AMPHIBIANS AND REPTILES OF NORTH AFRICA,(EE eraphic distribution CONTENTS page Foreword by Wolfgang Bohme Mt Preface Vv Introduction: 1 3 Systematic questions le 1.3. Biological aspects 4 1.4 Abbreviations 5 1.5 Additional information in the appendix 6 2 Amphibians and reptiles of North Africa = species list 6 3 Climate and Vegetation 3.1 Macroclimate ul 3.2 Microclimate 4 3.3 Vegetation zones 45 4 Geographie distribution 4.1 Zoogeographic affinities 4.1.1 Zoogeographic regions 22 4.1.2 Migration routes and radiations 2 4.2 Endemic forms 27 4.3 Disjunctions and relictary forms 28 441 Moyen Atlas: 33 4.4.2 Haut Atlas 34 45 Transsects 34,36-51 4.6 Herpetofauna of islands off the North African coast 32 4.7 Dubious records 33 48 Survey of regional species numbers 54 5 Ecological aspects 5.1 Biotopes 56 5.2 Relations to the ambient 5.2.1 Thermal biology 5.2.1.1 Thermal ecology 60, 5.2.1.2 Temperature-induced sex determination 62 5.22 Water regime 62 5.23 Psammophilous species 63 5.2.3.1 Locomotion on sand 64 5.2.3.2 Life in the sand 64 5.24 Activity cycles. 66 5.24.1 Diel activity cycles 66 5.2.4.2 _ Annual activity cycles 67 5.2.5 Reproductive cycles 1 5.3 Population density and biomass 74 S4 Trophic complexes 75 5.4.1 Role in food chains 7S5.4.2 55 55.1 55.2 5.6 5.6.1 5.6.2 563 71 72 81 82 83 O14 92 93 94 95 10 uw 12 16 751 52-54 55-62 63 1 ‘The region covered by this book comprises the Maghreb states - Morocco, Role in food webs Herpetological communities Lizard communities Landscape and biotope profiles Impact of man Direct impact Habitat change Protective measures Special part: Amphibians and reptiles of the Maghreb and Libya The Amphibian and Reptile species of Eeypt Systematic list Key to the Amphibians and Reptiles of Egypt Appendix Scientific terms Geographic and toponymic terms List of birds and mammals predating on North African Amphibians and Reptiles Index Scientific denominations English denominations French denominations German denominations Spanish denominations Literature References for text figures Addenda during press (genus Chalcides) Colour Plates Amphibians Repiiles Traces Biotopes Impact of man Introduction 7 81 82 88 89 558 565 S67 568 S75 581 584 587 590 S91 621 624 167 168 174 219 222 230 Algeria, Tunisia - and additionally Libya. For several reasons the species restricted to Egypt are only represented in a list and key (7), but not in thespecial part (6): We lack field experience in this country and, as many forms penetrated into Egypt via the Nile valley and Sinai Peninsula, the number of species would have to be greatly augmented. In the species list (2) these forms are listed with the remark ,EGY*. ‘The surprising diversity of the North African herpetofauna is caused by a complex of factors, primarily geological and climatological ones. The partition into a northern Mediterranean and a southern Saharian part is one of the cardinal points of the zoogeography in this region. The Mediterranean part roughly coincides with the geologically young mountain chains Rif and Atlas. The Saharian region begins with the relatively rigid African plate farther to the south which largely consists of a system of large basins. Climate and natural vegetation of these landscapes are outlined in part 3 as both create the prerequisites for animal life. Special adaptations to the diversified biotopes are shown in part 5. tage the surfacing and drowning of land bridges as well as drastic changes created a continually alternating scenario of evolutive factors as the breaking up of distribution areas, immigration of concurring species or changes in population sizes. The repeated fluctuation between high and low precipitations lowered and elevated selection pressures in different senses, inducing dramatic changes in sizes of areas and composition of gene pools. No wonder that the taxonomic relations of many species have been posing problems up to our days. LA Systematic questions A number of North African species was separated from their European relatives with the opening of the Gibraltar Strait after the Messinian crisis 5.7 million yrs b.p.: The Spadefoot toad Pelobates varaldii (from P. cultripes), the Ocellated lizard Lacerta pater (from L. lepida), the amphisbaenid Blanus tingitanus (from B. cinereus), Salamandra algira (from S. salamandra). Within Northern Africa, candidates for splitting into several species are the geckos Saurodactylus (mauritanicus) mauritanicus and Saurodactylus (mauritanicus) brosseti, the skinks Eumeces schneiderii aldrovandii and Eumeces (schneiderii) meridionalis and the widespread and very variable Thorny-eyelidded geckos (Quedenfeldtia). Still more intricate are the problems of classification in complexes where species are difficult to separate and the usual subspecies concept does not match. The Green frogs (provisionally called ,Rana saharica") still await a thorough study. With relations to the Iberian Rana perezi, the eastern Rana levantina, and additional triploid forms they promise to present at least the same degree of complexity as their European conterpart. ‘The Green Toads (Bufo viridis s.l.: sensu lato = in broad sense) appear homogeneous over much of their range, but appear in very different forms in SW-LIB (similar situations in Central Asia). ‘Two species groups of Fringe-toed lizards (the Acanthodactylus pardalis and scutellatus groups) have turned out to be real taxonomic labyrinths.‘Their recent thorough study by four renowned herpetologists resulted in four different ways of classification! Other cases of doubt are the Chalcides ocellatus group, Chalcides polylepis and the Psammodromus blanci/microdactylus complexes. ‘What has been called ,Podarcis hispanica vauchert™ seems to be rather misnamed and anything but homogeneous. Recently, two vipers, Macrovipera deserti and Macrovipera mauritanica, attained specific rank while they were formerly considered as forms of Macrovipera lebetina. A third form, Macrovipera lebetina trans- ‘mediterranea, is probably sympatric with at least one of them, but only one precise locality (Oran) is recorded. Another viperid genus, Echis, has been revised lately and this entailed a ‘number of name changes. Nevertheless the identification of NAF forms of this genus as well as of Macrovipera remains very problematic. ‘The recent discoveries of new species such as Tarentola bochmei, and Chalcides ghiarai show that paris of the Northern Africa are still terra incognita for herpetologists, and certainly more species (not to speak of subspecies) await their discovery in the field or in collections. 1.2 Geographic distribution (see also part 4) ‘The compilation of reliable distribution maps, even approximative ones, is an impossible task for many, if not all, species. In cultivated regions many forms, above all relictary ones, are dwindling away while in desert regions population densities are often thus low that even an expert observer discovers at most a few lizards per day. Taurau (1986) showed that a nocturnal species, considered to be extremely rare, only escaped detection by a secretive way of life. He found several specimens of the Night skink (Scincopus fasciatus) in deep ditches near Tozeur, Tunisia. They got trapped during their nocturnal excursions. The situation is similar in apparently relictary species, namely the dwarf gecko Tropiocolotes nattereri, and the Blind snake Leptotyphlops macrorhynchus. The most restricted area of a North African reptile seems to be that of Ebner’s skink, Chalcides ebneri, one specimen of which was caught near Fez while another of unknown provenience was known through an amateur’s photograph. Our knowledge on Saharan areals and their limitsis rudimentary and mostly restricted to oases and the vicinity of roads. Vast expanses between them, measuring thousands of square kilometers, are practically inaccessible. Fortunately the southern limits of many Saharan species fall outside our maps and we escape the thankless task to draw conjectural lines in a region where the desert steadily advances southward. In connection with distribution areas we try to give some idea of general distribution patterns, endemisms and disjunctions concerning the N-AFR herpetofauna.1.3 Biological aspects ‘The reasons why reptiles play a very important role in arid North African ‘ecosystems are for a considerable part of morphological and physiological nature: Many of them never drink, and evaporate relatively little water through lungs and keratinized skin. The excretion of nitrogenic wastes in form of solid uric acid (in contrast to dissolved urea in mammals) is another important factor in wa’ 2r economy, Considering the fact that endotherms (birds and mammals) use roughly 90% of their energy budget to generate heat, amphibians and reptiles as ektotherms use food energy more economically and are at advantage in warm surroundings where heat production is unnecessary and food is scarce. A short glance at the special part (6) will convince the disappointed reader of the fact that our knowledge of biological aspects consists of islets in an ocean of ignorance. Often we had to refer to extralimital (ext.) data from Europe (mostly Spain) or other parts of Africa. The best studied North African species seems to be the Dabb lizard, Uromastyx acanthinura (Grexor, 1976). It is typical that the studies made by Doumercue in 1901 remain a unique and invaluable source of knowledge to our days. ‘Many important data on behaviour and reproduction are from captive animals (See literature, 10). Only in recent years the maintenance in captivity of extremely heliophilous species such as the Common skink (Scincus scincus) and the Dabb lizard (Uromastyx acanthinura) became successful and resulted in breeding. The use of intense modern light sources helped to ‘overcome the difficulties of keeping these lizards in moderate latitudes (Grexor, 1967; Haerwanw, 1989b). A series of act systems cannot be released at low light intensities. In this regard geckos are less demanding with the exception of the Thorny-eyelidded Quedenfeldtia trachyblepharus. Extensive field studies are indispensable e.g. to learn about the densities, structures and dynamics of populations. Reproductive strategics at different elevations merit special attention as studies of European species (e.g. ‘Salamandra salamandra and Vipera sp.) have shown. An extreme case is Quedenfeldtia trachyblepharus again, living from sea level to a height of 4000 m. 1.4 Abbreviations Geographic names: AFR: Africa, NAF,N-Afr: North Africa, MAG: Maghreb (Morocco, Algeria, Tunisia), MED: Mediterranean, MOR: Morocco, ALG: Algeria, TU! ‘Tunisia, LIB: Libya, SAH: Sahara; EGY: Egypt, Dj: Djebel, Jebel (mount, ‘mountain(s)) Measurements: SVL: Snout-vent-length sec: second(s), h: hour(s), d: day(s), mth: month(s), yr: year(s). Etymology: ¢: English, f: French, g: German, a: Arabic, b: Berber, lat: Latin, gr: GreekExt: extralimital data from regions outside NAF Max., min.: maximal, maximum; minimal, minimum. 1.5 Additional information in the appendix (8) Besides the explanation of scientific terms (8.1), some Arabic and Berber denominations of landscape forms are compilatcd (8.2). The predator list (8.3) serves two purposes: Species cited with their scientific denominations in the special part (6) are rendered with their vernacular names. The predator list also includes general data on prey items, e.g. ,lizards, snakes, small reptiles, frogs and toads", which lack precision and could not be taken into account in the special part (6). 