Abstract
Lissamphibians (frogs, salamanders, caecilians, and the extinct Albanerpetontidae) have a near global distribution. Africa, its associated islands (especially Madagascar and the Seychelles) and the Arabian Plate are home to about 27 families (including 15 endemic) and 1135 species of extant lissamphibians or about 38 and 15 %, respectively, of the global totals. The region also contains an extensive, but patchy and somewhat under-appreciated fossil record. Based on published and unpublished information, we provide here the most comprehensive review to date of the lissamphibian fossil record from the region. We also discuss the insights those occurrences provide into past distributions and diversities of lissamphibians in the region and the establishment of the modern fauna. Our review relies on occurrence data from 93 sets of localities of basal Triassic through Holocene age, distributed across 23 countries. As with the modern lissamphibian fauna of the region, the fossil record is dominated by frogs, but there also are notable occurrences of other lissamphibians, including several genera of enigmatic Cretaceous salamanders, one of two known stem caecilians, and the only Gondwanan records for albanerpetontids. Africa is one of only two continents (the other being North America) to have occurrences for all four lissamphibian clades. Twenty named and currently accepted fossil lissamphibian species are recognised from the region: one stem and 14 crown frogs (11 or possibly 12 of which are pipimorphs, 1 alytid, and 1 neobatrachian possibly referable to the otherwise exclusively South American families Ceratophryidae or Calyptocephalellidae); three salamanders; one stem caecilian; and one albanerpetontid. Additional and as yet unnamed taxa are represented in existing collections, and others undoubtedly remain to be discovered. Of the 27 extant lissamphibian families currently recognised from the region, 12 of 22 frog families (including five endemics: Brevicipitidae, Heleophrynidae, Hyperoliidae, Ptychadenidae, and Pyxicephalidae) and the sole salamander family (Salamandridae) have fossil records; at present, none of the known caecilian fossils can be assigned with confidence to any of the four extant families currently recognised in the region. The biogeographic histories of lissamphibians in Africa, its associated islands and the Arabian Plate are characterised by vicariant and dispersal events related to the complex palaeogeographic history of the region.
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Notes
For the purposes of this paper, the Arabian Plate includes the following countries (listed clockwise starting from the northwest corner of the Arabian Plate): Israel, Palestine, Lebanon, Jordan, Syria, Iraq, Kuwait, Saudi Arabia, Qatar, Oman and Yemen.
As summarised by Frost (2015), species of Silurana have been variously placed in a separate genus (i.e. Silurana) that is the sister taxon to a more restricted Xenopus (e.g. Evans et al. 2004; Evans 2008; Frost et al. 2006; Pyron and Wiens 2011; Bewick et al. 2012) or placed in the so-called Xenopus tropicalis group within a more inclusive Xenopus (e.g. Loumont and Kobel 1991; Frost 2015).
The sole exception may be the latest Cretaceous (late Maastrichtian) Palaeobatrachus occidentalis, which is reliably known only by several incomplete ilia from the western USA (Estes and Sanchíz 1982; Gardner 2008; Gardner and DeMar 2013). Here we follow Wuttke et al. (2012) in questioning whether that species can be assigned to Palaeobatrachidae.
A tetrapod skeleton from the Lower Cretaceous (Aptian) Crato Formation of Brazil was briefly reported by Martill et al. (2013) in a conference abstract as a caudate and potentially referable to Noterpetontidae. Subsequent study of that Brazilian skeleton indicates that it likely is from a squamate (D. Martill, personal communication 2013).
Many earlier classifications (e.g. Duellman and Trueb 1986; Wilkinson and Nussbaum 2006; Zhang and Wake 2009a) recognised just two caecilian families in Africa: the endemic Scolecomorphidae (identical in the sense used here) and a more inclusive Caeciliidae that contained two dozen genera distributed through tropical regions of South America and Africa and on the Seychelles and the Indian subcontinent. As then conceived, Caeciliidae was ‘a relatively heterogeneous and paraphyletic assemblage comprising all those caecilians that have never been removed to another family’ (Wilkinson and Nussbaum 2006, p. 48); in other words, a ‘waste basket taxon’. Wilkinson et al. (2011) subsequently restricted Caeciliidae to two extant Central and South American genera (Caecilia and Oscaecilia) and partitioned the remaining extant African and Seychelles genera into the families Dermophiidae, Herpelidae and Indotyphilidae; this arrangement has been widely accepted since (e.g. Kamei et al. 2012; San Mauro et al. 2014; Frost 2015) and is followed here.
