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History of batrachology in North Africa
CHAPTER 5
History of batrachology in North Africa The first references to the herpetofauna of the region (Egypt) date from Herodotus (484 425 BC), but consisted in general descriptions of conspicuous reptiles (crocodiles, turtles, chameleons, snakes), mixed with descriptions of mythological animals. Older depictions of frogs are found in Egyptian hieroglyphs from the Early Dynastic period (around 3000 BC), representing the fertility—goddess Heqet or Heqtit (Budge, 1904). Amphibians were usually ignored by early naturalists and explorers, despite that some large reptiles were mentioned in their works, for example, Pierre Belon (1517 64) “Observations de plusieurs singularités et choses mémorables trouvées en Grèce, Asie, Judée, Egypte, Arabie et autres pays étranges” (1554). During the colonial period (late 18th to early 20th centuries), Northern Africa was an important destination for naturalists and explorers, who described new taxa based on the recently established Linnaean nomenclature. Early expeditions had mainly focused on Algeria. Jean Louis Marie Poiret (1755 1834), a French clergyman, botanist, and explorer, was sent by Louis XVI to La Calle (El Kala; north-eastern Algeria) to study the local flora. In his extensive work “Voyage en Barbarie, ou Lettres écrites de l’ancienne Numidie pendant les années 1785 & 1786, sur la Religion, les Coutumes & les Múurs des Maures & des Arabes-Bédouins; avec un Essai sur l’histoire naturelle de ce pays” (1789), Poiret mentioned for the first time the presence of urodeles in the North African region, “Lézard des marais (vulgairement Salamandre aquatique) Lacerta palustris,” describing it: “Queue lancéolée, de moyenne grandeur; pieds sans ongles, quatre doigts. Cette espèce de Salamandre vit dans les étangs, où elle se nourrit de petits poissons” (Pleurodeles species) and the “Lézard Salamandre Lacerta Salamandra” (Salamandra algira). Martin Lichtenstein (1780 1857), a German botanist and zoologist, published “Verzeichniss der Doubletten des zoologischen Museums der Königl. Universität zu Berlin nebst Beschreibung vieler bisher unbekannter Arten von Säugethieren, Vögeln, Amphibien und Fischen” (1823) where he described the presence of bufonids in Egypt for the first time, Bufo viridis (Bufotes boulengeri), Bufo cinereus (Sclerophrys regularis), and Bufo fuscus (an undetermined species). Étienne Geoffroy Saint-Hilaire (1772 1844), Isidore Geoffroy Saint-Hilaire (1805 61), and Jean Victoire Audouin (1797 1841) collaborated in a broad series of publications about Egyptian natural history “Description de l’Égypte” (1809 29). In this work, for the first time in Egypt, Rana esculenta (Ptychadena nilotica) and Hyla savignyi were recorded. Claude Antoine Amphibians of North Africa DOI: https://doi.org/10.1016/B978-0-12-815476-2.00005-5
r 2019 Elsevier Inc. All rights reserved.
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Rozet (1798 1858) wrote “Voyage dans la régence d’Alger” (1833), mentioning for the first time the presence in Algeria of Rana esculenta (Pelophylax saharicus), Rana temporaria (Discoglossus pictus), and a toad species (Bufo sp.). Paul Gervais (François Louis Paul Gervaise, 1816 79) wrote “Énumération de quelques espèces de reptiles provenant de Barbarie” (1835), describing Triton poireti (Pleurodeles poireti), commemorated to Jean Louis Marie Poiret, and mentioning a species of toad Bufo arabicus (an undetermined species) from the region of Annaba. Gervais produced a second inventory “Sur les Animaux vertebres de I’Algerie” (1848), where he mentioned for the first time the presence of green toads Bufo variabilis (B. boulengeri) in Algeria. An English botanist, Philip Barker-Webb (1793 1854) published “Histoire Naturelle des Îles Canaries. Reptiles” (1836 54), mentioning the presence of Pelophylax perezi and Hyla meridionalis in the Canary Islands. Hermann Schlegel (1804 84) wrote “Bemerkungen über die in der Regentschaft Algier gesammelten Amphibien” (1841), where he produced a list of five species of amphibians in Algeria: Hyla arborea (H. meridionalis), R. esculenta (P. saharicus), Rana picta (D. pictus), Bufo mauritanicus (Sclerophrys mauritanica), and T. poireti. In his work, he also produced the first taxonomic description of S. mauritanica. Antoine Alphone Guichenot (1809 76) wrote “Exploration Scientifique de l’Algérie: Pendant les Années 1840, 1841, 1842. Histoire Naturelle des Reptiles et des Poissons” (1850) (Fig. 5.1), describing the “Triton nébuleux Triton nebulosus” (Pleurodeles nebulosus) based on specimens found in the surroundings of Algiers. He also mentioned for the first time the presence of Bufo vulgaris (Bufo spinosus) in Algeria. In 1855 “Reisen durch Syrien, Palästina, Phönicien, die Transjordan-Länder, Arabia Petraea und Unter-Aegytpten” by Ulrich Jasper Seetzen (1767 1811) was published, with the description of Rana nilotica (P. nilotica). Alexander Strauch (1832 93), the curator of the zoological museum at the Imperial Academy of Sciences in Saint Petersburg, wrote “Essai d’une erpétologie de l’Algérie” (1862). In that book, he compiled the knowledge about reptiles and amphibians in Algeria, mentioning three species of urodeles, Euproctus rusconii (P. nebulosus), Euproctus poireti (P. poireti), and Salamandra maculosa (S. algira), and six species of anurans, Bufo pantherinus (S. mauritanica), B. viridis (B. boulengeri), B. vulgaris (B. spinosus), D. pictus, H. arborea (H. meridionalis), and R. esculenta (P. saharicus). Shortly after, C. Lallemant provided a similar species account in “Erpétologie de l’Algérie ou catalogue synoptique et analytique des reptiles et amphibiens de la Colonie” (1867). Oskar Boettger (1844 1910) published the first inventories of the herpetofauna of Morocco: “Reptilien von Marocco und von den canarischen Inseln” (1874) and “Die Reptilien und Amphibien von Marocco. II” (1883). The first work contained the description of H. arborea var. meridionalis, based on specimens collected in Tenerife (Böettger, 1874) and the first record of Pleurodeles waltl in Morocco, in the surroundings of Tangier (Böettger, 1874). The second work (1883) included one urodele,
History of batrachology in North Africa
Figure 5.1 Illustrations of Pleurodeles nebulosus and Pleurodeles poireti. From “Exploration Scientifique de l’Algérie: Pendant les Années 1840, 1841, 1842. Histoire Naturelle des Reptiles et des Poissons” (1850).
