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Ríos Ibáñez, María
Morales Romero, Jorge (dir.); M. Sánchez, Israel (dir.) Departament de Geologia |
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Aquest document és un/a tesi, creat/da en: 2017 | |
The Giraffidae is a monophyletic group of pecorans diagnosed by the presence of a bilobed canine and a special type of epiphyseal, permanent, cranial appendage called ossicone [1, 2]. They are the only living survivors of the Giraffomorpha, a large clade of crown-pecoran ruminants that contains Propalaeoryx, the Palaeomerycidae, the Prolybitheriidae, the Climacoceratidae and the Giraffidae [3].
Giraffids are represented today by only two living African forms, Okapia johnstoni [4] and Giraffa (with four different species) [5-7], but they were very diverse during the Miocene, when the family was widespread throughout the Old World. The more ancient record of the Giraffidae are the early and middle Miocene fossils of Canthumeryx sirtensis [8] from Gebel Zelten (Libya), Moruorot and Rusinga (Kenya) [8, 9] and Georgiomeryx georgalasi [10] from Thymiana (Greece) [11]. However, giraffid rem...
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The Giraffidae is a monophyletic group of pecorans diagnosed by the presence of a bilobed canine and a special type of epiphyseal, permanent, cranial appendage called ossicone [1, 2]. They are the only living survivors of the Giraffomorpha, a large clade of crown-pecoran ruminants that contains Propalaeoryx, the Palaeomerycidae, the Prolybitheriidae, the Climacoceratidae and the Giraffidae [3].
Giraffids are represented today by only two living African forms, Okapia johnstoni [4] and Giraffa (with four different species) [5-7], but they were very diverse during the Miocene, when the family was widespread throughout the Old World. The more ancient record of the Giraffidae are the early and middle Miocene fossils of Canthumeryx sirtensis [8] from Gebel Zelten (Libya), Moruorot and Rusinga (Kenya) [8, 9] and Georgiomeryx georgalasi [10] from Thymiana (Greece) [11]. However, giraffid remains from the middle Miocene are scarce. The group experiences a diversification process during the late Miocene, producing a high variety of shapes and forms. Fossil remains of late Miocene giraffids are abundant all over Eurasia and Africa [1, 12-14].
Only two late Miocene giraffid species had been described from the Iberian Peninsula so far: Decennatherium pachecoi (Vallesian, MN9), present in the Calatayud-Teruel, Duero and Tagus basins and Birgerbohlinia schaubi (early Turolian, MN11) from Piera and Crevillente-2 [15, 16] but since their first publication [15] very few works followed on the subject [17-21]. Several non-cladistic phylogenetic hypotheses were proposed for Decennatherium pachecoi and Birgerbohlinia schaubi. Decennatherium pachecoi was aligned with giraffines [15], samotheres [18], sivatheres [19, 21] and bohlinines [1], whereas Birgerbohlinia schaubi was aligned
with either sivatheres [15] or bohlinines [1]. Decennatherium pachecoi and Birgerbohlinia schaubi were either considered directly related [18, 21], or, alternatively, belonging to two different lineages [15]. Neither of these
hypotheses was tested through a cladistic analysis and the relationships of these genera to each other and to other members of the Giraffidae remained unresolved. The difficulties of clarifying these relationships derive from the plesiomorphic and morphological stable dentition and post-cranial skeleton [9], as well as the lack of good skull remains on the fossil record.
The aim of this thesis is to resolve the phylogenetic position of the Iberian giraffids, exploring the rela-tionships between them and of them within the Giraffidae.
First we started characterizing the clade Giraffomorpha, a large group of crown-pecorans that includes two lineages: the Palaeomerycoidea (Propalaeoryx plus the Palaeomerycidae) and the Giraffoidea (with the Prolybitheriidae as the sister-group to the Climacoceratidae plus the Giraffidae). The Giraffomorpha is defined
as the least inclusive clade containing Giraffa and Triceromeryx, and is diagnosed by derived cranial features, derived navicular-cuboid with a particular plantar morphology, lack of supra-articular fossetes in the metatarsal
III-IV. Our phylogeny also propose the presence of ossicone-like cranial appendages as basal for giraffomorphs, although this has to be still fully corroborated by both the fossil record and a deep analysis of the anatomy and structure of the diverse giraffomorph cranial appendages. We also proposed an emended diagnosis of the
Palaeomerycidae based on cranial and postcranial characters, including several features of the cranium not described so far, and described the beautifully preserved remains of a new palaeomerycid from Spain, Xenokeryx amidalae gen. nov. sp. nov., (La Retama, Loranca Basin, middle Miocene, MN5, Cuenca Province, Spain).
Subsequently we revised all the Spanish material previously assigned to Decennatherium pachecoi (MN9) and Birgerbohlinia schaubi (MN11) including previously undescribed fossils and all the fossil giraffids
stored at the AMNH (New York, USA). We constructed the first character matrix for the Giraffidae (37 taxa and 47 characters), with the aim of reassessing the systematic and phylogenetic position of the two aforementioned Spanish species. This resulted in the first published cladistics analysis of the Giraffidae.