2 Amphibians and reptiles of Northern Africa - species list Species marked with EGY are restricted to Egypt and mentioned only in the list and key for this region (7). Systematics CLASS AMPHIBIA CAUDATA (salamanders and newts) Fam. Salamandridae (True salamanders) Pleurodeles poireti (Gervais, 1835) - wall (Michahelles, 1830) Salamandra algira Bedriaga, 1883 ANURA (frogs and toads) Fam. Discoglossidae Alytes obstetricans (Laurenti, 1768) Discoglossus pictus (Otth, 1837) Fam. Pelobatidae Pelobates varaldii Pasteur & Bons, 1959 Fam. Bufonidae Bufo brongersmai Hoogmoed, 1972 - bufo spinosus Daudin, 1803 ~ dodsoni Boulenger, 1895 EGY - mauritanicus Schlegel, 1841 = pentoni Anderson, 1893 EGY - regularis. Reuss, 1834 - viridis Laurenti, 1768 EGY= vittatus Boulenger, 1906 EGY ~ xeros Tandy, Tandy, Keith & Duff McKay, 1976 Fam. Hylidae Hyla meridionalis Boetiger, 1874 Fam. Ranidae Dicroglossus (Rana) occipitalis Giinther, 1858 Prychadena mascareniensis (Duméril & Bibron, 1841) EGY Rana levantina Schneider, Sinsch & Nevo, 1992 EGY Rana saharica Boulenger, 1913 sometimes abbreviated as Rana compl. (complex) CLASS REPTILIA TESTUDINES (Turtles, Terrapins, Tortoises) Fam, Emydidae Enmys orbicularis (Linnaeus, 1758) Mauremys leprosa (Schweiger, 1812) Fam, Testudinidae Testudo graeca (Linnaeus, 1758) Testudo (Pseudotestudo) kleinmanni (Lortet, 1883) Fam. Trionychidae Trionyx triunguis (Forskal, 1775) EGY Fam. Cheloniidae Chelonia mydas Linnaeus, 1758 Eretmochelys imbricata (Linnaeus, 1766) Caretta caretta Linnaeus, 1758 Lepidochelys kempii (Garman, 1880) Fam. Dermochelyidae Dermochelys coriacea Linnaeus, 1766 CROCODYLIA (Crocodiles) Fam, Crocodylidae Laurenti, 1768 Crocodylus niloticus Laurenti, 1768 EGY SAURIA (Lizards) Fam. Gekkonidae Cyrtopodion scaber (Heyden, 1827) EGY Geckonia chazaliae Mocquard, 1895 Hemidactylus flaviviridis Riippell, 1835 EGY = turcicus Linnaeus, 1758 Phyllodactylus europaeus Géné, 1838 Pristurus flavipunctatus Riippell, 1838 EGY Pryodactylus guitatus (Heyden, 1827) EGY+ hasselquisti Donndorft, 1798 EGY ~ oudrii Lataste, 1880 = ragazzii Anderson, 1898 Quedenfeldtia trachyblepharus (Boettger, 1874) = moerens (Chabanaud, 1916) (not treated separately) Saurodactylus fasciatus Werner, 1936 ~ (mauritanicus) brosseti Bons & Pasteur, 1957 = ( mauritanicus) mauritanicus (Duméril & Bibron, 1836) Stenodactylus petrii Anderson, 1896 ~ sthenodactylus (Lichtenstein, 1823) Tarentola annularis (Geoffroy St.-Hilaire, 1809) - boehmei Joger, 1984 ~ deserti Boulenger, 1891 - ephippiata (O’ Shaugnessy, 1875) ~ mauritanica (Linnaeus, 1758) - neglecta Strauch, 1887 Tropiocolotes nattereri Steindachner, 1901 - steudneri (Peters, 1869) = tripolitanus Peters, 1880 Fam. Agamidae Agama impalearis Boetiger, 1874 + spinosa Gray, 1831 EGY Laudakia stellio (Linnaeus, 1758) EGY Pseudotrapelus sinaitus (Heyden, 1827) Trapelus mutabilis (Merrem, 1820) - pallidus (Reuss, 1833) EGY ~ savignyi (Duméril & Bibron, 1837) EGY - tournevillei (Lataste, 1880) Uromastyx acanthinura Bell, 1825 - aegyptia (Forskil, 1775) EGY - geyri L. Miller, 1922 - ocellata Lichtenstein, 1823 EGY Fam, Chamaeleonidae Chamaeleo africanus Laurenti, 1768 EGY ~ chamaeleon (Linnaeus, 1825) Fam. Scincidae Chalcides chalcides (Linnaeus, 1758) ~ guentheri Boulenger, 1887 EGY ~ mauritanicus (Duméril & Bibron, 1839) ~ mionecton (Boettger, 1873) - ocellatus group: - ocellatus (Forskal, 1775) ~ colosii Lanza, 1967 - ebneri Werner, 1931 - ghiarai Caputo & Mellado, 1992 ~ lanzaiPasteur, 1967 - manueli Hediger, 1935~ montanus Werner, 1931 - polylepis Boulenger, 1890 = ragazzii Boulenger, 1890 Eumeces algeriensis Peters, 1864 ~ schneiderii (Daudin, 1802) = (Schneiderii) meridionalis Doumergue, 1900 Mabuya quinquetaeniata (Lichtenstein, 1823) EGY - vittata (Olivier, 1804) Scincopus fasciatus (Peters, 1864) Scincus albifasciatus Boulenger, 1890 ~ scincus (Linnaeus, 1758) ‘Sphenops boulengeri (Anderson, 1896) ~ delislei (Lataste, 1876) ~ sepsoides Audouin, 1829 ~ sphenopsiformis (Duméril, 1856) Fam, Lacertidae Acanthodactylus boskianus group: ~ boskianus (Daudin, 1802) - erythrurus group: ~ blanci Doumergue, 1901 ~ erythrurus (Schinz, 1838) ~ savignyi (Audouin, 1809) pardalis group: ~ pardalis (Lichtenstein, 1823) including - bedriagai Lataste, 1881 ~ busacki Salvador, 1982 ~ maculatus (Gray, 1838) - spinicauda Doumergue, 1901 scutellatus group: - aureus Giinther, 1903 = scutellatus (Audouin, 1809) including - dumerili (Milne Edwards, 1829) - longipes Boulenger, 1918 Lacerta andreanskyi Werner, 1929 Latastia longicaudaia (Reuss, 1834) EGY Mesalina guttulata (Lichtenstein, 1823) = olivieri (Audouin, 1829) ~ pasteuri Bons, 1960 ~ rubropunctata (Lichtenstein, 1823) - simoni (Boettger, 1881) Ophisops elbaensis Schmidt & Marx, 1957 EGY - elegans Ménétries, 1832 + occidenialis (Boulenger, 1891) Philochortus intermedius Boulenger, 1917 EGY - zolii Scortecci, 1934 Podarcis hispanica (Steindachner, 1870) - perspicillata (Duméril & Bibron, 1839) Psammodromus algirus (Linnaeus, 1766) blanci Lataste, 188010 - microdactylus (Boetiger, 1881) Pseuderemias mucronata (Blanford, 1870) EGY Fam, Anguidae Ophisaurus koellikeri (Giinther, 1873) Fam, Varanidae Varanus griseus (Daudin, 1803) - niloticus (Linnaeus, 1766) EGY Fam. Amphisbaenidae Blanus metetali Bons, 1963 - tingitanus Busack, 1988 Fam, Trogonophidae Trogonophis wiegmanni Kaup, 1830 SERPENTES (Snakes) Fam. Typhlopidae ‘Typhlops vermicularis Merrem, 1820 EGY Fam. Leptotyphlopidae Leptotyphlops cairi (Duméril & Bibron, 1844) EGY - macrorhynchus (Jan, 1862) Fam. Boidae Eryx colubrinus (Linnaeus, 1758) EGY ~ jaculus (Linnaeus, 1758) Fam, Colubridae Coluber algirus (Jan, 1863) - florulentus Geoffroy St.-Hilaire, 1827 EGY - hippocrepis (Linnaeus, 1758) = rhodorachis (Jan, 1865) - rogersi (Anderson, 1893) Coronella girondica (Daudin, 1803) Dasypeliis scabra (Linnaeus, 1758) Lamprophis fuliginosus (Boic, 1827) Lycophidion capense (A. Smith, 1831) EGY Lytorhynchus diadema (Duméril & Bibron, 1854) Macroprotodon cucullatus (Geoffroy St-Hilaire, 1827) Malpolon moilensis (Reuss, 1834) = monspessulanus (Hermann, 1804) Natrix maura (Linnaeus, 1758) - natrix (Linnaeus, 1758) - tessellata (Laurenti, 1768) EGY Psammophis aegyptius Marx, 1958 - schokari (Forskiil, 1775)i - sibilans (Linnaeus, 1758) Spalerosophis diadema (Schlegel, 1837) = dolichospilus (Werner, 1923) Telescopus obtusus (Reuss, 1894) Fam. Elapidac Naja haje (Linnaeus, 1758) - nigricollis Reinhardt, 1843 EGY Walterinnesia aegyptia Lataste, 1887 EGY Fam. Viperidae Bitis arietans (Mertem, 1820) Cerastes cerastes (Linnaeus, 1758) - vipera (Linnaeus, 1758) Echis arenicola H. Boie in F. Boie, 1827 = pyramidum (Geoffroy St.-Hilaire, 1827) Macrovipera deserti (Anderson, 1892) - ,lebetina transmediterranea Nilson & Andrén, 1988 - mauritanica (Duméril & Bibron, 1849) Vipera latastei Bosca, 1878 = monticola Saint Girons, 1953 3 Climate and Vegetation Amphibians and reptiles depend on the main climatic factors, temperature and precipitation, in many ways. Being ectothermal they can only deploy their activities in surroundings with appropriate temperatures within a relatively narrow range. Optimal body temperatures can be maintained by different means of behavioural temperature control, the most obvious of which is basking. While in moderate climates sufficient heat intake is the major problem, animals inhabiting hot deserts are often threatened by overheating and have acquired different ways to avoid lethal temperatures. Precipitations furnish the water for plant life which forms the base for all food chains and food webs in which amphibians and reptiles occur. Water is also the living milieu for all larvae of North African amphibians and the adults of newt and some frog species. ‘The struggle for water is one of the main problems of survival in deserts, and population densities directly correlate with the quantity of water disposible for plants and animals. Man has become a fierce competitor in this struggle as increasing quantities of water are needed for irrigation projects. With the intensive channelling of natural watercourses the number of suitable biotopes for some aquatic species decreases rapidly. 3.1 Macroclimate The ,climates* in which large and small animals of the same biocoenosis, live may differ substantially. As solar radiation warms the air not directly, but after heating the soil, animals living close to the ground are exposed to12 16.5° 35m Bartamedo 2008 18.0" 315a0 Fig. 1. Effect of elevation: Climatograms for 1982 from two stations in Kouf National Park about 150 km NE of Bengasi (nr:3 following page). ‘Temperature and precipitation scale see following page Kouf Station 500m Fig. 2. Temperature extremes for 1982 in two stations of Kouf National Park (Cyrenaica, LIB), Scale see following page. much greater temperature extremes than the head of a standing or sitting human, The data on the .macroclimate* furnished by a meteorological station (2 mabove ground) can therefore give only a faint idea of the microclimate in which small diurnal vetrebrates live. Nevertheless they are indispensable to characterize the climate of a region and the character of its vegetation. ‘The climatograms (p. 13) used here allow the recognition of arid and humid seasons at the first glance. This is afforded by an appropriate scale of temperatures (left margin of graphs) and precipitations (right margin) where 10°C correspond with 20 mm of precipitation. This relation was chosen because for every 10 degrees centigrade above zero of monthly mean temperature, about 20 mm of monthly precipitation evaporate. If the precipitation curve rises above the temperature curve (hatched part), a surplus of water remains and the climate is called humid, In the contrary case (dotted part) the climate is arid. The distance between both curves gives a measure of the degree of aridity and humidity. Handicaps of climatograms are the difficulties in representing high precipitations (which can rise up to several thousand mm per year) and the fact, that the influences of air humidity and wind are not taken into account. ‘The meteorological data normally do not include all types of precipitation, such as dew and mist which can support plant growth in regions without rain, Regions with an arid climate generally lack permanent watercourses, bear a scant vegetation and possess lakes without runoffs. The immission of salt solutions into these, even with a very low concentration, produces salines or, when all the water evaporates, salt flats. Pei NN Precipitation 10000 [DD] sereters tunte: 100 - 1000 seuthera Lisit: >150an southers lisit CD () tonger 15m |(2) tunis 3a |(3) Bengasi 240 ® ee Matruh 17.2° 895am 18,3 4150m 20.1° 268um 19.1° 158mm ostchors Sa |) tegsatse. 5200 (iQ) Bitm 35% Khartoum 380m4 mm precipitation 100 - | Fig.1. Irregular annual precipita- tions of Beni Abbés (ALG) 1947-1959, Average 58 mm Fig. 2. Temperatures during a summer day in the Haut Atlas ‘The climates of North African regions are principally influenced by three wind belts which heavily bear on the precipitation regime namely (from north to south): 1. The west wind belt bringing moist air from the Atlantic. 2. The belt of Trade winds blowing from the northeast. Their influence on precipitations is negative in a double sense: They blow towards the Equator, become warmer and their degree of relative humidity decreases because warmer air can incorporate a higher concentration of moisture. In addition to this they blow from dry continental regions where they cannot absorb humidity. 3. The intertropical convergence (ITC) which roughly coincides with the sun’s culmination. The heated air rises and induces heavy summer (.zenital*) rains. The region treated here is only marginally influenced by this belt as its southern limit touches the Tropic of Capricorn, ‘The hibernal southward shift of these three belts entails heavy consequences on the precipitation regimes creating three main types of climate. summer winter Mediterranean Trade winds - drought western winds - rain Saharan ‘Trade winds - drought ‘Trade winds - drought Sahelian ire- rain ‘Trade winds - drought ‘The climatic charts on p. 13 show these three types more or less distinctly. ‘The Mediterranean type is characterized by arid summers and humid winters. In the Saharan type precipitations are minimal throughout the year, and in the Sahelian zone farther to the south summer rains prevail. Besides latitudinal effects a general decrease of rainfall from west to east, with increasing distance to the Atlantic, is pronounced. In marginal Saharan regions where precipitation averages are still rather high, the annual amounts of rainfall are extremely irregular (fig. 1). 