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Pleurodeles Waltli (P. waltl ), and six anuran species, B. mauritanicus (S. mauritanica), B. viridis (B. boulengeri), B. vulgaris (B. spinosus), D. pictus (Discoglossus scovazzi), Hyla viridis var. meridionalis (H. meridionalis), and R. esculenta var. hispanica (P. saharicus). Böettger also published two papers on the herpetological fauna of the eastern Maghreb: “Liste der von Hrn. Dr. med. W. Kobelt in der Prov. d’Oran, Algerien” (1880 81) and “Liste der von Hrn. Dr. med. W. Kobelt in Algerien und Tunisien gesammelten Kriechtiere” (1885). Lorenzo Camerano (1856 1917) wrote “Osservazioni intorno agli anfibi anuri del Marocco” (1878), containing several species descriptions, including a new species of discoglossid, D. scovazzi and the first record of B. vulgaris (B. spinosus) in Morocco. In 1882, L. Camerano produced the description of R. esculenta var. bedriagae in “Recherches sur les variations de la Rana esculenta et du Bufo viridis dans le Bassin de la Méditerranée.” Fernand Lataste (1847 1934) described Bufo boulengeri (Bufotes boulengeri) based in north-eastern Algerian specimens, in “La zoologie descriptive et la zoologie geographique” (1879). He also provided numerous records of amphibians in Algeria in “Diagnoses de reptiles nouveaux d’Algérie” (1881). Jacques von Bedriaga (1854 1906) described Salamandra maculosa var. algira in his extensive work “Beiträge zur Kenntniss der Amphibien und Reptilien der Fauna von Corsika” (1883) about the herpetofauna of Corsica; Bedriaga compared the morphology of the Corsican fire salamander, with specimens from North Africa and mainland Europe, in order to assess the taxonomic status of the first species. George Albert Boulenger (1858 1937), a Belgian zoologist and assistant in the Department of Zoology in the British Museum, wrote “On the Reptiles and Batrachians obtained in Morocco by Mr. Henry Vaucher” (1889), where he first described the presence of S. algira in Morocco (“Benider hills, near Tangier”; Boulenger, 1889, 1891). In 1891 he published his extensive monograph “Catalogue of Reptiles and Batrachians of Barbary (Morocco, Algeria and Tunisia) based chiefly upon the notes and collections made in 1880 1884 by M. Fernand Lataste,” one of the most complete inventories of the amphibians of the northern Maghreb. This work contained very detailed morphological descriptions and comments on the species’ distribution. In that work, G.A. Boulenger recognized four species of urodeles, Molge poireti (P. poireti), Molge hagenmuelleri (P. nebulosus), Molge waltlii (P. waltl ), and S. maculosa (S. algira), and six anurans B. viridis (B. boulengeri), B. mauritanicus (S. mauritanica), B. vulgaris (B. spinosus), D. pictus (D. pictus/D. scovazzi), H. arborea var. meridionalis (H. meridionalis), and R. esculenta var. ridibunda (P. saharicus). At this moment the species inventory was close to being exhaustive, although the knowledge of the distribution ranges was very incomplete—for example, B. spinosus was not yet known in Tunisia and P. waltl was restricted to the northern extreme of the Tingitana Peninsula (Boulenger, 1891). Joseph Ernest Olivier (1844 1914), a French entomologist, published “Herpétologie algérienne ou Catalogue raisonné des Reptiles et des Batraciens
History of batrachology in North Africa
observés jusqu’à ce jour en Algérie” (1894) and “Matériaux pour la faune de la Tunisie” (1896), providing new distribution data and taxonomic keys. John Anderson (1833 1900), a Scottish anatomist and zoologist, published “On a new species of Zamenis and a new species of Bufo from Egypt” (1893), including the taxonomic description of Sclerophrys pentoni and “Zoology of Egypt. Volume first, Reptilia and Batrachia” (1898), the first comprehensive inventory of Egyptian herpetofauna (including territories of northern Sudan) (Fig. 5.2). In his work, he included a Salamandridae species ignotae and six anuran species: R. esculenta (Pelophylax bedriagae), Rana mascareniensis (P. nilotica), B. viridis (B. boulengeri), Bufo regularis (S. regularis), and Bufo pentoni (S. pentoni) (restricted to Sudan). François Doumergue (1858 1936) published in 1901 “Essai sur la faune erpétologique de l’Oranie: avec des tableaux analytiques et des notions pour la détermination de tous les reptiles & batraciens du Maroc, de l’Algérie et de la Tunisie” (Fig. 5.3). His work followed the nomenclature of Boulenger (1891) and contained new data on natural history and distribution of North African amphibians.
Figure 5.2 Illustrations of (A) Ptychadena nilotica, Bufotes boulengeri, Sclerophrys regularis, and Sclerophrys pentoni (clockwise from top) and (B) Hyla savignyi. From “Zoology of Egypt. Volume first, Reptilia and Batrachia” (1898).
