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Description of the advertisement call and release calls of Bufo (Epidalea) brongersmai Hoogmoed 1972, new distribution records and an attempt to a “total” bibliography

Poster · September 2009

DOI: 10.13140/RG.2.1.2439.2728

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Description of the advertisement call and release calls of Bufo (Epidalea) brongersmai Hoogmoed 1972, new
distribution records and an attempt to a “total” bibliography
Stefano DOGLIO(1,2), Daniele SEGLIE(2,3), Lahcen KABIRI(4) & Massimo Delfino (5,6)
(1) Zirichiltaggi S.W.C. - Strada vicinale Filigheddu, 62/c - 07100 Sassari (Italy)
(2) CESMAP - Prehistoric Study Center and Museum of Prehistoric Art of Pinerolo (Italy)
(3) Dipartimento di Biologia animale e dell’Uomo, Università degli studi di Torino -
Via Accademia Albertina, 13 - 10123 Torino (Italy)
(4) Laboratoire LFS [SCEEP], Département de Géologie, Faculté des Sciences et Techniques
Errachidia, Université My Ismail - BP 509 Boutalamine - 52 000 Errachidia (Maroc)
(5) Paläeontologische Institut und Museum, Universität Zürich, Karl Schmid-Strasse, CH-8006 (Switzerland)
(6) – Dipartimento di Scienze della Terra, Università di Firenze, via G. La Pira 4, 50121 Firenze (Italy)

Abstract

The advertisement and release calls of the Moroccan endemic Bufo (Epidalea) brongersmai (from here on we won’t use anymore the subgenus in the text) from the jbel Sarhro (Anti Atlas) are described. The median value of pulses for an advertisement call is 5 (range 3 to 8
pulses), and the fundamental frequency varies from 1380 to 1890 Hz. It is a short, harsh call with a broad resemblance to a release call, and strikingly different from the advertisement calls of both the closely related Bufo viridis and B. calamita. The distinctive advertisement
call of B. brongersmai is possibly linked with a sympatric origin of this species from a “proto green toad”. The release calls are superficially similar, but the fundamental frequency is usually higher even if there’s a substantial overlap (the range of the recorded calls varies
from 1670 up to 1940 Hz), while the length of the call and the number of pulses is very variable .
New records for B. brongersmai give a more detailed view of the distribution of this toad in the jbel Sarhro area. Finally, an annotated list of the literature published on this still poorly known toad is provided
Short Taxonomical Note

We’re using the subgenus Epidalea instead Pseudepidalea following the suggestion of Speybroek & Crochet (2007) and Epidalea as a “mere” subgenus in an attempt to both accept new evidence and the need for taxonomical stability

Introduction

Bufo brongersmai Hoogmoed 1972 (see the 2 photos in the centre of an adult, fully grown, male) is still a little known bufonid endemic of Morocco, even if during the last few years a few papers and researches have started to come
out (see our “total bibliography” lower down). Broadly speaking B. brongersmai is a “Bufo viridis type” toad, but more adapted to a dry environment, even if this close relatedness has been and is often doubted (see our “total
bibliography”)

Materials & Methods

The toads have been sampled in the field, at site 3 described further on in “New Records”. The calls have been recorded with a Sony TCM-900 DV cassette recorder, an external microphone and a TDK CDing-I “high precision
cassette” (normal position). Sounds have then been digitized with the built-in audio card of a PowerMac ® G4 at a sampling rate of 44.1 kHz at 16 bit, and then analyzed using Audacity ® 1.2.6 and Raven Lite version 1.0 Build 9
Update 17. On the waveform of calls we measured the call duration and the number of pulses, while from the power spectrum we measured the fundamental frequency (Hamming, FFT size 2048) loosely following Castellano et al.
(2002)