3.2 Microclimate »Official* climatic data are recorded in weather stations where air temperatures are registered in a height of two meters and in shade. ‘These conditions are of little meaning to small animals, as most reptiles are, moving, about in the sun, Microclimatology studies the special local conditions created by different radiation, exposure, soil type and heat transport of the ambient. Close to the ground surface temperature differences between sun and shade, night and day are extreme when compared with changes higher up in the air. Every shrub or rock offers a range of different microclimates (fig. 2). The highest and lowest temperature values are measured on sand surfaces (with low heat conductivity; fig. 15/1, 19/3) during ,,radiation weather" (no clouds, dry air, no wind). At these conditions solar radiation rises noon temperatures to 70°C while heat emission at night brings nearFig.l. Temperatures above and below the sand surface on a summer day JF 9.5 10: ry Depth surface 50 co 100 cm Fig. 2. Soil temperatures over cone year (1982) in calcareous clay. Kouf Station (Cyrenaica, LIB) 1s zero values. Measurings above and below the surface show different values for most hours of the day with temperature waves reaching only a few decimeters into the soil but several meters into the air (fig. 1). On heterogeneous ground with vegetation and large stones, sun radiation creates a mosaic of different microclimates. Temperatures differ drastically between the sunny and shaded faces of a rock. On its lower side they are markedly less extreme combined with a generally higher humidity. For a desert reptile choosing the right microclimate is not only a matter of, feeling well, but often a question of survival. On summer days vast stretches are real death traps for small lizards which die within a few seconds if they cannot find a cool spot. 3.3 Vegetation zones All amphibians and reptiles depend on plant life, even geckos living on bare rocks. As primary producers plants stand at the beginning of all terrestrial food chains, they furnish living space for climbing species (Hyla meridionalis, Tarentola neglecta, Chamaeleo chamaeleon and Agama impalearis) and shelter from excessive heat. From the multitude of environmental factors influencing plant growth we only select two important ones, namely precipitation and temperature, to demonstrate the basic distributional pattem of some important vegetation types in northern Africa. decreasing temperature decrease with increasing height humidity hot temperate cool cold Olea & Pistacia Quercus faginea humid (Portuguese oak, deciduous) on silicate soil: Quercus ilex (Holm oak) subhumid Quercus suber Cedrus atlanticus (Cork oak) Juniperus thurifera (Incense juniper) semiarid Juniperus phoenicea Lygeum spartium coastal regions (Halfa grass steppe) | Tetraclinis articulata | Gandarae) Artemisia herba-alba Pinus halepensis (Artemisia steppe) | (Aleppo pine) arid semidesert with thorn shrubs - Zizyphus lotus saharic dispersed vegetation of xerophytic trees and shrubs eg. Ephedra alata, Cupressus dupreziana, Phoenix dactylifera, Populus euphratica, Ficus salicifolia, Acacia spec., Pistacia atlantica, Tamarix spec., Olea laperrini contracted vegetation of xerophytic trees and shrubs, ephemeral annuals after sporadic rains16 100 200.km ac 2 H.de Tindouf CMe See aK Cushions ‘Thom Cf Rock Plants Wy Woodlands: gk s co de = a} [Oy a] [do | le g g 32 8 S Sz = |] fe] fo o| | Jee} | oFPlaine 4, des Daias 4 BD xp De mun Mz oF Steppes: Halfa Grass Artemisia Salines & Halophytes, Semidesert with Tademait ‘Thorn Shrubs ee eg with Argania Desert Erg r Hamada he 7] Oasis. 2-7 _|& Oued18 Depth Fig. 1. Change of soil humidity (in %) at two sites of Kour National Park (LIB) a: Kouf station, calcareous loam b: coastal sand, oviposition site of Caretta caretta As the natural plant cover was considerably altered or destroyed by pasturing and agricultural activities, the areas of the listed plants are much reduced today. Fig. 89/1 shows the shrinking of woodlands by human activities from antique times. The impact of cultivation was heaviest in the lowlands, but also high mountain forests have dwindled, and the area of the Atlantic cedar is steadily shrinking. The state of the Argan (Argania spinosa) woods in SW-MOR, surroundings of Agadir, is extremely precarious (MELLADo, 1989). This tree is a member of the tropical family Sapotaceae, and the biocenosis associated with it, especially in its lowland form, is heavily threatened by lumbering, irrigation projects, intensive pasturing and development of tourist projects. ‘The traveller from the north may also be surprised looking at what is called woodland in southern regions: Much of what has remained from the Argan forests in SW-MOR consists of small trees ten or more meters distant on bare stony soil. Low precipitations during the last years additionally enhanced forest degradation throughout this region. ‘There ate several characteristic traits identifying landscapes with arid climates: 1. Lack of autochthonous permanent watercourses (the Nile is a “foreigner"). 2. Lakes without superficial and subterraneous runofis. 3, Salt lakes and salt flats. Rain water with diluted salts collects in depressions and evaporates. For this reason salines are absent in very arid regions with almost no rain. ‘The exact methods to measure aridity are beyond the scope of this book. ‘They have to take into account all forms of precipitation (rain, mist and dew) as well as all factors influencing evaporation, as insolation, air temperature, wind and relative air humidity. ‘The scarcity of precipitation in arid regions might suggest that desert plants need almost no water. This erroneous point of view neglects the fact that they are eager water collectors. Their struggle for survival is primarily a subterraneous fight for water in which the root systems are involved. The soil volume penetrated by the roots must furnish the necessary water. With increasing aridity the evaporating plant mass visible above the surface decreases while the roots expand. Desert plants which seem to grow far apart contact in reality with their concurring root systems which may cover many square meters. Additionally the roots can grow several meters downward to tap deep water reserves. Different soil types have substantially different water economies (fig. 19.1), and sand merits special attention in this regard. Its capillary forces trap a high quantity ofrain while precipitations on rock or gravel are almost entirely lost by superficial or subterraneous runoff. Therefore the different desert types djebel (mountain slope), ham(m)ada (plane rock surface), reg or serir (gravel) and erg (sand) (fig. 19.2) house different forms of life even if they receive an equal sum of precipitation. The amount of plant matter gives some idea of the available water, In stone and rock deserts the vegetation is concentrated along the intermittent watercourses, in sand deserts along interdune valleys. Runoff or storage are absent if precipitations are minimal, but many desert19 rock pebbles sand clay (hanada) (reg) (erg) (daya, sebkha) Fig. 1. Runoff and evaporation on different desert soils Rock Plateau dilution salt water transport-> eraporaticn sebkha silt ablation clay Fig. 3. Temperature fluctuations Fig. 2. Desert landscapes and their interrelations by water and wind transport ona summer dayf Paerik ta"fouse (iauritania) 40 30 0 0 Fig, 1. Temperature extremes of a coastal and an inland (300 km) station near the Atlantic coast Fig. 2. Vegetation of Atlantic coast and western Sahara (for symbols s. also pp. 20,21) rains are accompanied by heavy thunderstorms. The runoff waters are collected in depressions and valleys. Plants growing there can dispose of the water from huge surfaces which may be many hundred times larger than their root systems. In addition to this the dry river valleys offer ground water from very distant regions, e.g. mountains with a higher precipitation rate. Under these conditions plant life is confined to a few favourable spots amidst wide stretches of bare ground. This type of vegetation is called contracted in contrast to the ,,diffuse one in less arid regions. Even the , abiotic desert can be transformed into a lush garden by a heavy shower. Seeds of plants which can perform their whole life cycle from germination to seed ripening within a few weeks may wait many years for this event. Local (Uromastyx, rodents), and wandering (antelopes) herbivores feast on this desert bloom (a: acheb). Atlantic climate - coast south of the Oued Souss (fig. 2). The vegetation of this region does not fit into the general scheme of precipitation and temperature because of peculiar climatic conditions which also bear strikingly on the herpetofauna. In contrast to the Mediterranean coast there are no mountain ridges parallel to the shoreline. Air masses can move inland, there are no abrupt changes in the landscape, and the climate slowly becomes more continental with increasing distance from the coast. [HE] coastat uccetene st Litoral seaidesert Sublitoral semidesert Lae] Cares wien Sumach [see] Piftuse shrubs: Salsolacese, Aizoon [SE] earns with Acacias TE] tect cone, 3008 high Displacement, Yow ridge21 Of special influence are the Trade winds blowing from northeast parallel to the coast, They are the driving force for the cool Canary current which causes the upwelling of cold water from ocean depths. The contact of relatively warm air and cold water evokes the formation of fog banks which hang over the coast and eventually move inland. The climatic consequences of these special constellations become evident in comparison with conditions 200 km inland (in brackets): 1, Temperatures are relatively low. 2. Temperature maxima and minima are less extreme: mean maximum 25°C, absolute maximum 40°C (38°C and 50°C resp.). 3. Smaller temperature differences between means of coldest and warmest ‘months: coast 6°; El Aioun, 20 km inland 10°; (Samara, 200 km inland, 14%). 4. Low daily temperature fluctuation: 10° (18°). 5. Fogs: 14 days per year (5 days). 6. Heat radiation about 10% less than inland. 7. High air humidity: Average 80-90% (50-60%). 8. High additional hidden precipitation in the form of fog: 50-100 mm against 75 mm rainfall. ‘The effect of these conditions on the herpetofauna are manyfold: 1. A few species are restricted to the Atlantic coast: Geckonia chazaliae and Acanthodactylus aureus. 2. Mediterranean species reach their southernmost limit here: Lacerta pater (2) and Chamaeleo chamaeleon. 3. Desert species which reach the Mediterranean coast in arid regions can only be found inland: Uromastyx acanthinura, Varanus griseus. A striking botanical speciality of the coastal strip from Cap Ghir (Rhir) down to Cap Blanc are the succulent Euphorbias (e.g. Euphorbia echinus, a: Dakhmus) which strongly remind of columnar cacti. But there are no indigenous desert cacti in Africa. The well-known Barbary figs (Opuntia) were imported from America. ‘Together with Rius tripartitum, a thomy bush related to the Sumach, Euphorbia echinus forms the graras (Va.verne, 1957) in the Seguiet el Hamra. This type of vegetation consists of concentric zones of different plants in a depression with ground water. From center to margin the availability of water decreases. The innermost part is a swamp, a meadow ora piece of arable land, then follow bushes of Rhus tripartitwm up to Sm high, a ring of low Salsolaceae, and one of Euphorbia echinus surrounded by bare gravel soil. Graras are 100 - 300m, exceptionally up to one km long and 50 - 100m broad. They may be separated by a half km of bare gravel ground, but can also fuse. They are a typical feature of the coastal plain up to 40 km inland,22 Fig. 2. Distribution patterns. a: European; b: Mediterranean; c: Saharo-Sindian; d: Ethiopian 4.1 Zoogeographic affinities 4.1.1 Zoogeographic regions Zoogeographic regions are characterized by relatively homogeneous faunae (mainly of mammals and birds) originating from long evolutive processes within ecological limits. The distribution of reptiles and amphibians does not only reflect geohistoric processes as the separation or fusion of land masses but also climatic zones that are characterized by particular herpetofaunae. ‘The denomination and delimitation of zoogeographic regions and provinces (fig. 1) is subject to dispute, and some authors consider the Saharo-Sindian (= eremial) Province as intermediary between the Palacarctic and Ethiopian Region. Members of the NAF herpetofauna show affinities to four zoogeographic provinces or parts of them: Palaearctic Region: 1, W-European and Mediteranean (fig. 2a): They are relictary, mostly restricted to mountain regions of the MAG (Bufo bufo). 