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Figure 5.3 Illustrations (A) from top to bottom Pelophylax saharicus, Bufotes boulengeri, and Bufo spinosus and (B) Pleurodeles waltl and Salamandra algira. From “Essai sur la faune erpétologique de l’Oranie: avec des tableaux analytiques et des notions pour la détermination de tous les reptiles & batraciens du Maroc, de l’Algérie et de la Tunisie” (1901).
Subsequent works at the beginning of the 20th century maintained Boulenger’s list—for example, Mayet (1903) (“Catalogue raisonné des reptiles et batraciens de la Tunisie”) and Pellegrin (1912, 1926) (“Reptiles, batrachiens et poissons du Maroc”; “Reptiles, Batraciens et Poissons du Maroc oriental recueillis par M. P. Pallary”). Henri Gadeau de Kerville (1858 1940) published “Voyage Zoologique en Khroumirie, Tunisie: Mai-Juin 1906” (1908), including the first observation of B. spinosus in Tunisia (Aïn Draham). In 1913, Boulenger described R. esculenta var. saharica (P. saharicus) based on central Algerian specimens (Boulenger in Hartert, 1913). Fernand Angel (1881 1950) published “Sur une collection de reptiles et de batraciens, recueillis au Soudan Français par le Mission, du Dr. Millet Horsin” (1922), describing Arthroleptis milleti-horsini (Tomopterna milletihorsini) and “Sur quelques formes nouvelles de reptiles et de batraciens du Sahara Central” (1936), containing new records and taxonomic accounts from the southern margins of the Sahara (Niger). In 1925 Jacques Pellegrin (1873 1944) added a new species to the Boulenger’s list, describing for the first time the presence of spade-foot toads in Morocco: “Sur la présence au Maroc du Pélobate
History of batrachology in North Africa
cultripède.” Shortly after other works were published, mainly dealing with for the eastern Maghreb region: Seurat (1930) (“Exploration zoologique de l’Algérie de 1830 à 1930”), Mosauer (1934) (“The Reptiles and Amphibians of Tunisia”), Blanc (1935) (“Reptiles et Batraciens. Faune Tunisienne”), and Zavattari (1922, 1937) (“Vertebrati di Cirenaica raccolti dal Generale Medico Prof. Francesco Testi”; “I Vertebrati della Libia. Festschrift für Prof. Dr. Embrik Strand”). Seurat (1930) mentioned “La Grenouille des Mascareignes, espèce appartenant à la faune éthiopienne, a été signalée par le Dr. Pellegrin dans la mare d’Ifédil, du Tassili des Ajjer,” possibly referring to a Ptychadena species, although it was not subsequently found in this region (Le Berre, 1989; Schleich et al., 1996). In 1931 the Spanish herpetologist, F. Galán discovered Alytes obstetricans (Alytes maurus) in northern Morocco, between the localities of Bab-Taza, Beni Seddat, and Ketama; these new records were described in “Batracios y reptiles del Marruecos español.” Stanley Smyth Flower (1871 1946) published “Notes on the Recent Reptiles and Amphibians of Egypt, with a List of the Species recorded from that Kingdom.” Flower added a new species to the Egyptian inventory, Bufo vittatus (Sclerophrys kassasii), based on several specimens collected near Alexandria by Adolf Andres between 1908 and 1912. In 1934 Hugh Bamford Cott (1900 87) published an exhaustive study on the ecology of H. meridionalis in the Canary Islands “On the ecology of Hyla arborea var. meridionalis in Gran Canaria, with special reference to predatory habits considered in relation to the protective adaptations of insects.” Between 1934 and 1937, Giuseppe Scortecci (1898 1973) published several works about Libya and south-eastern Algeria batrachofauna, including “Rettili raccolti nel Deserto Libico dal prof. Lodovico di Caporiacco” (1935) and “Relazione preliminare di un viaggio nel Fezzan sud orientale e sui Tassili” (1937), adding B. regularis (Sclerophrys xeros) and Rana occipitalis (Hoplobatrachus occipitalis) to the north-western Africa inventory. In 1950 J. Guibé published a list of amphibian species occurring in Mauritania “Batraciens. Contribution à l’Étude de l’Aïr,” mentioning B. regularis (S. regularis), B. pentoni (S. pentoni ), and B. mauritanicus (S. xeros). In 1956 P.L. Dekeyser and A. Villiers wrote “Contribution à l’étude du peuplement de la Mauritanie. Notations écologiques et biogéographiques sur la faune de l’Adrar” (1956), adding Pyxicephalus sp. delalandii (T. milletihorsini) and Dicroglossus occipitalis (H. occipitalis) to the Mauritanian list. In 1956 K.P. Schmidt and H. Marx published a species account of the Sinai Peninsula, “The herpetology of Sinai” mentioning two species: B. viridis viridis (B. boulengeri) and R. mascareniensis mascareniensis (P. nilotica). The same authors described for the first time the presence of S. dodsoni in Egypt, restricted to the extreme south-east, in the Gabal Elba region in “Results of the namru-3 southeastern Egypt Expedition, 1954” (1957). In 1959 the French herpetologists Georges Pasteur and Jacques Bons published “Les Batraciens du Maroc.” In the work, they revised the taxonomy and provided new data about the natural history of Moroccan amphibians, describing a new species endemic to Morocco (Pelobates varaldii). Shortly after, the same authors described
