi (Seoane, 1885) (Antunes et al.,

1989). From the Gruta da Aroeira (Almonda System) there are undetermined anurans and

salamandrines as well as one species of Pelodytes Bonaparte, 1838 and Bufo spinosus (Daudin, 1803)

(López-García et al., 2018). Also there are some undetermined amphibian remains come from the

Morgadinho site in Algarve (Antunes et al., 1986a).

8

Figure 1.9: Vertebra of Salamandra salamandra from Figueira Brava cave in dorsal view; scale bar 1mm.

Catalog number of the specimen not provided in the original (Crespo et al., 2000).

Reptiles have been recognized in at least 12 sites of the Pleistocene of Portugal: Gruta da Aroeira

(Almonda System), Mealhada, Furninha, Caldeirão, Almonda, Gruta Nova da Columbeira, Lagar

Velho, Portocovo, Figueira Brava, Escoural, Foz do Enxarrique and Gruta de Ibn Amar. All of the

reptile species recognized still exist in Portugal nowadays except one. Most of the remains found are

Testudines Linnaeus, 1758 probably because this group of animals has a lot of sturdy bony plates that

favor conservation, compared with the relative fragile bones of other reptiles, but there are also some

references of Squamata Oppel, 1811 like Lacertilia Gunther, 1867 (Figure 1.10), Amphisbaenia Gray,

1844 and Serpentes Linnaeus, 1758 (Crespo, 2002; Morales Pérez & Serra, 2009).

Timon lepidus Daudin, 1802 is the most common lacertid, appearing in Figueira Brava and Guia.

Psammodromus algirus (Linnaeus, 1758) is referred in Figueira Brava, although the remains probably

should be ascribed to Psammodromus manuelae Busack et al., 2006 (Busack et al., 2006). From the

same locality there is a reference to an undetermined Podarcis Wagler, 1830. In Guia (Algarve)

appears Acanthodactylus erythrurus (Schinz, 1833) and from Lagar Velho come other two

undetermined lacertids (Antunes et al., 1989; Crespo et al., 2000; Moreno-García & Pimenta, 2002).

Undetermined lacertids appear as well at Gruta da Aroeira (Almonda System) (López-García et al.,

2018).

Figure 1.10: Mandible from an indeterminate lacertid recovered at the site of Santa Margarida (Portugal).

FCT-UNL.

The Serpentes of the Quaternary of Portugal are mostly known from the cave of Figueira Brava, where

at least two species of Colubridae Oppel, 1811 are recognized, being them probably Rinechis scalaris

(Schinz, 1822) and/or Hemorrhois hippocrepis (Linnaeus, 1758), but probably more are present (like

Coronella girondica Daudin, 1803 or Vipera lastei Bosca, 1878). In the Gruta da Aroeira (Almonda

System), there are references to cf. Coronella girondica and Natrix cf. maura (López-García et al.,

2018). In Guia there is a reference to the presence of Natrix Laurenti, 1758. Also two undetermined

Serpentes are present at Lagar Velho (Antunes et al., 1989, Crespo et al., 2000; Crespo, 2002;

Moreno-García & Pimenta, 2002).

The Amphisbaenia group is represented by Blanus cinereus Vandelli, 1797 of Figueira Brava, where it

is known by a right dentary (Figure 1.11) and vertebrae (Crespo et al., 2000).

9

Figure 1.11: Right dentary of Blanus cinereus from Figueira Brava in lingual view, scale bar 1 mm. Catalog

number of the specimen not provided in the original (Crespo et al., 2000).

Testudines are represented in Portugal by three species during this time period: Testudo hermanni

Gmelin, 1789, Mauremys leprosa (Schweiger, 1812) and Emys orbicularis (Linnaeus, 1758). Testudo

hermanni, nowadays extinct in Portugal, is the most common one appearing in the literature under

different nomenclatures like Agrionemys in the sites of Mealhada, Furninha, Figueira Brava (Figure

1.12), Gruta Nova da Columbeira, Caldeirão, Escoural and the Gruta of Ibn Amar; being the later one

probably of Holocene age. Mauremys leprosa appears in Mealhada (being this the oldest appearance

of the species in Portugal), Caldeirão, Porto Covo (Cascais) and probably Gruta Nova da Columbeira.

Finally, Emys orbicularis appears in Figueira Brava and probably Gruta Nova da Columbeira. It

should be noted that Gruta Nova da Columbeira has yielded more than 80 specimens of turtles, being

this the most important locality for the group made (Jiménez Fuentes et al., 1998; Lapparent de Broin

& Antunes, 2000; Morales Pérez & Serra, 2009).

Figure 1.12: Hyoplastron of Testudo hermanni from Figueira Brava in ventral view; scale bar 1 cm. Catalog

number of the specimen not provided in the original (Lapparent de Broin & Antunes, 2000).

The avifauna of the Pleistocene of Portugal is known in around 15 different sites: Gruta Nova da

Columbeira, Gruta das Salemas, Gruta das Fontainhas, Gruta da Furninha, Gruta do Pego do Diabo,

Gruta da Casa da Moura, Gruta do Caldeirão, Lapa da Rainha, Lagar Velho, Lapa do Picareiro,

Galería Pesada/Almonda, Lapa do Suão, Escoural, Gruta da Figueira Brava and Vale Boi (Figueiredo,

2010; Manne et al., 2012; Pimenta et al., 2015). Practically all of them (if not all) are situated in

archaeological contexts. More than 78 species have been recognized in diverse works, the immense

majority of them are already present in the country (Figure 1.13) but five species are extremely rare in

Portugal, four of them no longer inhabit the territory and three are extinct (Figueiredo, 2010; Pimenta

et al., 2015). It is also worth mentioning that in the Portuguese islands of Açores and Madeira there are

more examples of extinct birds, but given their insular nature those will not be discussed here (Alcover

et al., 2015).

10

Figure 1.13: Right femur of Bubo bubo (Linnaeus, 1758) from Furninha cave in anterior view. Geological

Museum, Lisbon.

Between the species that have been discovered in the Quaternary fossil record of Portugal, there are

some (most of them being of cold climate) today considered “rarities” and whose sights are

homologated by the “Portuguese Committee of Rarities” belonging to the SPEA, “Sociedade

Portuguesa para o Estudo das Aves”. All of the following homologated cites come from them:

Falco rusticolus Linnaeus, 1758; is known by one left tarsometatarsus from Gruta da Moura that have

the characteristics of the genus Falco but is bigger than any other species. The gyrfalcon today lives in

the high Arctic and has only one homologated sight in Portugal in 1991 (Poole & Boag, 1988;

Figueiredo, 2010).

Anser albifrons (Scopoli, 1767) is known by one carpometacarpus from Pego do Diabo. This species

today nest in the Arctic and winters in Northern Europe. It has ten homologated sights in Portugal (Ely

et al., 2005; Figueiredo, 2010).

Somateria mollisima (Linnaeus, 1758) is recognized in Furninha cave, being particularly abundant in

this site (the second most common duck). Although this fact, today it only lives in Northern Europe

and has five homologated sights in Portugal (Swennen, et al., 1989; Brugal et al., 2012).

Mergus merganser Linnaeus, 1758 has been recognized in Escoural cave, but nowadays is rare in

Iberian Peninsula and has only been seen eight in Portugal (Pimenta et al., 2015).

Cygnus olor (Gmelin, 1789) is known by one right humerus from Gruta da Furninha (Figure 1.14) and

in Galería Pesada, Almonda cave system. It lives today mostly in Northern Europe and has been seen

at least 20 times in Portugal (Figueiredo, 2010; Pimenta et al., 2015).

Figure 1.14: Right humerus of Cygnus olor from Furninha cave in anterior view. Geological Museum, Lisbon.

11

Between the species that no longer live in Portugal (most of them being also linked to cold, like the

previous group) we have:

Pyrrhocorax graculus (Linnaeus, 1766) is probably the most common species of this category, being it

represented by hundreds of bones in Lapa da Rainha, Furninha, Gruta de Casa da Moura, Lapa do

Picareiro, Fontainhas, Caldeirão and Gruta Nova da Columbeira (Figure 1.12). Although its

widespread distribution an abundance, it no longer appears in the mountains of Portugal according to

the SPEA, probably due to climatic reason (Figueiredo, 2010; Smith et al., 2013).

Tetrao urogallus Linnaeus, 1758; known by one tarsometatarsus from Lapa da Rainha, that lived in

the area of Peneda-Gêres during the second half of the XVIII century and went extinct around 100

years ago in Portugal (Dantas da Gama, 1998; Figueiredo, 2010).

Perdix perdix Linnaeus, 1758; is known by several bone remains from Lapa da Rainha and Gruta

Nova da Columbeira. Recently it has appeared back in the Northeast of Portugal (Martínez Nistal et

al., 1986; Figueiredo, 2010).

Lagopus muta (Montin, 1781) has been recognized by one tarsometatarsus from Gruta Nova da

Columbeira. Today the closer populations to Portugal are in the Pyrenees (Figueiredo, 2010; Smith et

al., 2013).

Lagopus lagopus Linnaeus, 1758 was discovered in Portugal in Gruta Nova da Columbeira (two

tarsometatarsus) and probably Lapa da Rainha (one femur, one tarsometatarsus). There is also an

interesting reference of possible Holocene survival of this species in Portugal until Roman times in

Quinta das Longas, but nowadays it is extinct from the Iberian Peninsula as a whole (Figueiredo,

2010; Smith et al., 2013; Pimenta et al., 2015).

Then we have three fossil species of birds found in the Quaternary of Portugal now extinct: Grus

primigenia (Milne-Edwards, 1869), Puffinus holeae Walker et al., 1990 and Pinguinus impennis

(Linnaeus, 1758). Grus primigenia, the cave crane, was a larger crane than the Grus grus Linnaeus,

1758; almost as big as the Asiatic sarus crane, Grus antigone (Linnaeus, 1758). It inhabited Western

Europe during the Pleistocene and part of the Holocene, being known from France, Germany,

Mallorca, Ibiza and England, surviving until the Iron Age in that case. It is known in Portugal by one

left scapula from Figueira Brava cave. This species was sturdier than the sarus crane and probably had

a different way of living. The cause and precise date of its demise remains in mystery (Harrison &

Cowles, 1977; Northcote & Mourer-Chauviré, 1985; Northcote & Mourer-Chauviré, 1988; Mourer-

Chauviré & Antunes, 2000).

Puffinus holeae Walker et al., 1990; the Canary dune shearwater, whose remains have been recovered

from different levels of Figueira Brava (Coracoids, humerus, ulnas, etc). This was the first time that

the species was recorded outside its nesting grounds in the Canary Islands. The fact that it appears

until recent levels in the cave pointed towards that its demise was triggered by the human colonization

of the islands that led to the destruction of its colonies and not climate change at the end of

Pleistocene, as was later proved (Mourer-Chauviré & Antunes, 2000; Rando & Alcover, 2010).

Pinguinus impennis (Linnaeus, 1758), the great auk, has been recognized in two Portuguese sites

(besides the island of Porto Santo): In Figueira Brava by two humerus and in Furninha by one humerus

(Figure 1.15) that was recovered more than 100 years ago by Nery Delgado. Its demise was caused by

human persecution and it went extinct in the XIX century (Mourer-Chauviré & Antunes, 1991;

Pimenta et al., 2008).

12

Figure 1.15: Pinguinus impennis proximal humerus from Furninha in lateral view. Geological Museum, Lisbon.

Finally there is a reference of Corvus antecorax Mourer-Chauviré, 1975 (the ancestor of the crow) in

the Almonda Cave and other of Alectoris barbara (Bonnaterre, 1791) in Gruta Nova da Columbeira

that became restricted to North Africa after the Pleistocene and have been reintroduced in Portugal, in

contrast with the other regionally extinct species that are linked towards a colder environment

(Figueiredo, 2010).

The study of Quaternary micromammals in Portugal is less developed than in Spain. Most of these

studies are centered on relatively recent archeological contexts (Morales & Rodríguez, 2009), and

although other go into the Pleistocene almost always they revolve around the archaezoological

question, not usually going deep into the paleoclimatic reconstruction or the past distribution of the

species of micromammals (Manne et al., 2006).

The “insectivores” of Portugal during the Quaternary are represented by at least seven species. The

hedgehog, Erinaceus europaeus Linnaeus, 1758 is represented mainly by an extraordinary array of

321 bones, belonging to at least 40 individuals from Furninha (Figure 1.16), also it is known by two

teeth from Figueira Brava cave, and some more remains from Lagar Velho and Lapa do Picareiro

(Mein & Antunes, 2000; Moreno-García & Pimenta, 2002; Bicho et al., 2003; Brugal et al., 2012).

Crocidura russula Hermann, 1780 is the most common member of this group, that appears in Goldra,

Guia, Figueira Brava, Lapa do Picareiro, Lagar Velho and Bom Santo Cave (Antunes et al., 1986b;

Antunes et al. , 1989; Mein & Antunes, 2000; Moreno-García & Pimenta, 2002; Bicho et al., 2003;

Pimenta, 2014). Crocidura suaveolens Pallas, 1811 is present in Figueira Brava cave and Lapa do

Picareiro (Mein & Antunes, 2000; Bicho et al., 2003). Undetermined species of the genus Crocidura

Wagler and Sorex Linnaeus, 1758 are presented in the sites of Gruta da Aroeira (Almonda System)

and Lagar Velho (Moreno-García & Pimenta, 2002; López-García et al., 2018). Sorex araneus

Linnaeus, 1758 is identified in Goldra (Antunes et al., 1986b). Talpa occidentalis Cabrera, 1907 is

known from Figueira Brava cave, Lapa do Picareiro and Bom Santo Cave (Mein & Antunes, 2000;

Bicho et al., 2003; Pimenta, 2014), Talpa europea Linnaeus, 1758 is cited in Goldra (Antunes et al.,

1986b) and an unidentified Talpa Linnaeus, 1758 species is known from Lagar Velho, although this

last two probably belong to the first species. Finally Galemys kormosi (Shereuder, 1940) is the only

extinct insectivore that has been found during this time period in Portugal, concretely by one upper

third premolar from the site of Morgadinho (Antunes et al., 1986a), its relative Galemys pyrenaicus

Geoffroy, 1811 that still exists today in the North of Portugal has been discovered in Lapa do Picareiro

(Bicho et al., 2003).

13

Figure 1.16: Right hemimandible of Erinaceus europaeus from Furninha in lingual view. Geological Museum,

Lisbon.

The fossil Chiroptera Blumenbach, 1779 have mostly been studied in detail in Portugal in the Figueira

Brava cave, where seven species are recognized: Rinolophus ferrumequinun (Shereber, 1774),

Rinolophus hipposideros (Bechstein, 1800), Rinolophus euryale Blasius, 1853; Myotis myotis

(Borkhausen, 1797), Myotis blythi (Tomes, 1857), Myotis nattereri (Khul, 1818) and Miniopterus

schreibersi (Khul, 1819). From Gruta da Aroeira (Almonda System) five species are cited: Myotis

myotis, another undetermined Myotis Borkhausen, 1797; Rinolophus ferrumequinun and another

undetermined Rhinolophus Lacepede, 1799 plus possibly Miniopterus schreibersi. Then we have other

two species recognized in Lapa do Picareiro: Myotis myotis and Myotis mystacinus Khul, 1817. There

is also one indeterminate bat from Bom Santo Cave (Mein & Antunes, 2000; Bicho et al., 2003;

Pimenta, 2014, López-García et al., 2018).

The rodents (figure 1.17) are the most common micromammals from the Quaternary of Portugal, with

at least 20 species known in eleven localities across Portugal (Morgadinho, Gruta da Aroeira, Guia,

Vale Boi, Bom Santo Cave, Figueira Brava, Galería Pesada, Lagar Velho, Lapa do Suão, Lapa do

Picareiro and Caldeirão).

Figure 1.17: Rodenia incisor recovered at the site of Santa Margarida (Portugal). FCT-UNL.

The Muridae Illiger, 1811 are represented by at least four species. Apodemus sylvaticus (Linnaeus,

1758) is the most common one, being recognized in Goldra, Guia, Caldeirão, Bom Santo (Figure

1.15), Lagar Velho and Lapa do Picareiro; meanwhile the bigger Apodemus flavicollis (Melchior,

1834) is present in Figueira Brava Cave and Gruta da Aroeira. There are also two other species more

closely related to human activity; the house mouse, Mus musculus Linnaeus, 1758 that appears in Guia

and Figueira Brava; and the black rat, Rattus rattus Linnaeus, 1758 that is found in Bom Santo Cave

and Figueira Brava. Their presence in the latter site is due to the inclusion of more recent sediments in

the Pleistocene deposits. Finally there is an indeterminate Muridae in the site of Morgadinho (Antunes

et al., 1986a; Antunes et al., 1986b; Antunes et al., 1989; Póvoas et al., 1992; Jeannet, 2000; Moreno-

García & Pimenta, 2002; Bicho et al., 2003; Pimenta, 2014; López-García et al., 2018).

Sciuridae Fischer de Waldheim, 1817 is only represented by one species: Sciurus vulgaris Linnaeus,

1758 that appear in Figueira Brava and Lagar Velho (Jeannet, 2000; Moreno-García & Pimenta,

2002). Gliridae Muirhead in Brewster, 1819 is represented by two species: Eliomys quercinus

Linnaeus, 1766 that appear in Gruta da Aroeira (Almonda System), Goldra, Guia, Caldeirão, Bom

Santo, Figueira Brava and Lapa do Suão and Glis glis (Linnaeus, 1766) that only appear in Lapa do

Picareiro. There is also one indeterminate glirid from Lagar Velho (Antunes et al., 1986b; Antunes et

al., 1989; Póvoas et al., 1992; Jeannet, 2000; Moreno-García & Pimenta, 2002; Bicho et al., 2003;

Valente & Haws, 2006; Pimenta, 2014, López-García et al., 2018).

14

The Cricetidae Fischer, 1817 is the most common family of rodents in Portugal, being represented by

at least fifteen species: Allocricetus bursae Schaub., 1930 is an extinct species of rodent closely related

to actual hamsters that has been found in Gruta da Aroeira (Almonda System) and Caldeirão (Póvoas

et al., 1992; López-García et al., 2018). Complete teeth of another extinct rodent, a species of

Mimomys Forsyth-Major, 1902 has been located in the site of Morgadinho (Antunes et al., 1986 a).

Pliomys episcopalis Méhely, 1914 is yet another extinct member of this group cited in the middle

Pleistocene sites of Galería Pesada and Gruta da Aroeira, in the Almonda system (Marks et al., 2002a;

López-García et al., 2018) . There is also a strange reference of a Myodes Pallas, 1811 from an

unknown locality in Atouguia da Baleia (Nehring, 1899). Chionomys nivalis Martins, 1842 appears in

Caldeirão and Lapa do Picareiro, but is no longer present in Portugal since it is related to colder

conditions (Póvoas et al., 1992; Bicho et al., 2003). At least 3 species of water voles have been

recorded in Portugal: Arvicola sapidus Miller, 1908 is the most common one appearing in Caldeirão,

Figueira Brava and Lagar Velho. Arvicola amphibious (Linnaeus, 1758) (Formerly known as Arvicola

terrestris) and Arvicola cantiana Hinton, 1910, the probable ancestor of A. sapidus, appear in Lapa do

Suão and Figueira Brava respectively (Póvoas et al., 1992; Jeannet, 2000; Moreno-García & Pimenta,

2002; Valente & Haws, 2006).

The genus Microtus Schrank, 1798 and /or Iberomys Chaline, 1972 is represented by at least 6 species

from 9 localities in Portugal that will be treated in the Table 1.1.

Table 1.1: Presence of different species of Microtus/Iberomys in different Quaternary sites in Portugal, data

from Antunes et al., 1986b, Antunes et al., 1989; Póvoas et al., 1992; Jeannet, 2000; Moreno-García & Pimenta,

2002; Bicho et al., 2003; Valente & Haws, 2006; Pimenta, 2014, López-García et al., 2018).

Figueira Brava Caldeirão

Bom Santo

Guia Lapa do

Suão Lapa do Picareiro

Lagar Velho

Goldra Gruta da Aroeira

TOTAL

Microtus lusitanicus (Gerbe,

1879) X X X X

X

5

Microtus duodecimcostatus (de Selys-Longchamps,

1839)

X X X

X X X

6

Microtus arvalis (Pallas,

1778) X X

X

3

Microtus agrestis(Linnaeus,

1761) X

X

2

Iberomys brecciensis

(Forsyth Major, 1905) X X

X

X X 5

Iberomys cabrerae

(Thomas, 1906) X X

2

Microtus sp. X X

2

TOTAL 4 7 3 1 1 2 4 2 1

Several pairs of species of Microtus are difficult to tell apart from each other, being the pair M.

lusitanicus/M. duodecimcostatus the most common one. It should be also addressed that the most

common rodent taxa today (Mus, Apodemus, Rattus, etc) appear to be under-represented in the fossil

record, probably because of their “lack of scientific interest” that have caused them to be neglected

sometimes in the literature. To end with rodents, the beaver, Castor fiber also appear in Portugal,

namely in Caldeirão cave (Antunes, 1989a).

The lagomorphs are overwhelmingly represented in Portugal by the Oryctolagus cuniculus (Linnaeus,

1758), the European rabbit. It appears in all the major Quaternary sites including hundreds or

thousands of bones that are a testimony of its hunt by ancient human populations (Hockett & Bicho,

2000). The hares, Lepus Linnaeus 1758, seem to be way less abundant but are mentioned in Caldeirão

and Pego do Diabo (Davis, 2002; Valente, 2004) and some tracks in the eolianites of the South of

Portugal are attributed to hares, with a new ichnospecies being described: Leporidichnites malhaoi

(Neto de Carvalho, 2010). Also from Algarve, from the old site of Morgadinho (Almost 1 million

YBP) comes a Mediterranean pika of the genus Prolagus Pomel, 1853 probably P. calpensis Major,

1905 (Antunes et al., 1986a). In the nearby and probably contemporaneous site of Algoz is cited also

Oryctolagus lacosti (Pomel, 1853) (Antunes et al., 1986c).

15

The only non human primate known in the Pleistocene of Portugal is Macaca sylvanus (Linnaeus,

1758) that has been recognized in the Galería Pesada of the Almonda system during the middle

Pleistocene (Marks et al., 2002b).

Now we will start to discuss the big mammals of the Quaternary from Portugal, an area that has been

better covered than the microvertebrates or the bird fauna since the XIX century, as we have seen in

the previous section.

Besides an African elephant (Loxodonta africana Blumenbach, 1797) lamella and other ivory

elements brought to Portugal by humans from Africa since the middle Holocene (Schuhmacher et al.,

2009) two species of proboscideans have been cited in the Quaternary of Portugal: Palaeoloxodon

antiquus and Mammuthus primigenius.

The straight tusked elephant was cited already in the XIX century by Paul Choffat, who found a molar

(Figure 1.18) in the calcareous tuffs of Condeixa (Choffat, 1895). Most of the remains known until

today are dental remains, like the tusks known from Meirinha, Algés and Condeixa or the molars or

molar laminae found in Furninha, Almonda, Carregado, Mealhada, the before mentioned Condeixa

molar and Foz do Enxarrique; this last one was regarded as the most recent known remain for the

species with around 30.000 YBP; and although new radiometric datings (Cunha et al., 2019) have

pushed back this date to around 44.000 YBP it is still the most recent known remain for this species.

Other bone remains include several fragments of ribs, tibia and humerus from Mealhada, an unciform

from Santa Cruz and a set of bones coming from Santo Antão do Tojal (Femur, tibia, phalanx…etc)

that will be treated later in this work (Antunes & Cardoso, 1992; Figueiredo, 2012).

The Mammuthus primigenius presence is well more cryptic. There is a fragment of femur from Algar

de João Ramos that is badly damaged, but its C14 dating of 14000 YBP make its attribution only

possible to this species, being it also the most recent proboscidean remain from Portugal (Antunes &

Cardoso, 1992). Also two molar lamellas are known, one from Figueira Brava cave and another one

that was recovered by a fisherman from the ocean bottom in front of the coast of Oeiras (Cardoso &

Regala, 2002). Also there are some tracks of proboscideans conserved in the Pleistocene eolianites

from South Portugal, in Pessegueiro Island (Neto de Carvalho, 2010).

Figure 1.18: Molar of Palaeoloxodon antiquus (MG 3561) recovered by Paul Choffat in Condeixa during the

XIX century in lateral view.

16

Rhinoceroses were identified for first time in the Quaternary of Portugal by Nery Delgado, from the

material of Furninha cave (Delgado, 1884). Until now it seems that only one species is known in this

time period in Portugal: Stephanorhinus hemitoechus (Falconer, 1859), mostly thanks to several dental

specimens (Figure 1.19) but also some postcranial (two metatarsals, four astragalus and one

calcaneus). The species is known only from cave material and appears in the localities of Serra dos

Molianos, Furninha, Gruta Nova da Columbeira, Lapa da Rainha, Lorga de Dine, Correio-Mor,

Figueira Brava, Pedreira das Salemas and Escoural, only between 30000 and 20000 YBP (Veiga

Ferreira, 1975; Cardoso, 1993a).

Figure 1.19: Partial molar of Stephanorhinus hemitoechus from Gruta da Serra de Molianos. Geological

Museum, Lisbon.

Other Perissodactyla, Owen 1848, include three species of equids that have been recognized in the

Quaternary of Portugal: Equus hydruntinus, Equus ferus (Linnaeus, 1758) and Equus africanus

(Linnaeus, 1758). The later one arrived to Portugal thanks to human introduction around 4000 YBP,

1500 years earlier than previously thought (Cardoso et al., 2013) but was absent of the country for all

the Pleistocene.

Equus ferus has a native subspecies described in Portugal: Equus ferus antunesi; whose type specimen

(Figure 1.20) comes from Fointainhas cave. It was represented by a superbly preserved skull and some

other limb bones that allow reconstructing this subspecies like a slender and small horse of 140 cm to

the withers (Cardoso & Eisenmann, 1989). Other material that cannot be compared with the type of E.

f. antunesi appear in many Portuguese sites and has been simply attributed to Equus ferus appearing

quite commonly in the Portuguese records (mostly dental material), from the prewürmian sites of

Mealhada to the archaeological contexts of Furninha, Vale Boi and Foz do Enxarrique to the

relatively modern sites like Casais Robustos (Cardoso, 1993a; Raposo, 1995; Manne et al., 2012).

Equus hydruntinus appear at least in Pedreira das Salemas, Caldeirão cave and the site of Vale Boi

(Cardoso, 1995; Davis, 2002; Manne et al., 2012). If the “zebro” of medieval chronicles is this

species, it only disappeared in Portugal during the XV century (Antunes, 2006).

Figure 1.20: Type specimen of Equus ferus antunesi in lateral view (MG). Geological Museum, Lisbon.

17

There is no big associated or articulated cetacean skeleton from geological contexts during this period

in Portugal, but several bones collected in archaeological contexts have been recovered, some of them

only centuries old, that are testimony of the hunt of whales during medieval and modern times

(Teixeira et al., 2014). In similar contexts but way older we have a cetacean bone from the site of Vale

Boi, Algarve, with around 28000 years (Manne et al., 2012) a possible cetacean bone from Lagar

Velho (Moreno-García & Pimenta, 2002) and six vertebrae ascribed to Delphinus delphis Linnaeus,

1758 from Figueira Brava with around 30.000 years (Antunes, 2000).

The hippopotamuses are known in Portugal in three localities: Condeixa, Mealhada and Algoz.

The remains of Condeixa (Figure 1.21) are dental and mandibular remains (a fragment of inferior

mandible, a molar and an isolated canine) already collected by Paul Choffat in the XIX century

(Choffat, 1895). In Mealhada there is also a left cuboid (Antunes et al., 1988). Both have been

attributed to Hippopotamus incognitus Faure, 1884 by Cardoso, 1993 although some authors

considered it a synonym of Hippopotamus amphibious Linnaeus, 1758 (Petronio, 1995).

The remains from Algoz are the better preserved ones, including one femur head, a distal part of a

tibia, a calcaneum, an incomplete astragalus; a fragmentary cuboid and navicular, a fourth left

metatarsal and two phalanges that are regarded as belonging to the same animal. They had been

attributed to Hippopotamus major Cuvier, 1825 (Antunes et al., 1986c) although by priority rule they

should be named Hippopotamus antiquus Desmarets, 1822 (Magri et al., 2010).

Figure 1.21: Anterior mandibular fragment of Hippopotamus incognitus from Condeixa in dorsal view.

Geological Museum, Lisbon.

Most authors agree that nearly all of the suid remains on Europe from the Pleistocene belong to the

extant species Sus scrofa Linnaeus, 1758; although the presence of Sus strozzi Forsyth Major, 1881 in

the Early Pleistocene of Europe has been proved (Alcalá et al., 1985). Sus scrofa is divided nowadays

in five subspecies across Europe, but the distinction at sub specific level in the fossil record (mostly

based on dental remains and size, like for example the subspecies S. s. priscus) is specially

problematic (Cardoso, 1993a; Lister et al., 2010).

In Portugal, although this species is nowadays in expansion and quite common (Sáez-Royuela &

Tellería, 1986) its remains from the Upper Pleistocene and Holocene are rare. It is known in sites like

Fontainhas, Pedreira e Gruta das Salemas, Escoural, Lapa do Picareiro, Figueira Brava and Caldeirão

(Cardoso, 1993a; Hockett & Haws, 2009). The later ones have been suggested to belong to domestic

animals (Davis, 2002).

18

The red deer (Cervus elaphus Linnaeus, 1758) (Figure 1.22) is probably the most common big

mammal from the Pleistocene and Holocene of Portugal. It is known since the very first excavation of

Furninha (Delgado, 1884), spanning from the oldest sites of Mealhada (Antunes et al., 1988) to

Holocene sites in Archaeological contexts (Dean & Carvalho, 2011). It seems that there is a smaller

morphotype in the Würm of Portugal (Cardoso, 1993a) and genetic studies of recent deer have pointed

towards the genetic separation of European and Iberian red deer during the LGM (Queirós et al.,

2019); therefore given the abundance of materials, ancient DNA analysis of the red deer fossil

occurrences would be interesting.

The roe deer (Capreolus capreolus Linnaeus, 1758) is rare, being it known from the caves of

Caldeirão, Columbeira, Lapa da Rainha, Picareiro and Porto Covo (Cascais) the later ones probably

from Holocene (Cardoso, 1993a, Hockett & Haws, 2009).

The fallow deer (Dama dama Linnaeus, 1758) has been recognized in the Algar de João Ramos by an

inferior tooth, Gruta Nova da Columbeira with dental series and Pedreira das Salemas by a first

phalanx. The chronology of all the Pleistocene remains is younger than 30000 and older than 14000

YBP (Cardoso, 1989). There are not known remains of this species of Pre-Roman Holocene time,

coming the oldest of this period from the Roman sites of São Pedro da Fronteira and Torre de Palma,

so it is possible that Pleistocene remains belonged to a different species of the genus Dama (Davis &

MacKinnon, 2009). An extinct species of the genus; Dama vallonnetensis De Lumley et al., 1988, is

cited from the older middle Pleistocene deposits in the Almonda system (Marks, Monigal, et al.,

2002).

Eucladoceros sp. Falconer, 1868 has also been cited from the lowermost Middle Pleistocene of Algoz,

Algarve (Antunes et al., 1986c).

Figure 1.22: Skull of female Cervus elaphus from Gruta das Fontainhas in lateral view. Geological Museum,

Lisbon.