2. Mediterranean (fig. 2b): Largely represented in the MAG and to minor degree in the Cyrenaica (Hemidactylus turcicus). 3. Saharo-Sindian (fig. 2c): In desert regions, penetrating into steppes and savannas (Varanus griseus). Ethiopian Region (fig. 24): Nile Valley; relictary in mountains of the Central SAH and near the Atlantic coast (Lamprophis fuliginosus). 4.1.2 Migration routes and radiations ‘The opening and closing of passages between different regions can be caused by movements of continental plates and by climatic changes which can induce massive oscillations of the sea level (by accumulation or melting of polar ice caps) and vast shifts of the vegetation cover. Migrations play a major role in evolution, entailing isolation, new selective factors, adaptive radiation in a new ambient or increased possibilities for gene recombination if previously separated populations fuse. Key events for the development of today’s NAF herpetofauna: 1. The long separation of the African and Eurasian continents by the Tethys Sea (fig. 23/1below) allowed the evolution of autochthonous African forms (Tareniola groups, Geckonia). 2. The fusion of the African and Eurasian plates (fig. 23/1 middle) was followed by an intense faunal exchange between both continents. Note that the Arabian Peninsula was originally part of the African plate, Its separation started during the Early Miocene when the formation of the East African Rift Valley System including the Red Sea began.23 Salinity crisis ~ 6 Mio years Me ie - 11 Mio years Late Mic Fig. 1. Palacogeographic evolution of the Mediterranean during the Tertiary 3. The Messinian salinity crisis (fig. 1 above) during which parts of the Mediterranean fell dry and opened a passage across the Gibraltar strait and possibly between Tunisia and Sicily. 4, The flooding of the Mediterranean Basin separated European and NAF populations again. In spite of 2 multitude of available data all conclusions on the time and direction of migrations must remain speculative and the results of Palaeontology, Palaeogeography and Molecular biology are often difficult to match. The fossil records just document faunal displacements via land bridges in the late caenozoic. Looking at features like shell morphology, an evolutionary linkage between Testudo graeca-marginata-kleinmanni may be recognized (in contrary to a Testudo graeca-Agrionemys horsfieldi line; Scuveicn (1989) and still un- published data) that possibly realized after separation by the postmessinian filling of the Mediterranean about 5.7 million years ago. Fossil records of NAF herpetofaunas are rare and most findings are diffe- rent from the recent circummediterranean fauna. There are records of Uroma- sx in Southern Italy (Esres 1983) and probably of other NAF reptiles in the Neogene of Southeastern Spain that may be explained by migrations during the Messinian, Szywo1 ax (1984) favored the concept of Europe being invaded by snakes from North Africa. He (p.138) considered at least Coronella girondica and Natrix maura to originate from local speciation during the Pliocene/Pleistocene from the common ancestors of Coronella austriaca and Natrix tesselata respectively. Rogl & Steininger (1983) provided evidence that palaeogeographically, migrations could only have been successful in the early Miocene (20-17 Mio a) via a possible land bridge from East Africa to Asia Minor and in the upper Miocene (12-11 mio a), too. There might have existed several land bridges during the Messinian evaporation of the Mediterranean, which seems to be vouched by fossil herpetofaunas in postmessinian outcrops in Spain and Southera Italy. Recent findings of the youngest fossil records of crocodiles from Southern Spain are dated as Pliocene, but still await scientific description. They either represent postmessinian invaders from Africa, most probably in more humid Pliocene climates, or they were a last offshoot of the rather ubiquitary ‘Tertiary genus Diplocynodon. During late Pliocene (2-3 mio yrs) forms of the Agama group still lived in southern France (BaiLon, 1987). A fossil amphisbaenid dissimilar to all others found in Europe may also document a postmessinian invasion from NAF into South Spain were an agamid has also been recorded (Scuteicx; still unpublished data). Earlier fossil records of NAF herpetofaunal assemblages are typical for palaeptropical coenoses like pleurodiran turtles or several distinct crocodiles. Hecir (1987) men- tions for the Pliocene Sahabi formation of EGY findings of a 2-3 m long boid snake, a Crocodile up to 3 m and another long-snouted crocodilian Euthecodon. As with Trionyx and Geochelone only African origin can be presumed. ‘The Miocene findings from Beni Mellal (MOR) provide the most detailed24 Uromastyx [pore ‘geri spec. Mali | mactadyeni ocellata AR fornata AR aegypia AR microlepis AR, SW-Asia thomasi AR ‘benti AR SW-Asian forms loricata asmussi \nardwickii Asian origin pulcheltus bedriagai ther. pistaciae Iber. sexlineatus viridianus both on Canar. and Madeira il ‘eccidentalis Canar. colosii ebucci shiarai anzai it SW-EUR., SW-Asia fevitoni AR knowledge on a fossil herpetocoenosis (Bons, 1973; Race, 1976). It shows the following spectrum: Anura (Pipidae, Discoglossidae, Bufonidae-Bujo, Ranidae-Rana, Ptychadena), Reptilia (Gekkonidae, Scincidae, ?Lacertidae, Anguidae, Amphisbaenidae, Typhlopidae or Leptotyphlopidae, Boidae- ‘°Calabaria, Eryx, Colubridae-Coluber, Elaphe, Elapidae, Viperidae). A recentmost publication (Gaura, 1993) is reporting a fossil tortoise (Testudo Aenitrensis) from the Pleistocene of Kenitra, MOR. But for this comprehensive list, other localities with fossil records from NAF are scarce, and for migratory routes it seems to be appropriate to follow palaeo-biogeographical and geological deductions regarding pos- sible pathes for invaders. Bons (1974) discussed postglacial migration and the following speciation for Hyla meridionalis from S-Europe into NAF. He also discussed the situ- ation of amphisbaenians and mentioned their absence from south of the Sahara, At that time there was no idea of the many fossil findings that were to be described during the last decade. Scxteicx (1985) reported from the mid-Miocene of S-Germany an ancestral form of Blanus, clearly differ- ent from Palacoblanus of the lower Miocene. As discussed by SzyNpLax (1984) Coronella girondica, Natrix maura and also Blanus might be te- garded as being of European origin. Szywp.ar also acknowledges only one faunal exchange, at the end of the Miocene, as a possibility for a snake fauna invasion from Africa to Iberia. This must have also been possible vice versa, and Quaternary invasions during periods of low sea level, al- though less important, are to be reconsidered. Mauremys, also reported from localities south of the Sahara, is insufficiently studied, and we have no information whether the occurrences in far southern oases are anthropogeneous or relictary. The recent distributional pattern of Mauremy's might be due to Quaternary ecological differentiations followed by climatic deteriorations and separations. In Neogene times Blanus had a wide distri- bution in Europe and one could assume an invasion from Europe into NAF. Bons (1973) provided a rather comprehensive discussion on the origin and evolution of MOR herpetofaunas. Whether the fossil Varanids of Europe (Germany, Spain) are related to NAF forms is still not understood, a situa- tion similar to that of the fossil chameleons. In Central and Eastem Europe fossil chameleons were rather common during the mid-Miocene, but for Spain first fossil records were described by TaLavera & Sancutz (1983) from the preanthropogeneous Quaternary. Bons (1973) stressed the following migrations: Testudo graeca is relictary in Europe, and after its extinction from much of the continent it invaded NAF via Spain and possibly via Italy. Fossil Testudo graeca are known from the Quaternary of MOR (Bons). He votes for three subsequent migra- tion waves: -A first one via the Betikum-Rif passage: Lacerta pater, Podarcis hispanica, Psammodromus, Natrix natrix, N. maura, Malpolon monspessulanus, Vipera latastei, Coronella girondica, Testudo graeca, Emys orbicularis, Pleurodeles, Salamandra, Discoglossus, Bufo bufo. -Another, later one via the Betikum-Rif passage: Hyla, Acanthodactylus25 Agama in wide sense (groups largely incomplete) atricollis group adramitana AR yemenensis AR SS ‘cyanogasier E-AFR Pseudotrapelus sinaitus AR Trapelus agilis mustabilis pallidus AR sanguinolentus SE-EUR, C-Asia ruderatus SW-Asia — persicus AR, SW-Asia Lournevillei suvigay] AR Agama agama (paragon ‘soinosa Ae Lark sankaranica ‘numerous other ‘African species Lavdakia chernovi SW-Asia Iehmanni SW-Asia ‘caucasica SE-EUR, SW-Asia S nupta AR ‘stellio AR, SE-EUR, SW-Asia -Phrynocephalus SE-EUR to E-Asia ‘Asian origin erythrurus, Chalcides, Psammodromus, Tarentola (however we know of much earlier radiation in the Macaronesian Tarentolas), Chamaeleo, Coluber hippocrepis, Macroprotodon -A third one via Sicily-Tunisia or Suez: Rana ridibunda, Bufo viridis, ?Alytes, ?Pelobates. Other faunal radiations might have followed ecologically-climatologically stable pathways: first, the Atlantic coast strip, mountain chains within MOR-ALG-TUN, and various Saharan regions. Lamprophis, Dasypeltis and Bitis are of tropical origin, but Naja, also included by Bons (p.94), received a different explanation by SzynpLax & Rack (1990). The genus ‘Naja probably dates back to the end of the Paleogene with its origin lying in Africa where it is still in a state of speciation. According to the latter authors the ancestral stock of Naja entered Africa before the Miocene, per- haps during the Oligocene, via India and East Africa. A similar explana- tion could be feasable for the evolution of the genus Chamaeleo s.\. ‘The Egyptian black cobra Walterinnesia is regarded as an early offshoot of the genus Naja, Szynptak (1988) regards the endemic occurrence of Vipera mauritanica in NAF, far away from the Near East distribution center with the close akin Vipera lebetina, as proof for that common ancestors had a continuous ran- ge in the past. This would explain a former NAF invasion to Iberia as deduced from fossil records of a past Vipera "russelli” complex on the peninsula, Pleistocene sea level changes might also have favoured some herpetofaunal elements to cross narrowed sea passages by drifting. Recent migrations along irrigation channels may result in a mixing of genetically separated groups (phentotypically different and geographically long separated populations) e.g. in the case of terrapins and amphibians. Main hypothesis on the origin of the North African Herpetofauna: 1. The stocks of the African Agama, Uromastyx, Varanus and Echis groups immigrated into NAF from W-Asia during the Miocene, 2. The Naja stock originated in India and perhaps reached AFR via Mada- gascar, 3. Radiation on the African plate, mainly in its north, took place in the following groups: Chalcides, Tarentola, Ptyodactylus, Acanthodactylus, Mesalina, Psammodromus, the AFR stocks of Agamids, Chamaeleo, Varanus, Naja, Bitis, Echis, (subgen. Toxicoa CxERUN, 1990) and the Eumeces schneiderii subgroup. ‘The marginal drawings visualizing radiations (p, 24-26) are highly hypo- thetical and based on biochemical and morphological evidences. The start of African radiations is marked with a dark dot. African forms appear in bold print; NAF ones are additionally underlined. Species of the Arabian Peninsula are marked with AR.26 Mesalina olver! AR a suttulata AR watsoniana SW-Asia pastewri Vp subropunctate breviretris AR, SW-Asia Z adramitana AR = 10 species inS-AFR Varanus griseus AR, SW-Asia |yemenensis Ar albigularis iti Indo-Asian and Australian radiations ca. 35 species Asian origin Echis arenicola paamidum ocellats Jogert varia AR (— w w esau Spapraeean 34,5 (32), 4,5 graaare 4 oe ae aay oePhilochortus zolii (genus with Ethiopian affinities) see a. Dasypeltis scabra 1 Lamprophis fuliginosus 1 Bitis arietans 1 44 Altitudinal distribution With increasing height we find a decrease in thermophilous species, with N-MED (Coronella girondica) ot even palaearctic (Bufo bufo) elements remaining, besides endemites such as Quedenfeldtia trachyblepharus or Lacerta andreanskyi. The reproductive and thermal adaptations of reptiles extreme altitudes of the Haut Atlas is unknown, their study would certainly be very promising. 