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another endemic, A. maurus, in “Note préliminaire sur Alytes maurus: gemellarité ou polytopisme? Remarques biogéographiques, génétiques et taxinomiques” (1962). In 1968 the American herpetologist H. Marx published the “Checklist of the reptiles and amphibians of Egypt,” mentioning six species of anurans: R. mascareniensis mascareniensis (P. nilotica), Rana ridibunda (P. bedriagae), B. vittatus (S. kassasii), Bufo dodsoni (Sclerophrys dodsoni), B. regularis regularis (S. regularis), and B. viridis viridis (B. boulengeri). In 1972 M.S. Hoogmoed described a new species of green toad from southern Morocco, Bufo brongersmai (Barbarophryne brongersmai) in “On a new species of toad from southern Morocco.” In 1974 A. Salvador published “Guia de los anfibios y reptiles españoles” where the distribution of Canarian amphibians was revised; two species were mentioned for the Canary Islands: H. meridionalis and R. ridibunda (P. perezi). In 1975 A. Salvador and S. Peris described a new subspecies of green frog from southern Morocco R. ridibunda riodeoroi (P. saharicus) in “Contribución al estudio de la fauna herpetológica de Río de Oro.” In 1989 M. Le Berre produced a book summarizing the knowledge on the amphibians that inhabit the Sahara and the peripheral regions, in “Faune du Sahara. I. Poissons-Amphibiens-Reptiles.” In 1993 Sherif Baha El Din described the Egyptian populations of B. vittatus as a new species, Bufo kassasii (S. kassasii) in “A new species of toad (Anura: Bufonidae) from Egypt.” Later this author added two more species to the inventory of Egypt: Ptychadena schillukorum in “On the occurrence of Ptychadena schillukorum (Werner, 1907) in Egypt” (2005) and Rana saharica (P. saharicus) in “A Guide to the Reptiles and Amphibians of Egypt” (2006). In 1992 H. Schneider and U. Sinsch published “Mating call variation in lake frogs referred to as Rana ridibunda Pallas, 1771. Taxonomic implications,” suggesting that the populations of Pelophylax ridibundus from the south-east Mediterranean (Israel) should be considered a distinct species, Rana levantina (P. bedriagae). In 1996 three broad reviews were published on amphibians from the Northern Africa region: A. Salvador’s “Amphibians of northwest Africa”; J. Bons and P. Geniez’s “Amphibiens et reptiles du Maroc (Sahara Occidental compris) Atlas bio-géographique”; and H.H. Schleich, W. Kästle, and K. Kabisch’s “Amphibians and Reptiles of North Africa, Biology, Systematics, Field Guide.” Between 2001 and 2003, H. Nickel published two papers, “Biodiversité de la Mauritanie: Liste Comentée des Vertébrés Réellement ou Potentiellement Présents en Mauritanie, à l’Exception des Oiseaux” and “Ökologische Untersuchungen zur Wirbeltierfauna in südóstilichen Mauretanien. Zwei Fallstudien unter besonderer Berücksichtigung der Krokodile,” adding three new species to the Mauritanian list: Phrynobatrachus natalensis, Ptychadena cf. maccarthyensis (Ptychadena bibroni), and Pyxicephalus edulis. In 2003 D. Donaire and S. Bogaerts described S. algira tingitana in “A new subspecies of S. algira Bedriaga, 1883 from northern Morocco.” In 2004 José M. Padial and I. de la Riva published a revision of Mauritania’s inventory “Annotated checklist of the amphibians of Mauritania (West Africa),” adding Ptychadena trinodis and Kassina senegalensis. In 2004 S. Carranza
History of batrachology in North Africa
and E. Wade resurrected the name of P. nebulosus, restricting P. poireti to the Edough Peninsula (“Taxonomic revision of Algero-Tunisian Pleurodeles (Caudata: Salamandridae) using molecular and morphological data. Revalidation of the taxon Pleurodeles nebulosus (Guichenot, 1850)”). In 2004 I. Martinez-Solano in “Phylogeography of Iberian Discoglossus (Anura: Discoglossidae)” resurrected the name of D. scovazzi for the Moroccan discoglossids. In 2006 M. Stöck, C. Moritz, M. Hickerson, D. Frynta, T. Dujsebayeva, V. Eremchenko, J.R. Macey, T.J. Papenfuss, and D.B. Wake (“Evolution of mitochondrial relationships and biogeography of Palearctic green toads (Bufo viridis subgroup) with insights in their genomic plasticity”) demonstrated that North African green toads should be considered as a different species, B. boulengeri. In 2007 D. Escoriza and M.M. Comas described the microendemic S. algira spelaea in “Description of a new subspecies of Salamandra algira Bedriaga, 1883 (Amphibia: Salamandridae) from the Beni Snassen massif (Northeast Morocco).” In 2012, J. Ben Hassine and S. Nouira published a broad review of the distribution of amphibians in Tunisia in “Répartition géographique et affinités écologiques des Amphibiens de Tunisie.” In 2013, another taxonomic revision of the batrachofauna of Morocco by W. Beukema, P. de Pous, D. Donaire-Barroso, S. Bogaerts, J. GarciaPorta, D. Escoriza, O. Arribas, E.H. El Mouden, and S. Carranza (“Review of the systematics, distribution, biogeography and natural history of Moroccan amphibians”) was published, containing the description of S. algira splendens and a new genus of true toads, Barbarophryne. Also in the same year, J.W. Arntzen, E. Recuero, D. Canestrelli, and Í. Martínez-Solano (“How complex is the Bufo bufo species group?”) demonstrated that the Ibero-Maghrebian populations of B. bufo should be considered as distinct species, B. spinosus. In 2013 M. Dehling and U. Sinsch demonstrated that the populations of Ptychadena mascareniensis of the Nile Valley belong to a different species, P. nilotica [“Diversity of Ridged Frogs (Anura:Ptychadenidae: Ptychadena spp.) in wetlands of the upper Nile in Rwanda: morphological, bioacoustic, and molecular evidence”]. In 2017 P. schillukorum was added to the Mauritanian species inventory by A. Sánchez-Vialas, M. Calvo-Revuelta, and R. Márquez in “Ptychadena in Mauritania and the first record of Ptychadena schillukorum.” In 2017 J. Ben Hassine and D. Escoriza published a broad review of the distribution of amphibians in Algeria, providing also new data on their ecology and phenology, “Amphibians of Algeria: new data on the occurrence and natural history.”