The bovids in Portugal during the Quaternary are represented by the auroch, Bos primigenius Bojanus,

1827; the Iberian ibex, Capra pyrenaica Schinz, 1838; the Pyrenean chamois, Rupicapra pyrenaica

Bonaparte, 1845 and the sheep/mouflon Ovis orientalis Linnaeus, 1758; arriving the last one by

human hand to Portugal in the Holocene only 3000 thousands year after its domestication, 5450 YBP

(Davis & Simões, 2016).

The auroch, Bos primigenius, is common in Portugal, appearing in all major Quaternary sites. It is

necessary to remark that the difficulties to differentiate this species from members of Bison Hamilton

Smith, 1827 make the former´s presence in Portugal probable, although not proved, because not

19

enough well preserved material have been recovered (the easiest way to differentiate between them is

from cranial material, quite scarce), (Cardoso, 1993a).

The Iberian ibex, Capra pyrenaica, is an endemism of the Iberian Peninsula. The subspecies typical of

Portugal, Capra pyrenaica lusitanica went extinct at the end of the XIX century but nowadays the

species is again living in Portugal (Moço et al., 2006). It is known from sites like Fontainhas,

Caldeirão (Figure 1.23), Casais Robustos (that we will discuss later), Pedreira das Salemas, Figueira

Brava, Escoural, Pego de Diablo, Algar of João Ramos or Vale Boi (Cardoso, 1993a, Manne et al.,

2012).

The Pyrenean chamois, Rupicapra pyrenaica is known in Portugal from the localities of Gruta das

Salemas, Caldeirão, Pego do Diabo and Lapa do Picareiro (Cardoso, 1993a; Bicho et al., 2000). The

fossil population of Portugal was regarded as a subspecies of the chamois Rupicapra rupicapra

pyrenaica (Linnaeus ,1758) but further genetic investigation revealed that the animals living in the

Cantabric mountains, the Pyrenees and the Apennines have been separated for around 30000 years

from the main population in the Alps, that came from the East during the Würm, so the specimens

from Portugal probably belonged to Rupicapra pyrenaica (Pérez et al., 2002).

Figure 1.23: Distal part of a humerus of Capra pyrenaica in anterior view from Caldeirão cave. Catalog number

and scale of the specimen not provided in the original (Davis, 2002).

Concerning Carnivora Bowdich, 1821 the pinnipeds are represented by one right ulna from Figueira

Brava attributed to Pusa hispida (Schereber, 1775), (Antunes, 2000) that today lives in the high Arctic,

in locations like Svalbard (Krafft et al., 2007). There is also a part of a mandible at exposition in the

Geological Museum in Lisbon that is labeled as “Foca (Phocus monachus)”, but it does not appear to

be in the bibliography.

Mustelidae Fischer de Waldheim, 1817 is present in the Quaternary of Portugal. In the site of Vale Boi

there is a reference to the appearance of Mustela Linnaeus, 1758. Martes Pinel, 1792 appear in Vale

Boi and Furninha and the badger, Meles meles Linnaeus, 1758 appear in Furninha, Caldeirão and Lapa

do Suão (Davis, 2002; Valente & Haws, 2006; Brugal et al., 2012; Manne et al., 2012).

Three canid species have been recognized from this time period in Portugal: Canis lupus Linnaeus,

1758, Vulpes vulpes (Linnaeus, 1758) and Cuon alpinus Pallas, 1811. The first two are relatively

common, being known since the XIX century from diverse Portuguese caves, like Furninha,

Fontainhas or Caldeirão, but Cuon alpinus is only known in Portugal thanks to a fragment of

hemimandible from the Escoural cave (Figure 1.24) ( Cardoso, 1993a), although studies suggest that

many Cuon remains could be misidentified in late Pleistocene and Holocene contexts so it would be

risky to assess the relative abundance of that species compared to the other two (Ripoll et al., 2010).

20

Figure 1.24: Fragment of left hemimandible of Cuon alpinus with P4 and fragment of M1 from Escoural cave in

oclusal (left) and labial (right) views. Catalog number of the specimen or not provided in the original, scale do

not appear in the image (Cardoso, 1993a).

At least three works until this date agree in the conclusion that there is only one species of ursid

recognized from the Quaternary of Portugal: the brown bear, Ursus arctos Linnaeus, 1758 (Harlé,

1910; Torres, 1979; Cardoso, 1993a). There are bear remains known in most of major Portuguese

Quaternary sites: Furninha (Figure 1.25), Fontainhas, Molianos, Pedreira and Gruta das Salemas,

Algar de Cascais, Figueira Brava, Gruta de Almonda (Galería Pesada), Caldeirão and the hesperic

massif sites of Lorga de Dine and Escoural, although it is only abundant in the two first sites. This is

not a surprise given the affinity of this species for caves during winter periods (Cardoso, 1993a; Marks

et al., 2002a; Davis, 2002). Besides this locations it has been detected in recent Holocene

archaeological contexts, for example in Leceias (Pires, 2002). It is also worth to mention that the cave

bear Ursus spelaeus Rosenmüller, 1794 in known to have lived near the Portuguese territory in the

Galician cave of Rebolal, only 50 km away from Portugal (Grandal-d’Anglade et al., 2006) .

Figure 1.25: Left Ursus arctos mandible from Furninha cave in lingual view. Geological Museum, Lisbon.

Two species of hyenas have been cited in the Pleistocene of Portugal: Hyaena hyaena (Linnaeus,

1758) and Crocuta crocuta (Erxleben, 1777).

Hyaena hyaena prisca de Serres, 1828 is cited from Furninha cave with some abundance (Figure

1.26), which is curious given that the species is not cited in the Pleistocene of Spain, meanwhile it

appears in France. It seems that it survived longer in Portugal that in other locations of its distribution

(Cardoso, 1996; Brugal et al., 2012).

21

Crocuta crocuta spelaea (Goldfuss, 1832) is more common, appearing in at least ten sites in the

Pleistocene of Portugal including Caldeirão or Pego do Diabo (Cardoso, 1996; Valente, 2004).

There is also a mention to the presence of Crocuta crocuta intermedia, an older and smaller subspecies

of spotted hyena in Lorga de Dine (Cardoso, 1994).

Figure 1.26: Skull of Hyaena hyaena prisca from Furninha cave in dorsal view. Geological Museum, Lisbon.

Felidae Fischer, 1817 is represented in the Quaternary of Portugal by at least five species: Felis

silvestris Schreber, 1777, Panthera spelaea, Panthera pardus Linnaeus, 1758, at least one species of

Lynx Kerr, 1792, and Homotherium latidens (Cardoso, 1993a; Antunes, 1986).

Felis silvestris in known from at least ten sites of this chronology in Portugal (Villaluenga, 2016), in

more recent archaeological contexts (younger than 10000 YBP) the specimens could belong to

domestic cat (Baca et al., 2018).

The lynx is the most common felid in the Pleistocene of Portugal, appearing at least in more than

eleven sites (Villaluenga, 2016) including a new one that will be presented later in this work. If all that

remains are from one or various species of lynx is a question that is not clear. Until now all of them

have been attributed either to the extant endangered Lynx pardinus Temminck, 1827 or to Lynx

spelaea (Boulé, 1906), being the latter regarded as a junior synonym of the former by some (Boscaini

et al., 2016). The possible presence of European lynx, Lynx lynx (Linnaeus, 1758) cannot be ruled out

under the light of new works that have proved its presence in the Iberian Peninsula in the Holocene

(Rodríguez-Varela et al., 2016). There are also tracks attributed to lynxes in the eolianites of the South

of Portugal (Neto de Carvalho, 2014).

The leopard, Panthera pardus, is present in twelve sites of the Quaternary of Portugal (Cardoso &

Regala, 2006; Villaluenga, 2016) including one mostly complete and well preserved skull from the

Algar da Manga Larga (Figure 1.27).

The cave lion, Panthera spelaea is known by scarce material from seven localities in Portugal: Vale

Boi (non specified remains), Pedreira das Salemas (three canine teeth and one phalanx), Gruta de

Penacova (one canine teeth), Lorga de Dine (one fourth right metacarpal), Escoural (one third

premolar), Caldeirão (one phalanx) and Figueira Brava (one phalanx) (Antunes & Cardoso, 1986;

Cardoso, 1993a; Manne et al., 2012; Villaluenga, 2016).

Homotherium latidens has been identified in the Pleistocene of Portugal by one left astragalus in the

Mealhada site, probably older than 100000 YBP (Antunes, 1986).

22

Figure 1.27: The leopard (Panthera pardus) skull from Algar da Manga Larga in anterolateral view. Geological

Museum, Lisbon.

1.3 Aim, context and objectives of the present thesis.

This thesis will focus in Pleistocene tetrapods (with special emphasis on macro-mammals), although

some Holocene material will be included on it:

1. Paleodiversity census. Compile a state of the art census of the fossil Quaternary tetrapods

known in Portugal across the literature, with a list of species, localities and publications.

Quantify bias in the fossil record, the spatial distribution of the sites, extinction rates and

differences between groups (What groups suffered more severe extinctions? Which are the

better represented in the fossil record?) Establish the quality of the fossil record, i.e., what

percentage of biodiversity is represented in the fossil record.

2. The Palaeoloxodon antiquus of Santo Antão do Tojal: Estimate for first time with the most

up to date numerical methods the body mass of a Portuguese proboscidean and study its

implications. Perform a metric study with a dataset of more than 40 Eurasian Palaeoloxodon

antiquus and Mammuthus primigenius (including the Santo Antão do Tojal specimen) to

determine metric differences between the two species.

3. The bears of Algar do Vale da Pena: Excavation and description of the new locality and

recovered bear materials. Compare the skeletal measurements of the recovered remains to with

other Iberian populations. Provide a specific determination. Asses the implications of this new

information in the context of Portugal.

4. The microvertebrates of Santa Margarida: Describe the new locality. Elaborate a list of

fauna. Provide a chronology for the site.

Given this objectives the thesis will be separated in 4 sections according to each one of them.

23

2. PALEOBIODIVERSITY OF THE QUATERNARY FOSSIL

TETRAPODS IN CONTINENTAL PORTUGAL

2.1 Introduction and methods

On this chapter an inventory of localities and tetrapod species recognized in the Quaternary of

Portugal will be provided in order to give us an approach to the paleobiodiversity of the territory. For

each locality some general information will be given. For each group a table will be provided

including the species name, the localities where it has been recognized and the references to that

occurrence, also specifying if the species is found today in Portugal or not. Then a resume table with

some numerical considerations for each group will be given, like the number and percentage of extinct

species in the sample, the number and percentage of species locally extinct in Portugal and the number

and percentage of species of the group that are only known as fossils (Species extinct + Species locally

extinct). Also there will be the number and percentage of species detected and undetected in the fossil

record for each group, in comparison with the species that live today in Portugal. Then there will be a

general overview, providing some comparison between groups, some numerical examples about the

importance of the cave deposits in the Quaternary record and some inferences about the

paleobiodiversity of other periods. This section has been partially published in an abtract and

presented in the 1st Paleontological Virtual Congress (Estraviz-López & Mateus, 2018a).

2.2 Lists of localities and fossil tetrapods of the Quaternary of Portugal

In the following table 2.1 a list of the principal Portuguese Quaternary localities will be provided,

including name, coordinates, municipality, approximate age(s) and some references. It should be noted

that not all the sites are on it; some minor localities where isolated remains of elephants or other fauna

have been found are not listed, like Meirinha, Carregado or Porto Covo (Cascais). Also the coordinates

of some localities are not totally precise, since they were not explicitly pointed in the literature.

Table 2.1: Localities of the Quaternary of Portugal.

NAME

MUNICIPALIT

Y COORDINATES AGE REFERENCES

Lorga de Dine Vinhais

41° 54' 35" N

6° 55' 49" W

Late Pleistocene Pre

Würm (80.000 YBP?)

and mainly Würm

(20.000 YBP)

Cardoso, 1994;

Cardoso, 1996

Mealhada Mealhada

40° 22' 35" N

8° 27' 30" W

Middle Pleistocene

(interglacial Mindel-

Riss?)

Zbyszewski,

1977a; Antunes

et al., 1988

Condeixa

Condeixa-a-

Nova

40° 05' 49" N

8° 29' 59" W

Middle Pleistocene

(Mindel?)

Choffat, 1895;

Cardoso, 1993a

Caldeirão Tomar

39° 38' 59" N

8° 24' 54" W 30.000 - 5.700 YBP

Cardoso,

1993a; Davis,

2002

Foz do

Enxarrique

Vila Velha de

Ródão

39° 38' 58" N

7° 40' 11" W 140.000-35.000 YBP

Antunes &

Cardoso, 1989;

Raposo, 1995;

Cunha et al.,

2019

Lapa do

Picareiro Fátima

39° 32' 33" N

8° 38' 02" W 12.000 - 6.500 YBP

Bicho et al.,

2006

Almonda

System Torres Novas

39° 30' 19" N

8° 36' 58" W

Middle Pleistocene

(~250.000 YBP)

Marks et al.,

2002 a; López-

García et al.,

24

2018

Casais Robustos Alcanena

39° 30' 12" N

8° 40' 20" W Recent Würm?

Cardoso,

1993a;

Estraviz-López

& Mateus,

2018c. Gruta da Serra

de Molianos Alcobaça Unknown Recent Würm? Cardoso, 1993a Algar de João

Ramos Alcobaça

39° 23' 57" N

8° 56' 10" W ~15.000 YBP Cardoso, 1993a

Furninha Peniche

39° 21' 23" N

9° 24' 14" W

Aprox 80.000 YBP(But

with great uncertainty)

Brugal et al.,

2012

Casa da Moura Óbidos

39° 19' 36" N

9° 15' 14" W ~25.000 YBP Cardoso, 1993a Gruta Nova da

Columbeira Bombarral

39° 18' 06" N

9° 11' 58" W ~25.000 YBP Cardoso, 1993a

Lapa do Suão Bombarral

39° 17' 59" N

9° 12' 01" W 15.000 - 10.000 YBP

Valente &

Haws, 2006

Fontainhas Cadaval

39° 11' 37" N

9° 02' 39" W ~22.000 YBP Cardoso, 1993a

Lapa da Rainha Torres Vedras

39° 10' 51" N

9° 19' 38" W 25.000 - 20.000 YBP Cardoso, 1997

Bom Santo Cave Alenquer

39° 10' 29" N

9° 02' 02" W 5.800 - 5.400 YBP Pimenta, 2014 Pedreira e Gruta

das Salemas Loures

38° 52' 38" N

9° 11' 58" W 30.000 -20.000 YBP Cardoso, 1993a

Pego do Diabo Loures

38° 51' 50" N

9° 13' 00" W 28.000 - 23.000 YBP

Cardoso,

1993a; Valente,

2004 Santo Antão do

Tojal Loures

38° 50' 26" N 9°

8' 37" W 80.000 YBP Raposo, 1995

Correio-Mor Loures

38° 49' 44" N

9° 10' 50" W Recent Würm Cardoso, 1993a

Porto Covo Cascais

38° 45' 06" N

9° 25' 23" W Recent Würm Cardoso, 1993a

Algar de Cascais Cascais

38° 42' 10" N

9° 25' 08" W 18.000 YBP Cardoso, 1993a

Escoural

Motemor-o-

Novo

38° 32' 59" N

8° 08' 15" W 20.000-15.000 YBP Cardoso, 1993a

Figueira Brava Setúbal

38° 28' 23" N

8° 59' 04" W 30.000 YBP Antunes, 2000

Algoz Silves

37° 09' 07" N

8° 17' 51" W Nearly 1 million YBP

Antunes et al.,

1986c

Goldra Loulé

37° 07' 38" N

7° 59' 19" W Early Würm

Antunes et al.,

1986b

Guia Albufeira

37° 07' 13" N

8° 17' 43" W

Late Pleistocene-Early

Holocene

Antunes et al.,

1989

Morgadinho Tavira

37° 05' 48" N

7° 42' 43" W Nearly 1 million YBP

Antunes et al.,

1986a

Vale Boi Vila do Bispo

37° 05' 25" N

8° 48' 26" W

Mostly 27.000-17.000

YBP (8.000 YBP)

Manne et al.,

2012

25

Figure 2.1: Satellital picture showing the location of 30 sites with Quaternary fossil vertebrates in Portugal,

mentioned in the table 2.1. In black, the localities situated in the Lusitanian basin and in red the ones situated in

the Algarve basin. Credit Google Earth.

As we can see in figure 2.1.; practically all the localities are situated in two areas over de Mesozoic

basins of Portugal (Terrinha et al., 2006; Kullberg et al., 2014). The stretch of the Lusitanian basin

between Lisbon and Leiria (plus the Setúbal Peninsula) covers an area of 5000 square kilometers,

about 5,5% the surface of the country, yet it comprises 18 of the 30 localities considered (More than

50% of them). Most of the other localities are situated in the Algarve basin, although it is interesting to

note that most of them are situated in open air sites and not in caves, opposed to what happens in the

North. Only three localities are outside these main areas: Lorga de Dine (Northernmost point),

Escoural (Southernmost point outside the main areas) and Foz do Enxarrique (Between them). Those

two caves are the only ones whose fauna have been studied in detail in the Hesperian Massif of

Portugal and Foz do Enxarrique not only is outside the main areas of findings, but it is also an open air

site near the Tagus River. Only ten localities are situated in open air sites, most of them in Algarve.

26

Tables 2.2 (amphibians), 2.3 (reptiles), 2.4 (birds) and 2.5 (mammals) summarize the known

vertebrates from the Quaternary of Portugal. Some nomenclature is corrected, see previous chapter for

further information. Table 2.2: Amphibians of the Quaternary of Portugal.

TAXON LOCALITIES REFERENCES

EXTANT IN

PORTUGAL

?

Salamandra salamandra

(Linnaeus, 1758)

Lagar Velho, Figueira

Brava

Moreno-García &

Pimenta, 2002;

Crespo et al., 2000

YES

Triturus marmoratus

(Latreille, 1800) Lagar Velho

Moreno-García &

Pimenta, 2002

YES

Bufo bufo (Linnaeus, 1758) Lapa do Picareiro, Guia

Hockett & Haws,

2009; Antunes et al.,

1989

YES

Pelobates cultripes Cuvier,

1829

Lapa do Picareiro, Figueira

Brava, Guia

Hockett & Haws,

2009; Crespo et al.,

2000

YES

Pleurodeles waltl

(Michahelles, 1830) Guia Antunes et al., 1989

YES

Epidalea calamita Laurenti,

1758 Guia Antunes et al., 1989

YES

Pelophylax perezi (Seoane,

1885) Guia Antunes et al., 1989

YES

Table 2.3: Reptiles of the Quaternary of Portugal.


EXTANT IN

PORTUGAL?

Timon lepidus Daudin,

1802 Figueira Brava, Guia

Crespo et al., 2000,

Antunes et al., 1989

YES

Psammodromus manuelae

Busack et al., 2006 Figueira Brava Crespo et al., 2000

YES

Podarcis sp. Wagler, 1830 Figueira Brava Crespo et al., 2000

YES

Acanthodactylus

erythrurus (Schinz, 1833) Guia Antunes et al., 1989

YES

Rinechis scalaris (Schinz,

1822) Figueira Brava Crespo et al., 2000

YES

Hemorrhois hippocrepis

(Linnaeus, 1758) Figueira Brava Crespo et al., 2000

YES

Coronella girondica

Daudin, 1803

Gruta da Aroeira

(Almonda System),

Figueira Brava

Crespo et al., 2000;

López-García et al.,

2018

YES

Vipera lastei Bosca, 1878 Figueira Brava Crespo et al., 2000 YES

Natrix sp. Laurenti, 1758

Gruta da Aroeira

(Almonda System),

Guia

Antunes et al., 1989;

López-García et al.,

2018

YES

Blanus cinereus Vandelli,

1797 Figueira Brava Crespo et al., 2000

YES

Testudo hermanni Gmelin, Mealhada, Furninha, Lapparent de Broin &

27

1789 Figueira Brava, Gruta

Nova da Columbeira,

Caldeirão, Escoural,

Gruta de Ibn Amar

Antunes, 2000;

Morales Pérez &

Serra, 2009.

NO

Mauremys leprosa

(Schweiger, 1812)

Mealhada, Caldeirão,

Porto Covo (Cascais),

Gruta Nova da

Columbeira

Lapparent de Broin &

Antunes, 2000;

Morales Pérez &

Serra, 2009.

YES

Emys orbicularis

(Linnaeus, 1758)

Figueira Brava, Gruta

Nova da Columbeira

Lapparent de Broin &

Antunes, 2000;

Morales Pérez &

Serra, 2009.

YES

Table 2.4: Birds of the Quaternary of Portugal.


EXTANT IN

PORTUGAL?

Tetrao urogallus

Linnaeus, 1758 Lapa da Rainha Figueiredo, 2010

NO

Alectoris rufa

Linnaeus, 1758

Lapa do Picareiro, Furninha, Lapa

do Suão, Gruta Nova da

Columbeira, Escoural, Figueira

Brava, Bom Santo Cave Pimenta et al., 2015

YES

Alectoris barbara

(Bonnaterre, 1791) Gruta Nova da Columbeira Figueiredo, 2010

YES

Perdix perdix

Linnaeus, 1758

Lapa do Picareiro, Furninha, Gruta

Nova da Columbeira, Caldeirão

Figueiredo, 2010;

Pimenta et al., 2015

NO

Coturnix coturnix

Linnaeus, 1758

Furninha, Gruta Nova da

Columbeira Figueiredo, 2010

YES

Phasianus

colchicus Linnaeus,

1758 Gruta Nova da Columbeira Figueiredo, 2010

YES

Lagopus lagopus


NO

Lagopus muta

Montin, 1781 Gruta Nova da Columbeira Figueiredo, 2010

NO

Anas platyrhynchos

Linnaeus, 1758

Lapa do Suão, Escoural, Figueira

Brava Pimenta et al., 2015

YES

Anas acuta

Linnaeus, 1758 Gruta Nova da Columbeira Pimenta et al., 2015

YES

Anas crecca

Linnaeus, 1758 Furninha Pimenta et al., 2015

YES

Melanitta nigra

(Linnaeus, 1758) Furninha, Figueira Brava Pimenta et al., 2015

YES

Melanitta fusca

(Linnaeus, 1758) Figueira Brava Pimenta et al., 2015

YES

Clangula hyemalis

Linnaeus, 1758 Figueira Brava Pimenta et al., 2015

YES

Tadorna tadorna

(Linnaeus, 1758) Furninha Figueiredo, 2010

YES

Tadorna ferruginea

Pallas, 1761 Furninha Figueiredo, 2010

YES

Somateria Furninha Figueiredo, 2010

28

mollisima

(Linnaeus, 1758)

NO

Mergus albellus

(Linnaeus, 1758) Galería Pesada (Almonda) Figueiredo, 2010

YES

Anser albifrons

(Scopoli, 1769) Pego do Diabo Figueiredo, 2010

NO

Cygnus olor

(Gmelin, 1789) Galería Pesada (Almonda), Furninha Pimenta et al., 2015

NO

Scolopax rusticola

(Linnaeus, 1758)

Galería Pesada (Almonda), Figueira

Brava Pimenta et al., 2015

YES

Numeius phaeopus


YES

Calidris canutus

Linnaeus, 1758 Lapa do Picareiro, Figueira Brava Pimenta et al., 2015

YES

Larus fuscus


YES

Pinguinus impennis


NO

Alca torda


YES

Vanellus vanellus

(Linnaeus, 1758)

Gruta Nova da Columbeira,

Fontainhas Figueiredo, 2010

YES

Pluvialis

squatarola

(Linnaeus, 1758) Gruta Nova da Columbeira Figueiredo, 2010

YES

Pluvialis apricaria

(Linnaeus, 1758) Lapa do Picareiro Pimenta et al., 2015

YES

Podiceps nigricollis

Brehm, 1831 Figueira Brava Figueiredo, 2010

YES

Otis tarda

Linnaeus, 1758 Gruta de Casa da Moura Figueiredo, 2010

YES

Grus grus


YES

Grus primigenia

(Milne-Edwards,

1869) Figueira Brava Figueiredo, 2010

NO

Puffinus puffinus

(Brünnich, 1764)


Columbeira Pimenta et al., 2015

YES

Puffinus holeae

Walker et al., 1990 Figueira Brava Pimenta et al., 2015

NO

Phalacrocorax

aristotelis


YES

Phalacrocorax

carbo (Linnaeus,

1758) Figueira Brava Pimenta et al., 2015

YES

Morus bassanus


YES

Phoenicopterus

ruber Pallas, 1811 Furninha Figueiredo, 2010

YES

Gavia stellata

(Pontoppidan,

1763) Figueira Brava Pimenta et al., 2015

YES

29

Accipiter nisus


YES

Aquila chrysaetos

Linnaeus, 1758 Figueira Brava, Vale Boi Pimenta et al., 2015

YES

Aquila adalberti

Brehm, 1861 Lapa da Rainha Figueiredo, 2010

YES

Milvus migrans

(Boddaert, 1783) Figueira Brava Pimenta et al., 2015

YES

Falco tinnuncus

Linnaeus, 1758 Lagar Velho, Figueira Brava, Pimenta et al., 2015

YES

Falco rusticolus


NO

Aquila fasciata

(Viellot, 1822) Figueira Brava Pimenta et al., 2015

YES

Aegypius monachus

(Linnaeus, 1766) Caldeirão Figueiredo, 2010

YES

Gyps fulvus

Hablizl, 1783 Furninha, Caldeirão Pimenta et al., 2015

YES

Buteo buteo

(Linnaeus, 1758) Galería Pesada (Almonda) Figueiredo, 2010

YES

Haliaeetus albicilla

Linnaeus, 1758 Galería Pesada (Almonda) Pimenta et al., 2015

YES

Bubo bubo

(Linnaeus, 1758)

Galería Pesada (Almonda),

Caldeirão, Furninha, Figueira Brava Pimenta et al., 2015

YES

Athene noctua

Scopoli, 1769


Caldeirão, Lapa do Picareiro, Gruta

Nova da Columbeira, Lapa da

Rainha, Escoural, Figueira Brava Pimenta et al., 2015

YES

Strix aluco

Linnaeus, 1758 Gruta Nova da Columbeira Figueiredo, 2010

YES

Asio flammeus

(Potoppidan, 1763) Lapa do Picareiro, Furninha Pimenta et al., 2015

YES

Columba livia

Gmelin, 1789

Galería Pesada (Almonda), Lapa do

Picareiro, Furninha, Figueira Brava Pimenta et al., 2015

YES

Columba palumbus

Linnaeus, 1758


Caldeirão, Escoural, Figueira Brava Pimenta et al., 2015

YES

Coracias garrulus

Linnaeus, 1758

Lapa do Picareiro, Lapa do Suão,

Gruta Nova da Columbeira Pimenta et al., 2015

YES

Egretta sp. Forster,

1817 Galería Pesada (Almonda)

Figueiredo, 2010,


YES

Pyrrhocorax

pyrrhocorax

Linnaeus, 1758


Caldeirão, Lapa do Picareiro,

Furninha, Lapa do Suão, Gruta Nova

da Columbeira, Fontainhas, Lapa da

Rainha, Figueira Brava Pimenta et al., 2015

YES

Pyrrhocorax

graculus (Linnaeus,

1766)

Galería Pesada (Almonda), Lapa da

Rainha, Lapa do Picareiro, Furninha,

Gruta de Casa da Moura,

Fontainhas, Caldeirão, Gruta Nova

da Columbeira

Figueiredo, 2010,


NO

30

Corvus monedula

(Linnaeus, 1758)

Galería Pesada (Almonda), Lagar

Velho, Lapa do Picareiro, Furninha,

Lapa do Suão, Gruta Nova da

Columbeira, Fontainhas, Lapa da

Rainha, Figueira Brava Pimenta et al., 2015

YES

Corvus corax

(Linnaeus, 1758)

Caldeirão, Gruta Nova da

Columbeira, Lapa da Rainha, Gruta

de Casa da Moura

Figueiredo, 2010,


YES

Corvus corone

Linnaeus, 1758

Galería Pesada (Almonda), Lapa do

Picareiro, Furninha Pimenta et al., 2015

YES

Corvus frugilegus

Linnaeus, 1758 Furninha, Escoural Pimenta et al., 2015

YES

Corvus antecorax

Mourer-Chauviré,

1975 Galería Pesada (Almonda) Figueiredo, 2010

NO

Pica pica Linnaeus,

1758


Caldeirão, Furninha, Escoural Pimenta et al., 2015

YES

Garrulus

glandarius

Linnaeus, 1758

Lapa do Picareiro, Lapa do Suão,

Gruta Nova da Columbeira Pimenta et al., 2015

YES

Turdus iliacus

Linnaeus, 1766 Furninha Figueiredo, 2010

YES

Turdus pilaris

Linnaeus, 1758


Columbeira Figueiredo, 2010

YES

Turdus philomelos

Brehm, 1831 Gruta Nova da Columbeira Figueiredo, 2010

YES

Turdus merula

Linnaeus, 1758 Furninha Figueiredo, 2010

YES

Picus viridis

Linnaeus, 1758

Galería Pesada (Almonda), Gruta

Nova da Columbeira

Figueiredo, 2010,


YES

Cyanopica cyanus

(Pallas, 1776) Gruta Nova da Columbeira Figueiredo, 2010

YES

Sturnus vulgaris

Linnaeus, 1758 Lagar Velho, Lapa do Picareiro Figueiredo, 2010

YES

Carduelis carduelis

Linnaeus, 1758 Gruta Nova da Columbeira Figueiredo, 2010

YES

Ptyonoprogne

rupespris (Scopoli,

1769) Gruta Nova da Columbeira Figueiredo, 2010

YES

Motacilla alba


YES

Caprimulgus

europaeus

Linnaeus, 1758

Galería Pesada (Almonda), Gruta

Nova da Columbeira Pimenta et al., 2015

YES

Cuculus canorus

(Linnaeus, 1758) Furninha, Gruta de Casa da Moura Figueiredo, 2010

YES

31

Table 2.5: Mammals of the Quaternary of Portugal.


EXTANT IN

PORTUGAL?