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Acknowledgements
We are grateful to colleagues from the Université Montpellier II (France), the Universités of Tlemcen and Oran (both in Algeria) and the Office National des Mines of Tunis (Tunisia) for allowing us to include unpublished lissamphibian records from the Eocene (Chambi, El Kohol and Glib Zegdou) of North Africa; to Emmanuel Gheerbrant and Martin Pickford (Muséum National d’Histoire Naturelle, Paris, France) for allowing us to include unpublished anuran records from, respectively, the Palaeocene (Adrar Mgorn 1) of Morocco and the Miocene (Berg Aukas) of Namibia; and to Hillary Maddin (Carleton University, Ottawa, ON, Canada) for allowing us to mention her micro-CT study of the Miocene caecilian skull from Napak XV, Uganda.
For assistance with references, clarifying details about localities and occurrences, providing images, alerting us to occurrences and other help with this project we thank: Robert Asher (Cambridge University, Cambridge, UK); Margaret Avery, Thalassa Matthews and Roger Smith (Iziko South African Museum, Cape Town, South Africa); Jen Bancescu, Caleb Brown, Julien Divay, Dave Eberth, Don Henderson, Craig Scott, Patty Ralrick and Dan Spivak (Royal Tyrrell Museum of Palaeontology, Drumheller, AB, Canada); Daniella Bar-Yosef Mayer (The Steinhardt Museum of Natural History and Institute of Archaeology, Tel Aviv University, Tel Aviv, Israel); Paul Barrett and Zerina Johanson (Natural History Museum, London, UK); Aaron Bauer (Villanova University, Villanova, PA, USA); Rebecca Biton (The Hebrew University of Jerusalem, Jerusalem, Israel); Winand Brinkmann (Paläontologisches Institut und Museum, Zurich, Switzerland); Eric Bruth (Montrouge, France); Eric Buffetaut (Ecole Normale Supérieure, Paris, France); Anusuya Chinsamy-Turan (University of Cape Town, Cape Town, South Africa); Rich Cifelli (University of Oklahoma, Norman, OK, USA); Jim Clark (George Washington University, Washington, DC, USA); Dave DeMar, Jr. (University of Washington, Seattle, WA, USA); David Evans (Royal Ontario Museum, Toronto, ON, Canada); Susan Evans (University College London, London, UK); Amy Henrici (Carnegie Museum of Natural History, Pittsburgh, PA, USA); Robert Holmes and Alison Murray (University of Alberta, Edmonton, AB, Canada); Saida Hossini (Université Moulay-Ismaïl, Meknès, Morocco); August Ilg (Düsseldorf, Germany); Marc Jones (University of Adelaide, Adelaide, SA, Australia); Richard G. Klein (Stanford University, Stanford, CA, USA); Olivier Maridet (Jurassica Museum, Porrentruy, Switzerland); David Martill (University of Portsmouth, Portsmouth, UK); John Measey (Stellenbosch University, Stellenbosch, South Africa); Dick Moody (Kingston University, London, UK); Johannes Müller (Museum für Naturkunde, Berlin, Germany); Marianne Pearson (London, UK); Martin Pickford and Brigitte Senut (Muséum National d’Histoire Naturelle, Paris, France); Cory Redman (Des Moines University, IA, USA); Zbyněk Roček (Geological Institute, Czech Academy of Sciences Prague, Czech Republic); Nicolas Samuelian (Allée de l'Université, Nanterre, France); Gonen Sharon (Tel Hai Academic College, Upper Galilee, Israel); Hans-Dieter Sues (Smithsonian Institution, Washington, DC, USA); Steve Sweetman (Marvel Farm, Isle of Wight, UK); Nancy Stevens (Ohio University, Athens, OH, USA); Bob Sullivan (Albuquerque, NM, USA); and Miguel Vences (Technical University of Braunschweig, Braunschweig Germany). Although we did not pester Eduard Van Dijk (Stellenbosch University, South Africa) while working on this project, we wish to acknowledge his pair of 1995 review papers that provided a starting point for our updated review here. Finally, we are grateful to Amy Henrici and Márton Venczel for their constructive reviews of our manuscript, and to Peter Königshof, Tomáš Přikryl and Sinje Weber for their editorial help.