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History of batrachology in North Africa The first references to the herpetofauna of the region (Egypt) date from Herodotus (484 425 BC), but consisted in general descriptions of conspicuous reptiles (crocodiles, turtles, chameleons, snakes), mixed with descriptions of mythological animals. Older depictions of frogs are found in Egyptian hieroglyphs from the Early Dynastic period (around 3000 BC), representing the fertility—goddess Heqet or Heqtit (Budge, 1904). Amphibians were usually ignored by early naturalists and explorers, despite that some large reptiles were mentioned in their works, for example, Pierre Belon (1517 64) “Observations de plusieurs singularités et choses mémorables trouvées en Grèce, Asie, Judée, Egypte, Arabie et autres pays étranges” (1554). During the colonial period (late 18th to early 20th centuries), Northern Africa was an important destination for naturalists and explorers, who described new taxa based on the recently established Linnaean nomenclature. Early expeditions had mainly focused on Algeria. Jean Louis Marie Poiret (1755 1834), a French clergyman, botanist, and explorer, was sent by Louis XVI to La Calle (El Kala; north-eastern Algeria) to study the local flora. In his extensive work “Voyage en Barbarie, ou Lettres écrites de l’ancienne Numidie pendant les années 1785 & 1786, sur la Religion, les Coutumes & les Múurs des Maures & des Arabes-Bédouins; avec un Essai sur l’histoire naturelle de ce pays” (1789), Poiret mentioned for the first time the presence of urodeles in the North African region, “Lézard des marais (vulgairement Salamandre aquatique) Lacerta palustris,” describing it: “Queue lancéolée, de moyenne grandeur; pieds sans ongles, quatre doigts. Cette espèce de Salamandre vit dans les étangs, où elle se nourrit de petits poissons” (Pleurodeles species) and the “Lézard Salamandre Lacerta Salamandra” (Salamandra algira). Martin Lichtenstein (1780 1857), a German botanist and zoologist, published “Verzeichniss der Doubletten des zoologischen Museums der Königl. Universität zu Berlin nebst Beschreibung vieler bisher unbekannter Arten von Säugethieren, Vögeln, Amphibien und Fischen” (1823) where he described the presence of bufonids in Egypt for the first time, Bufo viridis (Bufotes boulengeri), Bufo cinereus (Sclerophrys regularis), and Bufo fuscus (an undetermined species). Étienne Geoffroy Saint-Hilaire (1772 1844), Isidore Geoffroy Saint-Hilaire (1805 61), and Jean Victoire Audouin (1797 1841) collaborated in a broad series of publications about Egyptian natural history “Description de l’Égypte” (1809 29). In this work, for the first time in Egypt, Rana esculenta (Ptychadena nilotica) and Hyla savignyi were recorded. Claude Antoine Amphibians of North Africa DOI: https://doi.org/10.1016/B978-0-12-815476-2.00005-5
r 2019 Elsevier Inc. All rights reserved.
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Rozet (1798 1858) wrote “Voyage dans la régence d’Alger” (1833), mentioning for the first time the presence in Algeria of Rana esculenta (Pelophylax saharicus), Rana temporaria (Discoglossus pictus), and a toad species (Bufo sp.). Paul Gervais (François Louis Paul Gervaise, 1816 79) wrote “Énumération de quelques espèces de reptiles provenant de Barbarie” (1835), describing Triton poireti (Pleurodeles poireti), commemorated to Jean Louis Marie Poiret, and mentioning a species of toad Bufo arabicus (an undetermined species) from the region of Annaba. Gervais produced a second inventory “Sur les Animaux vertebres de I’Algerie” (1848), where he mentioned for the first time the presence of green toads Bufo variabilis (B. boulengeri) in Algeria. An English botanist, Philip Barker-Webb (1793 1854) published “Histoire Naturelle des Îles Canaries. Reptiles” (1836 54), mentioning the presence of Pelophylax perezi and Hyla meridionalis in the Canary Islands. Hermann Schlegel (1804 84) wrote “Bemerkungen über die in der Regentschaft Algier gesammelten Amphibien” (1841), where he produced a list of five species of amphibians in Algeria: Hyla arborea (H. meridionalis), R. esculenta (P. saharicus), Rana picta (D. pictus), Bufo mauritanicus (Sclerophrys mauritanica), and T. poireti. In his work, he also produced the first taxonomic description of S. mauritanica. Antoine Alphone Guichenot (1809 76) wrote “Exploration Scientifique de l’Algérie: Pendant les Années 1840, 1841, 1842. Histoire Naturelle des Reptiles et des Poissons” (1850) (Fig. 5.1), describing the “Triton nébuleux Triton nebulosus” (Pleurodeles nebulosus) based on specimens found in the surroundings of Algiers. He also mentioned for the first time the presence of Bufo vulgaris (Bufo spinosus) in Algeria. In 1855 “Reisen durch Syrien, Palästina, Phönicien, die Transjordan-Länder, Arabia Petraea und Unter-Aegytpten” by Ulrich Jasper Seetzen (1767 1811) was published, with the description of Rana nilotica (P. nilotica). Alexander Strauch (1832 93), the curator of the zoological museum at the Imperial Academy of Sciences in Saint Petersburg, wrote “Essai d’une erpétologie de l’Algérie” (1862). In that book, he compiled the knowledge about reptiles and amphibians in Algeria, mentioning three species of urodeles, Euproctus rusconii (P. nebulosus), Euproctus poireti (P. poireti), and Salamandra maculosa (S. algira), and six species of anurans, Bufo pantherinus (S. mauritanica), B. viridis (B. boulengeri), B. vulgaris (B. spinosus), D. pictus, H. arborea (H. meridionalis), and R. esculenta (P. saharicus). Shortly after, C. Lallemant provided a similar species account in “Erpétologie de l’Algérie ou catalogue synoptique et analytique des reptiles et amphibiens de la Colonie” (1867). Oskar Boettger (1844 1910) published the first inventories of the herpetofauna of Morocco: “Reptilien von Marocco und von den canarischen Inseln” (1874) and “Die Reptilien und Amphibien von Marocco. II” (1883). The first work contained the description of H. arborea var. meridionalis, based on specimens collected in Tenerife (Böettger, 1874) and the first record of Pleurodeles waltl in Morocco, in the surroundings of Tangier (Böettger, 1874). The second work (1883) included one urodele,
History of batrachology in North Africa
Figure 5.1 Illustrations of Pleurodeles nebulosus and Pleurodeles poireti. From “Exploration Scientifique de l’Algérie: Pendant les Années 1840, 1841, 1842. Histoire Naturelle des Reptiles et des Poissons” (1850).