Erinaceus

europaeus

Linnaeus, 1758

Furninha, Figueira Brava, Lagar

Velho, Lapa do Picareiro

Mein & Antunes, 2000; Moreno-

García & Pimenta, 2002; Bicho et

al., 2003; Brugal et al., 2012

YES

Crocidura russula

Hermann, 1780

Figueira Brava, Lapa do

Picareiro, Lagar Velho, Guia,

Bom Santo Cave

Antunes et al., 1989; Mein &

Antunes, 2000; Moreno-García &

Pimenta, 2002; Bicho et al., 2003;

Pimenta, 2014

YES

Crocidura

suaveolens Pallas,

1811

Figueira Brava cave, Lapa do

Picareiro

Mein & Antunes, 2000; Bicho et al.,

2003

YES

Sorex araneus

Linnaeus, 1758 Goldra Antunes et al., 1986b

NO

Sorex sp. Linnaeus,

1758 Lagar Velho Moreno-García & Pimenta, 2002

YES

Talpa occidentalis

Cabrera, 1907


Picareiro, Bom Santo Cave

Mein & Antunes, 2000; Bicho et al.,

2003; Pimenta, 2014

YES

Talpa europaea

Linnaeus, 1758 Goldra Antunes et al., 1986b

NO

Galemys kormosi

(Shereuder, 1940) Morgadinho Antunes et al., 1986a

NO

Galemys

pyrenaicus

Geoffroy, 1811 Lapa do Picareiro Bicho et al., 2003

YES

Rinolophus

ferrumequinun

(Shereber, 1774)

Gruta da Aroeira (Almonda

System), Figueira Brava

Mein & Antunes, 2000; López-

García et al., 2018

YES

Rinolophus

hipposideros

(Bechstein, 1800) Figueira Brava Mein & Antunes, 2000

YES

Rinolophus euryale

Blasius, 1853 Figueira Brava Mein & Antunes, 2000

YES

Myotis myotis

(Borkhausen,

1797)

Gruta da Aroeira (Almonda),


Picareiro

Mein & Antunes, 2000, Bicho et al.,

2003; López-García et al., 2018

YES

Myotis blythi

(Tomes, 1857) Figueira Brava Mein & Antunes, 2000

YES

Myotis nattereri

(Khul, 1818) Figueira Brava Mein & Antunes, 2000

YES

Myotis mystacinus

Khul, 1817 Lapa do Picareiro Bicho et al., 2003

YES

Miniopterus

schreibersi (Khul,

1819) Figueira Brava Mein & Antunes, 2000

YES

Apodemus

sylvaticus

(Linnaeus, 1758)

Goldra, Guia, Caldeirão, Lagar

Velho, Lapa do Picareiro, Bom

Santo Cave

Antunes et al., 1986b; Antunes et

al., 1989; Póvoas et al., 1992;

Moreno-García & Pimenta, 2002;

Bicho et al., 2003; Pimenta, 2014

YES

32

Apodemus

flavicollis

(Melchior, 1834)


System), Figueira Brava

Jeannet, 2000; López-García et al.,

2018

NO

Mus musculus

Linnaeus, 1758 Guia, Figueira Brava Antunes et al., 1989; Jeannet, 2000

YES

Rattus rattus

Linnaeus, 1758 Figueira Brava, Bom Santo Cave Jeannet, 2000; Pimenta, 2014

YES

Sciurus vulgaris

Linnaeus, 1758 Figueira Brava, Lagar Velho

Jeannet, 2000; Moreno-García &

Pimenta, 2002

YES

Eliomys quercinus

Linnaeus, 1766

Goldra, Guia, Caldeirão, Figueira

Brava, Lapa do Suão, Bom Santo

Cave

Antunes et al., 1986b; Antunes et

al., 1989; Póvoas et al., 1992;

Jeannet, 2000; Valente & Haws,

2006; Pimenta, 2014

YES

Glis glis (Linnaeus,

1766) Lapa do Picareiro Bicho et al., 2003

YES

Allocricetus bursae

Schaub., 1930 Caldeirão Póvoas et al., 1992

NO

Mimomys sp.

Forsyth-Major,

1902 Morgadinho Antunes et al., 1986a

NO

Pliomys

episcopalis

Méhely, 1914

Galería Pesada and Gruta da

Aroeira (Almonda)

Marks et al., 2002a; López-García et

al., 2018

NO

Myodes sp. Pallas,

1811 ? Nehring, 1899

NO

Chionomys nivalis

Martins, 1842 Caldeirão, Lapa do Picareiro

Póvoas et al., 1992; Bicho et al.,

2003

YES

Arvicola sapidus

Miller, 1908

Caldeirão, Figueira Brava, Lagar

Velho

Póvoas et al., 1992; Jeannet, 2000;

Moreno-García & Pimenta, 2002

YES

Arvicola

amphibious

(Linnaeus, 1758) Lapa do Suão Valente & Haws, 2006

NO

Arvicola cantiana

Hinton, 1910 Figueira Brava Jeannet, 2000

NO

Microtus

lusitanicus (Gerbe,

1879)

Caldeirão, Lapa do Suão, Lagar

Velho, Guia, Bom Santo Cave

Antunes et al., 1989; Póvoas et al.,

1992; Moreno-García & Pimenta,

2002; Valente & Haws, 2006;

Pimenta, 2014

YES

Microtus

duodecimcostatus

(de Selys-

Longchamps,

1839)

Figueira Brava, Caldeirão, Lapa

do Picareiro, Lagar Velho,

Goldra, Bom Santo Cave


1992; Jeannet, 2000; Moreno-García

& Pimenta, 2002; Bicho et al., 2003;

Valente & Haws, 2006; Pimenta,

2014

YES

Microtus arvalis

(Pallas, 1778)

Figueira Brava, Caldeirão, Lagar

Velho

Póvoas et al., 1992; Jeannet, 2000;

Moreno-García & Pimenta, 2002

YES

Microtus agrestis

(Linnaeus, 1761)

Caldeirão, Lapa do Picareiro,

Lagar Velho

Póvoas et al., 1992; Moreno-García

& Pimenta, 2002; Bicho et al., 2003

YES

Microtus

brecciensis Forsyth


System), Figueira Brava,


1992; Jeannet, 2000; López-García

NO

33

Major, 1905 Caldeirão, Goldra et al., 2018

Microtus cabrerae

(Thomas, 1906) Caldeirão, Bom Santo Cave Póvoas et al., 1992; Pimenta, 2014

Castor fiber

Linnaeus, 1758 Caldeirão Antunes, 1989

NO

Prolagus calpensis

Major, 1905 Morgadinho Antunes et al., 1986a

NO

Lepus granatensis

Rosenhauer, 1856

Caldeirão, Pego do Diabo, Lapa

do Suão Davis, 2002; Valente, 2004

Oryctolagus lacosti

(Pomel, 1853) Algoz Antunes et al., 1986c

NO

Oryctolagus

cuniculus

(Linnaeus, 1758)

Pego do Diabo, Lapa do Suão,

Lapa do Picareiro, Caldeirão,

Vale Boi, Lagar Velho

Hockett & Bicho, 2000; Davis,

2002; Moreno-García & Pimenta,

2002; Valente & Haws, 2006;

Manne et al., 2012

YES

Macaca sylvanus

(Linnaeus, 1758) Galería Pesada (Almonda) Marks et al., 2002b

NO

Paleoloxodon

antiquus

(Falconer & Cautle

y, 1847)

Meirinha, Algés, Condeixa,

Furninha, Almonda, Carregado,

Mealhada, Foz do Enxarrique,

Santa Cruz, Santo Antão do Tojal

Antunes & Cardoso, 1992;

Figueiredo, 2012

NO

Mammuthus

primigenius

Blumebach, 1799

Algar de João Ramos, Figueira

Brava, Oeiras sea floor

Antunes & Cardoso, 1992; Cardoso

& Regala, 2002

NO

Stephanorhinus

hemitoechus

(Falconer, 1859)

Galería Pesada (Almonda), Serra

dos Molianos, Furninha, Gruta

Nova da Columbeira, Lapa da

Rainha, Lorga de Dine, Correio-

Mor, Figueira Brava, Pedreira

das Salemas, Escoural

Veiga Ferreira, 1975; Cardoso,

1993a, Marks et al., 2002a

NO

Equus hydruntinus

Regalia, 1905

Pedreira das Salemas, Caldeirão,

Vale Boi

Cardoso, 1995; Davis, 2002; Manne

et al., 2012

NO

Equus ferus

(Linnaeus, 1758)

Algar de João Ramos, Gruta das

Fontainhas, Pedreira das

Salemas, Algar de Cascais,

Escoural, Gruta Nova da

Columbeira, Furninha, Lapa da

Rainha, Figueira Brava,

Caldeirão, Vale Boi, Foz do

Enxarrique

Cardoso, 1993a; Raposo, 1995;

David, 2002; Manne et al., 2012

NO

Delphinus delphis

Linnaeus, 1758 Figueira Brava Antunes, 2000

Hippopotamus

amphibious

Linnaeus, 1758 Condeixa, Mealhada Choffat, 1895; Antunes et al., 1988

NO

Hippopotamus

antiquus Desmaret,

1822 Algoz Antunes et al., 1986b

NO

Sus scrofa

Linnaeus, 1758

Fontainhas, Pedreira das

Salemas, Gruta das Salemas,

Escoural, Lapa do Picareiro,

Figueira Brava, Caldeirão, Vale

Boi, Goldra, Lagar Velho

Antunes et al., 1989; Cardoso,

1993a; Davis, 2002; Moreno-García

& Pimenta, 2002; Hockett & Haws,

2009; Manne et al., 2012

YES

34

Cervus elaphus

Linnaeus, 1758


Mealhada, Furninha, Fontainhas,

Caldeirão, Pedreira das Salemas,

Escoural, Lorga de Dine, Casais

Robustos, Algar de Cascais,

Porto Covo (Cascais), Lapa da

Rainha, Figueira Brava, Pego do

Diabo Cardoso, 1993a

YES

Capreolus

capreolus

Linnaeus, 1758

Caldeirão, Columbeira, Lapa da

Rainha, Picareiro, Porto Covo

(Cascais), Lagar Velho

Cardoso, 1993a; Moreno-García &

Pimenta, 2002; Hockett & Haws,

2009

YES

Dama dama

Linnaeus, 1758

Algar de João Ramos, Gruta

Nova da Columbeira, Pedreira

das Salemas Cardoso, 1989

YES

Dama

vallonnetensis De

Lumley et al., 1988 Galería Pesada (Almonda) Marks et al., 2002a

NO

Eucladoceros sp.

Falconer, 1868 Algoz Antunes et al., 1986c

NO

Bos primigenius

Bojanus, 1827

Lorga de Dine, Fujaca, Gruta

Nova da Columbeira, Pedreira

das Salemas, Algar de Cascais,

Figueira Brava, Escoural, Lapa

da Rainha, Mealhada, Foz do

Enxarrique, Furninha, Vale Boi,

Guia, Lagar Velho

Antunes et al., 1989; Cardoso,

1993a; Moreno-García & Pimenta,

2002; Manne et al., 2012

NO

Capra pyrenaica

Schinz, 1838

Fontainhas, Caldeirão, Casais

Robustos, Pedreira das Salemas,

Figueira Brava, Escoural, Pego

de Diablo, Algar of João Ramos,

Vale Boi Cardoso, 1993a; Manne et al., 2012

YES

Rupicapra

pyrenaica

Bonaparte, 1845

Gruta das Salemas, Caldeirão,

Pego do Diabo, Lapa do

Picareiro Cardoso, 1993a; Bicho et al., 2000

NO

Pusa hispida

(Schereber, 1775) Figueira Brava Antunes, 2000

NO

Monachus

monachus

(Hermann, 1779) Furninha This work

NO

Canis lupus

Linnaeus, 1758

Furninha, Fontainhas, Casa da

Moura, Algar de João Ramos,

Lapa da Rainha, Pego do Diabo,

Gruta Nova da Columbeira,

Pedreira das Salemas, Gruta das

Salemas, Caldeirão, Escoural,

Lagar Velho, Vale Boi


Pimenta, 2002; Manne et al., 2012

YES

Cuon alpinus

Pallas, 1811 Escoural Cardoso, 1993a

NO

Vulpes vulpes

(Linnaeus, 1758)

Furninha, Casa da Moura,

Figueira Brava, Caldeirão,

Escoural, Vale Boi, Lagar Velho


Pimenta, 2002; Manne et al., 2012

YES

Ursus arctos

Linnaeus, 1758

Furninha, Fontainhas, Molianos,

Gruta das Salemas, Algar de

Cascais, Pedreira das Salemas,

Cardoso, 1993a; Marks et al.,

2002a; Davis, 2002

35

Figueira Brava, Galería

Pesada(Almonda), Caldeirão,

Lorga de Dine, Escoural

NO

Mustela sp.

Linnaeus, 1758 Vale Boi, Furninha

Brugal et al., 2012; Manne et al.,

2012

YES

Martes sp. Pinel,

1792 Vale Boi, Furninha

Brugal et al., 2012; Manne et al.,

2012

YES

Meles meles

Linnaeus, 1758

Furninha, Caldeirão, Lapa do

Suão

Davis, 2002; Valente & Haws, 2006;

Brugal et al., 2012

YES

Hyaena hyaena

(Linnaeus, 1758) Furninha Brugal et al., 2012

NO

Crocuta crocuta

(Erxleben, 1777)

Lorga de Dine, Caldeirão,

Fontainhas, Gruta Nova da

Columbeira, Lapa da Rainha,

Gruta das Salemas, Algar de

Cascais, Figueira Brava,

Escoural, Porto Covo Cardoso, 1996; Valente, 2004

NO

Felis silvestris

Schreber, 1777

Escoural, Figueira Brava, Pego

do Diabo, Gruta das Salemas,

Pedreira das Salemas,

Fontainhas, Furninha, Caldeirão,

Lapa da Rainha, Gruta Nova da

Columbeira Villaluenga, 2016

YES

Lynx pardinus

Temminck, 1827

Escoural, Pego do Diabo,

Pedreira das Salemas, Gruta das

Salemas, Furninha, Caldeirão,

Lapa da Rainha, Gruta Nova da

Columbeira, Casa da Moura,

Algar de João Ramos, Algar de

Cascais, Casais Robustos Villaluenga, 2016, This work

YES

Panthera pardus

Linnaeus, 1758

Escoural, Figueira Brava, Pego

do Diabo, Pedreira das Salemas,

Fontainhas, Furninha, Casa da

Moura, Galería Pesada

(Almonda), Caldeirão, Lorga de

Dine, Algar da Manga Larga

Cardoso & Regala, 2006;

Villaluenga, 2016

NO

Panthera spelaea

Goldfuss, 1810

Escoural, Figueira Brava,

Pedreira das Salemas, Lorga de

Dine, Vale Boi

Manne et al., 2012; Villaluenga,

2016

NO

Homotherium

latidens Owen,

1846 Mealhada Antunes, 1988

NO

36

2.3 Results

In the following resume table (Table 2.6) we can see some general numbers for the different groups. It

should be noted that only native species are counted in each case, not the ones introduced in recent

years, because those could not have been detected in the fossil record and therefore distort the

representativeness of it. The table has been arranged following the data provided for nowadays species

by Loureiro et al., 2008 for amphibians and reptiles; Assírio & Alvim, 2008 for birds; Rainho et al.,

1998 for bats and Bencatel et al., 2017 for the other mammals (The four Quaternary marine mammals

detected, as well as the modern ones are excluded). An extra column is added with the big mammals

(mammals that are bigger than Meles meles and Felis silvestris) in the Quaternary of Portugal, but

those are listed within the mammal category already.

Table 2.6: Resume table showing the number, percentage and status of extant and fossil species of the

Quaternary of Portugal.

Amphibians Reptiles Birds Mammals Total Big mammals

Extant 17 28 221 66 332 10

Known in the fossil

record 7 13 80 74 174 30

Extinct 0 0 3 17 20 9

Locally extinct 0 1 10 17 29 11

Fossil only locally 0 1 13 34 49 20

In the fossil record and

extant locally 7 12 67 40 125 10

Not known as fossil 10 15 154 26 207 0

% Extinct in the fossil

record 0,00% 0,00% 3,75% 22,97% 11,49% 30,00%

% Locally extinct in the

fossil record 0,00% 7,69% 12,50% 22,97% 16,67% 36,67%

% Fossil only locally 0,00% 7,69% 16,25% 45,95% 28,16% 66,67%

% of extant species

detected in the fossil

record 41,18% 42,86% 30,32% 60,61% 37,65% 100,00%

% of extant species

undetected in the fossil

record 58,82% 57,14% 69,68% 39,39% 62,35% 0,00%

% of each group in

extant species 5,12% 8,43% 66,57% 19,88% 100,00% 3,01%

% of each group in

fossil species 4,02% 7,47% 45,98% 42,53% 100,00% 17,24%

Looking at the general numbers at table 2.5, we can see that 174 species of fossil tetrapods have been

discovered in the Quaternary of Portugal. A total of 49 (28,16%) of them no longer appear in the area

either by local or complete extinction. 125 of the fossil species (that represent the 71,84% of them),

still live in Portugal nowadays, comprising the 37,65% of the 332 species of living tetrapods in the

country. We can presume therefore that 207 (or 62,35%) of them have not been detected in the fossil

record despite already living in Portugal.

We can compare the groups now. In figure 2.2 we can see graphically the percentage of extinct,

locally extinct and extant species in the fossil record for each group. We can see also the percentage of

extant species detected vs the undetected in each group in the fossil record and the percentage of the

total fossil and extant species that each group comprises.

37

Figure 2.2: Percentages of different parameters about the paleodiversity of the Quaternary in Portugal.

Of special interest is the case of the big mammals. In figure 2.3 we can see the status of the different

species known in the fossil record. It should be noted that 2/3 of them are extinct or locally extinct. All

ten species of them living nowadays in Portugal appear in the fossil record. Interestingly they

comprised 17,24% of the fossil species, meanwhile today they are only the 3,01% of the living

tetrapod species in Portugal (Table 2.5).

38

Figure 2.3: Status of the different big mammals that appear in the fossil record in Portugal.

Looking at the localities in the table 2.1 we can see that most of them are cave deposits; but precisely

how important are they to the understanding of the Quaternary fauna? In table 2.7 the species and

localities associated with cave deposits have been eliminated. The references are in the table 2.6.

Table 2.7: Table showing the tetrapods and localities known outside the cave deposits in the Quaternary of

Portugal.

TAXA LOCALITIES

Salamandra salamandra (Linnaeus, 1758) Lagar Velho

Triturus marmoratus (Latreille, 1800) Lagar Velho

Bufo bufo (Linnaeus, 1758) Guia

Pelobates cultripes Cuvier, 1829 Guia

Pleurodeles waltl (Michahelles, 1830) Guia

Epidalea calamita Laurenti, 1758 Guia

Pelophylax perezi (Seoane, 1885) Guia

Timon lepidus Daudin, 1802 Guia

Acanthodactylus erythrurus (Schinz, 1833) Guia

Natrix sp. Laurenti, 1758 Guia

Testudo hermanni Gmelin, 1789 Mealhada

Mauremys leprosa (Schweiger, 1812) Mealhada

Aquila chrysaetos Linnaeus, 1758 Vale Boi

Falco tinnuncus Linnaeus, 1758 Lagar Velho

Sturnus vulgaris Linnaeus, 1758 Lagar Velho

Crocidura russula Hermann, 1780 Lagar Velho, Guia

Sorex araneus Linnaeus, 1758 Goldra

Sorex sp. Linnaeus, 1758 Lagar Velho

Talpa europaea Linnaeus, 1758 Goldra

Galemys kormosi (Shereuder, 1940) Morgadinho

Apodemus sylvaticus (Linnaeus, 1758) Goldra, Guia, Lagar Velho

30,00%

36,67%

33,33%

% Extinct in the fossil record

% Locally extinct in the fossil record

%Extant

39

Mus musculus Linnaeus, 1758 Guia

Sciurus vulgaris Linnaeus, 1758 Lagar Velho

Eliomys quercinus Linnaeus, 1766 Goldra, Guia

Mimomys sp. Forsyth-Major, 1902 Morgadinho

Arvicola sapidus Miller, 1908 Lagar Velho

Microtus lusitanicus (Gerbe, 1879) Lagar Velho, Guia

Microtus duodecimcostatus (de Selys-Longchamps, 1839) Lagar Velho, Goldra

Microtus arvalis (Pallas, 1778) Lagar Velho

Microtus agrestis (Linnaeus, 1761) Lagar Velho

Microtus brecciensis Forsyth Major, 1905 Goldra

Prolagus calpensis Major, 1905 Morgadinho

Oryctolagus lacosti (Pomel, 1853) Algoz

Oryctolagus cuniculus (Linnaeus, 1758) Vale Boi, Lagar Velho

Paleoloxodon antiquus (Falconer & Cautley, 1847)

Meirinha, Algés, Condeixa, Carregado,

Mealhada, Foz do Enxarrique, Santa Cruz,

Santo Antão do Tojal

Mammuthus primigenius Blumebach, 1799 Oeiras sea floor

Equus hydruntinus Regalia, 1905 Vale Boi

Equus ferus (Linnaeus, 1758) Vale Boi, Foz do Enxarrique

Hippopotamus amphibious Linnaeus, 1758 Condeixa, Mealhada

Hippopotamus antiquus Desmaret, 1822 Algoz

Sus scrofa Linnaeus, 1758 Goldra, Lagar Velho

Cervus elaphus Linnaeus, 1758 Mealhada, Casais Robustos

Capreolus capreolus Linnaeus, 1758 Lagar Velho

Eucladoceros sp. Falconer, 1868 Algoz

Bos primigenius Bojanus, 1827

Pampilhosa do Botão, Mealhada, Foz do

Enxarrique, Vale Boi, Guia, Lagar Velho

Capra pyrenaica Schinz, 1838 Vale Boi

Canis lupus Linnaeus, 1758 Lagar Velho, Vale Boi

Vulpes vulpes (Linnaeus, 1758) Vale Boi, Lagar Velho

Mustela sp. Linnaeus, 1758 Vale Boi

Martes sp. Pinel, 1792 Vale Boi

Panthera spelaea Goldfuss, 1810 Vale Boi

Homotherium latidens Owen, 1846 Mealhada

We can see in the table 2.8 the new resume table for the data in table 2.7, with an extra column for big

mammals.

40

Table 2.8: Resume table showing the number, percentage and status of extant and fossil species in the

Quaternary of Portugal, excluding the cave deposits.

Amphibians Reptiles Birds Mammals Total

Big

mammals

Extant 17 28 221 66 332 10

Known in the fossil

record (No caves) 7 5 3 37 52 17

Extinct (No caves) 0 0 0 12 12 8

Locally extinct (No

caves) 0 1 0 4 5 3

Fossil only locally (No

caves) 0 1 0 16 17 11

In the fossil record and

extant (No caves) 7 4 3 21 35 6

Not known as fossil (No

caves) 10 24 219 45 297 4

% Extinct in the fossil

record (No caves) 0,00% 0,00% 0,00% 32,43% 23,08% 47,06%

% Locally extinct in the

fossil record (No caves) 0,00% 20,00% 0,00% 10,81% 9,62% 17,65%

% Fossil only locally (No

caves) 0,00% 20,00% 0,00% 43,24% 32,69% 64,71%

% of extant species

detected in the fossil

record (No caves) 41,18% 14,29% 1,36% 31,82% 10,54% 60,00%

% of extant species

undetected in the fossil

record (No caves) 58,82% 85,71% 99,10% 68,18% 89,46% 40,00%

% of each group in extant

species 5,12% 8,43% 66,57% 19,88% 100,00% 3,01%

% of each group in fossil

species (No caves) 13,46% 9,62% 5,77% 71,15% 100,00% 32,69%

If we exclude the cave deposits, the percentage of species detected overall falls from the previous

37,65% (125 species) to only a 10,54% (35 species). Also the total number of species falls from 174

to 52, a fall of the 70%. The cases of the birds and amphibians are especially interesting (As we see in

figure 2.3) the first nearly disappearing from the fossil record (falling from a 45,98% to a mere 5,77%

of the species) meanwhile the amphibians pass from the 4,02% to the 13,46% of all detected species,

which is not surprising since all the 7 species located appear outside the cave deposits. It is also

noteworthy that big mammal are over-represented comprising 32,69% of all the species when with the

cave deposits they are the 17,24% and nowadays only the 3,01%. This same phenomenon happens

with the relative number of mammal species inversely to the bird ones.

41

Figure 2.4: Percentage of species from each group in extant, fossil and fossil species (without cave deposits).

2.4 Discussion

An interesting feature of the data is the similarity between the numbers of amphibians and the reptiles.

Between both groups there is only one locally extinct species, Testudo hermanni, all other species

appear nowadays in Portugal. This feature is congruent with the proposed behavior for the

herpetofaunal communities across the Quaternary (Blain et al., 2016; Bisbal-Chinesta & Blain, 2018):

At the beginning of the Pleistocene, with most of the mountain ranges as barriers going from East to

West and the Mediterranean and the Black Sea in the South preventing movements towards warmer

areas, the “exotic” (and most vulnerable to climate cooling) taxa of Europe died out relatively quick,

for example the giant chelonians that appear since the Miocene in Europe (Pérez-García et al., 2017)

therefore only the most resilient species to cold conditions survived this first impact, enduring again

and again later glaciations without losing more species, although there are some exceptions, like the

varanids that only perished in the Middle Pleistocene (Georgalis et al., 2017) or relict thermophilic

species like Chioglossa lusitanica Bocage, 1864 that endured since the Neogene the cold moments

isolated in refugia (Alexandrino et al., 2001).

Being a higher metabolic rate related with the rate of evolution (Martin & Palumbi, 1993) the capacity

of the herpetofauna for speciation events was quite low during this period, even more taking into

account the impoverish starting number of species.

Both the percentage of species compared to the overall number of tetrapods and the representativeness

of the abundance of these groups are quite reliable measures of their diversity. The percentage of

species compared to the total is close to the same number in extant, fossil species and fossil species

without cave deposits as we see in figure 2.4. Also, with around 40% of the extant species represented

in the fossil record their representativeness is close to the average of all tetrapod species combined.

Now talking about birds we have to bear in mind that their mobility make the number of species

present in the Portuguese territory change constantly, either related to seasonality (As migratory

species displace North to breed and South to winter) or to mere chance (for example with American

species that cross the Atlantic). Therefore the number of nesting species seem the closest proxy to the

number of “resident” birds that we can found. Their importance relative to the total number of species

of tetrapods today is great; with 221 species, they comprise 2/3 of tetrapods. But this numbers are not

reflected in the fossil record, with only 80 species that represent less than half of the fossil tetrapods.

This is probably related to several factors, like the lack of interest of many researchers in their bones

until relative recent years, their fragility given the pneumaticity of them and the difficulties to identify

to the specific level (like for example a lot of Passeriformes Linnaeus, 1758, whose skeletons are

remarkable similar).

5,12% 8,43%

66,57%

19,88% 4,02% 7,47%

45,98% 42,53%

13,46% 9,62% 5,77%

71,15%

Amphibians Reptiles Birds Mammals

Percentage of species from each group in different contexts

% of each group in extant species

% of each group in fossil species

% of each group in fossil species (Without cave deposits)

42

If we remove the cave deposits the situation is even more pronounced. Only three species “survive”

comprising just 5,77% of all tetrapods. This situation could be explained for the accentuation of all the

after mentioned factors in the more difficult preservation environment out of the caves. It is interesting

to note that this situation is quite similar to what we see in the Cenozoic fossil bird record in Portugal

with only 6 sites with bird remains, most of them impossible to ascribe at specific or even generic

level, although in the Eocene of Silveirinha appears Fluviatilavis antunesi Harrison, 1983; in the

Miocene site of Olival da Susana appears Palaeoperdix media Milne-Edwards, 1871 (Figueiredo,

2010) and in Costa da Caparica, also with Miocene chronology, a large sternum assigned tentatively to

Pelagornis miocaenus Lartet, 1857 (Mayr et al., 2008). Therefore is not risky to assume that there is a

huge hidden diversity of birds across the Cenozoic, even in the Quaternary, given that the detected

species in the fossil record that nowadays live in Portugal only reach about the 30% of the total.

Mammal data have the most complex interpretation of all; they are extremely overrepresented in the

fossil record without cave deposits (More than 70%), if we include the later they still comprise about

half of the total tetrapod species, but nowadays they only represent 19% of the species (as we see in

figure 2.3). This fact could be explained with the intersection of three factors: More than the 45% of

the fossil mammal fauna is extinct or locally extinct and this large extinction lowered the overall

number of species today. Also between them we have the biggest land animals in the Quaternary of

Portugal, with dense and large bones that are more likely to be found even in less favorable conditions

outside the cave deposits, like in alluvial terraces. Finally they have been subject of great scientific

interest for 150 years in the country, since the first excavations of Carlos Ribeiro and Nery Delgado.

These factors could also explain why around 60% of the actual species have been detected in the fossil

record; with only small carnivores, “insectivores” and mostly bats being missing; although in the

realm of extinct species there is a very real possibility of finding some known megafauna in the

Iberian Peninsula but not in Portugal yet, like bison or cave bear.

Finally, if we exclude the cave deposits we can find a good picture of the importance of those in our

knowledge of the paleobiodiversity of the Quaternary. Without them we pass from 174 species to 52; a

fall of around the 70% if we do not count with the fountain of fossils that is the karst. This is

congruent with what we see in the older Cenozoic fossil record, for example in the Miocene

pseudokarstic infillings of Cerro Batallones, Madrid that produce a rich fauna otherwise impossible to

find (Morales & Baquedano, 2017). For contexts where we don´t have that type of deposits (for

example the Jurassic of the Lusitanian Basin) our sampling can be quite reduced.

2.5 Conclusions

- Most of the vertebrate fossil record of the Quaternary of Portugal (with emphasis in the

Pleistocene) is situated in the Lusitanian and Algarvian basins. Only 3 out of 30 sites are

outside those areas and 20 out of 30 sites are situated in caves.

- Quaternary fossil herpetofaunal communities in Portugal evolve across the period with

reduced numbers and only one locally extinct species in the area.

- ~70% of living bird species are undetected in the fossil record due to their fragile bones, the

difficulties in their identification and the lack of scientific interest.

- 46% of mammal species known in the Quaternary fossil record of Portugal became locally

extinct in the area or went totally extinct. They are also the better known group of Quaternary

fossil tetrapods in the country, with most of the living species (60%) appearing in the fossil

record.

- The importance of the cave deposits is capital in our understanding of the Quaternary

vertebrate fauna, without it we would not known about most of tetrapod species, not to

mention facts about their paleobiology or paleoecology (Like their denning behavior). The

tetrapod living species detected in the fossil record would fall from 37,65% (125 species) to

only a 10,54% (35 species) without them.

43

3.THE PALAEOLOXODON ANTIQUUS OF SANTO ANTÃO DO

TOJAL

3.1 Introduction and methods

The proboscidean fossil record of the Quaternary in Portugal is poor if we compare it with the record

of neighboring Spain (Ros-Montoya, 2010) and similar sized countries like Greece (Tsoukala et al.,

2011) or Serbia (Lister et al., 2012), or even with the Miocene record of Portugal itself (Antunes,

1989b). As we have seen before, this group is represented in the Quaternary of continental Portugal by

isolated dental remains and a handful of bones, none of them really well preserved (Antunes &

Cardoso, 1992). Between this absence of articulated (or even associated) skeletons, the occurrence of

Santo Antão do Tojal is by far the most complete of a proboscidean of this age in Portugal, including

an almost complete femur, a partial tibia, phalanx and neural spine, assigned to Elephas (Now

Palaeoloxodon) antiquus (Zbyszewski, 1943, 1977b). On this chapter, for first time on a Portuguese

proboscidean, an allometric equation (Larramendi, 2015) will be used in order to give an estimation of

the body mass of the animal when it was alive. Also those newly obtained measurements of this

specimen will be compared with an extensive sample of 47 femurs and 29 tibias of other European

proboscideans in order to investigate the metric differences in the hind limb of Palaeoloxodon

antiquus and Mammuthus primigenius. In base of these two new sets of data some conclusions

regarding this individual specimen and both species will be taken. This section has been partially

published in the “VII Congresso de Jovens Invertigadores em Geociências” (Estraviz-López &

Mateus, 2018d).

3.2 Geographical and geological settings

In February of 1941 during a visit to a recently dug channel in Loures, Georges Zbyszewski found

several bone fragments. Two days later an excavation was carried on the margin of the channel and

about 2/3 of a right femur, a partial tibia and neural spine of a proboscidean were dug; alongside

several lithic tools, a horse tooth and a probable hyena coprolite (Zbyszewski, 1943). In 1977 after the

cleaning of the vegetation of the channel, Zbyszewski returned to the site. He found a mutilated

phalanx of proboscidean in the other margin of the channel that he considered belonging to the same

animal (Zbyszewski, 1977b). All of them were ascribed by him to Palaeoloxodon antiquus. The exact

location of the findings is relatively unknown, but by the description provided (“1080m South, 27°

West of Santo Antão do Tojal Church”) the probable coordinates are 38° 50' 27" N, 9° 8' 38" W. In the

figure 3.1 we can see the approximate location of the site, alongside the channel side.