The calls

The calls were recorded the 19th March 2005 from 7pm to about 10.30pm; water temperature ranged between 14° C and 15° C during this time, with no wind during the recordings (it had been often moderately windy earlier in the
daylight) and partially cloudy sky. The toads were either in the streamlet or hiding under stones but never more than 1-1.5 m from the water; all the advertisement calls we heard were from males sitting in the water, usually facing
the bank, in a quite typical “green toad” posture.
As already noted by Bogaerts (2002) B. brongersmai advertisement calls are strikingly different from the calls of both B. viridis and B. calamita. Also Guillon et al. (2004) observed that B. brongersmai advertisements were similar Fig.1
to B. viridis release calls. Fig 1 shows one of the calls we recorded, on the waveform on top and the spectrogram down, to be compared with a Bufo calamita call (Fig. 2) and the enlargement of a much longer Bufo viridis call (Fig.
3). In our sample of 7 males every call is composed by a train of pulses, the median number of pulses for an advertisement call being 5 (range 3 to 8 pulses), and the fundamental frequency varies from 1380 to 1890 Hz. The calls
last from 0.53” to 3.11”, with an average value of 1.51” (n = 31 calls, st. dev. 0.58). The single pulses are quite short: from 0.03” up to 0.12” and an average value of 0.09” (n = 51 pulses, from 2 different males, 34 pulses lasted
either 0.09” or 0.10”). Calls are given with a rather irregular interval and are quite difficult to hear from a human perspective. The release calls are superficially similar, but the fundamental frequency is usually higher even if there’s
a substantial overlap (the range of the recorded calls varies from 1670 up to 1940 Hz, n = 13 calls from 5 different males), the length of the call and the number of pulses is rather variable: from 0.68” up to over 5”, and from 3 up to
16 pulses. The single pulse last from 0.1” to 0.11” with an average value of 0.06” (n = 16 pulses
A “total bibliography”
measured from 2 different males).
This is a first attempt to list every publication on this toad, not just the ones more or less focused on it, but also down to works (also for a general public) where the B. brongersmai is just mentioned. The idea is that with a limited range, and not particularly
“famous” animal like this one it’s actually possible to track down all (or most) such publication; quite obviously this is a work in progress and the authors would welcome any addition or correction! In the meantime we hope people working on B. brongersmai or
other Moroccan amphibians will find this “bibliography” useful.
A final warning
The following list is made up searching the Zoological Record database, last accessed late August 2009 (with 3 separate searches for Bufo brongersmai, Pseudepidalea brongersmai and Epidalea brongersmai), and the specialized search engines
scholar.google.com and scirus.com (both last accessed on the 24th September 2009) and by the personal archives of the authors.
It must be noted that only 14 of the following 32 records have been found on the Zoological Records database.
The list follows the order of comparison.
Apparently Gallix (1999) has already analyzed the B. brongersmai calls, but we don’t know if
Books
his results match or not with ours, as we’ve not been able to read his work
1 - van Gelder J. J., Strijbosch H., Bergers P. J. M. eds. (1987) Proceedings of the 4th Ordinary General Meeting of the Societas Europaea Herpetologica - Nijmegen, The Netherlands: Nijmegen Faculty of Sciences, 473 pages
In these proceedings a chapter is dedicated to B. brongersmai:
Grillitsch B, H Grillitsch - The larval development of Bufo brongersmai Hoogmoed 1972 (Amphibia: Anura: Bufonidae), pages 167-170
In English

2 - Le Berre M. (1989) Faune du Sahara, tome 1. Poissons, amphibiens, reptiles - Chabaud Editions, Paris, 332 pages
This book after an introduction to the ecology and geography of the Sahara describes every single species of the area. Even if in need of an update this book is still possibly the best overview of the regional herpetology. B. brongersmai is described from page 76
to 77, and mentioned in the introductory ecology chapter (at page 20, and not 19 as the index says) in the description of the river Draa basin, and also at page 71 in the key of the larvae.
In French

3 - Schleich H. H., Kästle W., Kabisch K. (1996) Amphibians and Reptiles of North Africa - Koeltz Scientific Books, 630 pages
Comprehensive monography on the herps of North Africa. B. brongersmai is described from page 115 to 117, mentioned a few more times in the introduction and the B. viridis pages of the chapter devoted to Bufonidae (pages 114 to 131) and possibly elsewhere
in the same book.
In English

4 - Bons J. & Geniez P. (1996) Amphibiens et reptiles du Maroc (Sahara occidental compris): atlas biogéographique - Asociación Herpetológica Española, Barcelona, 319 pages
Invaluable distributional atlas of the Moroccan herpetofauna. B. brongersmai’s distribution is discussed at pages 62-63 and then again at page 300, and it’s mentioned a few times in the introductory chapters (pages 3, 22, 23, 31, 297)
In French and Spanish through the book, with an English translation at the end