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Pleurodeles Waltli (P. waltl ), and six anuran species, B. mauritanicus (S. mauritanica), B. viridis (B. boulengeri), B. vulgaris (B. spinosus), D. pictus (Discoglossus scovazzi), Hyla viridis var. meridionalis (H. meridionalis), and R. esculenta var. hispanica (P. saharicus). Böettger also published two papers on the herpetological fauna of the eastern Maghreb: “Liste der von Hrn. Dr. med. W. Kobelt in der Prov. d’Oran, Algerien” (1880 81) and “Liste der von Hrn. Dr. med. W. Kobelt in Algerien und Tunisien gesammelten Kriechtiere” (1885). Lorenzo Camerano (1856 1917) wrote “Osservazioni intorno agli anfibi anuri del Marocco” (1878), containing several species descriptions, including a new species of discoglossid, D. scovazzi and the first record of B. vulgaris (B. spinosus) in Morocco. In 1882, L. Camerano produced the description of R. esculenta var. bedriagae in “Recherches sur les variations de la Rana esculenta et du Bufo viridis dans le Bassin de la Méditerranée.” Fernand Lataste (1847 1934) described Bufo boulengeri (Bufotes boulengeri) based in north-eastern Algerian specimens, in “La zoologie descriptive et la zoologie geographique” (1879). He also provided numerous records of amphibians in Algeria in “Diagnoses de reptiles nouveaux d’Algérie” (1881). Jacques von Bedriaga (1854 1906) described Salamandra maculosa var. algira in his extensive work “Beiträge zur Kenntniss der Amphibien und Reptilien der Fauna von Corsika” (1883) about the herpetofauna of Corsica; Bedriaga compared the morphology of the Corsican fire salamander, with specimens from North Africa and mainland Europe, in order to assess the taxonomic status of the first species. George Albert Boulenger (1858 1937), a Belgian zoologist and assistant in the Department of Zoology in the British Museum, wrote “On the Reptiles and Batrachians obtained in Morocco by Mr. Henry Vaucher” (1889), where he first described the presence of S. algira in Morocco (“Benider hills, near Tangier”; Boulenger, 1889, 1891). In 1891 he published his extensive monograph “Catalogue of Reptiles and Batrachians of Barbary (Morocco, Algeria and Tunisia) based chiefly upon the notes and collections made in 1880 1884 by M. Fernand Lataste,” one of the most complete inventories of the amphibians of the northern Maghreb. This work contained very detailed morphological descriptions and comments on the species’ distribution. In that work, G.A. Boulenger recognized four species of urodeles, Molge poireti (P. poireti), Molge hagenmuelleri (P. nebulosus), Molge waltlii (P. waltl ), and S. maculosa (S. algira), and six anurans B. viridis (B. boulengeri), B. mauritanicus (S. mauritanica), B. vulgaris (B. spinosus), D. pictus (D. pictus/D. scovazzi), H. arborea var. meridionalis (H. meridionalis), and R. esculenta var. ridibunda (P. saharicus). At this moment the species inventory was close to being exhaustive, although the knowledge of the distribution ranges was very incomplete—for example, B. spinosus was not yet known in Tunisia and P. waltl was restricted to the northern extreme of the Tingitana Peninsula (Boulenger, 1891). Joseph Ernest Olivier (1844 1914), a French entomologist, published “Herpétologie algérienne ou Catalogue raisonné des Reptiles et des Batraciens
History of batrachology in North Africa
observés jusqu’à ce jour en Algérie” (1894) and “Matériaux pour la faune de la Tunisie” (1896), providing new distribution data and taxonomic keys. John Anderson (1833 1900), a Scottish anatomist and zoologist, published “On a new species of Zamenis and a new species of Bufo from Egypt” (1893), including the taxonomic description of Sclerophrys pentoni and “Zoology of Egypt. Volume first, Reptilia and Batrachia” (1898), the first comprehensive inventory of Egyptian herpetofauna (including territories of northern Sudan) (Fig. 5.2). In his work, he included a Salamandridae species ignotae and six anuran species: R. esculenta (Pelophylax bedriagae), Rana mascareniensis (P. nilotica), B. viridis (B. boulengeri), Bufo regularis (S. regularis), and Bufo pentoni (S. pentoni) (restricted to Sudan). François Doumergue (1858 1936) published in 1901 “Essai sur la faune erpétologique de l’Oranie: avec des tableaux analytiques et des notions pour la détermination de tous les reptiles & batraciens du Maroc, de l’Algérie et de la Tunisie” (Fig. 5.3). His work followed the nomenclature of Boulenger (1891) and contained new data on natural history and distribution of North African amphibians.
Figure 5.2 Illustrations of (A) Ptychadena nilotica, Bufotes boulengeri, Sclerophrys regularis, and Sclerophrys pentoni (clockwise from top) and (B) Hyla savignyi. From “Zoology of Egypt. Volume first, Reptilia and Batrachia” (1898).