We dedicate this paper to Zbyněk Roček on the occasion of his 70th birthday. Zbyněk has been a generous and supportive colleague and friend, and we wish him all the best as he eases into semi-retirement.
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Appendix 1
Appendix 1
Summary of 46 Quaternary occurrences for Lissamphibia in Africa, Madagascar and the Arabian Plate. Names for families generally follow current taxonomies (e.g. Blackburn and Wake 2011; Frost 2015), whereas names for genera and species generally follow cited papers and, thus, may not reflect current taxonomies (e.g. Beukema et al. 2013; Frost 2015). Generic names used in an older and more inclusive sense (i.e., sensu lato) are designated with an asterisk (*). Citations behind ‘Anura’ and ‘Urodela’ refer to publications in which listed taxa were identified.
I. Morocco ( n = 10). See recent overview of Moroccan localities and their herpetofaunas by Bailon et al. (in press).
-
1)
Irhoud Ocre; age = early Pleistocene. For further details, see text account for locality 48.
Alytidae: Alytes sp. (originally reported as Baleaphryne sp.)
Bufonidae: Bufo* cf. regularis, Bufo* cf. viridis, and Bufo* sp. indet.
Hylidae: aff. Hyla sp.
?Pelodytidae: ?Pelodytes sp.
Ranidae: Rana* sp.
Urodela (Bailon et al. 2011)
Salamandridae: Pleurodeles cf. waltl
-
2)
Sidi Abdallah; age = early Pleistocene.
Anura (Hossini 2002)
Alytidae: Alytes sp. (originally reported as ?Baleaphryne sp. or Alytes sp).
Ranidae: Rana* sp.
Urodela (Bailon et al. 2011)
Salamandridae: Pleurodeles cf. waltl
-
3)
Grotte des Hominidés; age = middle Pleistocene. Locality incorrectly listed by Hossini (2002) as ‘Grotte des Rhinocéros’ (see Bailon et al. in press).
Anura (Hossini 2002)
Alytidae: Discoglossus sp.
Bufonidae: Bufo bufo spinosus, Bufo* mauritanicus, and Bufo* viridis
Hylidae: Hyla meridionalis
Ranidae: Rana* sp.
-
4)
Doukkala II; age = middle–late Pleistocene.
Anura (Ould Sabar and Michel 1996; Hossini 2002)
Alytidae: Discoglossus pictus cf. scovazzi
Bufonidae: Bufo bufo spinosus and Bufo* mauritanicus
Ranidae: Rana* ridibunda
-
5)
Doukkala I; age = late Pleistocene.
Anura (Ould Sabar and Michel 1996; Hossini 2002)
Bufonidae: Bufo bufo spinosus and Bufo* mauritanicus
-
6)
El Harhoura 1; age = late Pleistocene.
Anura (Bailon and Aouraghe 2002; Hossini 2002)
Bufonidae: Bufo bufo, Bufo* mauritanicus, and Bufo* sp. indet.
-
7)
El Harhoura 2; age = late Pleistocene–Holocene.
Anura (Stoetzel et al. 2010, 2012)
Alytidae: Discoglossus pictus
Bufonidae: Bufo bufo, Bufo* mauritanicus, and Bufo* sp. indet.
Hylidae: Hyla meridionalis
Pelobatidae: Pelobates cf. varaldi
Anura indet.
Urodela (Bailon et al. 2011)
Salamandridae: Pleurodeles cf. waltl
-
8)
Guenfouda; age = Holocene.
Anura (Aouraghe et al. 2010)
Alytidae: Discoglossus pictus
Bufonidae: Bufo* mauritanicus and Bufo* viridis
Ranidae: Rana* sp.
-
9)
Kaf-Taht-el Ghar (= Caf That el Ghar); age = Holocene.