We then proceeded to describe the fossil material from the late Miocene (MN10) fossil site of Batallones-10 (Madrid Basin, Madrid province, Spain; Fig 1), which can be considered the core of this thesis. The beautifully preserved and abundant giraffid material allowed us to describe the new species Decennatherium
rex sp. nov., based upon a complete sample of cranial, dental and postcranial remains. The amount of new morphological data given by Decennatherium rex fossils, especially that from the skull (we have four skulls in the
Batallones-10 sample) is highly relevant to study the systematics and evolution of the Giraffidae. Together with the information obtained from the fossil giraffid collections of the AMNH (New York, USA) NHM (London,UK) and MNHN (Paris, France) we built a new morphological dataset that included 111 characters. The analysis
revealed a scenario where Decennatherium appears as the basal off-shoot of a large clade that includes both samotheres and sivatheres (the sivathere-clade).The next research focused on the only and almost complete Decennatherium rex specimen
from Batallones-4, that helps us to better understand the cranial anatomy of this species, as well as its morphological development, sexual dimorphism and morphological variability.
We also have collaborated in the first comprehensive morphological analysis on the metapodials of the Giraffidae. Both the morphological features of the giraffid metapodials, and the limb length and proportions can be a useful tool for future phylogenetic studies.Esta tesis doctoral se centra en la filogenia y sistemática de los jiráfidos del Mioceno tardío español (Mammalia, Ruminantia, Pecora). Los jiráfidos son un grupo monofilético de rumiantes caracterizados por la presencia de caninos bilobulados y un tipo especial de apéndice craneal epifisario y permanente denominado osicono [1]. Asimismo son los únicos supervivientes del clado Giraffomorpha, un grupo de rumiantes pécoras que contiene a Propalaeoryx junto con los Palaeomerycidae, Prolybitheriidae, Climacoceratidae y Giraffidae [2,3].Actualmente la familia Giraffidae está representada por tan solo dos géneros actuales africanos, Okapia, con una
única especie, Okapia johnstoni, que habita las densas selvas tropicales de la República Democrática del Congo, y Giraffa (con cuatro especies diferentes), hallada en los hábitats de la sabana del África subsahariana [4]. Sin embargo los jiráfidos fueron mucho más diversos y abundantes durante el Mioceno, encontrándose dispersos a
través de toda África y Eurasia [1, 5-7]. El primer registro de la familia Giraffidae está representado por los fósilesdel Mioceno medio Canthumeryx sirtensis (Gebel Zelten (Libia) [8], Moruorot y Rusinga (Kenia) [9] y Georgiomeryx
georgalasi de Thymiana (Grecia) [10]. No obstante, los restos fósiles de jiráfidos del Mioceno medio son muy escasos y fragmentados [1]. Durante el Mioceno tardío el grupo experimentó un rápido proceso de diversificación, produciendo una alta variedad de formas que representan el mayor grado de variabilidad encontrado en la familia [1, 7].
Hasta la fecha tan solo dos especies de jiráfidos del Mioceno tardío han sido descritas en la Península Ibérica:
Decennatherium pachecoi (Vallesiense, MN9) [3, 11-13], presente en las cuencas de Calatayud-Teruel, Duero y Tajo y Birgerbohlinia schaubi (Turoliense temprano, MN11) [14-17] de Piera y Crevillente-2, pero desde que fueron publicadas por primera vez, han sido escasos los trabajos han continuado profundizando en el tema. Se han propuesto diversas hipótesis filogenéticas no clásticas para Decennatherium pachecoi y Birgerbohlinia schaubi.Decennatherium pachecoi fue asignado a los jirafinos, samoterinos, sivaterinos y bohlininos, mientras que Birgerbohlinia schaubi se relacionó con sivaterinos o bohlininos [13, 14, 16]. Decennatherium pachecoi y
Birgerbohlinia schaubi eran considerados parientes o bien pertenecientes a linajes diferentes. Ninguna de estas hipótesis fue testada mediante un análisis cladístico y las relaciones de estos dos géneros entre ellos y con las otras especies e jiráfidos permanecían sin resolver. Las dificultades a la hora de esclarecer las relaciones
existentes entre los Giraffidae residen en la morfología plesiomórfica del esqueleto craneal y postcraneal de esta familia, junto con la falta de restos de cráneos en buen estado de la mayoría de las especies.
Objetivos de la tesis doctoral
El objetivo central de esta tesis es el el de resolver las relaciones filogenéticas de los jiráfidos de la Peninsula Ibérica, explorando las relaciones entre ellos y con el resto de Giraffidae. Con este propósito hemos establecido los
siguientes objetivos:
1. Caracterizar el clado Giraffomorpha y describir un nuevo paleomerícido de España, Xenokeryx amidalae gen. et sp. nov., (La Retama, Cuenca de Loranca, Mioceno medio, MN5, Cuenca, España).
2. Revisar todo el material español previamente asignado a Decennatherium pachecoi (MN9) y Birgerbohlinia schaubi (MN11) incluyendo fósiles no descritos en trabajos anteriores. Construir de la primera matriz de caracteres morfológicos para la familia Giraffidae, con la intención de conocer la sistemática y posición filogenética de las
especies españolas.
3. Describir la nueva especie de jiráfido Decennatherium rex sp. nov., del Mioceno tardío (MN10) del yacimiento Batallones-10 (Cuenca de Madrid, Madrid, España). Mejorar y acrecentar el número de caracteres morfológicos de la matriz de la familia Giraffidae, refinándola e incluyendo la nueva especie española de jiráfido.
4. Describir un nuevo cráneo de Decennatherium rex sp. nov del yacimiento Batallones-4 (Cuenca de Madrid,Madrid, España, MN10), con la intención de ayudarnos a entender la anatomía craneal de esta especie, así como su desarrollo ontogenético, su dimorfismo sexual y su variabilidad morfológica.
5. Colaborar en el primer análisis morfológico en profundidad de los metápodos de Giraffidae con la intención de establecer nuevos caracteres morfológicos útiles para la filogenia y la caracterización de la familia.
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