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Figure 5.3 Illustrations (A) from top to bottom Pelophylax saharicus, Bufotes boulengeri, and Bufo spinosus and (B) Pleurodeles waltl and Salamandra algira. From “Essai sur la faune erpétologique de l’Oranie: avec des tableaux analytiques et des notions pour la détermination de tous les reptiles & batraciens du Maroc, de l’Algérie et de la Tunisie” (1901).
Subsequent works at the beginning of the 20th century maintained Boulenger’s list—for example, Mayet (1903) (“Catalogue raisonné des reptiles et batraciens de la Tunisie”) and Pellegrin (1912, 1926) (“Reptiles, batrachiens et poissons du Maroc”; “Reptiles, Batraciens et Poissons du Maroc oriental recueillis par M. P. Pallary”). Henri Gadeau de Kerville (1858 1940) published “Voyage Zoologique en Khroumirie, Tunisie: Mai-Juin 1906” (1908), including the first observation of B. spinosus in Tunisia (Aïn Draham). In 1913, Boulenger described R. esculenta var. saharica (P. saharicus) based on central Algerian specimens (Boulenger in Hartert, 1913). Fernand Angel (1881 1950) published “Sur une collection de reptiles et de batraciens, recueillis au Soudan Français par le Mission, du Dr. Millet Horsin” (1922), describing Arthroleptis milleti-horsini (Tomopterna milletihorsini) and “Sur quelques formes nouvelles de reptiles et de batraciens du Sahara Central” (1936), containing new records and taxonomic accounts from the southern margins of the Sahara (Niger). In 1925 Jacques Pellegrin (1873 1944) added a new species to the Boulenger’s list, describing for the first time the presence of spade-foot toads in Morocco: “Sur la présence au Maroc du Pélobate
History of batrachology in North Africa
cultripède.” Shortly after other works were published, mainly dealing with for the eastern Maghreb region: Seurat (1930) (“Exploration zoologique de l’Algérie de 1830 à 1930”), Mosauer (1934) (“The Reptiles and Amphibians of Tunisia”), Blanc (1935) (“Reptiles et Batraciens. Faune Tunisienne”), and Zavattari (1922, 1937) (“Vertebrati di Cirenaica raccolti dal Generale Medico Prof. Francesco Testi”; “I Vertebrati della Libia. Festschrift für Prof. Dr. Embrik Strand”). Seurat (1930) mentioned “La Grenouille des Mascareignes, espèce appartenant à la faune éthiopienne, a été signalée par le Dr. Pellegrin dans la mare d’Ifédil, du Tassili des Ajjer,” possibly referring to a Ptychadena species, although it was not subsequently found in this region (Le Berre, 1989; Schleich et al., 1996). In 1931 the Spanish herpetologist, F. Galán discovered Alytes obstetricans (Alytes maurus) in northern Morocco, between the localities of Bab-Taza, Beni Seddat, and Ketama; these new records were described in “Batracios y reptiles del Marruecos español.” Stanley Smyth Flower (1871 1946) published “Notes on the Recent Reptiles and Amphibians of Egypt, with a List of the Species recorded from that Kingdom.” Flower added a new species to the Egyptian inventory, Bufo vittatus (Sclerophrys kassasii), based on several specimens collected near Alexandria by Adolf Andres between 1908 and 1912. In 1934 Hugh Bamford Cott (1900 87) published an exhaustive study on the ecology of H. meridionalis in the Canary Islands “On the ecology of Hyla arborea var. meridionalis in Gran Canaria, with special reference to predatory habits considered in relation to the protective adaptations of insects.” Between 1934 and 1937, Giuseppe Scortecci (1898 1973) published several works about Libya and south-eastern Algeria batrachofauna, including “Rettili raccolti nel Deserto Libico dal prof. Lodovico di Caporiacco” (1935) and “Relazione preliminare di un viaggio nel Fezzan sud orientale e sui Tassili” (1937), adding B. regularis (Sclerophrys xeros) and Rana occipitalis (Hoplobatrachus occipitalis) to the north-western Africa inventory. In 1950 J. Guibé published a list of amphibian species occurring in Mauritania “Batraciens. Contribution à l’Étude de l’Aïr,” mentioning B. regularis (S. regularis), B. pentoni (S. pentoni ), and B. mauritanicus (S. xeros). In 1956 P.L. Dekeyser and A. Villiers wrote “Contribution à l’étude du peuplement de la Mauritanie. Notations écologiques et biogéographiques sur la faune de l’Adrar” (1956), adding Pyxicephalus sp. delalandii (T. milletihorsini) and Dicroglossus occipitalis (H. occipitalis) to the Mauritanian list. In 1956 K.P. Schmidt and H. Marx published a species account of the Sinai Peninsula, “The herpetology of Sinai” mentioning two species: B. viridis viridis (B. boulengeri) and R. mascareniensis mascareniensis (P. nilotica). The same authors described for the first time the presence of S. dodsoni in Egypt, restricted to the extreme south-east, in the Gabal Elba region in “Results of the namru-3 southeastern Egypt Expedition, 1954” (1957). In 1959 the French herpetologists Georges Pasteur and Jacques Bons published “Les Batraciens du Maroc.” In the work, they revised the taxonomy and provided new data about the natural history of Moroccan amphibians, describing a new species endemic to Morocco (Pelobates varaldii). Shortly after, the same authors described