Anura (Hossini 2002; Ouchaou and Hossini 2008)
Bufonidae: Bufo bufo spinosus and Bufo* mauritanicus
-
10)
Kef-el Baroud; age = Holocene.
Anura (Hossini 2002)
Bufonidae: Bufo* mauritanicus
II. Israel ( n = 19)
-
11)
Gesher Benot Ya‘aqov; age = early-middle Pleistocene boundary. For further details, see text account for locality 49.
Anura (Rabinovich and Biton 2011; Biton et al. 2013)
Alytidae: Latonia nigriventer
Bufonidae: Bufo* cf. viridis
Hylidae: Hyla savignyi
Ranidae: Rana* levantina
-
12)
Ubeidiya; age = early Pleistocene.
Anura (Haas 1963, 1966, 1968; Biton et al. 2013)
Alytidae: Latonia nigriventer
Hylidae: ?Hyla sp.
-
13)
Mugharet-el-Zuttiyeh; age = middle Pleistocene.
Anura (Bate 1927)
Bufonidae: Bufo* sp.
Ranidae: Rana* sp.
-
14)
Nesher Ramla; age = middle Pleistocene.
Anura indet. (Zaidner et al. 2014)
-
15)
Qesem Cave; age = middle Pleistocene.
Anura (Smith et al. 2015)
Ranidae indet.
-
16)
Revadim Quarry; age = middle Pleistocene.
Anura (Rabinovich et al. 2012)
Bufonidae: Bufotes cf. viridis (originally reported as Pseudepidalea cf. viridis)
Amphibia indet. (Rabinovich et al. 2012)
-
17)
Rantis Cave; age = middle–late Pleistocene.
Anura (Marder et al. 2011)
Bufonidae: ?Bufo* sp.
Ranidae: ?Rana* sp.
-
18)
Tabun Cave; age = middle–late Pleistocene.
Anura indet.: listed as ‘Frog or Toad’ (Bate 1937)
-
19)
Ain Mallha-Eynan; age = late Pleistocene.
Amphibia unspecified (Bouchud 1987; Valla et al. 1998, 2001, 2004)
Anura (Biton et al. 2013)
Alytidae: Latonia nigriventer
-
20)
Nahal Mahanayeen Outlet; age = late Pleistocene.
Anura (Biton et al. 2013)
Alytidae: Latonia nigriventer
-
21)
Geula; age = Pleistocene.
Anura (Haas 1967)
Pelobatidae: Pelobates cf. syriacus
-
22)
Iraq e Zigan; age = Pleistocene.
Anura (Heller 1978)
Pelobatidae: Pelobates syriacus
-
23)
Abu Usba Cave; age = Holocene.
Anura (Haas in Stekelis and Haas 1952)
Bufonidae: Bufo* viridis
Hylidae: Hyla arborea
Pelobatidae: Pelobates syriacus
Ranidae: ?Rana* sp.
-
24)
El-Wad Terrace (= El Ouad); age = Holocene.
Anura (Valla et al. 1986)
Pelobatidae: Pelobates sp.
Anura indet.
-
25)
Gilgal; age = Holocene.
Anura (Noy et al. 1980)
Bufonidae: Bufo* viridis
Ranidae: Rana* ridibunda
-
26)
Hayonim Cave; age = Holocene.
Anura (Bar-Yosef and Tchernov 1966)
Bufonidae: Bufo* viridis
Hylidae: Hyla arborea
Pelobatidae: Pelobates syriacus
-
27)
Netiv Hagdud; age = Holocene.
Anura (Tchernov 1994)
Ranidae: Rana* ridibunda
-
28)
Sefunim Cave; age = Holocene.
Anura (Tchernov 1984)
Bufonidae: Bufo* viridis
-
29)
Tel Ara (= Ara Cave); age = Holocene.
Anura (Delfino et al. 2007)
Bufonidae: Bufo* viridis
Pelobatidae: Pelobates cf. syriacus
Anura indet.
III. South Africa ( n = 4)
-
30)
Duinefontein 2; age = middle Pleistocene. For further details, see text account for locality 50.