5 - Geniez P., Mateo J. A., Geniez M. & Pether J. (2004) Amphibians and reptiles of the western Sahara. An atlas and field guide - Edition Chimaira, 230 pages
Monograph on the herpetofauna of the former Spanish Sahara, with discussion and description of the range of B. brongersmai in the area
In English Fig. 2
Articles with a strong focus on B. brongersmai

6 - Hoogmoed M. S. (1972) On a new species of toad from southern Morocco - Zoologische Mededelingen, 47 (5) : 49-64
The original article with the description of B. brongersmai. It’s a long paper with a detailed description of the holotype, a discussion of the ecology and distribution of the new species, description of the tadpoles and keys to the adult and larval Moroccan Bufo
spp.
In English

7 - Benhachem L. M., Benazzou T. & Ayachi S. (1989) Etude du polymorphisme morphologique dans une population de Bufo viridis Laurenti, 1768 de Mohammedia - Bulletin de l'Institut Scientifique (Rabat), 13 : 117-124
This paper is a morphological and karyological study of a B. viridis population north of Casablanca, with a revision of the diagnostic characters to disriminate between B. viridis and B. brongersmai ; it’s mentioned by Bons & Geniez (1996) at page 62 of their
Atlas and by Mellado & Dakki (1988)
In French with abstracts in English and Arabic

8 - Grillitsch, B., Grillitsch, H. & Splechtna, H. (1989) The tadpole of Bufo brongersmai Hoogmoed 1972 - Amphibia-Reptilia, 10 : 215-229
Description of the developmental stages, the oral disk and the buccopharyngeal cavity, with comparison to other Mediterranean toads
In English with abstract in German

9 - de la Riva, I. (1992) Bufo brongersmai (Brongersma's toad) - Herpetological Review, 23 : 24

Not read by us, mentioned by Oualidi & Jaziri (2001) and also elsewhere
In English

10 - Herrero P., Lopez-Jurado L. F., Arano B. & Garcia-Paris M. (1993) Karyotype analyses and nuclear DNA content of Bufo brongersmai Hoogmoed - Journal of Herpetology, 27 (4) : 463-465
Note that further confirms the specific status of B. brongersmai and its sister species relationship to B. viridis
In English

11 - El Oualidi J. & Jaziri H. (2001) Variabilités morphologiques et génétique du Crapaud vert Bufo viridis: valeurs systematiques - Bulletin de l'Institut Scientifique (Rabat), 23 : 1-8
This paper, although mainly focused on B. viridis, also argues that more research (on genetics) was needed to clarify the status of B. brongersmai
In French, with summary in English

12 - Bogaerts S. (2001) Breeding Brongersma’s Toad, Bufo brongersmai - Pod@rcis, 2 (3) : 81-88
Based on the captivity breeding experiences of the authors, one of the most useful and interesting papers on the ecology requirements and behaviour of the B. brongersmai
In English

13 - Guillon M., Le Liard G. & Slimani T. (2004) Nouvelles données sur la répartition et l’écologie de reproduction de Bufo brongersmai, B. viridis et B. mauritanicus dans le Jbilets centrales (Maroc) - Bulletin de la Société Herpetologique de France, 111-112 :
37-48
Paper on the ecology and distribution of the toad species in the dry hills north of Marrakech, with a special attention to B. viridis and B. brongersmai
In French, the summary also in English

14 - Schlüter U. (2006) Bufo brongersmai Hoogmoed, 1979 Brongersmas Kröte, Atlaskröte - Reptilia (D), Nr 58 : 51-54
Listed on Zoological Record but unknown to us

15 - Doglio S., Seglie D. & Kabiri L. (2008) First skeletochronology results of Bufo (Epidalea) brongersmai from the Moroccan Anti Atlas (pages 226 to 230) - In Corti C. (ed.) 2008. Herpetologia Sardiniae. SHI/Edizioni Belvedere, Latina “le scienze” (8) 504
pages
Skeletochronolgy results from a small B. brongersmai population from the jbel Sarhro
In English