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another endemic, A. maurus, in “Note préliminaire sur Alytes maurus: gemellarité ou polytopisme? Remarques biogéographiques, génétiques et taxinomiques” (1962). In 1968 the American herpetologist H. Marx published the “Checklist of the reptiles and amphibians of Egypt,” mentioning six species of anurans: R. mascareniensis mascareniensis (P. nilotica), Rana ridibunda (P. bedriagae), B. vittatus (S. kassasii), Bufo dodsoni (Sclerophrys dodsoni), B. regularis regularis (S. regularis), and B. viridis viridis (B. boulengeri). In 1972 M.S. Hoogmoed described a new species of green toad from southern Morocco, Bufo brongersmai (Barbarophryne brongersmai) in “On a new species of toad from southern Morocco.” In 1974 A. Salvador published “Guia de los anfibios y reptiles españoles” where the distribution of Canarian amphibians was revised; two species were mentioned for the Canary Islands: H. meridionalis and R. ridibunda (P. perezi). In 1975 A. Salvador and S. Peris described a new subspecies of green frog from southern Morocco R. ridibunda riodeoroi (P. saharicus) in “Contribución al estudio de la fauna herpetológica de Río de Oro.” In 1989 M. Le Berre produced a book summarizing the knowledge on the amphibians that inhabit the Sahara and the peripheral regions, in “Faune du Sahara. I. Poissons-Amphibiens-Reptiles.” In 1993 Sherif Baha El Din described the Egyptian populations of B. vittatus as a new species, Bufo kassasii (S. kassasii) in “A new species of toad (Anura: Bufonidae) from Egypt.” Later this author added two more species to the inventory of Egypt: Ptychadena schillukorum in “On the occurrence of Ptychadena schillukorum (Werner, 1907) in Egypt” (2005) and Rana saharica (P. saharicus) in “A Guide to the Reptiles and Amphibians of Egypt” (2006). In 1992 H. Schneider and U. Sinsch published “Mating call variation in lake frogs referred to as Rana ridibunda Pallas, 1771. Taxonomic implications,” suggesting that the populations of Pelophylax ridibundus from the south-east Mediterranean (Israel) should be considered a distinct species, Rana levantina (P. bedriagae). In 1996 three broad reviews were published on amphibians from the Northern Africa region: A. Salvador’s “Amphibians of northwest Africa”; J. Bons and P. Geniez’s “Amphibiens et reptiles du Maroc (Sahara Occidental compris) Atlas bio-géographique”; and H.H. Schleich, W. Kästle, and K. Kabisch’s “Amphibians and Reptiles of North Africa, Biology, Systematics, Field Guide.” Between 2001 and 2003, H. Nickel published two papers, “Biodiversité de la Mauritanie: Liste Comentée des Vertébrés Réellement ou Potentiellement Présents en Mauritanie, à l’Exception des Oiseaux” and “Ökologische Untersuchungen zur Wirbeltierfauna in südóstilichen Mauretanien. Zwei Fallstudien unter besonderer Berücksichtigung der Krokodile,” adding three new species to the Mauritanian list: Phrynobatrachus natalensis, Ptychadena cf. maccarthyensis (Ptychadena bibroni), and Pyxicephalus edulis. In 2003 D. Donaire and S. Bogaerts described S. algira tingitana in “A new subspecies of S. algira Bedriaga, 1883 from northern Morocco.” In 2004 José M. Padial and I. de la Riva published a revision of Mauritania’s inventory “Annotated checklist of the amphibians of Mauritania (West Africa),” adding Ptychadena trinodis and Kassina senegalensis. In 2004 S. Carranza
History of batrachology in North Africa
and E. Wade resurrected the name of P. nebulosus, restricting P. poireti to the Edough Peninsula (“Taxonomic revision of Algero-Tunisian Pleurodeles (Caudata: Salamandridae) using molecular and morphological data. Revalidation of the taxon Pleurodeles nebulosus (Guichenot, 1850)”). In 2004 I. Martinez-Solano in “Phylogeography of Iberian Discoglossus (Anura: Discoglossidae)” resurrected the name of D. scovazzi for the Moroccan discoglossids. In 2006 M. Stöck, C. Moritz, M. Hickerson, D. Frynta, T. Dujsebayeva, V. Eremchenko, J.R. Macey, T.J. Papenfuss, and D.B. Wake (“Evolution of mitochondrial relationships and biogeography of Palearctic green toads (Bufo viridis subgroup) with insights in their genomic plasticity”) demonstrated that North African green toads should be considered as a different species, B. boulengeri. In 2007 D. Escoriza and M.M. Comas described the microendemic S. algira spelaea in “Description of a new subspecies of Salamandra algira Bedriaga, 1883 (Amphibia: Salamandridae) from the Beni Snassen massif (Northeast Morocco).” In 2012, J. Ben Hassine and S. Nouira published a broad review of the distribution of amphibians in Tunisia in “Répartition géographique et affinités écologiques des Amphibiens de Tunisie.” In 2013, another taxonomic revision of the batrachofauna of Morocco by W. Beukema, P. de Pous, D. Donaire-Barroso, S. Bogaerts, J. GarciaPorta, D. Escoriza, O. Arribas, E.H. El Mouden, and S. Carranza (“Review of the systematics, distribution, biogeography and natural history of Moroccan amphibians”) was published, containing the description of S. algira splendens and a new genus of true toads, Barbarophryne. Also in the same year, J.W. Arntzen, E. Recuero, D. Canestrelli, and Í. Martínez-Solano (“How complex is the Bufo bufo species group?”) demonstrated that the Ibero-Maghrebian populations of B. bufo should be considered as distinct species, B. spinosus. In 2013 M. Dehling and U. Sinsch demonstrated that the populations of Ptychadena mascareniensis of the Nile Valley belong to a different species, P. nilotica [“Diversity of Ridged Frogs (Anura:Ptychadenidae: Ptychadena spp.) in wetlands of the upper Nile in Rwanda: morphological, bioacoustic, and molecular evidence”]. In 2017 P. schillukorum was added to the Mauritanian species inventory by A. Sánchez-Vialas, M. Calvo-Revuelta, and R. Márquez in “Ptychadena in Mauritania and the first record of Ptychadena schillukorum.” In 2017 J. Ben Hassine and D. Escoriza published a broad review of the distribution of amphibians in Algeria, providing also new data on their ecology and phenology, “Amphibians of Algeria: new data on the occurrence and natural history.”
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