Anura (Sampson 2003)
Bufonidae: Bufo* sp. (possibly B. angusticeps) and Bufo* sp. indet. (= ‘giant form’ of Sampson 2003, p. 551)
Brevicipitidae: Breviceps sp. (possibly B. rosei)
Hyperoliidae: Hyperolius sp. (possibly H. horstockii)
Pipidae: Xenopus sp.
Pyxicephalidae: Tomopterna sp.
Ranidae sensu lato: ‘Ranid A’ and ‘Ranid B’ (Sampson 2003, p. 551). The ilia of what Sampson (2003, p, 551) regarded as ‘a new and as yet unnamed [ranid] species’ instead are likely referable to the extant pyxicephalid Tomopterna delalandii (Van Dijk 2006).
-
31)
Klasies River Mouth; age = late Pleistocene.
Bufonidae: Bufo* sp.
Brevicipitidae: Breviceps sp.
Heleophrynidae: Heleophryne sp.
Hyperoliidae: Genus indet.
Pipidae: Xenopus sp.
Pyxicephalidae: Strongylopus sp. and Tomopterna sp.
?Ranidae sensu lato: Genus indet.
-
32)
Bushman Rock Shelter; age = late Pleistocene.
Anura (Badenhorst and Plug 2012)
Anura indet.
-
33)
Swartkans Cave; age = early–late Pleistocene.
Anura (Watson 2004)
Anura indet.
IV. Namibia ( n = 4). Note we exclude the Kumakrans 1 and 2 localities listed by Van Dijk (1995a, table 1; 1995b, table 1), because according to Cruz-Uribe and Klein (1983, p. 96) those localities are no more than 100 years old.
-
34)
Nosib Cave; age = Pleistocene.
Amphibia unspecified (Pickford and Senut 2010)
-
35)
Bremen; age = Holocene.
Anura (Cruz-Uribe and Klein 1983)
Pyxicephalidae: Pyxicephalus sp.
-
36)
Maguams ‘Andalusia’ and ‘Elefant’; age = Holocene.
Anura (Cruz-Uribe and Klein 1983)
Pyxicephalidae: Pyxicephalus sp.
-
37)
Zebrarivier Cave; age = Holocene.
Amphibia unspecified (Avery 1984)
V. Other localities ( n = 8). Listed alphabetically by country
-
38)
Skikda, Algeria; age = Pleistocene
Anura (Ginsburg et al. 1968)
Alytidae: Discoglossus sp.
Bufonidae: Bufo* sp.
-
39)
Drotsky’s Cave, Botswana; age = late Pleistocene.
Anura (Robbins et al. 1996)
Pipidae: Xenopus sp.
Pyxicephalidae: Pyxicephalus adspersus
-
40)
Lake Ngami, Botswana; age = Holocene.
Anura (Robbins et al. 2009)
Bufonidae: Bufo* sp.
Pyxicephalidae: Pyxicephalus adspersus
Anura indet.
-
41)
Matupi Cave, Democratic Republic of Congo; age = Holocene.
Anura (van Neer 1984)
Anura indet.
-
42)
Bir Tarfawi, Egypt; age = Pleistocene.
Anura (Kowalski et al. 1989)
cf. Ranidae indet.
-
43)
Gadeb 8, Ethiopia; age = early Pleistocene
Anura (J.-C. Rage, unpublished observation)
Pipidae: Xenopus and/or Silurana-like
-
44)
Ksâr’Akil, Lebanon; age = Pleistocene.
Anura (Hooijer 1961)
Hylidae: Hyla arborea
-
45)
Ampasambazimba, Madagascar; age = Holocene.
Anura (MacPhee et al. 1985)
?Microhylidae: Genus indet.
-
46)
Olduvai Gorge, Tanzania age = early Pleistocene.
Anura (Leakey 1967).
Bufonidae: Bufo* sp.
Pipidae: Xenopus sp.
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Gardner, J.D., Rage, JC. The fossil record of lissamphibians from Africa, Madagascar, and the Arabian Plate. Palaeobio Palaeoenv 96, 169–220 (2016). https://doi.org/10.1007/s12549-015-0221-0
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DOI: https://doi.org/10.1007/s12549-015-0221-0