16 - Delfino M., Doglio S., Rocek Z., Seglie D. & Kabiri L. (2009) Osteological Peculiarities of Bufo brongersmai and Their Possible Relation to Life in an Arid Environment - Zoological Studies, 48 (1) 108-119
Description of selected skeletal elements differentiating B. brongersmai from other Bufo s.l. species
In English
New Records Fig. 3
Other articles
The following sites haven’t been so far described or listed on published papers (except, in part, the 3rd). 4 of them are in the jbel Sarhro or next to this range, while the last one is on the Atlantic coast,
17 - Guillaume C. P. & Bons J. (1982) Nouvelles observations herpétologiques au Maroc. Bull. Soc. Herp. Fr., 23 : 47-53
Never seen by us, this paper is anyway mentioned by Bons & Geniez (1996) for the B. brongersmai sites south of the Atlas, also by Gallix (1999) and elsewhere north of the famous Khnifiss lagoon. These sites don’t extend the range of B. brongersmai but increase our knowledge of its distribution.
Presumably in French

18 - Schouten J. R. & Thevenot M. (1988) Amphibians and Reptiles of the Khnifiss La Youne region. In: M. Dakki and De Ligny (eds), The Khnifiss lagoon and its surrounding environment (province of La youne, Morocc). Trav. Inst. Mém. Hors serie, Rabat,
pages 100 to 114 1 - Aït Ouaazik (10-11/10/2003)
Never seen by us, but widely cited, e.g. in the Bufo brongersmai discussion in Bons & Geniez (1996)
Presumably in English N 30° 40.723’ W 5° 39.173’ (900 m asl)
19 - Mellado J. & Dakki M. (1988) Inventaire commenté des amphibiens et reptiles du Maroc - Bull. Inst. Sci., Rabat, 12 : 171-181 Small village about 25 kms southwest from Tazzarine in the jbel Rhart. Many B. brongersmai tadpoles and newly metamorphosed toadlets found by MD, DS & SD in deep and narrow gullies
Annotated checklist of the Moroccan herpetofauna, B. brongersmai in mentioned also regarding some problems in the morphological diagnostic characters
In French, with abstracts also in English and Arabic dissecting the slope West of the village and the archaeological site. These deep gullies manage to keep a substantial amount of water and a sort of milder micro climate, with more shade, in an
20 - Destre R., Roux P., Geniez P., Thevenot M. & Bons J. (1989) - Nouvelles observations sur l’herpétofaune marocaine. Bulletin de la Société Herpetologique de France, 51 : 19-26 otherwise very barren and rocky slope. B. mauritanicus tadpoles were also observed in the same site
Never seen by us, this paper is again mentioned by Bons & Geniez (1996) for the B. brongersmai sites south of the Atlas, and by Gallix (1999)
Presumably in French

