Abstract
Intensified exploration of sub-Saharan Africa during the 18th and 19th centuries led to many newly described giraffe subspecies. Several populations described at that time are now extinct, which is problematic for a full understanding of giraffe taxonomy. In this study, we provide mitochondrial sequences for 41 giraffes, including 19 museum specimens of high importance to resolve giraffe taxonomy, such as Zarafa from Sennar and two giraffes from Abyssinia (subspecies camelopardalis), three of the first southern individuals collected by Levaillant and Delalande (subspecies capensis), topotypes of the former subspecies congoensis and cottoni, and giraffes from an extinct population in Senegal. Our phylogeographic analysis shows that no representative of the nominate subspecies camelopardalis was included in previous molecular studies, as Zarafa and two other specimens assigned to this taxon are characterized by a divergent haplogroup, that the former subspecies congoensis and cottoni should be treated as synonyms of antiquorum, and that the subspecies angolensis and capensis should be synonymized with giraffa, whereas the subspecies wardi should be rehabilitated. In addition, we found evidence for the existence of a previously unknown subspecies from Senegal (newly described in this study), which is now extinct. Based on these results, we propose a new classification of giraffes recognizing three species and 10 subspecies. According to our molecular dating estimates, the divergence among these taxa has been promoted by Pleistocene climatic changes resulting in either savannah expansion or the development of hydrographical networks (Zambezi, Nile, Lake Chad, Lake Victoria).
Highlights
The Europeans saw for the first time a living giraffe in Roman times when Julius Caesar returned with a menagerie of exotic animals from Africa in 46 BC
The southeastern (SE) haplogroup comprises giraffes from southeastern Africa belonging to the subspecies G. g. giraffa, G. t. tippelskirchi and G. t. thornicrofti
Three subgroups are separated by more than 10 mutations: (1) the haplogroup namedSoutheast Africais represented by 13 haplotypes and 83 individuals found exclusively among members of the subspecies G. g. giraffa; (2) the haplogroup namedMasai Iincludes the single haplotype shared by all 34 representatives of the subspecies G. t. thornicrofti and three haplotypes of G. t. tippelskirchi giraffes, comprising 24 extant individuals sampled in northern Tanzania (Serengeti National Park (NP); Manyara NP) and a museum specimen (MNHN-1913-523) collected in Kenya; and (3) the haplogroup namedMasai IIincludes eight haplotypes of the subspecies G. t. tippelskirchi detected in 58 extant individuals collected over the whole sampling area and a museum specimen (RMCA-2128M) sampled in the Serengeti-Mara in southern Kenya
Summary
The Europeans saw for the first time a living giraffe in Roman times when Julius Caesar returned with a menagerie of exotic animals from Africa in 46 BC. During the years 1780–1785, the French explorer François Levaillant undertook two expeditions in the Cape of Good Hope (western part of present South Africa). During his second journey (1783–1785), he encountered several giraffes close to the Orange River (Levaillant 1797), of which he sent a skin to the Muséum national d’histoire naturelle (MNHN) in Paris (illustrated by von Schreber 1784) and a skeleton to the collection of the governor of Netherlands, Guillaume V, in The Hague, which was transferred to the MNHN in 1795 (Schickh 1828). Pierre-Antoine Delalande (1822) enriched the giraffe collection of the MNHN with three additional skulls and a skin from giraffes of the Cape Colony
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