4.4.1 Moyen Atlas (p. 33) 44.2 Haut Atlas (p. 34-35) 4.5 Transsects (pp. 36-51) We tried to present a synoptic view of the species over a limited number of transsects, crudely in N-S direction from the Atlantic or MED to the SAH. The amount of precipitation is represented in relation to the 100mm level which marks the desert limit and in many cases the distribution Mediterranean/Saharan species. Algeria Morocco se Middle, cag, West East West Libya East Fig. 1. Position of transsects,32 ‘Symbols for vegetation types on transsects: Woodlands: Steppes: 2002 Oaks awe Malfa Crass 2484 Cedar uuu Artemisia ‘2098 Pine Semidesert, L20@ Olive & Pistachio secex Thora shrubs oon Sandarac & Junipers Sand Desert (Erg) High Mountains: aa Thorn Cushions susan Salines & Halophytes see Rock Plants At Oases Signs for species distrib x records of widely distributed species R records for relictary species general area of rather frequent species general area of rare species In the general areas the species occur of course only in adequate biotopes, which can be very scattered (e.g. in deserts). Direction of Transsects (fig. 31/1) Morocco- West Atlantic - western Haut Atlas - Oued Draa (p. 36-37) Morocco - Middle Mediterranean - central Haut Atlas - Tafilalt (p. 38-40) Morocco- East Mediterranean - Hauts Plateaux - Erg Occidental (p. 41-43) Algeria - West Mediterranean - Hauts Plateaux - Sahara Atlas: (p. 44-46) Algeria - East Mediterranean - Hauts Plateaux - Chott Djerid (p. 47-49) Libya - East Mediterranean - Cyrenaica - Libyan desert (p. 50-51) Note the marked regression in species number from west 10 east.33 Moyen Atlas - altitudinal distribution Elevation in meters 1000 1500 Vegetation: Evergreen broad-leaved woods chiefly Quercus ilex with enclaves of Quercus faginea, 2000 2500 3000 originally Cedrus atlantica wood, degraded parts with Quercus ilex and Juniperus oxycedrus Quercus suber and Pinus pinaster Reptiles and Amphibians: Podarcis hispanica vaucheri- Natrix natrix~. ~Chalcides chalcides- Coronella girondica Discoglossus pictus: Vipera latastei? Psanmodromus algirus: Lacerta pater- - Rana saharica- Bufo viridis-- Hyla meridionali -Chalcides lanzai- Chalcides polylepis? Acanthodactylus erythrurus-- Bufo mauritanicus- Podarcis perspicillata Trogonophis wiegmanni- Agana impalearis~ Bufo bufo Malpolon monspessulanus~ Natrix maura-~ Ophisaurus koellikeri Tarentola mauritanice- Nauremys leprosa Emys orbiculari. Salamandra algira Alytes obstetricans: Macroprotodon cucullatus-- Blanus mettetali-- Psanmodromus microdactylus x Pleurodeles waltl~ Because of rare records the occurrence of Vipera latastei and/or Vipera monticola remains doubtful. The case is similar with Chalcides lanzai and Chalcides polylepis which can only be distinguished by exact scale counts.34 4000m 3500m 3000m 2500m 2000m 1500m 1000m Haut Atlas - climate and vegetation (data from Saint Girons, 1953) temperature average yr. Jan. July } Max. Min.) Snow cover +3° -3° «13° 22° 20° 6-7 months 7° 0.5° 17.5°] 25.5° 1-2 months 11.574,.5° 22° | 39,5° -11°| 2-3 weeks 15.5° 9° 27° some hours Vegetation very scant, chiefly in debris and rock crevices thorn cushion: Peilotrichum(Alyssum) spinosum Bupleurun spinosum (Thorny Hare's Ear) Cytisus balancac Scant bushes Juniperus thurifera (Incense Juniper) Ormenis scariosa Anacarpus anagyrifolia Genista florida Yerophilous conifers: Tetraclinis articulata (Sandarac tree) Juniperus phoenicea (Phoenician Juniper) poplar, walnut, olive wheat, corn, meadows35 ~---snue[nssedsuow uopodTey == SNINIYIA IA snqAqoepoyzueay ----STieaTeduy euedy ---U0aTaeUeYI oaTaeuEY) -PITUPITINEW Pad TAOIIBY T49{T]T90q snanestydg ~----StdeAyod sapratey) ~stder20ddyy saqnqop ---POTIBYES PUBY sanneaeo=-SIPLITA 070g oen=-BINEW XTIIEH -SeptoTeyo sapraqey) sniq8[@ snworpowmesy -POTUeITINEW BTOIUAIE] -SnuequoW saproTey)---—~ ee] TTIIds1ad sToTepog mennnemJaqed 211998] eennn snTue]yinew 0jng a-------snjoqd snssoj800s1q -ST[euorpriew eyAy ----BT0ITIUON BIAdT A OTpuoITs BT TauoIO)-——— ween eee ee Tuayonea eatuedsty sto1epog-—~ TAysuearpue e7.10007 -----. --snreydayqhyoes9 BT apTazuapany-——-—----- wOQSE MQD0E wons? m000c wOOST WOOT36 Morocco West SALAMANDRIDAE Pleurodeles walt! DISCOGLOSSIDAE Discoglossus pictus BUFONIDAE Bufo bufo Bufo mauritanicus Bufo viridis HYLIDAE Hyla meridionalis RANIDAE Rana saharica EMYDIDAE Mauremys leprosa ‘TESTUDINIDAE, Testudo graeca GEKKONIDAE Ptyodactylus oudrit Quedenfeldtia trachyblepharus Saurodactylus (m.) brosseti Stenodactylus pen Stenodactylus sthenodactylus Tarentola annularis Tarentola boehmei Tarentola ephippiata Tarentola mauritanica Tropiocolotes tripolitanus AGAMIDAE Agama impalearis Trapelus mutabilis Uromastyx acanthinura CHAMAELEONIDAE Chamaeleo chamaeleon SCINCIDAE, Chalcides chalcides Chalcides mionecton Chalcides montanus Chalcides ocellatus Chalcides polylepis Eumeces algeriensis Scincus scincus Sphenops boulengeriLACERTIDAE Acanthodactylus boskianus ‘Acanthodactylus erythrurus Acanthodactylus maculatus Acanthodactylus scutellatus Lacerta andreanskyi Lacerta pater Mesalina guttulata Mesalina olivieri ‘Mesalina rubropunctata Mesalina simoni Podarcis hispanica Podarcis perspicillata Psammodromus algirus Psa. microdactylus Ophisaurus koellikeri VARANIDAE Varanus griseus AMPHISBAENIDAE Blanus mettetali Trogonophis wiegmanni LEPTOTYPHLOPIDAE Leptoyphlops macrorhynchus COLUBRIDAE Coluber algirus Coluber hippocrepis Coronella girondica Dasypeltis scabra Lamprophis fuliginosus Lytorhynchus diadema Macroprotodon cucullatus Malpolon moilensis Malpolon monspessulanus Natrix maura Psammophis schokari Spalerosophis diadema Spalerosophis dolichospilus ELAPIDAE Naja haje VIPERIDAE Cerastes cerastes Macrovipera mauritanica Vipera monticola 3738 SALAMANDRIDAE Pleurodeles walt! Salamandra algira DISCOGLOSSIDAE Alytes obstetricans Discoglossus pictus BUFONIDAE, Bufo brongersmai Bufo bujo Bufo mauritanicus Bufo viridis HYIIDAE Hyla meridionalis RANIDAE Rana saharica EMYDIDAE Emys orbicularis Mauremys leprosa ‘TESTUDINIDAE Testudo graeca GEKKONIDAE Hemidactylus turcicus Pryodactylus oudrii Saurodactylus fasciatus Stenodactylus petrii Stenodactylus sthenodactylus Tarentola deserti Tarentola mauritanica Tropiocolotes tripolitanus AGAMIDAE Agama impalearis Trapelus mutabilis Uromastyx acanthinura (CHAMAE LEONIDAE Chamaeleo chamaeleon 100 Morocco MiddleSCINCIDAE Chaicides chalcides Chalcides colosii Chalcides ebneri Chaicides lanzai Chalcides montanus Chalcides polylepis Eumeces algeriensis Scincus scincus Sphenops boulengeri LACERTIDAE Acanthodactylus boskianus Acanthodactylus erythrurus Acanthodactylus maculatus Acanthodactylus scutellatus Lacerta andreanskyi Lacerta pater Mesalina guttulata Mesalina olivieri Mesalina rubropunctata Podarcis hispanica Podarcis perspicillata Psammodromus algirus Psammodromus microdactylus ANGUIDAE Ophisaurus koellikeri VARANIDAE Varanus griseus AMPHISBAENIDAE Blanus tingitanus Blanus mettetali Trogonophis wiegmanni 39 ove Morocco Middle XXXXXXAXXNAXAAXY : : : Po XeX-x-40 COLUBRIDAE Coluber algirus Coluber hippocrepis Coronella girondica Lytorhynchus diadema Macroprotodon cucullatus Malpoton moilensis Malpolon monspessulanus Natrix maura Natrix natrix Psammophis schokari Spalerosophis diadema ‘Spalerosophis dolichospilus Telescopus obtusus ELAPIDAE Naja haje VIPERIDAE Cerastes cerasies Cerastes vipera Macrovipera mauritanica Vipera latastei Precipitation tsa tH00 Morocco Middle Djs Ayaana Morocco East ‘Seciberra, DISCOGLOSSIDAE Discoglossus pictus BUFONIDAE Bufo brongersmai Bufo mauritanicus Bufo viridis HYLIDAE Hyla meridionalis RANIDAE Rana saharica EMYDIDAE Mauremys leprosa ‘TESTUDINIDAE Testudo graeca GEKKONIDAE Hemidactylus turcicus Pryodactylus oudri Saurodactylus (m) mauritanicus : : Stenodactylus petrit 3 : : : x Stenodactylus sthenodactylus : Tarentola deserti Tarentola ephippiata Tarentola mauritanica Tarentola neglecta Tropiocolotes tripolitanus AGAMIDAE Agama impalearis Trapelus mutabilis Trapelus tournevillei 3 : : Uromastyx acanthinura EX=XX MX = XXX CHAMAELEONIDAE Chamaeleo chamacleon2 ec SCINCIDAE Chalcides chalcides Chaicides mauritanicus Chaicides ocellatus Chaicides ghiarai Eumeces algeriensis Eumeces meridionalis Scincus scincus Sphenops boulengeri LACERTIDAE Acanthodactylus boskianus Acanthodactylus erythrurus Acanthodactylus maculatus Acanthodactylus seutellatus Lacerta pater Mesalina guttulata Mesalina oliviert Mesalina pasteuri ‘Mesalina rubropunctata Ophisops occidentalis Podarcis hispanica Podarcis perspicillata Psammodromus algirus Psammodromus blanci VARANIDAE Varanus griseus ‘TROGONOPHIDAE Trogonophis wiegmanniBOIDAE Eryx jaculus COLUBRIDAE Coluber algirus Coluber hippocrepis Coronella girondica Lytorhynchus diadema Macroprotodon cucullatus Malpolon moilensis Mapolon monspessulanus Natrix maura Psammophis schokari Psammophis sibilans Spalerosophis diadema Spalerosophis dolichospilus ELAPIDAE Naja haje VIPERIDAE Cerastes cerastes Cerastes vipera Echis arenicola Macrovipera (lebetina) transmed. Macrovipera mauritanica Ireanteaion ta Algeria WestSALAMANDRIDAE Pleurodeles poireti Salamandra algira DISCOGLOSSIDAE Discoglossus pictus BUFONIDAE Bufo bufo Bufo mauritanicus Bufo viridis HYLIDAE Hyla meridionalis RANIDAE Rana saharica EMYDIDAE Enmys orbicularis Mauemys leprosa TESTUDINIDAE Testudo graeca GEKKONIDAE Hemidactylus turcicus Stenodactylus petrii Stenodactylus sthenodactylus Tarentola deserti Tarentola mauritanica Tarentola neglecta Tropiocolotes tripolitanus AGAMIDAE Trapelus mutabilis Trapelus tournevillei Uromastyx acanthinura 7 Dj.0t oir Andaba Oued Hedjerda XX=2-2-48 sa Algeria East "bso ‘ane ta Sane theta hace Byers * v CHAMAELEONIDAE Chamaeleo chamaeleon SCINCIDAE Chalcides chalcides Chalcides occellatus Eumeces schneiderii Scincopus fasciatus Scincus scincus Sphenops boulengeri LACERTIDAE Acanthodactylus boskianus Acanthodactylus erythrurus Acanthodaciylus maculatus Acanthodaciylus scutellatus Lacerta pater Mesalina guttulata Mesalina olivieri Ophisops occidentalis Podarcis hispanica Psammodromus algirus Psammodromus blanci VARANIDAE Varanus griseus AMPHISBAENIDAE Trogonophis wiegmanni49 BOIDAE Eryx jaculus COLUBRIDAE Coluber algirus Coluber hippocrepis Lytorhynchus diadema Macroprotodon cucullatus Malpolon moilensis Malpolon monspessulanus Natrix maura Natrix natrix Psammophis schokari Psammophis sibilans Spalerosophis diadema Spalerosophis dolichospilus Telescopus obscurus ELAPIDAE Naja haje VIPERIDAE Cerastes cerastes Cerastes vipera Echis arenicola Macrov. deserti & maurit. Vipera latastei Oued Made Algeria East ‘cuisine ontexrut50 BUFONIDAE Bufo viridis RANIDAE Rana saharica TESTUDINIDAE Testudo (Pst,) kleinmanni Testudo graeca GEKKONIDAE Hemidactylus turcicus Stenodactylus petrii Stenod. sthenodactylus Tarentola mauritanica Tropiocolotes tripolitanus AGAMIDAE Trapelus mutabilis Uromastyx acanthinura CHAMAELEONIDAE Chamaeleo chamaeleon SCINCIDAE Chalcides ocellatus Eumeces schneiderii Scincus scincus Sphenops sepsoides LACERTIDAE Acanthodactylus boskianus Acanthodactylus pardalis Acanthodactylus scutellatus Mesalina guttulata Mesalina olivieri Mesalina rubropunctata Ophisops elegans VARANIDAE Varanus griseus Tose. Libya East i Xxx, XXX HX xxx,SI BOIDAE Enyx jaculus COLUBRIDAE Coluber algirus Coluber rogersi Lytorhynchus diadema Macroprotodon cucullatus Malpolon moilensis Malpolon monspessulanus Psammophis schokari Spalerosophis diadema ELAPIDAE Naja haje VIPERIDAE Cerastes cerastes Cerastes vipera Echis pyramidum ake —<—52 Arch, Galita: aliisne guslia fe Lieveccea See tress Herts 4.6. Islands off the North African coast References: Boutencer (1891); Lanza & Bruzzons (1959); Mateo (1990); Satvapor (1982). No