21- Salvador A. (1996) Amphibians of northwestern Africa - Smithsonian Herpetological Information Service, 109 : 1-43 2 - Anou Ichou (12/10/2004)
Often cited monograph, however we have never been able to read it
Presumably in English N 31° 14.114’ W 5° 20.000’ (1070 m asl)
22 - Geniez P., Mateo J.-A. & Bons J. (2000) A checklist of the amphibians and reptiles of Western Sahara - Herpetozoa, 13 (3/4) : 149-163 Valley in the Eastern jbel Sarhro, one adult seen by SD & DS in the dry river bed, hiding in a rock crevice, next to an apparently recently dried shallow pool. The bottom of Anou Ichou is a rather
A long and detailed paper on the herpetofauna of Western Sahara. Bufo brongersmai had yet to be confirmed for the area when this paper came out
“lush” environment for B. brongersmai with many gardens, in the same crevices were hiding also many young B. mauritanicus
23 - Stöck M., Moritz C., Hickerson M., Frynta D., Dujsebayeva T., Eremchenko V., Macey J. R., Papenfuss T. J. & Wake D. B. (2006) Evolution of mitochondrial relationships and biogeography of Palearctic green toads (Bufo viridis subgroup) with insights in
their genomic plasticity. Molecular Phylogenetics and Evolution, 41 : 663-689
Long paper on the phylogeography of the “green toads”, also arguing that B. brongersmai does not belong to the B. viridis subgroup
In English 3 - jbel Sarhro - Boukhanjij Douar (16 & 19/3/2005)
24 - Frost D. R, Grant T., Faivovich J, Bain R. H., Haas A., Haddad C. F. B., De Sá R. O., Channing A., Wilkinson M, Donnellan S. C., Raxworthy C. J., Campbell J. A., Blotto B. L., Moler P., Drewes R. C., Nussbaum R. A., Lynch J. D., Green D. M. & N 31° 2’ 18.9” W 6° 10’ 14.6” (1680 m asl)
Wheeler W. C. (2006) The Amphibian Tree of Life - Bulletin of the American Museum of Natural History, 297 : 1-370
A truly revolutionary paper, moving Brongersma’s toad from Bufo to Pseudepidalea This is not a completely new site, as it’s the one MD & SD sampled and later we described in Doglio et al. (2008) and Delfino et al. (2009), about 20 kms south of the town of El Kelaa M’Gouna,
25 - Gassó Miracle M. E., van den Hoek Ostende L. W. & Arntzen J. W. (2007) Type specimens of amphibians in the National Museum of Natural History, Leiden, The Netherlands. Zootaxa, 1482 : 25-68 northwest jbel Sarhro. It’s also where the calls analyzed here were recorded. In this site B. brongersmai is sympatric with B. viridis, B. mauritanicus and Rana (Pelophylax) saharica
List of the 775 type specimen held in Leiden Museum, B. brongersmai is one of them

26 - Harris D. J., Carretero M. A., Brito J. C., Kaliontzopoulou A., Pinho C., Perera A., Vasconcelos R., Barata M., Barbosa D., Carvalho S., Fonseca M. M., Pérez-Lanuza G. & Rato C. (2008) Data on the distribution of the terrestrial herpetofauna of Morocco:
records from 2001-2006 - Herpetological Bulletin, 103 : 19-28 4 - jbel Sarhro - Akhamsi (17/3/2005)
List of new records of the Moroccan herps, B. brongersmai is found at page 20 with 2 new sites connecting “the locations of Souss basin and south of the High Atlas with the isolated observations in the Marrakech plains”
In English N 31° 4’ 11.6” W 5° 46’ 41.7” (1540 m asl)
27 - Van Bocxlaer I., Biju S. D., Loader S. P. & Bossuyt F. (2009) Toad radiation reveals into-India dispersal as a source of endemism in the Western Ghats-Sri Lanka biodiversity hotspot - BMC Evolutionary Biology, 9:131 This site is actually a roadside stop in a strikingly barren and dry environment on the way down down from Tizi-n’Tazazert toward N’kob. One adult B. brongersmai was spotted on the ground by SD
According to this new paper B. brongersmai (Pseudepidalea brongersmai here) has an “ambiguous phylogenetic position” and is the sister taxon to a big clade comprising the other several “Bufo”, Bufo s.s., Ansonia, Pelophryne, Pseudepidalea (thus non
monophyletic) and others. & MD; the nearest apparent water point was a small pool located about 2 kms away (where R. saharica was found)
Others