Figure 3.1: The location of the Santo Antão do Tojal Palaeoloxodon antiquus is marked by the pushpin, in the

Lisboa Peninsula (left) and at local level (right). Credit Google Earth.

44

According to Zbyszewski, (1943) the fluvial terrace where the specimen was found was subject to the

regular floodings from the estuary of the Tagus River, until the construction of channels to drain the

area. The log of the locality appearing in the same work is the following, from top to bottom. No

measurements of each layer were given:

“4 — Pink sandy silts with in-situ Upper Paleolithic and reworked Mousterian industries

.

3 — Pink or red sands with Mousterian industries.

2 — Alternation between greenish clays and yellow-orange sands with iron levels, Mousterian

industries and bones, among which those from Elephas antiquus.

1— Gravel and reddish claystone (Oligocene layers reworked).”

The material of Santo Antão do Tojal was stored in the Geological Museum of Lisbon, but no catalog

number was provided in any of the works referring to it.

In Zbyszewski (1943) several measurements of the femur were taken and then compared to six

specimens ascribed to Mammuthus primigenius, one to Palaeoloxodon antiquus and two to

Mammuthus meridionalis (Nesti, 1825). Meanwhile the tibia was also measured and compared with

five specimens ascribed to Mammuthus primigenius, one to Palaeoloxodon antiquus and one to

Mammuthus meridionalis. In base of these metric comparisons and the assumed age of the terrace

(“…belongs undoubtedly to a lower grimaldian terrace (Riss-Würm interglacial). It shows evidence of

local backfilling dating of this time and posterior to the digging contemporary from the rissian glacial

epoch…”) these remains were ascribed to Palaeoloxodon antiquus.

In Zbyszewski, (1977b) the mutilated phalanx was presented and its measurements given.

In Cardoso, (1993a) new measurements of the femur were presented.

Raposo, (1995) mentioned that the Palaeoloxodon antiquus remains of Santo Antão do Tojal were

dated by the method of Uranium-Thorium or Uranium series, yielding a datation of 81.900 +4000 -

3800 YBP.

Finally this proboscidean has been cited sometimes in reviews about the Quaternary Portuguese

proboscideans like Antunes & Cardoso, (1992) or Figueiredo, (2012) that agreed with the previously

given identification.

3.3 The material

As we have seen before all the materials (Femur, tibia, vertebral apophysis and phalanx) is part of the

exhibit of the Geological Museum in Lisbon, where it is situated in the archeological part of the

museum.

Right femur (MG 5788)

The proboscidean femur of Santo Antão do Tojal (Figure 3.2) is the most complete of all the bones of

the site, being it relatively well preserved compared to the other material. Only its distal part, including

the condyles, is missing. It is displayed in a standing position with the distal broken part integrated

into stand and two metallic rings securing it, all of this preventing the specimen of being removed

from the display structure. The trochanteric fossa is deep and well marked, character that point

towards P. antiquus, given its robustness and marked muscular insertions in bones (Larramendi et al.,

2017).

45

Figure 3.2: General view of the Palaeoloxodon antiquus femur of Santo Antão do Tojal (MG 5788) at display in

the geological museum in posterior view. Scale bar 7 cm. The chalk mark signals a cut in the bone where a stone

tool was found, being it regarded by Zbyszewski as a proof that the animal was butchered by Neanderthals.

Right tibia (MG 5793)

The tibia is in worse shape than the femur, being its distal end totally missing. The proximal one has

been clearly restored with plaster and has lost fragments. The tibia is displayed the same way than the

femur, with the distal and more damaged part embedded into the stand and a metal ring securing the

bone in an upright position that makes it impossible to take the piece out. In figure 3.3 we can see the

tibia in anterior view. Given the position of the medial articular surface (the best preserved feature of

the proximal part) relative to the tibial depression (marking the anterior part of the bone); the tibia was

a right exemplar (Shil et al., 2013). In Figure 3.4 we can see those features in the tibia.

46

Figure 3.3: Right tibia of the Palaeoloxodon antiquus of Santo Antão do Tojal (MG 5793) at display in the

Geological Museum in anterior view. Scale bar 7 cm.

Figure 3.4: Proboscidean tibia of Santo Antão do Tojal (MG 5793) in left, anterior and right proximal views.

The principal anatomical features are marked, in white the tibial depression and in red the medial articular

surface. Scale bar 7 cm.

47

Vertebral spinal process (MG 8081)

The incomplete spinal process of a vertebra from the proboscidean is 390 mm tall, despite that the tip

of it is broken. Meanwhile, the sides of the neural canal are better preserved, being it 165 mm wide. It

is impossible to be totally sure without the whole vertebra, but given the height of this process the

vertebra was a thoracic one (Larramendi et al., 2017). In figure 3.5, we can see it.

Figure 3.5: The spinal process of the proboscidean of Santo Antão do Tojal (MG 8081) in A) anterior and B)

posterior views.

48

First phalanx (MG 8080)

The phalanx of the Proboscidean of Santo Antão do Tojal is lightly damaged, as we can see in figure

3.6. In the table 3.1 we can see its measurements. It is a first phalanx according to Zbyszewski,

(1977b).

Figure 3.6: The phalanx from the proboscidean of Santo Antão do Tojal in A) dorsal; B) lateral; C) ventral and

D) lateral views.

Table 3.1: Measurements of the phalanx of the proboscidean of Santo Antão do Tojal. “Thickness” refers to

anteroposterior diameter of the bone and “width” to the mediolateral diameter of it. All the measurements are in

millimeters.

Work Zbyszewski, 1977, this work

Site Santo Antão do Tojal

Specimen Santo Antão do Tojal

Species attributed Palaeoloxodon antiquus

Number First

Maximum length 100

Proximal width 75

Proximal thickness 67

Distal width 50

Distal thickness 47

3.4 Measurements of femur and tibia

The measurements taken of the femur and tibia were the ones used by Larramendi et al. ( 2017); which

are similar to the ones of Von den Driesch, (1976) for the hindlimb of mammals. On them “width”

refers to medio-lateral diameter of the bone and “thickness” to the anteroposterior diameter of it. We

can see the measurements graphically in the figures 3.7 and 3.8. In the tables 3.2 and 3.3 we have the

measurements of the Santo Antão do Tojal material as well a collection of measurements that have

been taken from literature for other femora and tibiae of Palaeoloxodon antiquus and Mammuthus

primigenius.

49

Figure 3.7: Measurements of the femur of mammals by Von den Driesch, 1976.

Table 3.2: Measurements of 48 individual femurs of Palaeoloxodon antiquus and Mammuthus primigenius taken

from the literature. The specimen of Santo Antão do Tojal is marked. All measurements are in millimeters.

Work Site Species

attributed Maximum

length Proximal

width Caput width

Caput thickness

Minimum

diaphysis width

Minimum

diaphysis thickness

Minimum

diaphysis circumference

Distal width

Distal

articular width

Larramendi

et al., 2017

Neumark-

Nord 1

Palaeoloxodon

antiquus 1320 425 205 210 156 121 470 318 -

Larramendi

et al., 2017

Neumark-

Nord 1

Palaeoloxodon

antiquus 1398 - 216 205 - - - - -

Larramendi

et al., 2017

Neumark-

Nord 1

Palaeoloxodon

antiquus 1040 - 155 159 116 95 340 229 -

Larramendi

et al., 2017

Neumark-

Nord 1

Palaeoloxodon

antiquus 1070 338 146 159 121 86 332 244 192

Tsoukala et

al., 2011 Sotiras

Palaeoloxodon

antiquus - - - - - - - 330 290

Tsoukala et

al., 2011 Xerias

Palaeoloxodon

antiquus - - - - 124 289 - - -

Kevrekidis

& Mol, 2016 Amyntaio

Palaeoloxodon

antiquus - - 215 217 - - - - -

Mazo, 1998 Buelna

Palaeoloxodon

antiquus ~ 1320 - - - 175 130 460 320 -

Mazo, 1998 Torralba

Palaeoloxodon

antiquus 1210 - - - ~175 ~130 - ~320 -

Konidaris et

al., 2018

Marathousa

1

Palaeoloxodon

antiquus 1330 - - - - - - - -

Marra, 2009;

Beuval et al., 1998 Gröbern

Palaeoloxodon antiquus 1402 - 224 - 173 - - 308 -

Beauval et

al. 1998 Gröbern

Palaeoloxodon

antiquus - - 179 - - - - 234 -

Marra, 2009 Crumstad Palaeoloxodon

antiquus 1040 - 155 - 116 - - 208 -

Marra, 2009 Riano Palaeoloxodon 1290 - - - 155 - - 283 -

50

antiquus

Marra, 2009 Viterbo

Palaeoloxodon

antiquus 1370 - 200 - 192 - - 247 -

Marra, 2009; Larramendi

et al., 2015 Viterbo

Palaeoloxodon

antiquus 1440 - 242 - 242 - - 277 -

Andrews &

Cooper, 1928 Upnor

Palaeoloxodon antiquus ~1500 467 214 - - - - - -

Jakubowski,

1988 Konin

Palaeoloxodon

antiquus 1429 470 - - 157 130 - 323 275

Melentis, 1963 Megalopolis

Palaeoloxodon antiquus 1372 476 - - 170 117 478 298 -

Jakubowski,

1996 Grabsch tz

Palaeoloxodon

antiquus 1130 - - - - - - - -

Christiansen,

2004 ?

Palaeoloxodon

antiquus 1075 - - - 136 102 373 - -

Christiansen,

2004 ?

Palaeoloxodon

antiquus 1169 - - - 128 94 348 - 224

Zbyszewski,

1943; this

work

Santo

Antão do

Tojal

Palaeoloxodon

antiquus - 400 180 190 165 125 470 - -

Harington et al., 2012 Muirkirk

Mammuthus primigenius 1002 - - - 111 - 313 280 -

Harington et

al., 2012 Muirkirk

Mammuthus

primigenius 1010 - - - 121 - 327 - 185

Kirillova et al., 2012 Taymir

Mammuthus primigenius 880 - 132 132 108 70 - 202 -

Kirillova et

al., 2012 Taymir

Mammuthus

primigenius - - - - 110 74 - 209 -

Maschenko et al., 2006 Svesk

Mammuthus primigenius 790 - - - - - 158 - -

Maschenko

et al., 2006 Svesk

Mammuthus

primigenius 1010 - - - - - 210 - -

Maschenko et al., 2006 Berelekh


Maschenko

et al., 2006 Berelekh

Mammuthus

primigenius 1130 - - - - - 150 - -

Álvarez-Lao et al., 2009 Padul

Mammuthus primigenius 1060 314 147 152 142 73 - - -

Álvarez-Lao

et al., 2009 North Sea

Mammuthus

primigenius 810 245 110 120 95 60 - 165 -

Álvarez-Lao et al., 2009 North Sea

Mammuthus primigenius 1400 408 190 183 176 109 - 275 -

Christiansen,

2004 Hebior

Mammuthus

primigenius 1203 - - - 160 80 377 - 229

Christiansen,

2004 Brussels

Mammuthus

primigenius 1273 - - - 158 108 419 - 287

Christiansen,

2004 Paris

Mammuthus

primigenius 1071 - - - 128 88 339 - -

Christiansen, 2004 FMNH

Mammuthus primigenius 1111 - - - 126 88 337 - 195

Christiansen,

2004

FMNH

PM26267

Mammuthus

primigenius 1014 - - - 151 98 391 - 227

Larramendi, 2015 Siegsdorf


Larramendi,

2015 Rottweil

Mammuthus

primigenius 945 - 140 - 126 - 323 - 98

Lister, 2009 Condover Mammuthus primigenius 1165 - 161 - - - - 234 211

Lister, 2009 Condover

Mammuthus

primigenius 1190 - 158 - - - - 242 207

Demay,

2014

Changis-

sur-Marne

Mammuthus primigenius 1100 349 150 - 142 - - 198 -

Demay,

2014

Changis-

sur-Marne

Mammuthus

primigenius 1200 320 156 - 122 - - - -

Petrova, et al., 2017 Ahlen

Mammuthus primigenius 1240 - 183 - - - - - -

Petrova, et

al., 2017 Khoma

Mammuthus

primigenius 970 - 150 - 125 - - 177 -

Petrova, et al., 2017 Berezovka

Mammuthus primigenius 1050 - 153 - 122 - - 226 -

51

Figure 3.8: Measurements of the tibia of mammals by Von den Driesch, 1976.

Table 3.3: Measurements of 30 individual tibias of Palaeoloxodon antiquus and Mammuthus primigenius taken

from the literature. The specimen of Santo Antão do Tojal is marked. In order to fit the table, abbreviations have

been used: ML, maximum length; AL, articular length; PW, proximal width; PT, proximal thickness; PAW,

proximal articular width; PAT, proximal articular thickness; MDW, minimum diaphysis width; MDT, minimum

diaphysis thickness; MDC, minimum diaphysis circumference; DW, distal width; DT, distal thickness; DAW,

distal articular width; DAT, Distal articular thickness. All measurements are in millimeters.

Work

Species

attributed ML AL PW PT PAW PAT MDW MDT MDC DW DT DAW DAT

Larramendi et al., 2017

Palaeoloxodon antiquus 795 775 278 - 263 163 151 136 440 232 180 231 62

Larramendi et al.,

2017

Palaeoloxodon

antiquus 638 625 217 171 210 152 94 80 280 183 130 169 52


Palaeoloxodon antiquus 842 820 286 234 274 183 140 139 430 252 182 215 67

Larramendi et al.,

2017

Palaeoloxodon

antiquus 914 794 312 - 274 164 152 - - 237 - 205 63


Palaeoloxodon antiquus 576 490 208 150 188 123 98 83 - 170 132 153 47

Larramendi et al.,

2017

Palaeoloxodon

antiquus 624 598 205 143 192 139 89 87 288 167 133 145 67

Kevrekidis & Mol, 2016

Palaeoloxodon antiquus - - - - - - 114 105 335 - - - -

Konidaris et al.,

2017

Palaeoloxodon

antiquus 792 - - - - - - - 360 - - - -

Andrews & Forster-Cooper,

1928

Palaeoloxodon

antiquus 1020 - 281 183 - - 150 - - 230 - - -

Van Kolfschoten,

1993

Palaeoloxodon

antiquus ~775 - ~265 183 245 - 115 113 - ~200 - - -

Van Kolfschoten,

1993

Palaeoloxodon

antiquus 602 - 208 - - - - - - - - - -

Van Kolfschoten, Palaeoloxodon 600 - 198 - - - - - - - - - -

52

1993 antiquus

Larramendi, 2015

Palaeoloxodon

antiquus 880 - - - - - - - 435 - - - -

Athanassiou, 2001 Palaeoloxodon

antiquus - - - - - - 108 85 - 177 136 - -

Zbyszewski,

1943; this work

Palaeoloxodon

antiquus - - 270 190 - - 110 125 400 - - - -

Christiansen, 2004

Mammuthus primigenius 546 - - - - - 80 66 230 - - - -

Christiansen,

2004

Mammuthus

primigenius 499 - - - - - 81 70 237 - - - -

Christiansen, 2004

Mammuthus primigenius 548 - - - - - 87 74 252 - - - -

Larramendi, 2015

Mammuthus

primigenius 800 - - - - - - - 341 - - - -

Larramendi, 2015

Mammuthus

primigenius 540 - - - - - - - 248 - - - -

Kirillova et al.,

2012

Mammuthus

primigenius 501 - 174 149 - - 78 70 - 139 117 - -

Kirillova et al., 2012

Mammuthus primigenius 500 - 182 148 - - 78 69 - 141 117 - -

Lister, 2009

Mammuthus

primigenius 680 - 228 180 - - - - - 180 147 - -

Lister, 2009 Mammuthus primigenius 685 - 227 181 - - - - - 182 141 - -

Harington et al.,

2012

Mammuthus

primigenius 588 - 197 - - - 90 - 259 123 - - -

Harington et al., 2012

Mammuthus primigenius 595 - - - - - 90 - - - - - -

Adams, 1881

Mammuthus

primigenius 597 - 266 - - - - - 266 - - - -

Adams, 1881

Mammuthus primigenius 508 - - - - - - - - - - - -

Adams, 1881

Mammuthus

primigenius 558 - - - - - - - 272 - - - -

Demay, 2014 Mammuthus primigenius 640 - 217 159 - - 102 - - 153 144 - -

3.5 Body mass estimation

The body mass of an organism is one of the most important parameters for understanding its biology,

being it closely related to metabolic rate, reproductive strategy, development and many other traits.

But reconstructing the body mass of dead animals, sometimes of extinct species is a challenge.

Traditionally, allometric equations based on the living proboscideans have been used, but they are not

reliable for inferring the masses of long gone species, with different proportions and much larger or

smaller masses. Recently the volumetric methods have started to be recognized as more accurate for

reconstructing the masses of such animals; although this method has its own shortcomings: Full body

and precise restorations based on well preserved and complete specimens are needed to achieve

reliable results (Larramendi, 2015; Jukar et al., 2018). Considering the incompleteness of the Santo

Antão do Tojal proboscidean, the total length for both bones discovered was inferred from the

equation developed by Christiansen (2007). The shoulder height of the animal was calculated in base

of the specific ratios for Palaeoloxodon antiquus, developed by Larramendi (2015). Finally in base of

a specific allometric equation developed from volumetric reconstruction of better preserved specimens

presented in that work, a body mass is given.

To estimate the body mass of the Santo Antão do Tojal proboscidean the total length for both bones

should be inferred first thanks to the equation developed by Christiansen, (2007):

log Y = α + β log X

53

“Y” represents the maximum length of the bone in mm, “X” its least circumference in mm and “ α ”

and “β” represent two constants, empirically parameterized either for Elephantidae Gray, 1821

(femur) or for proboscideans in general (tibia). Considering the measurements of the least

circumference for both bones (470 mm for the femur and 400 for the tibia), the above equation

provides an estimation for the lengths of the femur (1388 mm) and tibia (865 mm). To test how

accurate these estimations are we can look to table 3.4 were they are compared with other similarly

sized specimens in the dataset (ratio of maximum length vs minimum diaphysis circumference of the

hind limb bones). In table 3.5 we have the ration between femur and tibia total lengths.

Table 3.4: Maximum length vs minimum diaphysis circumference ratios of femur and tibia, for selected

Palaeoloxodon antiquus specimens. All measurements in mm.”E” stands for “estimated”.

FEMUR

Work Specimen Maximum length

Minimum

diaphysis

circumference

Maximum length vs diaphysis

circumference ratio

Larramendi et al., 2017 152-E9 1320 470 2,81

Mazo, 1998 ? 1320 460 2,87

Melentis, 1963 AMPG 1960/32 1372 478 2,87

This work

Santo Antão do

Tojal 1388 E 470 2,95

TIBIA

Work Specimen Maximum length

Minimum

diaphysis circumference

Maximum length vs diaphysis circumference ratio

Larramendi et al.,

2017 152-E9 795 440 1,81

Larramendi et al., 2017 167-E43A 842 430 1,96

Konidaris et al., 2017

MAR-1-940/675-

50 792 360 2,20

Larramendi, 2016 Konin 880 435 2,02

This work

Santo Antão do

Tojal 865 E 400 2,16

Table 3.5: Femur vs tibia lengths for selected Palaeoloxodon antiquus specimens. All measurements in mm.”E”

stands for “estimated”.

Work Specimen

Maximum length

femur

Maximum length

tibia

Maximum length of femur vs Maximum length of

tibia ratio

Larramendi et al., 2017 152-E9 1320 795 1,66

Larramendi et al.,

2017 167-E43A 1398 842 1,66

Konidaris et al., 2017 MAR-1-940/675-

50 1330 792 1,68

Andrews & Cooper,

1928 Upnor 1500 E 1020 1,47

Larramendi, 2015 Konin 1429 880 1,62

This work

Santo Antão do

Tojal 1388 E 865 E 1,60

Looking to table 3.6 we can see that the estimated maximum length of femur is slightly larger than the

other specimens (greater length for the same circumference) but does not fall far from them. The tibia

is also in the upper part of the estimation but inside the variation of proportions of the species. If we

look at table 3.7 we can see that the tibia is larger for the same length of the femur than other similarly

sized animals, but again it falls in the variation of proportions for the species.

54

According to Larramendi, (2015) the shoulder height of a given individual Palaeoloxodon antiquus is

about 2,56 times the femur length and 4,15 the tibia length, which yields 3553 mm and 3589 mm of

skeleton shoulder height respectively. The average of this estimation gives a result of 3571 mm or 3,57

meters of shoulder height for the skeleton. According to Larramendi et al., (2017) for animals with

similar proportions to the Santo Antão do Tojal specimen the shoulder height with flesh for the species

should be about 20 cm larger than the skeletal one, therefore the shoulder height in life is estimated in

377 centimeters or 3,77 meters. This estimated value is used in the equation presented in the same

work for the body mass of average specimens of Palaeoloxodon antiquus:

Body mass in kilograms = 3,63 × 10-4

× (Shoulder height in cm)2,903

Once solved, the equation yields a result of 10940 kg or about 11 metric tons. On table 3.6 we can see

a comparison of these measurements with other specimens of Palaeoloxodon antiquus from different

sites. It must be noted that NOT all the shoulder heights and body mass estimations in the table have

been obtained by the same methods.

Table 3.6: Shoulder height and body masses of different Palaeoloxodon antiquus specimens. The specimens

below to the Santo Antão do Tojal one had not been calculated using the same methods as presented. Possible

males are marked in blue.

Work Site Specimen Shoulder height (cm) Body mass (metric tons)

Larramendi et al., 2017 Neumark-Nord 1 152-E9 363 9,5/9,8

Larramendi et al., 2017 Neumark-Nord 1 167-E43A 385 11,4/11,6

Larramendi et al., 2017 Neumark-Nord 1 151-E8 264 3,9

Larramendi et al., 2017 Neumark-Nord 1 171-E34A 295 4,5/5,4

Marra, 2009; Larramendi et al., 2016 Viterbo Fontana Campanile 381 11,3

This work Santo Antão do Tojal Santo Antão do Tojal 377 11

Kevrekidis & Mol, 2016 Amyntaio PHP AME 046/006 355 9,2

Konidaris et al., 2017 Marathousa 1 MAR-1-940/675-50 370 9

Jakubowski, 1996 Grabsch tz Grabsch tz 301 5,7

The Santo Antão do Tojal specimen seems to be average sized for a big male of this species, only

three centimeters below the expected average of 380-420 cm shoulder height (Larramendi, 2015). It is

almost as big as the Fontana di Campanile specimen of Viterbo and the specimen 167-E43A from

Neumark-Nord; being the age of the last one estimated in 47 years (Larramendi et al., 2017).

3.6 Analysis of metrical data

Completeness of the sample

There are two reasons why the sample is incomplete: First, some bones (like the Santo Antão do Tojal

specimen) are incomplete themselves and second, not all measurements considered here are presented

in the literature, even in the case of entire specimens. The completeness of the sample can be quite

problematic for its representativeness. In the case of the femur (Table 3.2) of 384 possible individual

measurements there are 198 represented, or 51,56% of them. For the tibia (Table 3.3) of 360 possible

individual measurements there are 163, representing 45,27% of them.

But not all measurements are equally represented, for example some (like the maximum length of the

tibia) appear in 90% of the specimens, meanwhile others (like the proximal articular thickness of the

tibia) appear only in 20% of the specimens. In the table 3.7 we can see the number of individual

measurements of the bone and the % of specimens in which they appear.

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Table 3.7: Number of individual measurements taken for proboscidean femurs and tibias of the considered

sample. Percentage of specimens where the measurement has been taken compared to the total is also presented.

The measurements that have been taken in 50% or more of the specimens are marked.

FEMUR

Measurement Number of individual measurements

% of specimens were the measurement has been

taken

Maximum length 42 87,50

Proximal width 11 22,92

Caput width 24 50

Caput thickness 10 20,83

Minimum diaphysis width 33 68,75

Minimum diaphysis thickness 21 43,75

Minimum diaphysis circumference 21 43,75

Distal width 24 50

Distal articular width 12 25

TIBIA

Maximum length 27 90

Articular length 6 20

Proximal width 18 60

Proximal thickness 12 40

Proximal articular width 7 23,33

Proximal articular thickness 6 20

Minimum diaphysis width 19 63,33

Minimum diaphysis thickness 14 46,67

Minimum diaphysis circumference 16 53,33

Distal width 15 50

Distal thickness 11 36,67

Distal articular width 6 20

Distal articular thickness 6 20

Univariate statistical analysis

After identifying the most promising measurements for analyzing the data, those were investigated in

R, version 3.2.2 of 64 bits; running in a computer Toshiba Satellite L655-1JV of 64 bits with an Intel

Core i5 and Windows 7 Home Premium as operative system. The measurements (table 3.4) are well

distributed across the bone, with at least one measurement in the proximal and distal epiphysis and one

in the diaphysis. In the table 3.8 we have the mean of each measurement and species for femur and

tibia, with a % in difference between those averages.

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Table 3.8: Mean for selected measurements in the hind limb of Palaeoloxodon antiquus and Mammuthus

primigenius, with percentage of difference between the species for each measurement and bone. All

measurements are given in millimeters.

FEMUR

Maximum length Caput width

Minimum diaphysis

width Distal width

Mammuthus primigenius 1.075,17 152,50 130,76 220,80

Palaeoloxodon antiquus 1.272,50 194,25 156,31 281,36

% Difference 15,51 21,49 16,34 21,52

% Overall difference 18,72

TIBIA

Maximum length Proximal width

Minimum diaphysis

circumference

Minimum diaphysis

width Distal width

Mammuthus primigenius 585,67 213 263,13 85,75 153

Palaeoloxodon antiquus 754,83 248 371 120,09 205,33

% Difference 22,41 14,11 29,08 28,60 25,49

% Overall difference 23,94

Looking at this table (table 3.5) we can see that on average the femur of Palaeoloxodon antiquus is

18,72% larger than the one of Mammuthus primigenius and the tibia is 23,94% larger. In figures 3.8

(Femur) and 3.9 (Tibia) there is the plot of means for each selected measurement according to the

bone and species; with an interval of confidence of 95%.

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Figure 3.9: Plot of means for the previously selected measurements of the femur of Mammuthus primigenius

(left) and Palaeoloxodon antiquus (right) with an interval of confidence of 95%. All measurements are in

millimeters.

In figure 3.9 we can see that there is no overlap in the 95% confidence intervals in all the selected

measurements of the femur except in the minimum diaphysis width.

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Figure 3.10: Plot of means for the previously selected measurements of the tibia of Mammuthus primigenius

(left) and Palaeoloxodon antiquus (right) with an interval of confidence of 95%. All measurements are in

millimeters.

59

As we see in figure 3.10, we can detect a reliable difference in all selected measurements of the tibia

except in the proximal width of the bone. Therefore, we are left with measurements for each bone that

are keen to distinguish between the two species. In the femur: maximum length, caput width and distal

width. In the tibia: maximum length, minimum diaphysis circumference, minimum diaphysis width

and distal width.

Of those seven measurements (Three in the femur and four in tibia) it is possible to record three in the

Santo Antão do Tojal specimen; one in the femur (Caput width) and two in the tibia (Minimum

diaphysis circumference and width). In figure 3.11 we can see the measurements of the Santo Antão

do Tojal specimen compared with the means of both bones for those measurements, falling it into the

variability of Palaeoloxodon antiquus in each case.

Figure 3.11: Measurements of Santo Antão do Tojal specimen (in red) compared to the means for them in

Mammuthus primigenius (left) and Palaeoloxodon antiquus (right) with a 95% confidence interval.

In some cases it seems that the range of variability of Palaeoloxodon antiquus is greater than in

Mammuthus primigenius, so now we will investigate it by using the density estimation of the two most

numerous measurements in each bone.

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Figure 3.12: Estimation of density for maximum length and minimum diaphysis width of the femur of

Mammuthus primigenius and Palaeoloxodon antiquus.

For the femur (Figure 3.12) we can see that the maximum length for Mammuthus primigenius follows

an almost perfect normal distribution around the mean of 1075 mm of length, meanwhile

Palaeoloxodon antiquus follows a bimodal distribution with peaks at 1100 mm and 1400 mm of total

length. Looking at the minimum diaphysis width we can see that in Mammuthus primigenius it does

not have neither a perfect bimodal or normal distribution, but a grouping of measurements at 120 mm

and 150 mm meanwhile in Palaeoloxodon antiquus the bimodal distribution is clearer, with one slope

around the 125mm and another around 175mm.

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Figure 3.13: Estimation of density for maximum length and minimum diaphysis width of the tibia of Mammuthus

primigenius and Palaeoloxodon antiquus

For the tibia (Figure 3.13) we can see for the maximum length in Mammuthus primigenius a relatively

weak bimodal distribution grouped mostly around 500 mm of total length, with fewer measurements

somewhat around 600 mm. Meanwhile for Palaeoloxodon antiquus, the bimodal distribution is again

clear, with the measurements grouped around 600 and 800 mm. Looking at the minimum diaphysis

width, the pattern is more complex: Mammuthus primigenius has three peaks of density around 80, 90

and 100 mm, when the mean is 85 mm. Palaeoloxodon antiquus bimodal distribution has peaks at 110

and 150 mm with the mean at 120 mm.

This bimodal distribution is interpreted therefore as proof of the more marked sexual dimorphism in

Palaeoloxodon antiquus.

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Multivariate statistical analysis

To investigate the sample in multivariate statistic the program Past (version 3.14) was used, running in

the same hardware as previously specified. This program allows running PCA (Principal Component

Analysis) without all data by replacing the missing data with the mean of the column. The results

reached by this technique should be therefore taken with caution.

First we will investigate the femur dataset. We can see that the three first principal components

account for more than 92% of the variation, with the first one alone amassing 80% of it. Then we can

see that the PC1 is related strongly with the maximum length of the bone and PC2 with its minimum

diaphysis circumference, meanwhile PC3 is a mix of different measurements with little differences

between them, as shown by the bootstrapping of N=100000 (Figure 3.14).

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Figure 3.14: From upper to bottom: Percentage of variation accounted by the first three Principal Component

of the femur; PC1, PC2 and PC3. In PC1 and PC2 the highest contribution comes from the maximum length and

the minimum diaphysis circumference respectively.

Plotting the PC1 versus the PC2 (Figure 3.15) we can see that for the same values of PC1 the

Palaeoloxodon antiquus specimens tend to score higher in PC2, therefore having greatest

circumferences and overall robustness; meanwhile the lower PC1 scores of all are obtained by

Mammuthus primigenius individuals (shortest length of the femur).

Figure 3.15: PC1 vs PC2 of the femur of selected proboscideans. The Santo Antão do Tojal specimen is marked

with the red square, Mammuthus primigenius in blue and Palaeoloxodon antiquus in black.

64

In the tibia dataset we can see a fairly similar situation (Figure 3.16). PC1 and PC2 account for most of

the variation. The measurements that contribute the most to them are respectively the same as in the

femur: The maximum length for the PC1 and the minimum diaphysis circumference in the PC2. The

PC3 is again a mix of different measurements without a clear meaning.

65

Figure 3.16: From upper to bottom: Percentage of variation accounted by the first three Principal Component

of the tibia; PC1, PC2 and PC3. In PC1 and PC2 the highest contribution comes from the maximum length and

the minimum diaphysis circumference respectively.

Plotting the PC1 vs the PC2 produces a situation relatively similar to the one in the femur (Figure

3.17). The Palaeloxodon antiquus specimens tend to have more robust tibia (higher PC2) for the same

lengths (PC1). It is also interesting to note the greatest variance of the tibia of Palaeloxodon antiquus

that could be reflecting the previously detected bimodal distribution in the univariate statistic.