data were available for islands mentioned in the map, but not in the following list. Archipelago Island surface distance species (dimension from or diameter) continent 1 Alhucemas Isla Tierra 0.3 ha 65m Tarentola mauritanica Isla Mar 0.2 ha 200m Tarentola mauritanica Pefion Alhucemas 1.5 ha 700 m_ Hemidactylus turcicus Podarcis hispanica Saurodact. mauritanicus Tarentola mauritanica "Chaicides ocellatus” Trogonophis wiegmanni Coluber hippocrepis 2 Chafarinas, The recently described Chalcides ghiarai lives on Congreso and Rey. It is doubtful that Chalcides ocellatus really occurs on the archipelago Congreso 22ha 4,5 km Tarentola mauritanica *Chalcides ocellatus" Podarcis hispanica Trogonophis wiegmanni Isabel I 1,3 ha 45 km Tarentola mauritanica "Chalcides ocellatus” Podarcis hispanica Trogonophis wiegmanni Rey 12ha 45 km Tarentola mauritanica Chalcides ocellatus Podarcis hispanica Trogonophis wiegmanni 3 Rachgoun 2 km Trogonophis wiegmanni 4 Habibas 7 km Macroproted. cucullatus 5S Galita Galitone 750 m 36 km Phyllodactyl. europaeus Chalcides ocellatus Psammodromus algirus53 Aguglia 500 m 36 km Phyliodact. europaeus Chalcides ocellatus Psammodromus algirus Galita 5.2x28km — 38 km Discoglossus pictus Testudo graeca Hemidactylus turcicus Phyllodactyl. europaeus Chalcides ocellatus Lacerta pater Macroprotod. cucullatus Natric maura Gallina 300m 39 km Chalcides ocellatus Gallo 450m 40 km Chalcides ocellatus 6 Cani 11 km Phyliodactyl. europaeus Chalcides ocellatus 7 Kuriate 17 km Caretta caretta nesting 8 Kerkenna Hemidactylus turcicus (Gharbi 15x 7km20 km Stenodactylus sthenod. Chergui 42.x8km26 km) Mesalina oliviert Coluber algirus 9 Djerba 514 km? 1.9 km Stenodactylus sthenod. Chalcides ocellatus Acanthodact. boskianus Mesalina guttulata Coluber algirus 10 Purple Islands off Essaouira (MOR, Atlantic coast) In 1933 H. Hepicer visited these islands (Hepicer, 1935) and found the following species: Pleurodeles wall, Testudo graeca, Tarentola mauritanica, Chaleides mionecton, Acanthodactylus erythrurus lineomaculatus. 4.7 Dubious records Bufo regularis - Ghat, SW-LIB: Scoxrecct (1935) states the presence of this toad in the Ghat region. ,,Si tratta senza alcun dubbio di Bufo regularis* while Jocer (1981) supposes that Scorrecc1 found Bufo xeros, which was unknown at that time and described much later (Tanpy & al. 1979). Its continuous southern distribution range is much closer than that of Bufo regularis. Prychadena mascareniensis. The presence of this frog was mentioned by Pritam (1911) for the pond of Ifedil (Mare d’fedil, Tassili Mts. close to54 the famous crocodile habitat in the Ouadi Mihero), but could not be reconfirmed (Rousset to Le Berne, 1984). Agama agama does not occur in the Tessili (SE-ALG). Old records from is mountain area are obviously based on wrongly determined Agama impalearis. The northernmost records forA gama agama are from Mauritania (Atar) and Niger (JoceR, 1981). Mabuya quinquetaeniata was recorded for the Cyrenaica (NE LIB) by CConporeLt FRANCAVIOLIA (1886) under the name ,.£uprepes savignii D.B." based on a collection of Prof. Paxcext from ,Egitto ¢ regioni vicine“. As there are no recent records of M. quinquetaeniata for LIB it may be that the specimens refered to by ConporeLi FRANcAvIGUA were from EGY. Sphenops delislei and Sphenops sphenopsiformis: The records of both species from the Hoggar mountains and the W-SAH are inexact and need to be reconfirmed. Anguis fragilis was believed to occur in ALG ,rare et localisé“, e.g. in the surroundings of Bone and Aumale (Seurar, 1930), and Perzotp (1971) stated without any further comment: ,,In Nordwestafrika bewohnt die Blindschlei- che einige begrenzte Gebiete Algeriens.“ None of these records was ever confirmed and we suppose that Chalcides chaleides was mistaken for this species. Walterinnesia aegyptia was recorded from the Cyrenaica (NW-LIB) by Ruwotta (1923) as frequent“. Geixomt (1926) and Francuint (1929) repeated this assertion uncritically, but there is no proof that this snake was ever found in LIB. 4.8 Survey of regional species numbers ‘The following compilation is based on our more , splitting concept, counting, Satvaor’s Acanthodactylus species as well as the mostly Moroccan Chalcides endemites which were formerly regarded as subspecies. Especially critical are dubious forms like Rana saharica, Bufo viridis, the Chalcides ocellatus group, Podarcis hispanica, and the genera Echis and Macrovipera. MOR ALG TUN LIB AMPHIBIA CAUDATA Salamandridae Pleurodeles 1 2 1 - Salamandra 1 1 - - ANURA, Discoglossidae —Alytes: 1 - -5S MOR ALG TUN LIB Discoglossus 1 an Pelobatidae Pelobates 1 - os Bufonidae Bufo 44 1 Hylidae Hyla 1 toa Ranidae Rana 1 144 Dicroglossus 1 CAUDATA 2 3 10 ANURA 9 7 6 3 AMPHIBIA total nu 0 7 3 REPTILIA ‘TESTUDINES Emydidae Emys 1 nr Mauremys 1 144 Testudinidae Testudo 1 1 1 2 TESTUDINES total 300303 SAURIA Gekkonidae Geckonia 1 - ose Hemidactylus 1 111 Phyllodaciylus - 1 - Pryodactylus 1 11 Quedenfeldtia 1 - oe Saurodactylus 3 1. Stenodactylus 2 2 2 2 Tarentola 5 5 3 4 Tropiocolotes 1 201 3 Agamidae Agama 1 toe Pseudotrapelus. - 2 1 Trapelus 1 202 1 Uromastyx 1 2 1 1 Chamacleonidae — Chamaeleon 1 1 14 Scincidae Chaleides mo 5 2 2 Eumeces 2 3 411 Mabuya - 144 Scincopus : 14. Scincus 1 1 414 Sphenops 2 11 ot Lacertidae Acanthodactylus 8 9 4 5 Lacerta 2 14 - Mesalina 4 4 33 Ophisops 1 2 410156 Philochortus - - ed Podarcis 2 201 - Psammodromus 3 202 ~- Anguidae Ophisaurus 1 se Varanidae Varanus 1 144 Amphisbaenidae —_Trogonophis 1 tobe Blanus 2 - ee SAURIA total 58 533532 SERPENTES Leptotyphlopidae Leptotyphlops 1 14 ot Boidac Enx 1 road Colubridae Coluber 2 30023 Coronella 1 Dasypeltis 1 se Lamprophis 1 - ee Lytorhynchus 1 1414 Macroprotodon 1 »o41o4 Malpolon 2 2 2 2 Natrix 2 2 21 Psammophis 2 2.02 3 Spalerosophis 2 20201 Telescopus 1 »o41o4 Elapidac Naja 1 » 1 ot Viperidae Bitis 1 ee Cerastes 2 2 2 2 Echis - 1 41 2 Macrovipera 2 300301 Vipera 2 14 - SERPENTES total 2 SH REPTILIA total 87 Bl 56 5 Ecological aspects 5.1 Biotopes Data mainly from Bows (1959), Doumexave (1901), Gauriner (1966), Grenor (1976), Lawpert (1984), MELLADO & OLMEDO (1991), SAINT-GiRONS (1953), ‘Scteicn (1987), StemMLeR (many articles, see literature), StemMLer & Horz (1972), Verwer & Grenor (1972), Wenner, Y. (1982).S7 Symbols used for 5 Ecologic “further ecological data": 5.1 Biotopes terrestrial aquatic The last two columns contain further ecological data: Relation to substrate semiaquatic and diel activity. fossorial subfossorial limbit id j climbing an mediterraneansot.steppes Sahara climbing diurnal nocturnal (and crepuscular) nocturnal with occasional diurnal activities fn nocturnal with regular diurnal activities & diurnal and nocturnal activities depending on weather /life phase / activity type spects arte 2 2700 fields rocks and walls coastal sand oasis grara scrub and thornbush daya succulent steppe erg low vegetation reg bushes epen ground, standing water running water thorn cushions rocks Artemisia Halfa grass hamada dry oued jebel guelta Q o o ba w SALAMANDRIDAE Pleurodeles poiret Pleurodeles walt. Satamandra algiré DISCOGLOSSIDAE. Alytes obstetricans PELOBATIDAE : Pelobates varaldit.. Ix: BUFONIDAE : Bufo brongersmai Bufo bufo .. Bufo mauritanicus Bufo viridis HYLIDAE Hyla meridionalis RANIDAE, Dicroglossus occipitalis Rana saharica. EMYDIDAE Emys orbicularis... Mauremys leprosa.. ‘TESTUDINIDAE Testudo graeca Testudo kleinmai GEKKONIDAE Geckonia chazaliae . Hemidactylus turcicus... Phyllodactylus europacu Ptyodactylus oudrii. Pryodactylus ragazzii.. SS ZRZ SSbSS RR RF RRRR 2 2 RRR58 Quedenfeldtia trachyblepharus| Sidbeyls (re) Broce Saurodactylus fasciatus .. Tarentola annularis. Tarentola boehmei. Tarentola deserti Tarentola ephippiata Tarentola mauritanica. Tarentola neglecta Tropiocolotes nattereri. Tropiocolotes steudner Tropiocolotes tripolitanus . AGAMIDAE Trapelus mutabilis 0.0.0. Trapelus tournevillei Uromastyx acanthinure Chamaeleo chamaeleon.. SCINCIDAE Chalcides chalcides Chalcides mauritanicus. Chalcides mionecton Chalcides ocellatus. Chalcides colosii... Chalcides ebneri Chalcides. ghiar Chalcides. foncat evens Chalcides. montanus.. Mabuya vittata Scincopus fasciatus, Scincus scincus... Sphenops spec .(Agadirt).. LACERTIDAE Acanthodactylus boskianus Acanthodactylus erythrurus Acanthodactylus blanci ‘Acanthodactylus savignyi Acanthodactylus pardalis. Acanthodactylus spinicauda .. Acanthodactylus scutellatus .. Acanthodactylus aureus... Lacerta andreanskyi Lacerta pater . Mesalina guttulata. ‘Mesalina olivierMesalina pasteuri Mesalina rubropunctata Mesalina simont Ophisops elegans. Ophisops occidentali Philochortus zol Podarcis hispanica. Podarcis perspicillata Psammodromus algiru Psammodromus blanci Psammodromus microdactylus| AMPHISBAENIDAE Blanus mettetali. Blanus tingitanus Trogonophis wiegmanni.. ANGUIDAE, Ophisaurus koellikeri.. VARANIDAE, Varanus griseus BOIDAE Eryx jaculus LEPTOTYPHLOPIDAE Lepto. macrorhynchus COLUBRIDAE pid Coluber algirus. xX Coluber hippocrepis : Coluber rhodorachis Coluber rogersi.. Coronella girondica Dasypeliis scabra . Lamprophis fuliginosus Lytorhynchus diadema.. ‘Macroprotodon cucullatus Malpoton moilensis : Malpolon monspessulanus . Natrix maura . Natrix natris Psammophis aegyptius Psammophis schokari.. Psammophis sibilans Spalerosophis diadema. Spalerosophis dolichospilus.. Telescopus obtusu: . ELAPIDAE, Naja haje VIPERIDAE, Bitis arietans. Cerastes ceraste: Cerastes vipera. Echis pyramid Macrovipera deserti Macrovipera mauritanica Vipera latastei . Vipera monticol TREE SSSeS SSS S5 BeEE5.2 Relations to the ambient 5.2.1 ‘Thermal biology 5.2.1.1 Thermal ecology The standardisized terms used here are defined according to Poust & Gans (1982) and are especially selected for dealing with amphibians and reptiles. These animals are ectotherms as they chiefly rely on exterior sources of heat energy. They are capable of thermoregulation (temperature regulation), chiefly by behavioural means. Heliotherms bask in the sun, thigmotherms gain heat from warm surfaces, but many of them apply both methods at the ‘same time or alternately. Thermoconformers have body temperatures which change with the ambient temperature. ‘The important body temperature set points are symmetrically arranged above and below activity temperature. In brackets we add the numeric values for Uromastyx acanthinura °C (after GreNor, 1976), one of the few species for which exact data are available. lethal maximum (49,3°)__a short exposure causes irreversible physiological damage critical maximum (48°) __causes heat narcosis, bringing the animal eventually into lethal danger voluntary maximum (46°) active animals withdraw into shelter activity temperature range body temperature at which an animal shows (30-44°) its routine activities mean activity temperature arithmetical mean of the body temperatures (38-3959) of active animals selected body temperature body temperature maintained in a laboratory G9-41°) temperature gradient basking temperature range releases activities which increase the body temperature to the activity temperature ran- ge voluntary minimum (21°) active animals withdraw into shelter critical minimum (13.5*) causes cold narcosis, bringing the animal eventually into lethal danger lethal minimum a short exposure causes irreversible physiological damage. There are two ways to study temperature preferences of ectotherms: Measuring body temperatures of active animals in the field, orin alaboratory temperature gradient. Preferred temperatures which guarantee optimal performance are a rarity in the natural ambient. Ectotherms cannot always stay at such spots, even if they exist, and often thermoregulate by shuttling between heat sources and heat sinks. In many species a rich behavioural repertoire allows to maintain body temperatures within a few degrees.61 Others, the thermoconformers, which mostly stay in the shade, save the energy and risks involved with steady locomotion. They stay on the spot and tolerate the oscillation of their body temperature with ambient conditions. Exapmples are found in many toads ot geckos (Saurodactylus, ‘Stenodactylus sthenodactylus, Geckonia chazaliae) which hide under stones. ‘Thermoregulators