28 - Gallix T. (1999) Premieres données sur la biologie de Bufo brongersmai Hoogmoed 1972 Endemique du Maroc, Suivi éco-éthologique d’une population en sympatrie avec Bufo viridis Laurenti 1768 pendant l’année 1999 (Amphibia: Anura: Bufonidae) 5 - oued Laaguig (23/2/2008)
(Oued Oudoudou; S.O.Maroc) - Ecole Pratique des Hautes Etudes, Montpellier, France
Unpublished thesis on the biology and ecology of B. brongersmai (and B. viridis) in southwest Morocco (Souss Massa National Park & Tiznit). N 28° 15’ 9.3” W 11° 37’ 48.7” (sea level)
In French
This site is a deeply cut wadi starting next to the route national about 45 kms north of Sidi Akhfennir and drying in the Atlantic Ocean about 1.5 km from the road. The 23rd February 2008 there were
29 - Berrahmouni N. & Burgess N. (2001) - North Saharan steppe and woodlands (PA1321) - accessed the 15th September 2009 at: http://www.worldwildlife.org/wildworld/profiles/terrestrial/pa/pa1321_full.html
Description of the ecoregion on the WWF website, B. brongersmai is mentioned as the only amphibian of “particular interest” several pools at the upper end of the wadi and running water in the lower half of it. A dead, adult B. brongersmai was found by SD & MD in a pool at the start of the wadi
In English
Bibliography (Poster)
30 - Dakki M., Qninba A., El Agbani M.-A., Bayed A. & Fekhaoui M. (2003) Fiche descriptive sur le zones umides Ramsar (FDR) Baie de Khnifiss - downloaded from www.wetlands.org, accessed the 24th September 2009
Description of the Khnifiss lagoon in the south of Morocco, B. brongersmai is mentioned as one of the reason this area needs Ramsar protection Bogaerts S. (2001) Breeding Brongersma’s Toad, Bufo brongersmai - Pod@rcis, 2 (3) : 81-88 Acknowledgments
In French
Castellano S., Tontini L., Giacoma C., Lattes A. & Balletto E. (2002) The evolution of release and advertisement calls in green toads (Bufo viridis complex) - Biological Journal of the Linnean Society, 77 : 379-391

31 - Jaziri A., El Oualidi J., & Beaubrun P. (2003) Fiche descriptive sur le zones umides Ramsar (FDR) Archipel et dunes d’Essawira - downloaded from www.wetlands.org, accessed the 24th September 2009 Delfino M., Doglio S., Rocek Z., Seglie D. & Kabiri L. (2009) Osteological Peculiarities of Bufo brongersmai and Their Possible Relation to Life in an Arid Environment - Zoological Studies, 48 (1) 108-119
The recordings for this paper have been realized in the frame of a cooperation project between INSAP (Institut
Description of the Essawira area along the Moroccan Atlantic coast, B. brongersmai is mentioned as part of the area “remarkable fauna” National des Sciences de l'Archéologie et du Patrimoine, Rabat, Morocco), and CeSMAP (Study Centre and
Doglio S., Seglie D. & Kabiri L. (2008) First skeletochronology results of Bufo (Epidalea) brongersmai from the Moroccan Anti Atlas (pages 226 to 230) - In Corti C. (ed.) 2008. Herpetologia Sardiniae. SHI/Edizioni Belvedere, Latina “le scienze” (8) 504 pages
In French Museum of Prehistoric Art, Pinerolo, Italy) to set up a new National Park in the jbel Sarhro area, project
Gallix T. (1999) Premieres données sur la biologie de Bufo brongersmai Hoogmoed 1972 Endemique du Maroc, Suivi éco-éthologique d’une population en sympatrie avec Bufo viridis Laurenti 1768 pendant l’année 1999 (Amphibia: Anura: Bufonidae) (Oued Oudoudou; S.O.Maroc) - Ecole Pratique des Hautes Etudes, Montpellier, France supported by both the Moroccan and Italian governments. The B. calamita and B. viridis calls are from the cd
32 - Salvador A., Donaire-Barroso D., El Mouden El H., Slimani T., Geniez P., Mateo J. 2006. Pseudepidalea brongersmai. In: IUCN 2009. IUCN Red List of Threatened Species. Version 2009.1. <www.iucnredlist.org>. (Downloaded on 24 September 2009)
Guillon M., Le Liard G. & Slimani T. (2004) Nouvelles données sur la répartition et l’écologie de reproduction de Bufo brongersmai, B. viridis et B. mauritanicus dans le Jbilets centrales (Maroc) - Bulletin de la Société Herpetologique de France, 111-112 : 37-48 “Guide sonore des Amphibiens” attached to the book “Les Amphibiens de France, Belgique et Luxembourg”,
Brongersma’s toad is listed and briefly described in the IUCN Red List as “Near Threatened” under the name Pseudepidalea brongersmai. It was already listed in possibly all previous editions of the Red List
Duguet et al., editions Biotope, 2005. B. calamita recording comes from Pas de Calais and the B. viridis from
Speybroek & Crochet (2007) Species list of the European herpetofauna - a tentative update. Pod@rcis, 8 (1/2) : 8-34
the Balearic islands

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