Figure 3.17: PC1 vs PC2 of the tibia of selected proboscideans. The Santo Antão do Tojal specimen is marked

with the red square, Mammuthus primigenius in blue and Palaeoloxodon antiquus in black.

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3.7 Conclusions

- The remains of the Santo Antão do Tojal proboscidean (Palaeoloxodon antiquus) which is

around 80.000 YBP old; comprise a first phalanx, a neural arch and a right femur and tibia.

- The Santo Antão do Tojal specimen is estimated to have measured alive about 3,8 meters in

the shoulder and weighted nearly 11 tons, being it in the average for a male Palaeoloxodon

antiquus. Despite being one of the most recent and Southwestern occurrences of this species,

it is nearly identical to specimens that lived several thousand years before in Central Europe or

the Italian Peninsula.

- The size difference between Mammuthus primigenius and Palaeoloxodon antiquus is 18% on

average for the femur and 24% in the tibia. The best measurements for differentiating between

both species are in the femur maximum length, caput width and distal width; and in the tibia

maximum length, minimum diaphysis circumference, minimum diaphysis width and distal

width.

- We agree with previous works (Zbyszewski, 1943; Antunes & Cardoso, 1992) in that the

Santo Antão do Tojal proboscidean is a Palaeoloxodon antiquus in base of the caput width of

the femur and the minimum diaphysis circumference and width of the tibia, combined with a

deep trochanteric fossa in the femur and the overall size of the bone and its robustness.

- Palaeoloxodon antiquus measurements follow a clearer bimodal distribution than Mammuthus

primigenius ones, indicating a more pronounced sexual dimorphism.

- The hind limb anatomy of Palaeoloxodon antiquus reflects more robust bones than in

Mammuthus primigenius, even for similar sized animals, as other works have shown

(Larramendi et al., 2017).

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4.ALGAR DO VALE DA PENA: A NEW FOSSIL BEAR SITE FROM

PORTUGAL

4.1 Overview

On this chapter a new locality with Ursus remains will be presented. First we will talk in the

introduction about the location, geological settings, history of the cave and its morphology and

physical characteristics. Then we will talk about the works carried until this moment and the

techniques used in the expeditions. In the third part of the chapter we will talk about the bear remains

recovered in detail, including comments about the mode of preservation. Then comparisons with the

literature (Morphological and metrical) will be done in order to characterize the population of the

cave. Next, the claw marks found in the walls of the cave will be presented; being this the first time

that such markings are described in Portugal. Finally, there will a discussion about the data and some

conclusions about the bear population of the cave as well as some comments about further work to be

done in the cavity and with the material recovered.

4.2 Introduction

Location

The Algar do Vale da Pena is located near the village of Moita do Poço, parish of Turquel,

municipality of Alcobaça, district of Leiria. It is inside of the natural park “Parque Natural das Serras

de Aire e Candeeiros”. In the figure 4.1 we can see its location graphically.

Figure 4.1: Location of the Algar do Vale da Pena. Upper left, at peninsular level, it is situated in the

Portuguese Estremadura. Bottom, view at regional level, the closest city is Alcobaça. Upper right, view at local

level. Credit Google Earth.

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Geological settings

The cave is located in the Northwestern slope of the Candeeiros range, a mountain range that appear

between Rio Maior in the Southwest and Porto de Mos in the Northeast; spanning for about 30

kilometers and reaching about 600 meters ASL.

This range is part of the system “Estrela-Montejunto”, a series of ranges that divide Portugal from

Southwest to Northeast. It is formed in calcareous rocks of the Maciço Calcário Estremenho, limited

between the thrust of Arrife in the East and the fault of Candeeiros in the West, which runs parallel to

the Candeeiros range. The rocks that form the cave are part of the Candeeiros Formation, Middle

Jurassic in age, concretely Bajocian as we can see in figure 4.2 (Carvalho, 1996; Crispim, 2008).These

calcareous rocks are not fossiliferous and are either oolitic or micritic in texture (Figure 4.3). Finally,

we should note that there are some detritic deposits of Pliocene age that are located in the lower part of

the slope and correspond to a marine transgression (Zbyszewski & de Matos, 1959).

Figure 4.2: Litostratigraphic log of the “Maciço Calcário Estremenho”, with the Candeeiros Formation marked

(Carvalho, 1996).

Figure 4.3: Location of the cave (red and orange dot) in the corresponding sheet (26 D, Caldas da Rainha) of

the geologic map (1:50000) of Portugal, including legend (Zbyszewski & de Matos, 1959).

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History of the cave

This version of the history of the cave has been put together according to personal communications by

Manuel Pinto Soares, the first professional speleologist in Portugal, who was also one of the first

people to enter the cave, Octávio Mateus, of the FCT-UNL who contacted the park authorities for the

authorization to recover the bones and Sérgio Medeiros, president of the GPS (Grupo de Protecção

Sicó).

The calcareous rock of the Candeeiros range is exploited by several enterprises in open air quarries

inside the natural park since many decades ago (Carvalho, 1996); like the one already active that can

be seen south of the site in the figure 4.1. In one of such quarries, at the end of the decade of 1980,

there was a rock fall that connected the outside with the cave. The quarry workers thought that maybe

they could fill the gap and continue working as nothing happened, so they proceeded to throw several

tons of big rock blocks into the newly opened cavity; but after throwing as many blocks as they could

into the hole it became evident that they could not fill it.

They contacted with Olimpio Martins, worker and speleologist of the natural park, who told to them to

stop the exploitation and realized the first descend into the cave; then the beginning of the decade of

1990 (1991 or 1992) Manel Pinto Soares accompanied Olimpio and installed the nails in the walls of

the cave to descend safely. Olimpio told him about the bones at the end of the cave (“bear number

four” in this work, as we will discuss later) and after he took a picture, Olimpio said to him to keep the

secret about them. In that visit Manuel did not notice any other bone remains despite he passed nearby

the main bone assemblage (“bear” number one in this work).

As the years passed the difficult vertical entrance of the cave became popular within the speleological

community, as well as the claw marks and bone remains that were inside the cave and earned it the

nickname of “Algar dos Ursos” or “Cave of the bears”. Manuel even heard other people talk about

“three bears” or “three bear skulls”. During this time the cave was visited by groups of firefighters

doing rescue practices and a dog fell into the cave, where its bones can be seen nowadays.

In 2011 some cavernicolous oniscidea (a type of crustacean isopods) were recovered in the cave and

later described (Reboleira et al., 2015). In 2012 the GEM (Grupo de Espeleología e Montanhismo)

started the works in order to map the previously uncharted cave; this map was completed in august

2015 and will be used later in this work, they also put some security tape to prevent people from

stepping over sand deposits and the fossil remains (Amendoeira et al., 2015). Around that time (2013)

the scientist of the CAVE project visited the site; it was an international project aiming to understand

the cave evolution from an integrative perspective and its use for paleoenvironmental reconstruction

from the point of view of several disciplines like archaeology, isotopic studies and geomorphology;

but no study centered on this cave came to light (http://caveportugal.wixsite.com/cave/homeengl).

Sérgio Medeiros contacted with the Museu da Lourinhã to make them know about the fossil remains

and Octávio Mateus prepared the first of the visits to the cave that would lead to the work presented

here.

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Morphology of the cave

The cave is not fully mapped yet and we do not have a side view of it. The cartography that will be

used was produced by members of the GEM (Grupo de Espeleología e Montanhismo) during 2014 and

2015. In the figure 4.4 we have marked in black numbers some landmarks of the cave like claw marks,

small conducts or sand deposits. The location of the bone assemblages is marked with red numbers,

they will be referred as individual “bears” in the text but the MNI will be discussed in the next section.

Figure 4.4: Topography of the Algar do Vale da Pena. In black, some landmarks of the cave are marked and in

red, the bone findings (Modified from Amendoeira et al., 2015).

71

The entrance of the cave is not very clearly drawn in the topography, but we have some pictures of it

in the figure 4.5. It includes a first ramp of about ten meters of different pending, then a short three

meters vertical step and then another ramp of about the same length before the final vertical well of

about fifteen meters.

Figure 4.5: A) Entrance of the cave showing the first ramp and the top of two meters step. B) Picture taken from

the base of the two meters step, showing the second ramp. C) Final vertical well of the entrance from the bottom.

D) Picture from the bottom of the cave showing the final well (white line), the second ramp (green line), the two

meter step (red line) and part of the firs ramp (blue line). Pictures by Carlos Ferreira.

The final landing is made over a pile of blocks and dirt thrown by the quarry workers into the cave in

their unsuccessful attempt to fill it (1 in the figure 4.4). Once properly in the cave, without the need of

ropes anymore, we descend the pile of blocks until we see a bifurcation. If we go right we will reach

quickly some bone remains (“bear” number 5 in the figure 4.4) and also a small conduct with claw

marks on it and the recent dog skeleton at the end that we can see in the figure 4.6 (2 and 3 in map).

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Figure 4.6: Claw marks (left) and dog skeleton (right) in the small conduct at the right of the cave.

Following the main right tunnel of the cave we can go down in the left part of it until we reach a lake

or going up sticking to the right wall until we find more bone remains near a bottle neck and the

beginning of a new conduct (“bear” number 2 in figure 4.4).

Meanwhile taking the left main tunnel we will pass by a big room with a natural “railing” formed by

speleotems that pass to the left of a big depression and then starts climbing. As soon as we turn left

from it we will find a panel with even more claw marks and following the easiest path another mount

of clay with many claw marks on it (number 4 and 5 in the map). Then going down right from that

mount we will find the main fossil findings and six meters at its left another bone assemblage

(numbers 1 and 3 in the figure 4.4). We can see sands near these findings (Figure 4.7 and number 6 in

the map). Finally after crossing another narrow “railing” we will find the best claw marks in the cave,

with the number 7 (Figure 4.7) and the last skeletal remains in the cave (“bear” number 4).

Figure 4.7: Thick deposit of sands below the calcareous crust (left) and claw marks (right).

After this last bone remains the cave starts to climb until it ends in a “cul-de-sac”. The location of the

original entrance of the cave is a mystery, and it might be impossible to find it if the path to it was

buried by the pile of tons of rocks that opened the current entrance.

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4.3 Methods, expeditions to the cave and preparation of the material

Up to the moment of delivering this thesis (March 2019) there has been three expeditions to the cave.

In all of them the entrance has been possible thanks to the support of the GPS (Grupo de Protecção

Sicó) that offered to the author and accompanists the material and technical support to descend and

climb safely by means of vertical progression through the 30 meters depth entrance. To descend, two

pieces of equipment called the “shunt” and the “descensor” (Figure 4.8) have to be used. The

technique consists in letting the rope pass through the “descensor” controlling the descend velocity

with the position of the hand, up to go slow, down to go faster. The “shunt” is a security measurement,

if the weight of the person is too far from it, it will lock. Therefore it has to be close to the “descensor”

being pulled down with two fingers at the same time that the body of the climber goes down. To

climb, the “ventral croll” and the “climbing fist” (Figure 4.8) have to be used at the same time pulling

the body up with the help of the later and letting the rope to pass by the first one, then it will lock in

the new position, so the climb is done in “steps”. Either going down or up, when the end of a rope is

found the climber has to secure himself in the nail of the wall and change the equipment to the new

rope, keeping in all moments two elements of security. There are five of these changes in the way up

and down of the cave.

Figure 4.8: Equipment needed to descend (in black) and climb out of the cave again (in red). Left; A) Shunt, B)

Descensor, C) Climbing fist. Right; D) Ventral croll and security knot to stop descending, note how the fingers

secure the shunt.

It has to be noted that since its opening the cave has become an important refuge for bats, so the visit

is not allowed by the Natural Park authorities between November to April because the disturbance to

hibernating bats can cause them to exhaust their energy reserves and die. Every visit to the cave has

been communicated to the corresponding authorities and in April 2017 an agreement was reached

between the municipality of Alcobaça, the Natural Park and the FCT-UNL to take out the fossils of the

cave and deliver them prepared to the Alcobaça municipality three years after they are collected.

First expedition (15 of October, 2016)

During the first expedition (Figure 4.9) the most distinguishable claw marks were recognized, as well

as the bones of the “bear” number four and the main finding (the “bear” number one) was started to be

dug. Soon it became evident that it could not be taken out with the small tools brought in this first

expedition that could not pierce through the thick layer of calcareous crust in the side of the block.

Also it was impossible to take the skull alone without a serious risk of destroying it, so the block had

to be taken entirely.

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Figure 4.9: First excavation around the skull of “bear number one” in October 2016.

Second expedition (14 of October, 2017)

During the second expedition (Figure 4.10) more claw marks were located in several panels of the

cave, besides the remains of the “bear number five” and the dog that fell into the cave. In this

expedition it also became clear that in order to take out the block a big expedition should be assembled

and many people specialized in cave speleology would be required. Also Manuel Pinto Soares was

interviewed about his knowledge of the cave. Rui Guerra, a documentalist and camera started the

elaboration of a documentary about the digging in the cave filming the bear site and the surroundings.

Up until this moment no bone remains where taken out of the cave.

Figure 4.10: The main block as in October 2017, during the second expedition, no new diggings where carried

out this time. The skull is near the scale of seven centimeters,

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Third expedition (16 and 17 of June, 2018)

This time a big expedition was set with 13 participants in the first day. The jeep of the FCT-UNL as

well as its powerline and circular saw specially bought for the task were used, also the drill and stone

splitters of Octávio Mateus and the power generator and illumination of the GPS. First the clay and

mud was dug around the block of the “bear number one” with relative ease (more about it in the next

section). Then the bones were protected with a first layer of plasters and the block with the main bones

was successfully cut thanks to the drill and the stone splitters where the source of light was in Figure

4.10. Then the block was put in a net. In order to move it some members of the team went out of the

cave, cut an eucalyptus tree of about 10 meters tall, removed the branches and descended it safely into

de cave to be used. Even thanks to the tree, the block weighed several hundred kilograms and could

only be taken up to the first point where the path goes to the right, since is not possible for so many

people to maneuver in the confined space of the “railing”. Finally it was conveniently plastered with

several extra layers. In the figure 4.11 we have a series of pictures about that process.

Figure 4.11: A) Digging of the main block of “bear one” (note the camera of Rui Guerra). B) First layer of

plaster after the block was cut. C) Putting the block in the net. D) Moving the block with the tree trunk. E) Final

plastering of the block. F) Current state of the block. Pictures by Carlos Ferreira.

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At the same time that the block was receiving its final plastering Octávio Mateus moved alongside the

walls of the cave counting all the claw marks that he could find and discovering the remains of the

“bears” number two and three. As much bones as possible were collected, including three small

fragments from “bear number four”. The remains of “bears” two and three that could not be taken

away were covered with plaster. In the next sections we will discuss more precisely which remains

were taken, which ones stay in the cave and what was the position and preservation of the elements, as

well as the some notes about the taphonomy and a comparison of the remains with other bears.

The remains (a total of 37 items) were stored during summer in the Museu da Lourinhã and were

prepared in the same institution in October of 2018. Use of glues (Paraloid B72, at 5% of

concentration in acetone for consolidation and 40% for gluing) was reduced to the minimum possible

and the preparation was carried out only in a mechanical manner with several air-scribes in order to

improve the possibilities for hypothetical isotopic dating, ancient DNA and other isotopic diet studies.

In the figure 4.12 we can see the “before and after” of some bones recovered.

Figure 4.12: Before ( left) and after( right) for some ursid remains recovered in Algar do Vale da Pena: a)

distal radius, b) mandibular ramus and c) basicranium.

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4.4 The bone assemblages

Bear number one

It appears as 1 in figure 4.4 and is the main finding in the cave.

Items still in the cave: Main block with a nearly complete skull, scapula and humerus (Figure 4.13).

The process to remove this block and its current status appear in figure 4.11, also numerous bone

fragment and smaller bones still in the mud.

Figure 4.13: The main block of “bear” one during the second expedition. Scapula and protruding long bones

are marked with black arrows. In red, line where the block was cut, main skull with the scale of 7 cm and muddy

area with numerous smaller bone remains.

Comments: This finding seems to have been subject to little if any transportation of the bones. The

skull posses the articulated mandible and this is highly unlikely to be conserved if it was subject to the

movement. The bones seem to have been covered first with thin layer of calcareous crust, and then the

skull and the bones in the main block were covered (At least partially) with several thick layers of

calcareous crust, meanwhile that the bones in the muddy area were covered by sands and mostly clays.

Bear number two

It appears as 2 in figure 4.4, was discovered in the third expedition and is the second best finding in

the cave for now.

Items still in the cave: Plaster jacket in situ including at least some partial remains.

78

Figure 4.14: Remains of the “bear number two” before (left) and after (right) removal of the block that was

hiding them.

Comments: This assemblage of bones seems to have been moved by subterranean waters from their

original position and eroded in the process. In the figure 4.15 we can see how there is muddy sediment

below the calcareous crust in the hemimandible during the preparation, meanwhile in the previous

assemblage it was the opposite. Also the anterior part of the hemimandible and other remains show

signs of breakage and erosion before being covered by the calcareous crust.

Figure 4.15: Muddy sediment below the calcareous crust, during the preparation of the hemimandible (ADVP

2.1). Scale 7 cm.

Bear number three

This bone assemblage is marked with the number 3 in the figure 4.4. It is only situated six meters

below the first finding, and it could not be ruled out yet that it belongs to the same individual as the

first one.

Items still in the cave: Partial tibia with on isolated block plaster jacket and other remains (proximal

femur and possible fibula) in plaster jacket in situ.

79

Comments: This assemblage of bones is the worst preserved of all. There is not a single intact bone,

being all of them heavily eroded and quite fragile. This fact contributes to the hypothesis that these

bones were transported from the first site by water.

Bear number four

This assemblage is situated at the end of the cave, the number 4 in the map of figure 4.4.

Items still in the cave: Humerus, femur, and part of a pelvis totally encrusted in calcareous matrix

plus several smaller bones equally encased in the crust (figure 4.16).

Figure 4.16: Material of bear number four encased in the calcareous crust. Left, humerus; right, femur and

pelvis.

Comments: Is quite difficult to know if these bones are as well preserved as it seems, because they are

completely embedded into the calcareous matrix. The recovery of more material of this assemblage

seems nearly impossible. Even cut in blocks with a circular saw and successfully transported outside

the cave, the preparation has so many setbacks that the best option is probably to let them in the place.

Bear number five

This assemblage is near the bear number two; it appears with the number 5 in the map of figure 4.4

Items still in the cave: Distal humerus (figure 4.17), and possible tibia encased in calcareous crust.

Figure 4.17: Distal humerus of the bear number five.

Comments: Despite being also encased in the crust this bones could be recovered easier than the ones

of the previous assemblage, since the crust is thinner here.

80

4.5 Results: The recovered bone remains

In this section we will list the 37 recovered items; each of them will have the field number that

consists in the cave (ADVP stands for “Algar do Vale da Pena”) the number of the bear and the

number of item in particular. They are stored at FCT-UNL. Then measurements and/or pictures of the

most relevant items (marked with an asterisk) will be presented. These measurements will be given

according to the system for ursid measurements provided by Tsoukala & Grandal-d’Anglade (2002).

Then they will be compared with other brown and cave bears as well as with Asiatic black bear in

some specimens.

Items recovered for “bear” one:

- ADVP 1.1: Partial left scapula *.

- ADVP 1.2: Scapula fragments.

- ADVP 1.3: Right scapholunate bone*.

- ADVP 1.4: Right hamate bone*.

- ADVP 1.5: Highly damaged metatarsal.

- ADVP 1.6: Second phalanx*.

- ADVP 1.7: Proximal epiphysis of third right metatarsal *.

- ADVP 1.8: Distal epiphysis of right radius*.

- ADVP 1.9: Distal part of diaphysis of left ulna*.

- ADVP 1.10: Fragments of rib.

- ADVP 1.11: Undifferentiated bone fragments.

- ADVP 1.12: Calcareous matrix pieces in contact with bone.

- ADVP 1.13: Bone fragments recovered in the first excavation.

- ADVP 1.14: Big calcareous crust.

Items recovered for the “bear” two:

- ADVP 2.1: Posterior part of right hemimandible *.

- ADVP 2.2: Parietal and occipital *.

- ADVP 2.3: Temporal bone*.

- ADVP 2.4: Cranial fragments.

- ADVP 2.5: Heavily damaged canine *.

- ADVP 2.6: Left upper fourth premolar *.

- ADVP 2.7: Left upper first molar *.

- ADVP 2.8: Right lower second molar *.

- ADVP 2.9: Left lower third molar *.

- ADVP 2.10: Partial third phalanx*.

- ADVP 2.11: Bone fragments.

- ADVP 2.12: Calcareous crust in contact with bones.

Items recovered for the “bear” three: - ADVP 3.1: Partial distal epiphysis of right femur*.

- ADVP 3.2: Vertebral centrum *.

- ADVP 3.3: Highly damaged vertebral centrum.

- ADVP 3.4: Fragment of rib.

- ADVP 3.5: Bone laminae (Part of the proximal femur or the tibia).

- ADVP 3.6: Possible fragment of sacrum.


- ADVP 3.8: Calcareous matrix.

Items recovered for the “bear” four: - ADVP 4.1: Proximal epiphysis of second right metatarsal*.

- ADVP 4.2: Damaged proximal epiphysis of fifth right metatarsal*.


81

ADVP 1.1: Partial left scapula

This bone (Figure 4.18) can only be recognized as a scapula thanks to the calcareous crust that is

holding it together. During preparation it was clear that removing the calcareous crust would damage

irreversibly the already compromised bone.

Figure 4.18: Damaged scapula (ADVP 1.1) from “bear number one” (Ursus arctos) of Algar do Vale da Pena.

ADVP 1.3: Right scapholunate bone

This carpal bone (Figure 4.19) is nearly in pristine condition, and one of the best preserved in “bear

one”. Its measurements are presented in the table 4.1. The orientation and angle of the apophysis rule

out an Ursus thibetanus Cuvier, 1823 origin (Torres Pérez-Hidalgo, 1984).

Figure 4.19: Right scapholunate bone (ADVP 1.3) from “bear number one” (Ursus arctos) of Algar do Vale da

Pena in A) proximal; B) distal; C) anterior and D) exterior views.

82

Figure 4.20: Measurements for the scapholunate bone in ursids according to the system of Tsoukala & Grandal-

d’Anglade (2002). All of them could be recorded for the specimen.

Table 4.1: Measurements from the right scapholunate bone (ADVP 1.3) from “bear number one” (Ursus arctos)

of Algar do Vale da Pena compared to a set of measurements of other brown bears (García Vázquez, 2015) and

cave bears (unpublished own data from Cova da Ceza population). Measurements are in mm according to the

system of Tsoukala & Grandal-d’Anglade (2002).

Species Specimen L DT DAP DTart.prox. DAPart.prox. DTart.dist. DAPart.dist.

U. arctos? ADVP 1.3 24 43 44 39 24 22 37

U. arctos LCF-137 26,79 49,43 47,12 42,89 26,82 27,74 41,23

U. arctos SIPA-52 26,27 43,59 43,02 39,54 25,75 23,12 38,45

U. arctos SIPA-118 31,3 51,12 49,55 47,33 30,13 28,4 42,7

U. arctos PUR-80 27,69 49,08 47,78 46,86 28,81 29,4 44,2

U. arctos SH5-98-S28-089 33,14 50,86 54,39 49,46 30,75 30,13 48,41

U. arctos SH5-98-S28-100 33,23 51,51 53,1 50,66 31,8 30,93 47,45

U. arctos SH5-97-T29-25 25,83 42,62 41,15 37,37 24,87 23,67 33,31

U. arctos SH5-97-U29-079 24,34 42,35 40,64 37,05 24,87 23,95 33,17

U. arctos TA-Lu-c-34 26,65 44,33 42,01 39,28 23,71 23,04 36,33

U. arctos

42,42 45,89 41,56 27,61

U. arctos

42,81 44,57 43 33,02

U. arctos El Cuervo 25,31 39,95 37,11 39,24 24,97 25,23

U. spelaeus CEZ 42 36,49 58,8 57,67 57,27 36,34 32,48 56,13

U. spelaeus CEZ 171 25,45 48,24 51,17 47,17 29,05 23,68 42,37

U. spelaeus CEZ 729 28,9 49,84 50,19 48,75 30,39 27,22 46,22

U. spelaeus CEZ 730 28,42 49,8 49,88 47,24 30,3 28,64 45,97

U. spelaeus CEZ 731 40,36 64,61 68,71 65,44 42,99 38,34 61,69

According to García Vázquez, (2015) the transverse vs anteroposterior diameters of the scapholunar

are the best measurements to distinguish between males and females of brown bear; so in figure 4.21

we have a XY plot of those measurements.

83

Figure 4.21: Transverse diameter vs anteroposterior diameter of the scapholunate of brown bears (García

Vázquez, 2015) as brown dots and cave bears (unpublished own data from Cova da Ceza population) as green

squares; plus the scapholunate ADVP 1.3, marked as a yellow star.

Looking at the graph we can say that the dimensions of the scapholunar are similar to the expected on

brown bears so in figure 4.22 we exclude the cave bears.

Figure 4.22: Transverse diameter vs anteroposterior diameter of the scapholunate of brown bears (data from

García Vázquez, (2015)) as brown dots; plus the scapholunate ADVP 1.3, marked as a yellow star.

So ADVP 1.3 is classified as the scapholunate of a female brown bear in base of the ratio between

transversal and anteroposterior diameters plus their overall dimensions.

Putative females

Putative males

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ADVP 1.4: Right hamate bone

This carpal bone (Figure 4.23) is only slightly damaged and is still one of the best preserved of this

bear, but taking into account the morphology of this bone and the difficulties to measure it properly

(many of the views looks similar) only one measurement has been recorded. This measurement is

presented in the table 4.2 and we can graphically see them at figure 4.24.

Figure 4.23: Right hamate bone (ADVP 1.4) from “bear number one” (Ursus arctos) of Algar do Vale da Pena.

Figure 4.24: Measurements for the scapholunate ursid bone according to the system of Tsoukala & Grandal-

d’Anglade (2002).The recorded measurement is marked.

85

Table 4.2: Length of the right hamate bone (ADVP 1.4) from “bear number one” (Ursus arctos) of Algar do

Vale da Pena compared to the length of the same bone of other brown bears (García Vázquez, 2015) .


Individual Light

ADVP 1.4 27

LCF-141 33,07

LCF-142 27,1

SIPA-109 33,1

SIPA-111 32,17

Pur-Lu-33 35,37

SH5-98-S28-064 35,54

SH5-97-T29-084 25,75

TA-Lu-c-33 30,74

Figure 4.25: Length of the right hamate bone (ADVP 1.4) from “bear number one” of Algar do Vale da Pena

compared to the length of the same bone of other brown bears (García Vázquez, 2015) . Measurements are in

mm according to the system of Tsoukala & Grandal-d’Anglade (2002). Two groups (males and females) are

classified according to this measurement.

As we can see in figure 4.25, ADVP 1.4 has the second smallest length of the hamate in the sample of

brown bears; therefore it is classified as belonging to a female or juvenile brown bear.

20

22

24

26

28

30

32

34

36

Putative males

Putative females

86

ADVP 1.6: Second phalanx

This phalanx (Figure 4.26) has been classified as a second phalanx due to its morphology compared to

the work of Torres Pérez-Hidalgo, (1984). Some parts of it are slightly damaged, but overall the

condition is good. In the table 4.3 we have the longitude and distal transversal diameter the of the

second phalanx compared with the same measurements of the second phalanxes of a recent brown

bear, from which we know if the bones are from hindlimb or forelimb (García Vázquez, 2015). In

figure 4.27 we can see the measurements graphically. In figure 4.28 we have an XY plot elaborated

with the data of table 4.3.

Figure 4.26: Second phalanx (ADVP 1.6) from “bear number one” (Ursus arctos) of Algar do Vale da Pena in

A) dorsal; B) ventral; C) proximal; D) distal views.

Figure 4.27: Measurements for the second phalanx of ursids according to the system of Tsoukala & Grandal-

d’Anglade (2002). All of them could be recorded.

87

Table 4.3: Length vs distal transverse diameter of the second phalanx (ADVP 1.6) from “bear number one”

(Ursus arctos) of Algar do Vale da Pena compared to the length of the same bone of a recent brown bear

(García Vázquez, 2015) . Measurements are in mm according to the system of Tsoukala & Grandal-d’Anglade

(2002).

Anterior/posterior Number L DTdist.

? ADVP 1.6 25 11

Anterior LE1 28,97 14,53

Anterior LE1 29 14,87





Posterior LE1 21,55 12,11




Figure 4.28: Length vs distal transverse diameter of the second phalanx (ADVP 1.6, marked as blue square)

from “bear number one” (Ursus arctos) of Algar do Vale da Pena compared to the same ration of a recent

brown bear, data from García Vázquez, (2015) . The 95% ellipses are drawn in the graphs. Measurements are in

mm according to the system of Tsoukala & Grandal-d’Anglade (2002).

Therefore, according to the figure 4.28 the second phalanx is more likely to come from the pes of the

bear.

88

ADVP 1.7: Proximal epiphysis of third right metatarsal

This bone has been classified as the proximal epiphysis of the third right metatarsal in base of its

morphology (Figure 4.29). Only its proximal transverse and anteroposterior diameters could be

recovered. They are presented in the table 4.4 alongside the same measurements of brown bears from

NW Spain (García Vázquez, 2015). In Figure 4.30 we see the measurements. In Figure 4.31 we can

see graphically the comparison of table 4.4.

Figure 4.29: Proximal epiphysis of the third right metatarsal (ADVP 1.7) from “bear number one” (Ursus

arctos) of Algar do Vale da Pena in A) exterior; B) dorsal; C) interior and D) ventral views.

Figure 4.30: Measurements for the third metatarsal of ursids according to the system of Tsoukala & Grandal-

d’Anglade (2002).The recorded measurements are marked.

89

Table 4.4: Proximal transverse diameter and proximal anteroposterior diameter of the third metatarsal (ADVP

1.7) from “bear number one” (Ursus arctos) of Algar do Vale da Pena compared to same measurements of fossil

brown bears from NW Spain (García Vázquez, 2015) . Measurements are in mm according to the system of

Tsoukala & Grandal-d’Anglade (2002).

Specimen Dtprox. DAPprox.

ADVP 1.7 11 21

LCF-129 13,05 21,44

LCF-130 17,13 24,68

CGLL-028 17,37 27,87

SIPA-6 18,66 26,45

SH5-97-T29-42 16,13 23,05

SH5-98-V30-016 16,04 26,57

SH5-98-R29-001 17,06 25,03

SH5-97-T29-59 14,8 23,73

SH5-98-S28-075 20,26 30,67

SH5-98-T31-002 16,71 25,57

TA-Lu-c-22 15,61 23,37

Figure 4.31: Proximal transverse diameter vs Proximal anteroposterior diameter of the third metatarsal (ADVP

1.6, marked as blue square) from “bear number one” (Ursus arctos) of Algar do Vale da Pena (As green

triangle) compared to the same ration of fossil brown bears from NW Spain, data from García Vázquez, (2015)

. Measurements are in mm according to the system of Tsoukala & Grandal-d’Anglade (2002).

As we can see in figure 4.30, the metatarsal from Algar do Vale da Pena is the smallest from the entire

sample, almost as it belonged to a juvenile, taking into account how slender it is (DT prox), although it

has to be taken into account that it could be slightly wear down.