are able to rise and lower their body temperatures chiefly by posturing and locomotion, Heat gain from the ambient is effected in two principal ways: Heliotherms bask in the sun, absorbing radiating hi thigmotherms seek contact with warm objects, absorbing heat by conduction. Many baskers are able to flatten their bodies, most of them dorsoventrally (all Lacertidae and Agamidae, the gekkonid genera Tarentola, Ptyodactylus, Quedenfeldtia) or laterally (Chamaeleonidae). Species capable of colour change become dark. Snakes, Scincidae and serpentiform lizards such as Ophisaurus koellikeri, simply expose themselves to the sun rays at favourable locations. They are able to regulate heat absorption by exposing, different lenghts of their body . In flattened forms the same effect is attained by tilting the body in relation to sun rays. With rising temperatures heliotherms move into half or full shade. Thigmotherms can absorb heat with the entire body surface (in digging forms like Blanus or Trogonophis), with their backs (species resting under stones like Blanus, Trogonophis, Saurodactylus, Stenodactylus, Tropio- colotes) or their venters. The latter method is often used by heliotherms when they bask on sand, stones or logs. Different substrate temperatures (dark substrates gain more heat than bright surfaces) release different postures during thigmothermal behaviour (see Uromastyx acanthinura). In biotopes with a mosaic of microclimates, active reptiles move around to remain within their activity temperature range. The most vital problem in this regard is to escape lethal heat, which means to keep away from the irradiated soil surface. This is effected by three ways: moving into shade (most species), digging into the ground (Scincus scincus, Sphenops) and climbing on rocksor plants (Agama, Trapelus, Pseudotrapelus). Large forms like the latter even gain some advantage by lifting the body off the substrate in the elevated posture (fig. 291/1). Ultimate methods to lower body temperatures are the elevation of the respiratory rate and, in lizards, panting. Both methods are very risky for desert reptiles because of a considerable loss of water. Retiring to a subterraneous shelter has the double advantage of lower temperature and higher humidity, but excludes the performance of vital activities such as feeding, territorial behaviour and mating. (One strategy for minimizing predation by ectotherms isto avoid their activity temperature range. Desert ants of the genus Cataglyphis are an important prey item for the Fringe-toed lizard Acanthodactylus dumerili, As field studies by Weaver (1992) showed, the ants are active in a ,heat window" between 46.5 and 53.6°C to scavenge for dead insects, mostly heat victims, on the scorching sand, Despite the fact that the Fringe-toed lizard’s voluntary maximum is 46°, behavioural thermoregulation permits them to remain active at much higher temperatures of exposed soil and catch single ants.62 ‘On the other hand this strategy reduces the danger of being caught by returning to their subterraneous nests on a direct way. 5.2.1.2 Temperature-induced sex determination (Ravwau & Preau, 1985, Roser, 1986) In many amphibians and reptiles, besides sex chromosomes (genotypic sex), ambient temperature, probably in the first three weeks of incubation, plays an important role for the differentiation of the genital system in a developing ‘embryo (phenotypic sex). In amphibians this mechanism is known e.g. from Pleurodeles (fig. 95/2) and Bufo bufo. In reptiles at least 4 different cases can be distiguished: 1, Almost no apparent temperature influence. 2. Low temperatures induce more (or only) males, high temperatures more (or only) females. 3. The inverse. 4. High and low temperature induce more females, intermediate ones more males. To make things still more complicated these mechanisms are not linked to certain families, but occur across different groups. Apparently they have not been discovered in snakes. In the gecko Eublepharis macularius (mechanism 4), the ,coldfemales react normally, but the ,,hot“ones react like males and are very aggressive. They produce a higher rate of androgens and a lower rate of estrogens than normal females. As far as N-AFR reptiles are concerned, temperature-induced sex determination (type 2) was observed in: Turtles: Emys orbicularis (fig. 144/1), Testudo graeca, Chelonia mydas, Caretta caretia Lacertilia: Chamaeleo chamaeleon. This phenomenon still awaits discovery among the geckos and agamids of this region. 5.2.2 Water regime (Duxson, 1976, Bextiey, 1976) Water loss through respiration, evaporation via skin, defecation and exeretion has to be compensated by water intake with food and water production during the catabolism of organic matter. Some species can survive 4 considerable water loss during dry periods. ‘The total water content of a well-fed Uromastyx acanthinura is 75.4%, the extracellular water weighs 33% of the lizard. During times of water deficiency, only extracellular reserves are reduced while cellular water is kept at its normal level. Respiration causes a considerable percentage (57% in Avicenna’s viper,Fig. 1. Nasal gland (black) of Uromaswy acanthinura Cerastes cerastes) of the total water loss, and there are at least two methods to reduce this amount: 1. Relatively moist and coo! refuges help to diminish the respiratory loss as the air contained in them possesses a high relative humidity and cuts down ‘evaporation from skin and lung surfaces. 2. In warm surroundings the rate of oxygen extraction from the air rises (in Cerastes cerastes from 11% at 25° to 15% at 35°). For this reason the respiratory rate can be somewhat reduced and the higher evaporation at elevated temperatures is thus compensated to a certain extent. Nasal salt glands Desert plants have generally high osmotic values, which facilitate water extraction from the soil and_help to diminish evaporative loss. Herbivorous desert lizards inevitably ingest a considerable amount of salts, above all potassium compounds, which threaten to disturb the animal’s osm: balance. Many lizard families (Agamidae, Scincidae, Lacertidse, Varanidae, but not Gekkonidae and Anguidae), some turtle families (Emydidae, Cheloniidae), and also snakes (Hydrophiidae, Acrochordidae), possess na- sal salt glands which secrete a concentrated solution of salts (potassium, sodium, chloride and carbonate ions). In NAF reptiles these glands are best known from the Dabb lizard (Uromastyx acanthinura, fig. 1). They empty into the nasal chamber (vestibulum nasi) and the salty liquid is sneezed out from time to time, Nevertheless the sodium concentration in the body liquids rises during the dry period, but does not affect body functions. No salt ¢glands have been found in tortoises and land snakes. ‘The kidneys supplement nasal glands in osmoregulation 5.2.3 Psammophilous species ‘There are three main types of sandy biotopes in our region: Litoral sands, desert sands and sections of oucds. Their herpetofauna is listed in the biotope survey (5.1). The vast ,,sand seas“ of the erg with chains of dunes over 100 m high are bare of any vegetation and therefore lack the food base for the existence of animal life. For this reason the herpetological community of the erg (5.5) chiefly inhabits marginal regions with low dunes, annual precipitation and easy access to water reserves, The biotope list (5.1) shows that some species are largely restricted to litoral sands as are the skinks Chalcides mauritanicus, Chalcides ghiarai and the Fringe-toed lizards Acanthodactylus aureus, A. blanci and A. savignyi. Typical desert forms inhabiting mobile sands are Petrie’s gecko (Stenodactylus petrii), the Common skink (Scincus scincus), species of the scincid genus Sphenops, the Erg agama (Trapelus tournevillei), the Awl- headed snake (Lytorhynchus diadema), and Avicenna’s viper (Cerastes vipera). Many species ate rather tolerant as to substrate and live on diffe- rent soils, e.g. the Desert monitor (Varanus griseus), the Red-spotied desert racer (Mesalina rubropunctata), Bosk’s fringe-toed lizard (Acanthodactylus boskianus), the Sand snakes (Psammophis), or the Horned viper (Cerastes cerastes). Some digging species inhabit sand and other loose substrates as well: The Sand boa (Eryx jaculus), lives in sand and loose soil, the skink1, Snake tracks on sand; a: Colubrid on firm flat ground, b: Colubrid along slope (down- hill at right), c: Viperid sidewinding Chalcides mionecton also hides in dry leaf litter. Desert sand is a decay product of siliceous rock accumulated by water and wind owing its colour to ferric oxides. 91-98% of its grains belong to the size class of fine grain (between 0.125 - 0.5mm) while silt (below 0.063mm) and coarse sand (over 0.5mm) are restricted to amounts of about 1% each (Haxrmany, 1989a). Several of its physical peculiarities are essential for reptile life on and in the sand and are referred to later. 5.2. 1 Locomotion on sand ‘The digit pectination of Fringe-toed lizards (genus Acanthodactylus) is one of the classic adaptations of reptiles living on desert sand. In this genus one lateral series of toe scales can bear elongated appendages which in extre- me cases surpass the actual digit’s diameter. These flat and pointed (fig. 396/2) spines or ,,fringes attain their maximal lengths on the posterior border of the 4th toe and are much smaller along the anterior toe margins. They are entirely lacking on the fingers which are flattened below and bear sharp lateral keels. Pectinate toes are a key character of all Acanthodactylus species, but there is only a very general correlation between psammophily and the degree of pectination. In Acanthodactylus erythrurus and A. blanci, which belong to the same species group, the spines are of about equal length, but the first species mostly inhabits hard soils and the second one coastal sands. Another desert dweller with pectinate toes is the Common skink (Scincus scincus) in which both marginal rows of finger and toe scales are enlarged though not spinose (fig. 22/3). The species of the gekkonid genus Stenodactylus also bear lateral digital fringes. They are shorter than in Acanthodactylus and form a seam on the interior and exterior margins of fingers and toes (fig. 259/1). Among reptile traces on sand (pl. 52-54) those of sidewinding vipers (p.472) are very typical. They are copied by undulating colubrids (fig. 1b) moving along a slope. 