90

ADVP 1.8: Distal epiphysis of right radius

This distal epiphysis of the right radius, ADVP 1.8 (Figure 4.32) is the third most complete postcranial

bone recovered until this point in the whole site. It shows a totally fused epiphysis; therefore it is

interpreted as belonging to an adult or nearly adult bear. In table 4.5 we can see its measurements

compared with brown bears from NW Spain (García Vázquez, 2015), as well as some measurements

from Portuguese caves (only maximum and minimum transverse diameters of the bone) as presented

in Cardoso, (1993). In figure 4.33 we have graphically the measurements of the radius according to


Figure 4.32: Distal epiphysis of the right radius (ADVP 1.8) from “bear number one” (Ursus arctos) of Algar do

Vale da Pena in A) anterior; B) exterior; C) Posterior; D) Interior and E) Distal views.

Figure 4.33: Measurements for the radius of ursids according to the system of Tsoukala & Grandal-d’Anglade

(2002). Recorded measurements are marked.

91

Table 4.5: Measurements from the radio (ADVP 1.8) from “bear number one” of Algar do Vale da Pena

compared to same ratio of fossil brown bears from NW Spain (García Vázquez, 2015) as well as some of

Portuguese caves. Measurements are in mm according to the system of Tsoukala & Grandal-d’Anglade (2002).

DTdist. DAPdist. DTart.dist. DAPart.dist.

ADVP 1.8 52 32 33 27

CB-004 54,45 34,98

LCF-008-1996 66,13 41,14

LCF-009-1996 66,84 40,65

LCF-96-037 65,68 39,59

LCF-96-038 53,98 31,58

CGLL-043 55,91 35,42

SIPA-3 55,25 32,19

Pur-Lu-19 62,82 38,25

Pur-Lu-20 58,73 40,43

1996-SH5-001 52,76 32,56

SH5-97-V28-001 52,49 32,05

SH5-98-S28-035 65,81 41,75

SH5-98-S29-011 55,29 32,95

SH5-98-U30-012 57,61 33,88

TA-Lu-c-6 55,41 32,77

56,53 36,19

El Cuervo 50,34 32,48

Furninha max 62

Furninha min 56,2

Fontainhas max 63,5

Fontainhas min 58,7

Figure 4.34: Transverse diameter of distal radius from ADVP 1.8 and fossil and recent brown bears from NW

Spain and Portugal (Cardoso, 1993; García Vázquez, 2015). Specimens with more than 60 mm are classified as

male and less than 58 mm as females.

92

Figure 4.35: Transverse diameter vs anteroposterior diameter of distal radius from ADVP 1.8 (Green in the

graph) and fossil and recent brown bears from NW Spain (García Vázquez, 2015). Two clusters of individuals

are classified as females or males.

As we can see in figures 4.34 and 4.35 ADVP 1.8 can be classified according to its measurements as

the distal part of the radius of a small female brown bear.

ADVP 1.9: Distal part of diaphysis of left ulna

This distal part of the ulnar diaphysis (Figure 4.36) has its epiphysis highly damaged; therefore only

one measurement could be taken. In figure 4.37 we have graphically the measurements of the ulna

according to Tsoukala & Grandal-d’Anglade (2002). In table 4.6 we have the measurement of ADVP

1.9 together with equivalent measurements of brown bears from NW Spain (García Vázquez, 2015).

Figure 4.36: Distal distal part of the ulnar diaphysis (ADVP 1.9) from “bear number one” (Ursus arctos) of

Algar do Vale da Pena in A) anterior; B) exterior; C) Posterior and D) Interior views.

93

Figure 4.37: Measurements for the ulna of ursids according to the system of Tsoukala & Grandal-d’Anglade

(2002). Taken measurement is marked with a circle.

Table 4.6: Minimum anteroposterior diameter of the ulnar distal diaphysis (ADVP 1.9) from “bear number

one” (Ursus arctos) of Algar do Vale da Pena compared to the same measurement of fossil and recent brown

bears from NW Spain (García Vázquez, 2015). Measurements are in mm according to the system of Tsoukala &

Grandal-d’Anglade (2002).

Specimen DAPdia.min.

ADVP 1.9 20

CB-001 22,37

CB-002 22,81

LCF-010-

1996 28,04

LCF-163 27,42

LCF-164 22,13

EIX-009 22,82

CGLL-053 26,44

SIPA-4 21,82

SIPA-63 21,57

Pur-Lu-18 27,72

SH5-97-T29-

11 20,81

SH5-97-T29-

27 20,67

TA-Lu-c-4 22,55

El Cuervo 23,32

94

Figure 4.38: Minimum anteroposterior diameter of the ulnar distal diaphysis (ADVP 1.9) from “bear number

one” (Ursus arctos) of Algar do Vale da Pena compared graphically to the same measurement of fossil and

recent brown bears from NW Spain (García Vázquez, 2015). Suggested males and females are marked.

Acording to the data in table 4.6 and the figure 4.38 ADVP 1.9 is classified as the ulna of a small

female brown bear.

ADVP 2.1: Posterior part of right hemimandible

This is the best preserved cranial remain recovered in the cave so far. As we can see in figure 4.39, it

has been eroded in the anterior part and lost all its teeth; the mandibular condyle is also lightly

damaged. In figure 4.40 we can see the measurements taken to it and in table 4.7 their values

compared to other bears of NW Spain.

Figure 4.39: Posterior part of right hemimandible (ADVP 2.1) from “bear number two” (Ursus arctos) of

Algar do Vale da Pena in A) dorsal; B) posterior; C) exterior; D) anterior; E) interior and F) ventral views.

95

Figure 4.40: Measurements for the mandible of ursids according to the system of Tsoukala & Grandal-

d’Anglade (2002). Taken measurements are marked with a circle.

96

Table 4.7: Measurements from the partial hemimandible (ADVP 2.1) from “bear number two” (Ursus arctos)

of Algar do Vale da Pena compared to same measurements of fossil brown bears from NW Spain (García

Vázquez, 2015) as well as some cave bears from the same region (Unpublished own data from Cova de A Ceza).


Species Specimen L cond.-Mp H pro.ang.-Cor Hmd(M3)(la) Hcond DTmd(M2,M3) Hmd(max)

? ADVP 2.1 70 86 53 15 19 103

U. arctos LCF-003-1996 92,46 102,41 57,34 21,59 23,31 134,25

U. arctos LCF-002-1996 93,51 102 56,92 21,16 23,86 134,25

U. arctos LCF-155

44,98

19,47

U. arctos LCF-167 98,03 101,72 60,91 19,7 22,23 132,85

U. arctos CCV-003 96,02 102,8 57,9 25,05 25,03 141,25

U. arctos CCV-004 95,89

59,53 23,38 22,8 132,45

U. arctos GP-1 81,08

44,21 15,07 16,07

U. arctos CGLL-012

44,07

18,39

U. arctos CGLL-011

46,13

18,16

U. arctos SIPA-15

48,77

18,69

U. arctos SIPA-53 80,02

52,33 15,12

U. arctos SIPA-128

54,94

22,05

U. arctos PVR-013 89,79 95,87 54,77 22,65 20 129,85

U. arctos PVR-014 88,56 95,4 49,82 23,35 19,57 125,35

U. arctos SH5-97-U29-7 79,01 83,01 42,39 16,37 14,35 106,15

U. arctos SH5-97-U29-6 81,07 82,49 42,25 16,48 14,25 105,55

U. arctos SH5-98-V29-7 97,65 78,54 52,59 19,87 23,26 117,7

U. arctos SH5-98-S30-6

54,11

22,49

U. arctos SH5-97-N19-4 82,48 83,81

15,29 17,4 104,55

U. arctos SH5-98-U28-010 74,88

45,68 14,33 18,45 105,4

U. arctos TA-109 100,37 81,97 45,66 18,22 19,95 112

U. arctos TA-108

45,96

18,76

U. arctos Cráneo 001 57,23 68,66 36,25

16,16 85,91

U. arctos Cráneo 001 58,17 65,88 35,91 16,23 16,07 84,26

U. arctos Cráneo 002 67,08 60,55

14,83 14,4 84,45

U. arctos Cráneo 002 64,52 61,43 38,35 15,39 14,99 88,2

U. arctos ? 90,54 86,84 55,76 15,64 19,93 142,89

U. arctos ? 83,46 87,59 55,28 15,54 18,85 142,1

U. spelaeus 141 102,03

66,52 26,7 25,88

U. spelaeus 142 52,98 65,33

17,4 23,82 82,2

U. spelaeus 144 55,79 57,29

16,27 22,4 66,47

U. spelaeus 145 53,18 54,24

15,9

68,03

U. spelaeus 146

57,36

16,35 22,74 76,97

U. spelaeus 191

23,57

U. spelaeus 518 59,68 63,31

18,9 25,9 75,71

U. spelaeus 664

30,09

U. spelaeus 801 50,3

16,63 19,8 75,22

97

With this complete set of data a PCA was performed in the program Past (version 3.14), although the

results were not informative, they pointed towards good measurements to use in biplot graphs, which

can be seen in figures 4.41, 4.42 and 4.43.

Figure 4.41: Total height of the mandible vs length from the mandibular condyle to the third molar from ADVP

2.1 (blue square in the graph), fossil and recent brown bears from NW Spain as black dots (García Vázquez,

2015) and cave bears as green triangles (unpublished data from Cova da Ceza). Two clusters of individuals are

classified as females or males.

Figure 4.42: Total height of the mandible vs height of the mandibular condyle from ADVP 2.1 (blue square in

the graph) and fossil and recent brown bears from NW Spain as black dots (García Vázquez, 2015) and cave

bears as red inverted triangles (unpublished data from Cova da Ceza).

98

Figure 4.43: Diameter of the mandible between M2 and M3 vs height of the mandibular corpus labially at M3

from ADVP 2.1 (blue square in the graph) and fossil and recent brown bears from NW Spain as black dots

(García Vázquez, 2015) and a cave bear as a green diamond (unpublished own data from Cova da Ceza).

In the figure 4.41 we can see that the Portuguese specimen falls in the variability of the female brown

bears. In the figure 4.42 we can see that the cave bears tend to be more robust (taller) mandibular

condyles even when the total height of their mandible is smaller, ADVP 2.1 falls into the smaller

brown bears. In the figure 4.43 the measurements are centered in the mandibular corpus, the only cave

bear included in the analysis poses a sturdier mandibular corpus than any of the 24 confirmed brown

bears used.

This data suggest that the mandible belongs to a fairly average sized female brown bear.

ADVD: 2.2 Parietal and occipital

This item (Figure 4.44) comprises partial occipital and parietal bones. The foramen magnum and

cranial condyles are totally gone and the remaining part of the cranial cavity is obscured by a layer of

calcite that cannot be removed without compromising the stability of the whole bone. Interestingly,

despite that the interparietal suture is pretty much fused; the nucal crest is almost nonexistent, as well

as the external occipital protuberance and parietal crest, even taking into account that they might be

slightly damaged. Not a single measurement according to the system of Tsoukala & Grandal-

d’Anglade, (2002) could be recorded in the specimen.

99

Figure 4.44: Partial occipital and parietal bones (ADVP 2.2) from the “bear two” (Ursus arctos) of Algar do

Vale da Pena in A) dorsal; B) posterior and C) anterior views.

ADVP 2.3: Temporal bone

This specimen (Figure 4.45) comprises an almost complete temporal bone and part of a parietal one.

The zygomatic apophysis (the start of the zygomatic arch in the temporal bone) is broken and partially

covered with calcareous crust. The post-glenoid apophysis (the place where the mandibular condyle

articulates with the cranium) is broken as well; being the broken part separated and covered by

calcareous crust which prevents it to be glued back in place. The ootic channel and the petrosal part of

the temporal are lightly damaged. Both the temporal crest (between the base of the zygomatic

apophysis and the posterior part of the petrosal part of the temporal) and the lambda crest (that runs

along the posterior margin of the temporal) are well preserved. The squamosal part of the temporal is

intact and articulates with a partial parietal bone. In the part of the cranial cavity that remains we can

see the foramina of several cranial nerves, and the inner ear canal. Again, not a single measurement

according to the system of Tsoukala & Grandal-d’Anglade, (2002) could be recorded in the specimen.

Figure 4.45: Temporal and partial parietal bones (ADVP 2.3) from the “bear two” (Ursus arctos) of Algar do

Vale da Pena in A) exterior and B) interior views.

100

ADVP 2.5: Heavily damaged canine

This canine tooth is so heavily damaged that cannot be measured, since not only is eroded, but also

partially covered in calcareous matrix, that would compromise heavily the bone if removed (Figure

4.46).

Figure 4.46: Partial canine teeth (ADVP 2.5) from the “bear two” (Ursus arctos) of Algar do Vale da Pena in

lateral view.

ADVP 2.6: Right upper fourth premolar

This last premolar of the dental series has been found in a nearly pristine condition (Figure 4.47). It

has a marked metastyle near the metacone as well as a shallow cingulum near the paracone. In oclusal

view it has a strong triangular shape, compared to the Furninha teeth depicted by Torres Pérez-

Hidalgo, (1984), that are broader in the mesial part of the tooth. We can see its measurements

graphically in figure 4.48 and numerically in table 4.8, as well as the breadth and length of several of

these teeth of brown bears of NW Spain. If we look at the figure 4.49 that it is the longest tooth, but

one of the narrowest. It is difficult to assign it to a possible age or sex in Ursus arctos.

Figure 4.47: Fourth upper premolar (ADVP 2.6) from the “bear two” (Ursus arctos) of Algar do Vale da Pena

in A) oclusal, B) lingual and C) labial views.

101

Table 4.8: Measurements from the fourth upper premolar (ADVP 2.6) from “bear number two” of Algar do

Vale da Pena compared to broad an length of the same tooth of fossil and recent brown bears from NW Spain

(García Vázquez, 2015). Measurements are in mm according to the system of Tsoukala & Grandal-d’Anglade

(2002).

Specimen Species L B Hpa Hme Hde Lpa-me Lpa-de Lme-de

ADVP 2.6 ? 18 11 8 5 4 4 5 4

ARLU-38 U. arctos 16,9 13,62 10,37 8,81 5,29 7,02 7,61 7,26

ARLU-38 U. arctos 17,47 13,65 10,43 8,74 7,08 7,54 8,97 8,18

CB-010 U. arctos 17,61 14,55 12,38 8,85 8,61 5,36 7,62 6,17

CB-010 U. arctos 17,49 14,6 12,41 8,95 8,4 6,03 6,77 6,66

LCF-001-1996 U. arctos 15,5 13,31 8,27 7,24 7,72 7,48 8,07 6,75

LCF-001-1996 U. arctos 16,31 13,03 9,5 7,4 6,82 6,94 8,52 5,38

LCF-169 U. arctos 14,92 11,96 7,22 6,36 7,35 5,67 6,49 5,68

CGCH-015 U. arctos 16,7 14,37 11,83 7,37 6,6 5,54 7,85 5,73

CGCH-016 U. arctos 17,18 13,92 11,34 5,98 7 6,03 8,18 6,15

SIPA-16 U. arctos 14,8 12,22 9,29 6,15 4,93 6,14 6,88 5,81

SIPA-56 U. arctos 16,2 12,3 6,82 5,55 5,39 6,53 7,17 5,67

SIPA-57 U. arctos 16,27 12,92 7,94 4,98 5,41 6,59 7,07 6,29

SIPA-135 U. arctos 16,83 12,82 9,01 6,38 6,43 6,25 7,68 6,16

SIPA-140 U. arctos 17,57 13,39 9,49 7,52 6,04 6,1 7,46 6,04

RT-001 U. arctos 17,43 10,95 8,53 7,02 4 6,41 8,65 4,52

Pur-Lu-47 U. arctos 16,32 12,8 9,18 6,91 6,32 6,95 8,05 6,97

Pur-Lu-48 U. arctos 16,67 12,91 9,09 6,72 6,36 7,21 7,5 5,95

SAB-82 U. arctos 17,17 12,33 9,7 7,6 5,81 6,88 9,76 6,54

SH5-98-T28-013 U. arctos 15,02 9,45 10,05 6,45 6,6 4,86 7,16 3,83

SH5-97-N20-2 U. arctos 15,05 12,17 9,51 6,19 6,99 4,8 6,09 5,64

SH5-97-N20-2 U. arctos 15,33 11,81 8,87 7,59 6,9 4,41 7,82 6,51

SH5-98-U30-001 U. arctos 14,55 12,37 10 8,3 8,48 6,62 8,15 6,7

SH5-98-U30-001 U. arctos 14,34 13,12 10,57 8,86 8,22 6,23 7,38 7,42

? U. arctos 14,98 13,65 8 5,07 4,94 6,87 9,64 6,84

? U. arctos 14,1 12,59 8,14 5,35 6,19 6,58 9,22 5,89

- U. arctos 14,2 11,7

- U. arctos 14,9 11,8

MZB 82-7005 U. arctos 14,23 13,21 8,25 7,1 7,15 5,56 6,6 5,52

MZB 82-7005 U. arctos 13,15 12,64 8,54 6,47 6,94 5,74 6,61 5,03

102

Figure 4.48: Measurements for the fourth upper premolar of ursids according to the system of Tsoukala &

Grandal-d’Anglade (2002).

Figure 4.49: Breadth vs length of the fourth upper premolar ADVP 2.6 (blue square in the graph) and the same

measurements of recent and fossil brown bears from NW Spain as black dots (García Vázquez, 2015).

ADVP 2.7: Partial left upper first premolar.

As we can see in the figure 4.50, this molar is partially eroded in its lingual side, therefore only the

parastyle, paracone, metacone and metastyle are visible. The metastyle of Ursus thibetanus should

protrude more from the tooth and a continuous more marked cutting ridge should run over paracone

and metacone in a tooth of this species (Torres Pérez-Hidalgo, 1984). The space between the metacone

and the metaconule (That can be partially seen) does not have any ornamentation, as in cave bears. We

can see its measurements in figure 4.51 and table 4.9, as well as the breadth and length of several of

these teeth of brown bears of NW Spain (García Vázquez, 2015) and cave bears from Liñares cave,

Galicia, NW Spain (González & Martelli, 2003). If we look at the figure 4.52 we can see that even the

smallest cave bears have more robust teeth than the bigger brown bears. ADVP 2.7 falls into the lower

part of the variation of U. arctos.

103

Figure 4.50: First upper molar (ADVP 2.7) from the “bear two” (Ursus arctos) of Algar do Vale da Pena in A)

oclusal, B) lingual and C) labial views.

Figure 4.51: Measurements for the first upper molar according to the system of Tsoukala & Grandal-d’Anglade

(2002). Taken measurements are marked with a circle.

104

Table 4.9: Length and breadth from the first upper molar (ADVP 2.7) from “bear number two” of Algar do Vale

da Pena compared to breadth an length of the same tooth of fossil and recent brown bears from NW Spain

(García Vázquez, 2015) and cave bears from Liñares cave, Galicia, NW Spain (González & Martelli, 2003).


Specimen Species L B

ADVP 2.7 ? 21 15

273 U. spelaeus 28,41 19,85

272 U. spelaeus 26,88 19,46

273 U. spelaeus 27,96 20,31

272 U. spelaeus 27,07 20

271 U. spelaeus 26,86 19,87

1 U. spelaeus 28,16 20,06

1100 U. spelaeus 26,6 19,1

841 U. spelaeus 28,7 20,8

271 U. spelaeus 26,84 19,2

1 U. spelaeus 28,2 19,84

1100 U. spelaeus 27,2 18,3

841 U. spelaeus 28,6 20,8

ARLU-38 U. arctos 21,99 17,07

ARLU-38 U. arctos 21,67 17,23

CB-010 U. arctos 25,62 19,46

CB-010 U. arctos 25,14 19,81

LCF-001-1996 U. arctos 25,05 18,38

LCF-001-1996 U. arctos 25,07 18,18

LCF-169 U. arctos 22,54 16,48

CGCH-015 U. arctos 23,83 18,2

CGCH-016 U. arctos 23,56 18,52

Pur-Lu-49 U. arctos 22,61 17,23

RT-001 U. arctos 23,33 18,29

RT-001 U. arctos 23,09 18,12

SIPA-16 U. arctos 21,27 15,96

SIPA-56 U. arctos 23,26 16,43

SIPA-33 U. arctos 25,21 17,87

SIPA-136 U. arctos 23,85 17,71

SIPA-141 U. arctos 24,18 17,9

SH5-98-T28-013 U. arctos 20,43 15,01

SH5-97-N20-2 U. arctos 21,39 16,17

SH5-97-N20-2 U. arctos 21,68 15,94

SH5-98-U30-001 U. arctos 22,68 16,21

SH5-98-U30-001 U. arctos 22,57 16,12

SH5-97-AO34-39 U. arctos 21,19 15,08

SH5-97-AO34-36 U. arctos 20,79 14,68

TA-196 U. arctos 22,02 17,92

? U. arctos 23,11 17,25

? U. arctos 22,98 17,25

- U. arctos 21,85 15,9

- U. arctos 21,5 16,2

MZB 82-7005 U. arctos 21,74 15,14

105

Figure 4.52: Breadth vs length of the first upper premolar ADVP 2.7 (blue square in the graph) and the same

measurements of recent and fossil brown bears from NW Spain as black dots (García Vázquez, 2015) and cave

bears as orange triangles from Liñares cave, Galicia, NW Spain (González & Martelli, 2003).

ADVP 2.8: Right lower second molar

This molar (Figure 4.53) was slightly damaged in its posterior part around the entoconid, but overall it

was well preserved. In the table 4.10, we can see its measurements of breadth and length together with

those of several teeth of cave bears A Ceza cave, Galicia, NW Spain (Unpublished own data) and

brown bears of NW Spain (García Vázquez, 2015) . In figure 5.54 we have the measurements taken

and in figure 5.55 a biplot that shows that ADVP 2.8 falls into the variability of brown bears (between

the sizes of modern and fossil brown bears) and that it is well outside the range of U. spelaeus.

Figure 4.53: Second lower molar (ADVP 2.8) from the “bear two” (Ursus arctos) of Algar do Vale da Pena in

A) oclusal, B) lingual and C) labial views.

106

Table 4.10: Breadth and length of the second lower molar (ADVP 2.8) from the “bear two” of Algar do Vale da

Pena compared to breadth an length of the same tooth of fossil and recent brown bears from NW Spain (García

Vázquez, 2015) and cave bears from A Ceza cave, Galicia, NW Spain (Unpublished own data). Measurements

are in mm according to the system of Tsoukala & Grandal-d’Anglade (2002).

Specimen Species L B

ADVP 2.8 ? 22 14

del 518 U. spelaeus 32,79 18,48

del 144 U. spelaeus 31,7 18,44

del 142 U. spelaeus 31,38 17,71

del 518 U. spelaeus 32,92 19,1

del 517 U. spelaeus 32,32 18,68

del 801 U. spelaeus 30,88 18,03

LCF-003-1996 U. arctos 27,26 15,72

LCF-002-1996 U. arctos 27,89 15,79

LCF-154 U. arctos 23,44 14,71

LCF-155 U. arctos 23,43 15,22

EIX-018 U. arctos 24,59 14,9

CGLL-011 U. arctos 24,74 14,59

CGLL-004 U. arctos 25,19 15,32

SIPA-15 U. arctos 22,67 14,11

SIPA-129 U. arctos 26,03 16,42

SIPA-133 U. arctos 25,93 16,64

Pur-Lu-52 U. arctos 25,53 15,91

Pur-Lu-53 U. arctos 25,09 15,17

SH5-97-U29-6 U. arctos 23,11 14,51

SH5-98-V29-7 U. arctos 26,15 15,62

SH5-98-S30-6 U. arctos 26,77 15,95

SH5-97-N19-4 U. arctos 23,56 13,62

SH5-98-U30-

014 U. arctos 23,23 13,85

SH5-98-U30-

025 U. arctos 23,36 13,98

TA-109 U. arctos 25,28 14,09

TA-192 U. arctos 24,31 16,15

TA-195 U. arctos 24,62 16,03

VA87/16F/37 U. arctos 25,37 15,42

? U. arctos 22,7 15,2

? U. arctos 22,71 16,01

QU U. arctos 20,8 13,1

QU U. arctos 21,5 12,75

MZB 82-7005 U. arctos 21,05 12,28

MZB 82-7005 U. arctos 21,2 12,91

107

Figure 4.54: Measurements for the second lower molar according to the system of Tsoukala & Grandal-

d’Anglade (2002).

Figure 4.55: Breadth vs length in mm of the second lower molar ADVP 2.8 (blue square in the graph) and the

same measurements of recent and fossil brown bears from NW Spain as green diamonds(García Vázquez, 2015)

plus cave bears from A Ceza cave as black dots, Galicia, NW Spain (Unpublished own data).

108

ADVP 2.9: Left lower third premolar

This tooth (Figure 4.56) is the best preserved dental remain in the entire sample and has a fairly similar

morphology to the teeth depicted for U. arctos from Furninha by Torres Pérez-Hidalgo, (1984). In

figure 4.57 we have graphically its measurements and in table 4.11 the length and breadth of several of

those teeth from U. arctos and U. spelaeus, in figure 4.58 we can see that those measurements that can

tell apart the species in an almost perfect way. ADVP 2.9 falls into the variability of U. arctos. In

figure 4.58 we can see the notch in the posterior part of the tooth that is typical from most U. spelaeus

as depicted by Torres Pérez-Hidalgo, (1984).

Figure 4.56: Third lower molar (ADVP 2.9) from the “bear two” (Ursus arctos) of Algar do Vale da Pena in A)

oclusal, B) lingual and C) labial views.

Figure 4.57: Measurements for the third lower molar according to the system of Tsoukala & Grandal-d’Anglade

(2002).

109

Table 4.11: Breadth and length of the third lower molar (ADVP 2.9) from the “bear two” of Algar do Vale da

Pena compared to breadth an length of the same tooth of fossil and recent brown bears from NW Spain (García

Vázquez, 2015) and cave bears from A Ceza, Liñares and Eirós caves, Galicia, NW Spain (Grandal-d’Anglade,

1993; González & Martelli, 2003; unpublished own data). Measurements are in mm according to the system of


Specimen Species L B Specimen Species L B

ADVP 2.9 ? 18 15 995 U. spelaeus 26,2 18,9

LCF-003-1996 U.arctos 17,96 15,21 1310 U. spelaeus 24,2 19,2

LCF-002-1996 U.arctos 18,38 15,5 15 U. spelaeus 24 17

LCF-147 U.arctos 18,51 14,49 52 U. spelaeus 22,5 15

LCF-155 U.arctos 19,32 14,76 1588 U. spelaeus 24,7 19,2

EIX-018 U.arctos 19,25 14,6 1273 U. spelaeus 25,6 18,1

CGLL-006 U.arctos 20,52 15,68 526 U. spelaeus 25,5 19,4

CGLL-011 U.arctos 20,24 15,59 1580 U. spelaeus 25,4 17,5

SIPA-23 U.arctos 15,21 13,45 842 U. spelaeus 26,5 19,9

SIPA-53 U.arctos 19,59 15,68 1579 U. spelaeus 30 19,9



SH5-97-U29-7 U.arctos 19,42 14,39 843 U. spelaeus 30,3 20,6

SH5-97-U29-6 U.arctos 19,37 13,88 1589 U. spelaeus 30,6 21,1

TA-192 U.arctos 19,73 15,26 996 U. spelaeus 30,5 21,3

TA-200 U.arctos 19,52 14,68 1581 U. spelaeus 30,3 19,9

? U.arctos 16,25 13,77 1108 U. spelaeus 26,8 20

? U.arctos 16,71 13,93 1590 U. spelaeus 26,7 18,9

QU U.arctos 12,85 11,6 1169 U. spelaeus 26,5 19,6

QU U.arctos 13,2 11,4 1591 U. spelaeus 27 17,4

MZB 82-7005 U.arctos 16,99 14,35 1295 U. spelaeus 28,1 19,5

MZB 82-7005 U.arctos 17,27 14,36 841 U. spelaeus 27,4 19,3

153 U. spelaeus 25,31 17,91 19 U. spelaeus 27 20

163 U. spelaeus 23,87 17,8 270 U. spelaeus 29,96 20,4

del 146 U. spelaeus 26,4 20,3 269 U. spelaeus 30,27 20,14




489 U. spelaeus 26 18,6 3 U. spelaeus 27,12 18,64

1585 U. spelaeus 26 18 843 U. spelaeus 26,7 20,5


1102 U. spelaeus 26 20,3 848 U. spelaeus 25,4 18,7



110

Figure 4.58: Breadth vs length in mm of the third lower molar ADVP 2.9 (black dot in the graph) and the same

measurements of recent and fossil brown bears from NW Spain as blue diamonds (García Vázquez, 2015) plus

cave bears from A Ceza, Liñares and Eirós caves, Galicia, NW Spain (Grandal-d’Anglade, 1993; González &

Martelli, 2003; unpublished own data) as purple triangles.

ADVP 2.10: Partial third phalanx

This partial claw (Figure 4.59) is the only postcranial remain recovered within the “bear 2”

assemblage. It has a strongly marked base, a rectilinear profile and a relatively thin bone lamina

compared to the base; all these features are closer to the claw morphology of brown bears than to cave

bears (Figure 4.59 and figure 4.60) (Torres Pérez-Hidalgo, 1984). In figure 4.60 we can see the

measurements graphically (on a cave bear claw) and in table 4.12 we can see the height and diameter

of the base of the phalanx ADVP 2.10 compared to several of those phalanxes from U. arctos and U.

spelaeus. We can see the biplot with those measurements at figure 4.61 where despite the great

overlapping; the brown bear claws tend to have smaller heights for similar diameters, indicating

broader claws at the cave bears.

Figure 4.59: Third phalanx (ADVP 2.10) from the “bear two” (Ursus arctos) of Algar do Vale da Pena in A)

Distal; B) and D) Lateral; C) Proximal view.

111

Figure 4.60: Measurements for the third phalanx of ursids according to the system of Tsoukala & Grandal-

d’Anglade (2002). Taken measurements are marked with a circle.

Table 4.12: Height and diameter of the third phalanx (ADVP 2.10) from the “bear two” of Algar do Vale da

Pena compared to the height and diameter of the same phalanx of fossil and recent brown bears from NW Spain

(García Vázquez, 2015) and cave bears from A Ceza, Liñares and Eirós caves, Galicia, NW Spain (Grandal-

d’Anglade, 1993; González & Martelli, 2003; unpublished own data). Measurements are in mm according to the

system of Tsoukala & Grandal-d’Anglade (2002).

Specimen Species H DT Specimen Species H DT Specimen Species H DT

ADVP 2.10 U. arctos 18 11 SH5-98-S28 U. arctos 24,35 16,54 1194 U. spelaeus 29,3 15,0

CB-017 U. arctos 19,75 14,03 SH5-97-T29-30 U. arctos 19,84 13,02 1201 U. spelaeus 28,0 18,3

CB-018 U. arctos 21,22 15,24 SH5-98-U30-044 U. arctos 21,12 13,71 1329 U. spelaeus 29,0 17,5

CB-019 U. arctos 21,88 14,45 SH5-98-S28-105 U. arctos 21,69 14,12 1233 U. spelaeus 25,5 13,7

LCF-103 U. arctos 23,57 13,38 SH5-98-S28 U. arctos 26,48 16,35 1294 U. spelaeus 26,3 16,3

LCF-107 U. arctos 19,25 13,18 SH5-97-U29 U. arctos 19,74 13,01 390 U. spelaeus 22,1 12,6

SIPA-147 U. arctos 20,92 12,39 SH5-97-T29-20 U. arctos 19,01 11,87 145 U. spelaeus 22,4 13,8

SIPA-148 U. arctos 19,9 13,25 TA-Lu-c-17 U. arctos 17,87 13,05 89 U. spelaeus 22,5 14,2

SIPA-151 U. arctos 23,16 14,48 VA87/14D/74 U. arctos 26,09 16,21 188 U. spelaeus 21,5 13,0

SIPA-152 U. arctos 19,59 12,98 VA88/16E/231 U. arctos 24,23 15,39 1853 U. spelaeus 28,7 17,3

Pur-Lu-36 U. arctos 22,87 15,39 El Cuervo U. arctos 19,86 14,59 1854 U. spelaeus 26,1 16,2

Pur-Lu-37 U. arctos 24,94 16,26 El Cuervo U. arctos 23,33 14,43 1856 U. spelaeus 26,0 14,7

SH5-97-U29-24 U. arctos 21,83 12,73 El Cuervo U. arctos 22,73 15,33 1857 U. spelaeus 23,0 14,2

SH5-98-S28-079 U. arctos 22,76 14,56 El Cuervo U. arctos 19,55 14,64 1858 U. spelaeus 26,7 15,7

SH5-98-T30-003 U. arctos 22,58 13,27 1 U. spelaeus 17,1 17,92 1859 U. spelaeus 21,8 13,0

SH5-98-S28-103 U. arctos 26,3 16,54 2 U. spelaeus 14,79 15,18 1860 U. spelaeus 22,4 12,9

SH5-97-U29 U. arctos 21,37 13,3 743 U. spelaeus 13,38 14,52 1861 U. spelaeus 29,1 15,9

SH5-98-S28 U. arctos 21,15 15,31 651 U. spelaeus 23,0 14,5 1862 U. spelaeus 26,4 14,8


SH5-97-T29-16 U. arctos 21 12,64 805 U. spelaeus 25,3 14,2 1865 U. spelaeus 25,7 14,0

SH5-97-T29 U. arctos 22,19 13,17 806 U. spelaeus 25,9 14,7 1867 U. spelaeus 24,3 14,7


SH5-97-T29-19 U. arctos 22,3 13,18 901 U. spelaeus 26,3 14,7 1870 U. spelaeus 29,9 16,7

SH5-98-U28-012 U. arctos 19,95 13,92 1022 U. spelaeus 22,5 14,9 262 U. spelaeus 24,0 14,6

SH5-98-S28-045 U. arctos 22,8 15,02 1106 U. spelaeus 26,5 14,2 953 U. spelaeus 22,5 13,5

SH5-98-U30-009 U. arctos 23,75 14,09 1152 U. spelaeus 24,3 15,2 952 U. spelaeus 31,5 17,0

112

Figure 4.61: Height and diameter of the third phalanx (ADVP 2.10) (black dot in the graph) and the same

measurements of recent and fossil brown bears from NW Spain as blue squares (García Vázquez, 2015) plus

cave bears from A Ceza, Liñares and Eirós caves, Galicia, NW Spain (Grandal-d’Anglade, 1993; González &

Martelli, 2003; unpublished own data) as crimson diamonds. 95% ellipses are drawn.

ADVP 2.10 is precisely in the limit for the ellipse for brown bears and is the smallest of all claws

shown, possibly attributable to a juvenile.

ADVP 3.1: Partial distal epiphysis of right femur

The assemblage from whom this item comes (“bear 3”) is the worst preserved of all. Only one

measurement according to the system of Tsoukala & Grandal-d’Anglade (2002) could be recorded.

Only the most distal epiphysis is preserved as we see in figure 4.62. We can see the recorded

measurement as well as a collection of the same measurements from other fossil and recent brown

bears in table 4.13. They are presented graphically in figure 4.63, where we can see that it is difficult

to classify ADVP 3.1 as belonging to a male or female, being it a nearly average individual between

the biggest and the smallest in the sample.

Figure 4.62: Distal epiphysis of femur (ADVP 3.1) from the “bear three” of Algar do Vale da Pena in A) distal

B) lateral views.

113

Table 4.13: Anteroposterior diameter of the distal epiphysis of the femur (ADVP 3.1) from the “bear three” of

Algar do Vale da Pena compared to the same measurement of fossil and recent brown bears from NW Spain

(García Vázquez, 2015). Measurements are in mm according to the system of Tsoukala & Grandal-d’Anglade

(2002).

Specimen DAPdist.

ADVP 3.1 75

CB-008 80,55

LCF-006-1996 81,59

LCF-007-1996 82,74

LCF-173 69,97

LCF-174 72,15

EIX-010 65,36

CGLL-047 86,77

SIPA-2 99,07

SIPA-106 98,39

Pur-Lu-21 76,93

Pur-Lu-22 82,65

SH5-97-T28-12 106,3

SH5-98-T30-008 63,66

SH5-97-T29-58 62,18

SH5-98-S28-096 73,6

TA-Lu-c-7 69,57

TA-Lu-c-8 71,41

59,16

62,81

El Cuervo 62,22

El Cuervo 61,59

Figure 4.63: Anteroposterior diameter of the distal epiphysis of the femur (ADVP 3.1) and the same

measurement of recent and fossil brown bears from NW Spain (García Vázquez, 2015).

75

80,55 81,59 82,74

69,97 72,15

65,36

86,77

99,07 98,39

76,93

82,65

106,3

63,66 62,18

73,6 69,57

71,41

59,16 62,81 62,22 61,59

58

68

78

88

98

108

AD

VP

3.1

CB

-00

8

LCF-

00

6-1

99

6

LCF-

00

7-1

99

6

LCF-

17

3

LCF-

17

4

EIX

-01

0

CG

LL-0

47

SIP

A-2

SIP

A-1

06

Pu

r-Lu

-21

Pu

r-Lu

-22

SH5

-97

-T2

8-1

2

SH5

-98

-T3

0-0

08

SH5

-97

-T2

9-5

8

SH5

-98

-S2

8-0

96

TA-L

u-c

-7

TA-L

u-c

-8

El C

uer

vo

El C

uer

vo

DAPdist.

114

ADVP 3.2: Vertebral centrum

This vertebral centrum (Figure 4.64) is partially covered by calcareous crust, which prevented it to be

heavily eroded like the other centrum recovered (ADVP 3.3). The neural arch is nearly gone and one

the cranial/caudal sides of the vertebra is highly damaged. The better preserved part of the vertebra is

heart-shaped centrum.

Figure 4.64: Vertebral centrum (ADVP 3.2) from the “bear three” of Algar do Vale da Pena in A) dorsal and B)

anterior views.

ADVP 4.1: Proximal epiphysis of second right metatarsal

This metatarsal fragment (Figure 4.65) is slightly worn in its proximal part; we have its measurements

graphically in figure 4.66 and numerically in table 4.14. The erosion apparently caused a similar

phenomenon tan the one observed in ADVP 1.7; the metatarsal is smaller in size than the ones of fossil

and recent brown bears from NW Spain (Figure 4.67), almost like a juvenile individual.

Figure 4.65: Proximal epiphysis of second right metatarsal (ADVP 4.1) from the “bear four” of Algar do Vale

da Pena in A) Interior; B) Ventral; C) Exterior and D) Dorsal.

115

Figure 4.66: Measurements for the second metatarsal according to the system of Tsoukala & Grandal-

d’Anglade (2002).Taken measurements are marked with a circle.

Table 4.14: Measurements of the second metatarsal (ADVP 4.1) from the “bear four” of Algar do Vale da Pena

compared to the same measurement of fossil and recent brown bears from NW Spain (García Vázquez, 2015).


L DTprox. DAPprox. DTdia.min. DAPdia. DTdist. DAPdist. DTart.dist.

ADVP 4.1

9 20 9 8

SIPA-92

16,15 22,73 13,5 12,3

SIPA-95

16,38

13,25 12,18

SH5-98-97-S29-009 76,25 13,53 22,78 12,92 9,04 18,19 15,02 14,27

SH5-97-T29-7 71,99 11,31 20,1 12,3 8,14 17,11 14,04 14,46

SH5-98-S28-044 83,73 16,7 27,17 13,91 11,3 21,08 18,5 16,88

SH5-97-T29-088 71,84 13,32 20,64 12,13 8,28 17,51 14,31 14,12

SH5-98-S28-006 75,59 15,17 22,39 12,88 8,82 18,52 14,81 14,1

TA-Lu-c-23 69,28 13,06 21,63 11,01 7,75 18,07 15,25 10,95

TA-Lu-c-24 69,74 12,68 21,98 11,16 8,05 18,38 14,96 11,52

116

Figure 4.67: Anteroposterior vs transversal diameter of the proximal epiphysis (upper) and diaphysis (bottom)

of the second metatarsal, ADVP 4.1 (blue square) and the same measurements of recent and fossil brown bears

from NW Spain as black dots (García Vázquez, 2015).

ADVP 4.2: Damaged proximal epiphysis of fifth right metatarsal

This proximal part of the metatarsal (Figure 4.68) was reconstructed from several fragments and could

not be entirely repaired; we have its measurements graphically in figure 4.69 and numerically in table

4.15. In figure 4.70 we can see a graphical comparison with brown bears from NW Spain,

demonstrating a similar situation as with the other metapodial bones (ADVP 1.7 and ADVP 4.1), the

Portuguese specimen is by far the less robust.

Figure 4.68: Proximal epiphysis of fifth right metatarsal (ADVP 4.2) from the “bear four” of Algar do Vale da

Pena in A) Interior; B) Ventral; C) Exterior and D) Dorsal views.

117

Figure 4.69: Measurements for the fifth metatarsal according to the system of Tsoukala & Grandal-d’Anglade

(2002). Taken measurements are marked with a circle.

Table 4.15: Measurements of the fifth metatarsal (ADVP 4.2) from the “bear four” of Algar do Vale da Pena

compared to the same measurement of fossil and recent brown bears from NW Spain (García Vázquez, 2015).


DAPprox. DTdia.min.

ADVP 4.2 21 10

LCF-136 24,29 11,42

SH5-97-T29-36 24,17 11,57

SH5-97-T29-087 24,36 11,82

SH5-98-S28-076 31,56 14,51

TA-Lu-c-20 24,58 11,18

TA-Lu-c-19 25,74 11,09

VA87/LIMPIEZAEXTERIOR/4 26,43 12,75

24,79 11,37

25,85 11,23

Figure 4.70: Anteroposterior proximal diameter vs transversal diameter of diaphysis of the fifth metatarsal,

ADVP 4.2 (blue square) and the same measurements of recent and fossil brown bears from NW Spain as black

dots (García Vázquez, 2015).

118

4.6 The claw marks

In several walls of the cave, where there is an accumulation of clays, we can find several parallel

markings that appear mostly in groups of four or five grooves. Octávio Mateus has counted 189 of

those groups alongside the walls of the cave. They are interpreted as claw marks left behind by bears

that were navigating its way across the darkness of the cave or digging beds for hibernation, the

location of the largest concentrations of them can be seen as the numbers 2, 4, 5 and 7 at the cave map

of figure 4.4. In figure 4.71 we can see some of the best examples. They vary greatly in size, being

mostly between five to ten centimeters in width. Without the calcareous flowstones that cover many

clay areas of the cave the number of tracks would probably be in the hundreds. Given that U. arctos

tend to not let behind nearly as many of this marks as U. spelaeus (Fosse et al., 2004); this numbers

point towards the use of the cave repetitively during a long period of time. These are the first of this

type of marks described in Portugal (Neto de Carvalho, 2018).

Figure 4.71: Claw marks in the walls of the Algar do Vale da Pena. Some of them are today partially covered by

calcareous crusts. Scale bar 7 cm.

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4.7 Discussion

The cave was sealed until recently, so we do not know its original entrance (or entrances), but the

claw marks in the walls point towards that the bears entered the cave by their own means (probably for

hibernation) and died of natural causes inside, so the cave did not acted like a natural trap. The bones

of most assemblages also point towards this hypothesis, since they do not show signs of important

transport and are still articulated or closely associated; with the exception of “bear 2” which shows

clay below the calcareous crust and eroded bones and teeth, and “bear 3” whose bones are in the worst

condition of all groups of remains. Those exceptions could be explained by water flowing and eroding

in situ the bones or lightly transporting them. The number of tracks point to the use of the cave as den

by bears, maybe during generations.

Talking about the characterization of the bear population, first we have to address that the minimal

number of individual is two, in base of the recovered cranial parts and the articulated skull that is still

in the cave. But as pointed earlier it is likely that the real number is between four and five, in base of

the separation between assemblages, being it of several dozens of meters and in the absence of

important transportation.

Now we will discuss each studied assemblage and bone in the sections 4.4 and 4.5:

“Bear” one: Does not present signs of transportation, with some articulated parts. It can be assumed to

be a single animal.

- ADVP 1.1: Partial left scapula:

Not measured. Its morphology is too distorted by conservation.

- ADVP 1.3: Right scapholunate bone:

Morphology ruling out U. thibetanus, measurements are not congruent with U. spelaeus; the

most likely affinity is to an average female U. arctos.

- ADVP 1.4: Right hamate bone:

Only measurement is congruent with the hamate of a female U. arctos.

- ADVP 1.6: Second phalanx.

It is likely a bone of hind limb. Not useful for specific or sex determination.

- ADVP 1.7: Proximal epiphysis of third right metatarsal.

Smallest of all the considered bones, juvenile sized U. arctos.

- ADVP 1.8: Distal epiphysis of right radius.

Slender morphology is ruling out U. spelaeus. The epiphysis is solidly fused, so it belongs to a

mature individual. It is also less robust than the other Portuguese specimens; measurements

are congruent with a small female U. arctos.

- ADVP 1.9: Distal part of diaphysis of left ulna.

Slender morphology is ruling out U. spelaeus. Only recovered measurement is the smallest of

the sample, congruent with a small or young female U. arctos.

Putting together all this lines of evidence we can attribute this assemblage to a small, mature female of

U. arctos.

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“Bear” two: Presents some signs of erosion and could have been subject to some minor transport. It

could include the remains of more than one specimen (all are cranial elements except one third

phalanx).

- ADVP 2.1: Posterior part of right hemimandible.

The width of the mandibular ramus relative to its height is typical for a female U. arctos. The

height of the mandibular condyle relative to height of the mandibular ramus is also typical of a

below average U. arctos and different to the taller condyles in similarly sized ramus of U.

spelaeus. The robustness of the mandibular corpus is above the average, but well below the

values of U. spelaeus. Therefore this mandible could be attributed to a female U. arctos.

- ADVP 2.2: Parietal and occipital.

The piece is not very diagnostic at specific level, but the grade of fusion of the sutures points

towards an adult specimen. Although this, the nucal and parietal crests are not developed

which points rule out a big mature male.

- ADVP 2.3: Temporal bone.

Not diagnostic, although the calcareous crust overlaying the broken parts of the bone is one of

the best examples of the taphonomical processes in the cave.

- ADVP 2.5: Heavily damaged canine.

This piece is too fragile and damaged to be measured.

- ADVP 2.6: Left upper fourth premolar.

Long, yet slender, compared to the sample this specimen is so particular that it is difficult to

assign to a certain species.

- ADVP 2.7: Left upper first molar.

Its morphology differs from U. thibetanus and U. spelaeus; therefore it can be classified

confidently to U. arctos. The measurements point towards a small U. arctos.

- ADVP 2.8: Right lower second molar.

Its dimensions are far from the ones of U. spelaeus, they are in the lower spectrum from U.

arctos.

- ADVP 2.9: Left lower third molar.

This piece has the typical dimensions of an average U. arctos, far from what we can expect

from a cave bear.

- ADVP 2.10: Partial third phalanx.

Its morphology and measurements are totally coincident with the claw of a small sized brown

bear

This assemblage could belong to more than one specimen yet, it will be tentatively attributed to a

mature and small to average female brown bear, similar to what we see in the first assemblage.

“Bear” three: Worst preserved assemblage of all, it could belong to weathered and moved remains of

“bear” one.

- ADVP 3.1: Partial distal epiphysis of right femur.

Only one measurement could be recorded, being it almost perfectly into the average for a

brown bear.

- ADVP 3.2: Vertebral centrum.

It is partially covered by calcareous crust not possible to classify.

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“Bear” four: Most of the remains of this assemblage are closely associated and covered by thick

layers of calcareous crust. It is unlikely that they suffered meaningful transportation.

- ADVP 4.1: Proximal epiphysis of second right metatarsal.

This piece is noticeably smaller than any brown bear specimen sampled in NW Spain for any

measurement considered.

- ADVP 4.2: Damaged proximal epiphysis of fifth right metatarsal.

This piece is noticeably smaller than any brown bear specimen sampled in NW Spain for any

measurement considered.

Both measured metatarsals of this assemblage are surprisingly small. The humerus, femur and hip

bones that comprise the rest of it do not look proportionally smaller to other bears, but given that they

have not been recovered from the cave, neither measured nor studied, this is only an appreciation. This

assemblage cannot be attributed to any species with certainty according our current data.

“Bear” five: Is still encased in layers of calcareous crust and could not be studied for this work.

Therefore the fossil bear population in the cave is formed by brown bears of smaller size that the ones

recorded in Northwestern Spain (García Vázquez, 2015) and most likely dominated by adult female

animals.

It is curious to compare these facts with the works of Torres Pérez-Hidalgo, (1979) and Cardoso,

(1993) that concluded that the Portuguese brown bears (mainly from Fontainhas, Furninha and some

of Serra de Molianos) were more robust that the Spanish bears of comparable age.

Given the absence of a reliable stratigraphy for the cave, the age of the fossil remains could only be

addressed thanks to absolute methods like isotopic or ESR (Electron Spin Resonance) datings.

4.8 Conclusions

- The Algar do Vale da Pena is a cave recently reopened and is a new fossil brown bear locality

in Portuguese Estremadura.

- 37 items have been recovered from the cave, including 20 informative fossil remains.

- The fossil remains probably belong to five individuals considering its distribution in the cave,

althought the minimal number of individuals in the cave is two, in base of the cranial remains.

- The findings can be ascribed to Ursus arctos in base of morphology (A first upper molar with

clearly individualized metacone-paracone and protocone-hypocone, but without a marked and

big cingulum; mandible with a low condyle compared with the mandibular ramus, slender

long bones without marked muscle insertions, occipital bone with a shallow nucal crest and

straight third phalanxes with robust proximal parts) and metrical comparisons (In several

measurements of 20 bones from the assemblage).

- Two of them are congruent with the measurements and characteristics of mature, yet below

the average sized, female bears.

- The population is formed by relatively small animals (compared to their Spanish counterparts)

in contrast with the observed in other Portuguese localities by Torres Pérez-Hidalgo, (1979)

and Cardoso, (1993).

- The claw marks in the walls of the cave are the first of its type documented in Portugal.

- The great numbers of the claw marks, the morphology of the cave and the taphonomy of the

remains point toward repetitive occupation of the cave by bears during a long time period.

- The presence of Ursus arctos and the thickness of the calcareous mantles over the bones point

towards a Pleistocene age, although the absence of informative stratigraphy prevents the

precise dating of the assemblage for the moment.

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4.9 Further work

Neither the works in the cave, nor the study of the fossil findings is over.

As presented in the section 4.4, there are two in situ jackets containing remains of the “bears” two and

three, and two separated blocks including material of “bear” three and the main skull of “bear” one

already in the cave. This material as well as some other samples is scheduled to be removed from the

cave with the help of the GPS between May and November of 2019.

In December of 2018 samples of two rib fragments (ADVP 1.10 and ADVP 3.4) belonging to the

assemblage were analyzed in the SAI (Servizos de Apoio a Investigación) of the University of A

Coruña by mass spectrometry in order to determine the amount of Nitrogen present in them. If the

amount of this element was above 0,05% it could be assumed that the samples had enough collagen or

other proteins in them to be used in a 14C radiometric analysis; but as we can see in table 4.16, it was

not the case.

Table 4.16: Nitrogen and Carbon percentages in the samples of ADVP 1.10 and ADVP 3.4 from Algar do Vale

da Pena (Portugal).

Sample N.º SAI

Weight

(mg) % N % C

AVDP 3.4 2018/45957 4,965 < 0,05 1,85

AVDP 1.10 2018/45958 5,096 < 0,05 3,24

Therefore the petrosal part of the temporal bone of ADVP 2.3 could be used next in order to prove if

some proteins survived, or another absolute dating technique (ESR) could be used in other to date the

fossils, although this second path of action would negate the possibility of isotopic diet studies or

ancient DNA analysis.

Also, once all the material of the cave is recovered, a larger and more complete study of the

Portuguese Ursus material (including at least the specimens of Fontainhas, Furninha and Serra de

Molianos) with measurements taken according the system of Tsoukala & Grandal-d’Anglade (2002),

would be useful for a better understanding of the fossil bear population from the Algar do Vale da

Pena in its immediate geographical context.

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5. SANTA MARGARIDA: A NEW MICROVERTEBRATE LOCALITY

FROM ALGARVE

5.1 Overview

On this chapter a new and abundant microvertebrate assemblage from Algarve will be described for

first time. Special emphasis will be given to the discovery and preparation methods, taking into

account the peculiar mode of preservation of the remains (Bone breccias). Then a taxonomic

identification will be provided for some of the specimens, alongside a list of groups present. Inferences

will be made about the age and origin of the association. Finally some conclusions will be made as

well as some remarks about further work that could be done in the locality.

5.2 Introduction: Location, discovery, geological settings and nature of the

site

The site of Santa Margarida is located near the village of the same name in the freguesia of Alte,

municipality of Loulé, Algarve region, South Portugal, 300 m above the sea level (Figure 5.1).

Figure 5.1: Location of the site of Santa Margarida. Figure by Cátia Ribeiro. Credit of the maps, google maps.

The discovery of the site happened following this series of events: During a conference in Algarve in

2016 Octávio Mateus was approached by a local photographer, Jorge Graça, who informed him of the

presence of multitude of bones in the blocks of a wall near Santa Margarida. Despite the efforts for

finding the site that year it was not possible. In 2017 the team went back again to search for the wall

and this time they were able to find it, photograph several blocks and take other back to the

Department of Earth Sciences at FCT-UNL (Figure 5.2) that where prepared during the next months to

extract the microfossils. In May of 2018 a team of the FCT went back again to the place and

discovered that the source of the blocks was just adjacent to the wall. The cemented terra rossa blocks

with breccified bones and dental remains were found only in a span of 20 meters of the wall, with the

point of maximum concentration being found in 2017, plus other blocks with bones scattered inside

the area limited by it. More blocks were collected.

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Figure 5.2: Maximum concentration point of fossil bearing blocks in the wall of the site as it was discovered in

2017. Note the difference of color between the bone bearing breccified blocks of cemented terra rossa and the

grey tone of the calcareous rocks. Photo by Octávio Mateus.

The locality of Santa Margarida is situated in the karst of the Picavessa Formation (Figure 5.3), Early

Jurassic in age and described as formed by calciclastic and micritic rocks, with rare arrecifal and

oolitic limestones (Terrinha et al., 2006). As the successive expeditions to the area have shown, the

Quaternary microvertebrate fossils are conserved in heavily breccified deposits of cemented terra

rossa that is covered by a crystalline crust of calcareous material and a thin clayish soil (Figure 5.4).

Both layers and the soil are overlaying the limestone rocks of the Picavessa Formation and vary

greatly in thickness and quantity of fossils across the area, to fully understand its fine stratigraphy a

much bigger excavation should be done.

Figure 5.3: Location of Santa Margarida site. In blue the calcareous Picavessa Formation. Modified by Cátia

Ribeiro from the geologic map of Algarve by LNEG.

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Figure 5.4: Left; comparison of freshly cut cemented terra rossa block and the underlying grayish calcareous

rock of the Picavessa formation. Right; calcareous crust covering the “terra rossa”.

The wall where the fossils were found is a traditional algarvian wall. These walls were made with in

situ materials (usually limestone) and depending of their orientation and the topography they have

been traditionally used for a) forming terraces in steppe terrain, in order to reduce the soil erosion and

retain the soil water for cultivation and b) To separate proprieties. The walls of the first type can reach

three meters in height. They are perpendicular to the pending of the slope in order to retain the soil

(Gonçalves et al., 2018). The wall where the discovery was made runs parallel to the pending of the

slope, not perpendicular, making it a representative of the second type. These “property walls” are

usually between 0,6 to 1,5 m wide and 0,6 to 1,2 m in height (Gonçalves et al., 2018)., which is

congruent with the observed. In the terrain delimited by it there are some deteriorated and old fruit

trees that are indicative of an abandoned traditional cultivation field in Algarve: Fruit trees, and

between them legumes planted to fix Nitrogen in the soil (Gonçalves et al., 2018).

The abundance of small vertebrates in the blocks was clear. For every square centimeter of exposed

surface, the richest blocks could have several fossils on it. Also, some blocks with larger bones of

centimeters of diameter could be recovered (Figure 5.5).

Figure 5.5: A) Abundant microfossils in rich block; B) Rabbit humerus in situ; C) and D) larger long bones

protruding from block.

126

Using the Dino-lite™ (A device designed for taking pictures of small fossils) it was possible to take

some pictures to the microfossils in situ, before any preparation (Figure 5.6) which gave a first idea of

the taxonomic groups that could be identified. Those groups included rodents, “insectivores”, lizards

and even gastropods (not shown in the picture).

Figure 5.6: Upper left, possible leporid phalanx in dorsolateral view; upper right, arvicoline mandible in

lingual view; bottom left, shrew mandible in labial view; bottom right, lizard mandible in labial view.

5.3 Preparation methods

In spite of the good condition in which several microfossils could be observed alongside the surface of

the blocks it was necessary to extract them from the matrix to properly study them. First, the

mechanical preparation was experimented in order to recover as many of the largest remains as

possible. They were consolidated using PARALOID B72™ and extracted with the help of a air-scribe

(Figure 5.7). Some arvicoline and squamate mandibles were extracted using this method, but it was

not useful for the recovery of the small fossils found in the blocks (Most of them).

Figure 5.7: Left, consolidated arvicoline mandible with PARALOID B72™; center, extraction of the mandible

using a air-scribe and right the extracted mandible.

127

Therefore, the chemical preparation was started. The first step was to choose the chemical product to

be used. A test using five products was prepared, including: water (as control), acetic acid at 5% of

concentration in water, formic acid at 5% of concentration in water, Hydrogen peroxide in a

commercial concentration of 30% and a commercial anti-calcareous product (Water softener) at 50%

in water. Samples of cemented terra rossa were weighed before and after 24 hours in the respective

products (therefore discounting the dissolved and disaggregated rock) and two days of drying. The

results can be seen in table 5.1.

Table 5.1: Results of the tests for determining the product to be use in the chemical preparation of Santa

Margarida samples.

Test Initial weight (g) Final weight (g) % of weight lost

water (control) 103,102 104,504 + 1%

acetic acid at 5% 51,228 50,036 2%

formic acid at 5% 38,846 34,872 10%

Hydrogen peroxide at 30% 48,793 45,864 6%

Commercial anti-calcareous product at

50% 39,231 24,343 38%

After this test the water softener was chosen. A second minor qualitative test was conducted to proof

the best times for the product to be used without affecting too much the fossils, but maximizing the

amount of dissolved and disaggregated rock. Samples were submerged in the product for 12, 24 and

48 hours. The best results (less fragile and dissolved bones observed, but with good amount of matrix

dissolution) were the ones of the 24 hours test.

Therefore the processing of the maximum amount of sediment in the span of three months

(September-December of 2017) was carried in the Museum of Lourinhã using the facilities of the lab.

In the figure 5.9 we have a workflow of how the process was running. At every given time two

kilograms of rocks were being processed, one of them in the product and other one being de-acidified

in water, to prevent that product trapped in the pores of the rock and bones kept reacting all the time.

After the entire preparation process and the picking of the fossils, they were classified in broad groups

(postcranial bones, “insectivores”, lizards, gastropods… etc) and then the most promising of them

were further cleaned by ultrasound. Not all of them were subject of this preparation, because some

were already too fragile. Finally the teeth were mounted in plastic boxes over thin strands of Blu-

Tack™ in oclusal view and then photographed with the binocular lenses in the department of Earth

Sciences of the FCT-UNL (Figure 5.8) for further identification. This was a complex task given the

fragmentation of the sample.

Figure 5.8: Right, rodent teeth mounted in the plastic box and other fossils stored in similar settings (Scale

7cm); left, the binocular lenses used to take the pictures (Photo by Alexadre Guillaume).

128

Figure 5.9: Workflow followed in the preparation of the Santa Margarida material (Portugal) in the Museum of

Lourinhã.

129

5. 4 Results

A total of around 0,5 kg of sediment was disaggregated from the blocks subject to treatment and

picked for fossils. We can see the amount of recovered sediment in figure 5.10.

Figure 5.10: Amount of processed sediment from Santa Margarida (Portugal) at Museum of Lourinhã.

On this section a taxonomic identification of some of the microvertebrate and gastropod remains will

be given, as well as the number of dental remains recovered for broader groups that we have in table

5.2. It is noteworthy that 90% of all remains belong to rodents.

Table 5.2: Number of specimens from broad groups recovered from the processed sediment of Santa Margarida.

Gastropoda Cuvier, 1797 1 complete shell, 3 smaller fragments, 1 bisected shell in situ.

Lacertidae Gray, 1825 1 semi-complete mandible, 3 mandibular fragments

Chiropera Blumenbach, 1779 4 unicusp teeth, 1 molar

Soricidae Fischer, 1817 20 isolated teeth or mandible parts with attached teeth

Rodentia Bowdich, 1821 273 tooth fragments, isolated teeth, and mandibles with teeth

Lagomorpha Brandt, 1855 1 distal humerus.

In spite of this abundance of material (~0,6 fossils for gram of sediment) most of the remains became

impossible or too difficult to identify at generic or specific level, so only the better preserved ones

were subject to a more precise identification presented here.

130

Gastropoda Cuvier, 1797

In spite of the aggressive treatment with water softener, one nearly pristine snail shell was recovered

during the picking (Figure 5.11), as well as other three fragments of similar sized shells. This shell was

identified as belonging to the snail Paralaoma servilis (Shuttleworth, 1852) in base of the marked

umbilicus, the presence of ribs and the overall geometry of the shell (Callapez, personal

communication, 2019).

Figure 5.11: Paralaoma servilis from Santa Margarida in A) lateral, B) ventral and C) dorsal views.

Another gastropod was found and photographed in situ in one of the blocks in the field (Figure 5.12).

This was identified by Pedro Callapez in base of the whorl morphology as Pomatia elegans (Müller,

1774).

Figure 5.12: Bisected shell of Pomatia elegans from Santa Margarida.

131

Lacertidae Gray, 1825

A relative big lizard mandible was separated mechanically from one of the blocks (Figure 5.13). It

bears the classic lacertid features: pleurodont condition of the teeth, which are cylindrical and

monocuspid; plus a Meckelian groove opened on its whole length (Blain et al., 2007). The fragility of

the mandible prevented the usage of ultrasounds for cleaning it; therefore it was not possible to take

reliable measurements of the piece that could have allowed further identification. Another three

mandibular fragments that bear similar characteristics were picked from the disaggregated sediments.

In figure 5.14 we can see the most complete of those fragments.

Figure 5.13: Lacertid mandible from Santa Margarida (Portugal) in lingual view.

Figure 5.14: Lacertid mandible from Santa Margarida (Portugal) in A) lingual and B) oclusal views.

The teeth of the mandible were measured with the help of the software ImageJ, following the method

presented by Blain et al., 2007. The obtained measurements and ratios are presented in the table 5.3

Table 5.3: Measurements and ratios of the most complete lacertid mandible fragment picked from the

disaggregated sediments of Santa Margarida, Portugal, following the methods of Blain et al., 2007: h, is height

of the teeth; d, its diameter and a, the height of the teeth that protrudes over the dental crest. All measurements

are in mm.

Teeth h d a d/h a/h

1 0,638 0,347 0,266 0,544 0,417

2 0,729 0,323 0,287 0,443 0,394

3 0,756 0,26 0,301 0,344 0,398

4 0,860 0,301 0,364 0,350 0,423

5 0,945 0,364 0,371 0,385 0,393

6 1,037 0,357 0,364 0,344 0,351

Average 0,828 0,325 0,326 0,393 0,393

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Comparing these ratios with the ones presented by Blain et al., 2007 for the genera Podarcis Wagler,

1830 (d/h=0,269; a/h= 0,439); Acanthodactylus Daudin, 1802 (d/h=0,280; a/h= 0,330) and

Psammodromus Hallowell, 1852 (d/h=0,301; a/h= 0,327) it is possible to conclude that the Santa

Margarida lacertid (d/h=0,393; a/h= 0,393) does not belong to either of those genera.

Chiroptera Blumenbach, 1779

Five unicusp teeth and one molar assigned to bats have been recovered in the disaggregated sediments

of Santa Margarida. The unicusp teeth are ascribed to Chiroptera in base of the marked cingulum

(Hillson, 2005) (Figure 5.15).

Figure 5.15: Unicusp teeth ascribed to a bat from Santa Margarida (Portugal) in meso-lingual view (Image by

Cátia Ribeiro).

The recovered molar (Figure 5.16) bears some typical characters of the upper molars of bats, like

being dilambdadont, with tall “W” -shaped ridges and a low lingual shelf (Hillson, 2005). The

parastyle is curved, which is typical for European bats and the parastyle/paracone length is shorter

than the metacone/metastyle length. All the main cusps are quite robust. There is a thick lingual/distal

cingulum. All these are congruent with a right M2; second upper molars are generally more elongated

than first upper molars, which are more compressed mesio-distally (Piskoulis, personal

communication, 2019). The morphology is similar to the M2 some species of Myotis Kaup, 1829 with

a strong and marked postprotocrista (Gunnell et al., 2011).

Figure 5.16: Right upper second bat molar from Santa Margarida (Portugal) in oclusal view. The rectangle

marks the w shaped labial ridge and the circle the lingually recurved hypocone shelf and the strong

postprotocrista. Metastyle/paracone (1,3 mm) and metastyle/lingual (2 mm) lengths are marked. Drawing by

Pavlos Piskoulis.

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Soricidae Fischer, 1817

A total of 20 mandibular and dental remains are ascribed to this group. In the table 5.4 we can see the

different groups in which they are classified.

Table 5.4: Classification of the different Soricidae remains recovered from Santa Margarida (Portugal).

Soricinae indet. 3 tooth fragments and 1 lower molar

Sorex sp. 1 right upper incisive

Crocidura sp. 5 mandible parts with attached teeth, 5 lower molars, 1 upper molar, 4 upper incisors

In the figure 5.15 we can see three upper incisors. The first one (Figure 5.17 A) can be attributed to

Sorex Linnaeus, 1758, in base of the fissident (Having a small medial cupule) upper incisor with red

pigmented enamel, plus the size of the specimen (Reumer, 1984). The second specimen (Figure 5.17

B) can be assigned to the genus Crocidura Wagler, 1832 in base of the size, the unpigmented enamel

and the marked and slightly undulated cingulum in the posterior bucal part (Reumer, 1984). The third

specimen (Figure 5.17 C) bears similar characteristics than the previous one, but with a less robust

morphology. According to Raquel Moyá-Costa (Personal communication, 2019) it probably belongs

to an older individual that could also be representative of a different species of the genus.

Figure 5.17: Soricidae upper incisors recovered from Santa Margarida (Portugal) in labial view. A) Sorex, B)

and C) Crocidura.

In figure 5.18 we can see an upper molar that broke during the process of taking the pictures. Yet, it is

possible to distinguish some features that allow it to be classified, like a hypocone situated in a

postero-lingual position relative to the protocone and poorly individidualized of it. There is also a deep

valley between the protocone and the metacone and a crest connecting it with the paracone. It´s

trapezoidal shape is congruent with a second upper molar. Given the absence of enamel coloration, it´s

size and the previously commented characters it is assigned to the genus Crocidura (Reumer, 1984).

Figure 5.18: Broken second upper molar of Crocidura recovered from Santa Margarida (Portugal) in oclusal

view. Labial part to the right.

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Some teeth recovered from the locality were still attached to parts of the mandible, but only one

mandible part conserved with more than one tooth (Figure 5.19). It presents the P4, M1 and M2

(Moyá-Costa, Personal communication, 2019). The P4 is tetrahedral shaped; the M1 and M2 have the

valley openings well above the cingulum in labial view. The cingulum is well marked and undulated in

the labial part and weak in the lingual. The mandible also poses a small mandibular foramen situated

in labial view in the transition of the P4 to the M1. All this characteristics allow us to attribute this

mandible fragment to the genus Crocidura (Reumer, 1984).

Figure 5.19: Fragment of Crocidura mandible (bearing P4, M1 and M2) recovered from Santa Margarida

(Portugal) in A) oclusal, B) lingual and C) labial views.

Rodentia Bowdich, 1821

The rodents comprise the immense majority of the fossil remains recovered in the locality of Santa

Margarida (+90%). In the next table (Table 5.5) we have the breakage of the rodent dental remains

from Santa Margarida.

Table 5.5: Classification of the different rodent dental remains recovered from Santa Margarida (Portugal).

Rodentia indet. 152 incisors or parts of incisors, including 41 incisor tips

Allocricetus

bursae 1 upper left first molar

Eliomys

quercinus 1 lower premolar, 1 second lower molar, 2 second upper molars, 1 indeterminate molar

Apodemus cf.

sylvaticus

6 third lower molars, 2 second lower molars, 15 first lower molars, 5 third upper molars, 9

second upper molars, 5 first upper molars, 8 indeterminate teeth

Arvicolinae

indet. 46 teeth and 22 fragments

Victoriamys

chalinei 1 right first lower molar

Iberomys

huescarensis 1 Left hemimandible with first and second molar

Iberomys

brecciensis 1 Left hemimandible with first molar and 1 right first lower molar

Iberomys

cabrerae 1 left first lower molar

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The rodent chisel-like incisors are highly characteristic, only lagomorphs are similar in European

Quaternary faunas and its size mostly prevents confusion. In Figure 5.20 we have an incisor of a

rodent from Santa Margarida, 152 incisor parts (including 42 tips) have been recovered during the

picking. These teeth are mostly similar in all rodent species and are not usually used in the fossil

identifications when finding isolated (Hillson, 2005), although new techniques are developing quickly

in that direction and they might not be considered nearly useless anymore soon (Paine et al., 2019).

Figure 5.20: Tip of a rodent incisor from Santa Margarida (Portugal).

The molars and premolars of the rodents are much more characteristic of their different species and

allow its specific identification (Hillson, 2005). In the Santa Margarida sample a total of 51 teeth (6

third lower molars, 2 second lower molars, 15 first lower molars, 5 third upper molars, 9 second upper

molars, 5 first upper molars and 8 indeterminate teeth) whose morphology is congruent with

Apodemus Kaup, 1829 were recovered ( Hillson, 2005). Moreover, in the first lower molars we can see

six low principal cusps, which is a characteristic of Apodemus (López-García, 2008). All the first

upper molars have a developed tubercle 7 (t7) on them as well as a t7–t4 connection in almost all cases

(Figure 5.21) (Piñero et al., 2015). In all the lower first molars the anterocentral cusp is connected to

the anterolabial and anterolingual cusps, meanwhile in the Apodemus mystacinus (Danford and Alston,

1877) from the Quibas site (Spain) around 1/3 of them have this cusp totally isolated (Piñero et al.,

2015). All these characteristics allow us to classify the teeth as belonging to Apodemus cf. sylvaticus.

Figure 5.21: A) third lower molar and B) first upper molar of Apodemus cf. sylvaticus from Santa Margarida

(Portugal) in oclusal view.

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Another group of rodents present in Santa Margarida are the dormice; Gliridae Muirhead, 1819. These

rodents are characterized by their cheek teeth: “brachydont, low and table-like in most genera, with

multiple ridges running buccolingually across the occlusal surface” (Hillson, 2005). There are 1 lower

premolar, 1 second lower molar, 2 second upper molars and 1 indeterminate molar present in the

sample. The teeth of Santa Margarida (Figure 5.22) present highly curved and continuous ridges

without extra cusps besides the four main ones and strongly triangular shaped premolars, which allow

us to ascribe them to Eliomys quercinus (Linnaeus, 1766) (Hillson, 2005; Piñero et al., 2015).

Figure 5.22: Left upper first or second molar of Eliomys quercinus from Santa Margarida (Portugal) in oclusal

view.

There is at least another non arvicoline rodent present in the sample, a cricetine rodent represented by

an upper first molar (Figure 5.23) that bears the characteristic low crown with six cusps aligned in two

rows (Hillson, 2005). It has not anterostyle, the anterolophule is clearly bifurcated and the lingual side

(judging for the position of the anterolophule relative to the protocone) is longer than the labial one.

These characteristics are typical of the species Allocricetus bursae Schaub, 1930 according to Cuenca-

Bescós, (2003).

Figure 5.23: Left upper first molar of Allocricetus bursae from Santa Margarida (Portugal) in oclusal view.

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The arvicolines are the most diverse group of rodents in the sample of Santa Margarida with 73

mandibles, isolated teeth and teeth fragments. Their teeth are highly characteristic with multiple

prismatic elements arranged in a very long oclusal surface with characteristics folds that allow its

identification to specific level (Hillson, 2005). In the figure 5.24 we have a general schematic draw

with the characteristics of the teeth of this group. In the Santa Margarita sample, most of the teeth

were too broken to be useful for identification, but at least four species were recorded:

Figure 5.24: Nomenclature and measurements for an arvicoline first lower tooth according to López-García et

al., (2015). ABBREVIATIONS: a: length of the anteroconid complex; ACC, anteroconid complex; AC, anterior

cap; BRA, buccal reentrant angle; BSA, buccal salient angle; c: width of the opening of triangles T4-T5; L,

length; La, mean width of T4; Li, mean width of T5; LRA, lingual re-entrant angle; LSA, lingual salient angle;

PL, posterior lobe; TTC, trigonidtalonid complex, T1-T7, triangles 1-7; W, width.

The first tooth is a right first lower molar, whose frontal part has been worn down (Figure 5.25).

Morphologically it posses three closed triangles (T1-T3) but the T4-T5 are very confluent, with a wide

connection to the anterior cap, that translates in broad angles of the BRA3 and LRA4. The anterior cap

itself is wide but short. The labial and distal triangles are similar in size. There is not LRA5. These

characters are diagnostics of Victoriamys chalinei (Alcalde et al., 1981) (López-García et al., 2012;

Martin, 2012).

Figure 5.25: Right lower first molar of Victoriamys chalinei from Santa Margarida (Portugal) in oclusal view.

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The second tooth is a first lower molar (Figure 5.26), attached to a hemimandible which also bears the

second molar and was mechanically recovered from the block. It posses lingual triangles that are

larger than the labial ones. The T4-T5 connection is broad and evident but its connection to the

anterior cap is narrow. The BRA4 is absent and so is the T6. The LRA5 is small, therefore the T7 is

almost non-separated of the anterior cap, but both are present. The anterior cap is short and triangular.

These characteristics are congruent with Iberomys huescarensis (Ruiz-Bustos, 1988) according to

López-García et al., (2012, 2015).

Figure 5.26: Left lower first molar of Iberomys huescarensis from Santa Margarida (Portugal) in oclusal view.

The next two teeth (Figure 5. 27) are one left and one right first molars; being the left one attached to

part of its hemimandible. They have similar characteristics to the previous one, but the lingual

triangles are better developed, while the labial ones are smaller. The connection of the T4-T5 is

narrow, as well as the connection to the anterior cap. The LRA5 is marked and separates a T7 from the

anterior cap. The T6 is also visible in the left one. According to López-García et al., (2015) these

characteristics are congruent with Iberomys brecciensis (Giebel, 1847).

Figure 5.27: Right and left (respectively) lower first molars of Iberomys brecciensis from Santa Margarida


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Finally the last recognizable arvicoline teeth in the sample (Figure 5.28) is a first lower left molar that

has lost its posterior lobe. Despite that, it is possible to distinguish on it extremely elongated lingual

triangles with acute angles at their tips. All the T1-T5 triangles are well individualized. The T7 is

greatly developed, reaching the size of the lingual triangles. The LRA5 is greatly developed and well

marked as well. There characters allow it to be referred to Iberomys cabrerae (Thomas, 1906)

according to López-García & Cuenca-Bescós, (2012).

Figure 5.28: Left lower first molar (without the posterior lobe) of Iberomys cabrerae from Santa Margarida


The measurements and ratios for all the identified arvicoline teeth were recorded (Using the program

ImageJ) following the indications showed in the figure 5.22. We have them in the next table (table

5.6).

Table 5.6: Measurements and ratios of the arvicoline first lower molars from Santa Margarida

Allophaiomys

chalinei

Iberomys

huescarensis

Iberomys

brecciensis 1

Iberomys

brecciensis 2

Iberomys

cabrerae

L 2,701 2,958 2,516 2,330

W 1,004 1,041 1,050 0,972 0,955

TTC 1,446 1,427 1,214 1,064

ACC 1,286 1,514 1,349 1,223 1,419

a 1,286 1,514 1,349 1,223 1,419

c 0,260 0,271 0,129 0,056 0,043

La 0,380 0,407 0,384 0,417 0,304

Li 0,493 0,657 0,694 0,584 0,688

A/L 0,476 0,512 0,536 0,525

C/W 0,259 0,260 0,123 0,058 0,045

La/Li 0,771 0,619 0,553 0,714 0,442

A/L x100 47,612 51,183 53,617 52,489

C/Wx100 25,896 26,033 12,286 5,761 4,503

La/Lix100 77,079 61,948 55,331 71,404 44,186

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These measurements and ratios where compared with those provided in López-García et al., (2012,

2015) for Allophaiomys lavocati Laplana & Cuenca-Bescós, 2000; Victoriamys chalinei, Iberomys

huescarensis, Iberomys brecciensis and Iberomys cabrerae for a collection of localities.

Figure 5.29: A/L x100 vs C/W x100 ratios for a collection of arvicoline teeth. Victoriamys chalinei from El

Chaparral locality appear as pink circles. Allophaiomys lavocati from El Chaparral appear as blue pluses

symbols. Iberomys huescarensis, from a collection of Iberian and Italian sites appear as black dots. Data from

López-García et al., (2012, 2015). The specimens from Santa Margarida (Portugal) appear marked with a red

circle.

In the figure 5.29 we can see a comparison between the ratios A/L (which measures how much of the

total length of the tooth is comprised by the anteroconid complex) and C/W (which measures how

much thick is the connection of the T4-T5 compared to its overall width) for the specimens of Santa

Margarida and other localities. We can see how the Santa Margarida specimens ascribed to

Victoriamys chalinei and Iberomys huescarensis fall into the variability of their respective species

meanwhile both Iberomys brecciensis fall further from them with lower values of C/W (narrower

connections between the T4 and T5) and greater values of A/L (Longer anteroconid complexes

compared to the total length).

Figure 5.30: A/L x100 vs La/Li x100 ratios for a collection of arvicoline teeth. Victoriamys chalinei from El

Chaparral locality appear as red inverted triangles. Allophaiomys lavocati from El Chaparral appear as blue

squares. Iberomys huescarensis, from a collection of Iberian and Italian sites appear as black dots. Data from

López-García et al., (2012, 2015). The specimens from Santa Margarida (Portugal) appear marked with a red

circle.

141


total length of the tooth is comprised by the anteroconid complex) and La/Li (which measures how

long is the T4 compared to the T5) for the specimens of Santa Margarida and other localities. The

Victoriamys chalinei of Santa Margarida falls between the variability of the V. chalinei and A. lavocati

from El Chaparral. The I. huescarensis and the two I. brecciensis from Santa Margarida fall into the

variability of the first. This indicates that the I. brecciensis of Santa Margarida do not have very

asymmetrical teeth despite that the length of the anteroconid complex relative to the total length is

greater than in most of I. huescarensis.

Figure 5.31: A/L x100 vs La/Li x100 ratios for Iberomys brecciensis (green squares) and I. chalinei (gold

inverted triangles) from a collection of Iberian and Italian sites. Data from López-García et al., (2015). The

specimens from Santa Margarida (Portugal) appear marked with a red circle.


total length of the tooth is comprised by the anteroconid complex) and La/Li (which measures how

long is the T4 compared to the T5) for the specimens of Santa Margarida and other localities.

Meanwhile one specimen of Iberomys brecciensis fall into the variability of its species the other have

extremely similar sizes of the T4 and T5, almost at the level of the Victoriamys chalinei from Santa

Margarida and surpassing the more archaic I. huescarensis.

Lagomorpha Brandt, 1855

One of the few relatively large sized bones recovered with an air-scribe from one of the blocks is a

distal left humerus (Figure 5.32). It has a narrow trochlea that is inclined towards the medial line of the

body; the coronoid fossa and the olecranon fossa are connected by a distinctive supratrochleal

foramen; the anterior half of the diaphysis is mostly straight in lateral view meanwhile the posterior

part is slightly curved caudally, making the proximal part of the diaphysis wider in lateral view.

Compared with actual remains of rabbit (Oryctolagus cuniculus) it is easy to ascribe it to this species,

given the mentioned characteristics. This is the animal species of largest size identified at this moment

in Santa Margarida.

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Figure 5.32: Left distal Oryctolagus cuniculus humerus from Santa Margarida (Portugal) in A) Anterior; B)

Exterior; C) Posterior and D) Medial views.

Diverse postcranial elements

A wide array of postcranial bones (Figure 5.33) has been recovered in the sample, likely in the range

of hundreds, but the great majority of them are too fragmentary to be identified to the species, or even

family level.

Figure 5.33: Diverse postcranial elements of microvertebrates recovered from the sample of Santa Margarida.

A) Distal femur in posterolateral view, B) Partial distal humerus? in anterior view and C) Distal radius in

lateral view.

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5.5 Discussion

The age

Starting with the gastropods, Pomatias elegans has been cited in Gelasian-Calabrian (2,5 – 0,781

million YBP) deposits from Italy (Bianchi et al., 2013) and it still lives today in Portugal (Platts et al.,

2003), therefore it is not useful for precise dating of the site.

Much more interesting is the appearance of Paralaoma servilis. Its presence in New Zealand and in

Australia dates from the Pleistocene and Holocene, but elsewhere this land snail is regarded as an

invasive species that has arrived in modern times, although it should have departed from New Zealand

soon after western colonization (Christensen et al., 2012). In spite of this, there is a record of this

species coming from soil samples from a remote location of the island of Gávdos (Greece) that is

regarded as unlikely to have been produced by a recent introduction (Welter-Schultes, 1998), therefore

opening the possibility that this species is not an alien in South Europe. In this context the new cite of

Santa Margarida can have two explanations: Either it belongs to a recent contamination (some of the

blocks could have caveats that could house modern snails) or it proves the presence of this species as

native from Southern Europe. Only further work and new shells (especially in situ in blocks) could

prove this point.

The indeterminate lacertid remains are not of value for age determination, since this group has been

present in Europe for dozens of millions of years until today (Bisbal-Chinesta & Blain, 2018).

Talking about bat remains, they are not age-informative; because the Myotis genus has been living in

Europe at least since the beginning of the Pleistocene up to today (Gunnell et al., 2011). Something

similar can be said about the shrews: The Crocidura shrews have been living in Europe for at least 5

million of years (Rofes & Cuenca-Bescós, 2011) and Sorex probably more (Reumer, 1984). Without a

specific determination and it is not possible to give a more precise age.

Talking about the rodents, Eliomys quercinus , that today lives in the Iberian Peninsula, appears on it

during the Early Pleistocene and overlaps with Eliomys intermedius Friant, 1953 during that time

period (Piñero et al.; 2015, 2017). Allocricetus bursae is present in Iberia since the late Early

Pleistocene until its extinction around 20.000 YBP, being specially common during the middle

Pleistocene (López-García, 2008; Sesé et al., 2011). Apodemus sylvaticus is a species that appears in

Iberia around the beginning of the Pleistocene (Sesé et al., 2011) but it becomes abundant only during

the Late Pleistocene (López-García, 2008). The arvicolines are the most interesting taxa for age

determination. Victoriamys chalinei is a typical species from the late Early Pleistocene of the Iberian

Peninsula, being it restricted to the Arvicoline Zone 5 according to Martin, (2012) that ranges

approximately between 1 million and 800.000 YBP. Iberomys huescarensis ranges broadly with the

same time period (López-García et al., 2015). Iberomys brecciensis is a typical species from the

Middle Pleistocene that spans between 800.000 and 120.000 YBP (López-García et al., 2015),

meanwhile Iberomys cabrerae only appears after around 120 YBP and broadly correspond to the Late

Pleistocene until today.

The three chronospecies of Iberomys (Figure 5.34) indicates that in Santa Margarida there are fossils

separated between them for at least 680.000 years; which separates the last occurrence of I.

huescarensis and A. chalinei from the first occurrence of I. cabrerae. Given this absence of

overlapping between those species and the presence of I. brecciensis (That could be present either at

the beginning or the end of the middle Pleistocene) a model with at least two separated time periods

during the Pleistocene is proposed:

1) A first moment around 800.000 YBP that includes broadly the transition between the Early

and Middle Pleistocene (End of the Calabrian). It is characterized by the presence of I.

144

huescarensis and V. chalinei and archaic forms of I. brecciensis. The A. bursae tooth could

come from this phase too.

2) A second moment around 120.000 YBP that includes the end of Middle Pleistocene and the

beginning of the Late Pleistocene. It is characterized by the appearance of late forms of I.

brecciensis and the presence of I. cabrerae. The relatively abundant Apodemus teeth might

come from this phase too.

The first chronology is similar to that of the oldest Quaternary sites known in Portugal: Morgadinho

(Antunes et al., 1986a) and Algoz (Antunes et al., 1986b). In none of those sites I. huescarensis and A.

chalinei have been cited, making Santa Margarida their first known occurrence in Portugal. The

second chronology is similar to the site of Mealhada (Zbyszewski, 1977a) and slightly younger than

some sites in the Almonda karstic system (Marks et al., 2002a; López-García et al., 2018).

Figure 5.34: Evolution of the first lower molar of the genus Iberomys with examples from the site of Santa

Margarida (Portugal).

Stratigraphy and type of the site

All the works that have been carried out (see “methods” section) assumed that given the way of

construction of a traditional algarvian wall (Gonçalves et al., 2018) and the similarity of the hand

samples, all the blocks of cemented terra rossa belonged to the most superficial level of the site and

therefore they were treated equally. As we have seen in the previous section the amount of time

represented in the fossils contained in those superficial blocks leave us with two possible scenarios:

1) The fossils were mixed during the preparation. The blocks collected belonged to different

layers (Figure 5.35 upper).

2) The fossils of the oldest chronology are reworked. The recovered fossils actually come from

the same layer. (Figure 5.35 lower).

145

Figure 5.35: The two possible stratigraphy scenarios that explain the faunal assemblage recovered from the ex

situ blocks from Santa Margarida (Portugal). Upper: Fossils from different but similar layers that became mixed

during preparation. Lower: Reworked fossils situated in the same layer as the younger ones.

This second scenario is unlikely, given the fragility of microvertebrate fossils, but not impossible. It is

not possible to test it with the fragmentation or erosion of the pieces, given the aggressive treatment

with water softener that all specimens have been subject to.

About the nature of the deposit; given the calcareous crusts that cover the terra rossa (that could be

interpreted as speleothems of flowstone type); the presence of bat remains and its geographical

location in the slope of a hill, the site is interpreted as a cave deposit similar to the one known as

“Cueva Des-Cubierta” in Pinilla del Valle, Madrid (Baquedano et al., 2016). The cave itself has been

eroded away, but the cemented and breccified infilling is preserved, with the fossils that it contained.

This could explain why the blocks were in the surface and there is not a trace of a cave nearby.

Finally the richness of microfossil vertebrates found in the locality has to be remarked. In the middle

Pleistocene site of Gruta da Aroeira (Almonda System), 20 kg of sediment yielded 362 specimens

(López-García et al., 2018); meanwhile in Santa Margarida ~0,5 kg yielded ~300 fossils, although

they were much more fragmentary.

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5.6 Conclusions

- Santa Margarida site is a new open-air microvertebrate locality from Algarve, municipality of

Loulé.

- It is known thanks to ex situ blocks of cemented terra rossa with fossils that were used in the

construction of a traditional algarvian wall. Due to the hard cemented sediment of the blocks

the preparation was difficult, but successful.

- About 300 dental and mandibular remains of vertebrates and gastropod shells were collected

from about 0,5 kg of disaggregated sediments, plus hundreds of bone fragments.

- The number of dental remains recovered makes Santa Margarida a remarkably rich site for

microvertebrates.

- At least two invertebrate taxa (Pomatias elegans and Paralaoma servilis) and 12 vertebrates

(Lacertidae indet., Chiroptera indet., Crocidura sp., Sorex sp., Eliomys quercinus,

Allocricetus bursae, Apodemus cf. sylvaticus, Victoriamys chalinei, Iberomys huescarensis,

Iberomys brecciensis, Iberomys cabrerae and Oryctolagus cuniculus) have been recognized.

- This is the first reported occurrence of I. huescarensis and V. chalinei in Portugal.

- There are least two time periods of the Pleistocene represented in Santa Margarida; the first

one about 800.000 YBP and the second around 120.000 YBP.

- Santa Margarida one of the three oldest Quaternary sites known from Portugal, together with

Algoz and Morgadinho, also in Algarve.

- The characteristics of the site are congruent with an eroded cave.

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6. CONCLUSIONS

Here is presented the compilation of conclusions replicated from previous chapters.

Chapter 2: Paleobiodiversity of the Quaternary fossil tetrapods in

continental Portugal

- Most of the vertebrate fossil record of the Quaternary of Portugal (with emphasis in the

Pleistocene) is situated in the Lusitanian and Algarvian basins. Only 3 out of 30 sites are

outside those areas and 20 out of 30 sites are situated in caves.

- Quaternary fossil herpetofaunal communities in Portugal evolve across the period with

reduced numbers and only one extinct species in the area.

- 70% of living bird species are undetected in the fossil record due to their fragile bones, the

difficulties in their identification and the lack of scientific interest.

- 46% of mammal species known in the Quaternary fossil record of Portugal became locally

extinct in the area or went totally extinct. They are also the better known group of Quaternary

fossil tetrapods in the country, with most of the living species (60%) appearing in the fossil

record.

- The importance of the cave deposits is capital in our understanding of the Quaternary

vertebrate fauna, without it we would not known about most of tetrapod species, not to

mention facts about their paleobiology or paleoecology (Like their denning behavior). The

tetrapod living species detected in the fossil record would fall from 37,65% (125 species) to

only a 10,54% (35 species) without them.

Chapter 3: The proboscidean of Santo Antão do Tojal

- The remains of the Santo Antão do Tojal proboscidean (Palaeoloxodon antiquus), which is

around 80.000 YBP old; comprise a first phalanx, a neural arch and a right femur and tibia.

- The Santo Antão do Tojal specimen is estimated to have measured alive about 3,8 meters in

the shoulder and weighted nearly 11 tons, being it in the average for a male Palaeoloxodon

antiquus. Despite being one of the most recent and Southwestern occurrences of this species,

it is nearly identical to specimens that lived several thousand years before in Central Europe or

the Italian Peninsula.

- The size difference between Mammuthus primigenius and Palaeoloxodon antiquus is 18% on

average for the femur and 24% in the tibia. The best measurements for differentiating between

both species are in the femur maximum length, caput width and distal width; and in the tibia

maximum length, minimum diaphysis circumference, minimum diaphysis width and distal

width.

- We agree with previous works (Zbyszewski, 1943; Antunes & Cardoso, 1992) in that the

Santo Antão do Tojal proboscidean is a Palaeoloxodon antiquus in base of the caput width of

the femur and the minimum diaphysis circumference and width of the tibia, combined with a

deep trochanteric fossa in the femur and the overall size of the bone and its robustness.

- Palaeoloxodon antiquus measurements follow a clearer bimodal distribution than Mammuthus

primigenius ones, indicating a more pronounced sexual dimorphism.

- The hind limb anatomy of Palaeoloxodon antiquus reflects more robust bones than in

Mammuthus primigenius, even for similar sized animals, as other works have shown

(Larramendi et al., 2017).

148

Chapter 4: Algar do Vale Da Pena: A new fossil bear site from Portugal

- The Algar do Vale da Pena is a cave recently reopened and is a new fossil brown bear locality

in Portuguese Estremadura.

- 37 items have been recovered from the cave, including 20 informative fossil remains.

- The fossil remains probably belong to five individuals considering its distribution in the cave,

althought the minimal number of individuals in the cave is two, in base of the cranial remains.

- The findings can be ascribed to Ursus arctos in base of morphology (A first upper molar with

clearly individualized metacone-paracone and protocone-hypocone, but without a marked and

big cingulum; mandible with a low condyle compared with the mandibular ramus, slender

long bones without marked muscle insertions, occipital bone with a shallow nucal crest and

straight third phalanxes with robust proximal parts) and metrical comparisons (In several

measurements of 20 bones from the assemblage).

- Two of them are congruent with the measurements and characteristics of mature, yet below

the average sized, female bears.

- The population is formed by relatively small animals (compared to their Spanish counterparts)

in contrast with the observed in other Portuguese localities by Torres Pérez-Hidalgo, (1979)

and Cardoso, (1993).

- The claw marks in the walls of the cave are the first of its type documented in Portugal.

- The great numbers of the claw marks, the morphology of the cave and the taphonomy of the

remains point toward repetitive occupation of the cave by bears during a long time period.

- The presence of Ursus arctos and the thickness of the calcareous mantles over the bones point

towards a Pleistocene age, although the absence of informative stratigraphy prevents the

precise dating of the assemblage for the moment.

Chapter 5: Santa Margarida: A new microvertebrate locality from Algarve.

- Santa Margarida site is a new open-air microvertebrate locality from Algarve, municipality of

Loulé.

- It is known thanks to ex situ blocks of cemented terra rossa with fossils that were used in the

construction of a traditional algarvian wall. Due to the hard cemented sediment of the blocks

the preparation was difficult, but successful.

- About 300 dental and mandibular remains of vertebrates and gastropod shells were collected

from about 0,5 kg of disaggregated sediments, plus hundreds of bone fragments.

- The number of dental remains recovered makes Santa Margarida a remarkably rich site for

microvertebrates.

- At least two invertebrate taxa (Pomatias elegans and Paralaoma servilis) and 12 vertebrates

(Lacertidae indet., Chiroptera indet., Crocidura sp., Sorex sp., Eliomys quercinus,

Allocricetus bursae, Apodemus cf. sylvaticus, Victoriamys chalinei, Iberomys huescarensis,

Iberomys brecciensis, Iberomys cabrerae and Oryctolagus cuniculus) have been recognized.

- This is the first reported occurrence of I. huescarensis and V. chalinei in Portugal.

- There are least two time periods of the Pleistocene represented in Santa Margarida; the first

one about 800.000 YBP and the second around 120.000 YBP.

- Santa Margarida one of the three oldest Quaternary sites known from Portugal, together with

Algoz and Morgadinho, also in Algarve.

- The characteristics of the site are congruent with an eroded cave.

149

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Quaternary fossil vertebrates from continental Portugal: … Lopez... · 2019. 7. 10. · para a detecção de espécies vivas, resultando em que 38% dos tetrápodes terrestres vivos