5.2.3.2 Life in the sand Dry sand behaves like a semiliquid. It collapses immediately behind a digging animal, but - in contrast to a real liquid - does not penetrate into hollow spaces from below. This fact is used by undersand dwellers which find relief from sand pressure under flat stones or the root system of plants and during undersand respiration (see below). Sand swimming‘ is the rapid form of locomotion by lateral undulation in the Sand fish (Common skink, Scincus scincus) and the genus Sphenops. In contrast to normal snake undulation on firm ground the loops are not locally stable, but move backward at a speed slower than the skink’s progression (fig. 471/1). ‘The term ,,swimming* suggests great case, but the friction resistance of fine grain sand is considerable even if the grains are rounded, and the low65 percentage of silt entails a loose stacking. With all its adaptations to drag and sand pressure the Sand fish deploys most of its activities on or near the sand surface. Diving into the sand is mainly evoked by predators, rivals, prey or the need for thermoregulation. ‘The most evident morphological specialisations for sand-swimming (p. 359 ff. in the Common skink are the cuneiform snout, the stout fusiform and muscular body with a short tail, tightly imbricate smooth scales which are reinforced by a pavement of osteoderms. All openings of the head are protected against sand: The nostrils can be closed by means of a valve-like mechanism, the mouth is countersunk behind the prominent snout, and the ear-opening is protected by several fringed scales. The presence of a translucent palpebral disk allows light perception with the eyelids closed, but this character can be found in many skinks with more conventional ways of life. A sand-swimming skink moves by undulation and folds its limbs backwards along the body. In the more elongated skinks of the genus Sphenops similar adaptations are found, but the limbs are greatly reduced in size as well as digit number. ‘Sand-diving in snakes ‘Snakes living in the sand surface show a pronounced tendency to shift their eyes upward as is the case in the Awl-headed snake (Lyiorhynchus diadema, p. 494) and Avicenna’s viper (Cerastes vipera, p. 537). Lytorhynchus digs into loose substrate head first while sand-diving in Avicenna’s viper works totally differently. The viper literally sinks down into the sand shovelling sand on its back by undulating motions of its flattened body. Lying at ambush only the eyes and the dark tail tip remain visible. Undersand respiration ‘Two properties of desert sand play a key role in this process: The scarcity of silt components leaves relatively wide interstices between the grains, allowing a relatively free passage of air. The inter-grain friction of sand is too high to fill air cavities from below, During undersand respiration reptiles chiefly move their ventral sides up and down and during exspiration a groove forms below the thorax (PoUGH, 1969), Ventrolateral ridges (and the adpressed front limbs in Scincus scincus) prevent the adjacent sand to penetrate into this air-filled space and facilitate thoracic extension during the following inspiration phase. Some of the water contained in the exspiration air condenses in the sand close to the nostrils. Grains cohere to a small clot about Imm thick which may form a barrier against penetrating sand Captive sand dwellers forced to live in crushed sand may suffer badly. The sharp edges inflict wounds, and the high amount of silt damages the lungs. A viable substitute for desert sand is washed river sand sterilized by heat.su 6 12 18 Tie al Fig. 1. Activity of lizards from the Haut Atlas in captivity: a: Quedenfeldtia trachy. blepharus b: Podarcis perspicillata : Lacerta andreanskyi Share and distribution of different activities are specific! 5.24 Activity cycles Diel and annual activity cycles are a basic behavioural phenomenon in amphibians and reptiles on which macro- and microclimate exert a strong influence. For this reason most data on this subject are generalized and may differ with populations. Closely related species show marked differences, and within herpetological communities different species are active at different times (fig. 1). 5.2.4.1 Diel activity cycles The simple differentiation between diurnal or nocturnal activity can be applied to many, but not all species. Strictly diurnal are Testudines, Agamidae, Chamaeleonidae, Varanidae and Lacertidae; strictly nocturnal are some Gekkonidae; the rest follows a mixed program. In most amphibians diel cycles are less rigid. Strictly nocturnal are Pleurodeles walt! in their terrestrial phase,Alytes obstetricans and Pelobates varaldii. Salamandra algira is nocturnal but leaves its shelters during and after rain, The species of Bufo are usually nocturnal (B. bujo) or crepuscular (B.mauritanicus and viridis), but appear during moist weather. Discoglossus pictus can be observed around the clock. Hyla meridionalis is diurnal or nocturnal depending on the type of activity. Water frogs (Rana saharica) are diurnal with concerts lasting into the night. Experiments (Haxrmans, 1989) have shown that besides an adequate temperature a sufficient light intensity is indispensable for the performance of a normal activity cycle in diurnal reptiles. ForScincus scincus illumination by lamps must attain 800 W/m? which corresponds with full June midday sun at moderate northern latitudes. A reduction to 300 Wim? shortened the activity period from 10 to 5 hours while locomotion was reduced to 2/3 of its original value. ‘The usual diagrammatic presentation of diel cycles are graphs covering a 24h interval, but most of them are based on general impressions rather than on exact counts. While the number of basking specimens can be crudely estimated, things become extremely difficult with those resting in the shade or foraging under cover. A common phenomenon in diurnal species is the midday pause which increases in length and intensity according to the summer heat and may even cut the activity curve in two (bimodal curve). Another change evoked by summer heat is the total inversion from diurnal to nocturnal activity as was observed in Scincus scincus, Sphenops boulengeri, Chalcides ocellatus and the snakesLytorhynchus diadema, Spalerosophis diadema, Macrovipera species and Cerastes vipera. Ifthe periodicity really changes completely in these species remains dubious, nor do we know anything about the transition phase. Ophisaurus koellikeri from southern and northern Morocco (Inezgane and Ifrane) apparently have opposite activity cycles. One captive specimen from67 Inezgane observed by PetzoLn (1964) was strictly nocturnal in captivi while several of these lizards were seen basking in Ifrane and captivity obervations confirmed their diurnal activity. A surprising diversity in diel activity cycles is found in the Gekkonidae, ‘most of which are considered nocturnal. The following list shows that even closely related species may be different in this regard, and there are many intermediate stages between diurnal and nocturnal regimes. Tropiocolotes tripolitanus: strictly nocturnal Geckonia chazaliae: nocturnal with exceptional observations during day Stenodactylus petrié: strictly nocturnal Stenodactylus sthenodactylus: nocturnal with diurnal activities during cool weather Tarentola neglecta: nocturnal with very low diurnal activity Tarentola mauritanica: nocturnal hunting; extensive diurnal basking periods in most populations Hemidactylus turcicus: nocturnal hunting activity, frequent diurnal social interactions in refuges with vocalization ‘Saurodactylus (m.) brosseti: main locomotory activity at dusk, but continuing into early morning; sitting in ambush in shelter during the day Ptyodactylus oudrii: chiefly nocturnal with activities far into the forenoon hours and starting carly in the evening; diurnal in shady biotopes Quedenfeldtia trachyblepharus: almost strictly diurnal with low-ranking individuals remaining hidden by day and proceeding outside by night. 5.2.42 Annual activity cycles A continuous activity throughout the year is rather the exception than the rule in North African ectotherm animals and is chiefly observed in the mild maritime climates of Sous, Massa and Draa valleys (Atlantic MOR) and the region of Oran of the following list. Surprisingly enough, rock dwellers of the Haut Atlas at altitudes over 2500m (Quedenjeldtia trachyblepharus, Podarcis perspicillata) are also active in winter. Their microclimatic advantages are the efficient insolation of south exposed rock walls and the possibility to withdraw into deep rock crevices where temperatures are comparatively high in winter. In addition to this, these species have a very low basking temperature range (in Quedenfeldtia starting with about 4°C, in Padarcis perspicillata a few degrees higher) and can at once move out of their shelters to bask when the sun shines. Annual activity cycles can be interrupted by hibernation and estivation which allow to avoid unfavourable temperatures, lack of food and, especially in amphibians, lack of water. Things get extremely complicated as there are profound differences between different populations: Water frogs from the Moyen Atlas terminate hibernation earlier than those of the lowlands. Other important influences are weather and local climate, The onset of hibernation in Erg reptiles is released by the first winter rains, and all these species hibernate. The sand surface becomes very cold in winter as there is very little heat conductance from deep warmer layers.Asa tule, large reptiles hibernate longer than small ones. In the Grand Erg Occidental Varanus griseus and Cerastes vipera hibernate from October to March while small Lacertids are inactive from November to February. Body temperatures of hibernating Erg repiiles lie between 14 und 22° (Grenot & Verner, 1973). In several species, e.g. Agama impalearis, Chalcides ocellatus, Acantho- dactylus erythrurus, Acanthodactylus savignyi, Mesalina guttulata, Podareis hispanica, and juveniles do not follow the hibernation or estivation regime of adults as the following table shows; we tend to interprete this as, advantage from predators as most of these are also in dormancy or hibernating. Remarkable is the fact that several amphibians (Pleurodeles walil, Pelobates varaldii, Hyla meridionalis) reproduce during winter. Hibernation and estivation in North African amphibians and reptiles Data are mainly fromDoumercvt (1901) for the region of Oran, Bows (1957) for the Souss and Draa regions, Mr.apo & OtMeno (1987) for Morocco, many from Qued Massa S of Oued Souss, Pastzur & Boxs (1956) for Moroccan amphibians, Verwer & Gaexor for Beni Abbés (abbreviated Beni Abb.) in the Grand Erg Occidental. Notes include locality, duration and remarks. For species abbreviations see p. 10ff. Winter activities ‘Summer activities: Salamandridae Ple walt reproduction! estivation if dayas dry up Sal algira hibernation in high regions Discoglossidae Aly obstetricans N-MOR probably estivation Dis pictus estivation during drought Pelobatidae Pel varaldii reproduction! estivation Bufonidac Buf bufo hibernation Buf maurit. hibernation in cold regions _estivation during drought, Buf "regularis" probably B. xeros S-LIB estivatiow Buf viridis hibernation Hylidae Hyl meridionalis. low regions: reproduction _estivation high regions: hibernation _ reproduction Ranidae Ran saharica —_N-MOR hibernation S-MOR none or short69 Testudinidae Tes graeca Gekkonidae Gee chazaliae Py ragazzii Que trachybl Sau brosseti Ste sthenodact. Tar mauritanica Tar neglecta Tro tripolitanus Agamidae Aga impalearis Tra mutabilis Tra tournevilli Uro acanthinura Chamaeleonidae Cam chamaeleon Scincidae Cha chalcides Cha mionecton Cha ocellatus Cha polylepis En algeriensis Sci scincus Sph boulengeri Sph sphen.? Lacertidae Aca aureus Winter activities hibernation Sous, active Tademait, rarely active ‘Haut Atlas, active Sous, active Oran, active generally active Oran, rare activities Beni Abbés hibernation Dra, November-March common Sous, predom. juv. ‘Tademait, active Beni Abbés hibernation Ougarta near Beni Abbés short hibernation ‘Zemmour (W-SAH), active ‘Tahrit (C-SAH), rare Massa, hibernation Oran, hib. until Feb Sous, active Massa, hibemation Oran, hibernation until Feb Sous, rare activities hibernation Nov-Mar Massa, hibemation Beni Abbes: hibernation Oct-Mar Beni Abb. hibernation Massa, hibemation Sous, active Summer activities estivation possible locally less active in summer rarer predominantly juveniles estivation possible Draa, Feb - May active, June only pregnant fem. August estivation Oran, rare in June; yearlings & newly hatched active Draa, estivation rare activities70. Aca boskianus Aca erythrurus Aca ‘pardalis” Aca savignyi Aca scutellats Lac pater Mes guttulata Mes olivieri Mes pasteuri Pod hispanica Pod perspicillata Psd algirus Anguidae Oph koellikeri Varanidae Var griseus Amphisbaenidae Trg wiegmanni Colubridae Lyt diadema Mal moilensis Pso schokari Viperidae Cer vipera Winter activities Massa, hibernation Oran, hibernation until Feb Tademait, active Oran, hib. Nov-Feb predom. juveniles active Sous, active Sous, active rare Jul-Aug Oran, hibernation. until Feb Jul - Aug only juveniles active Oran, hibernation until May Ghardaia, active Beni Abbés hibern. Nov-Feb Oran, much reduced Oran, frequent from Sep Oran, Ghardaia, active tive predom. juv. Sous, Oran, active Oran, much reduced Beni Abbés hibern. Nov-Feb most localities active Oran only yearlings and Oran, hibernation newly hatched active Oran, active Haut Atlas, active Oran, hibernation Nov-Mar Moyen Atlas, hibernation Beni Abbés hibernation _see special part ‘Oct-Mar hibemation at high altitudes Massa, hibernation Sous, rare activities Oran hibernation Dec - Feb Beni Abbés hibernation Beni Abbés hibernation Beni Abbés hibernation Beni Abbés hibernation7 5.2.5 Reproductive cycles ‘The most favourable timing for egg cell and sperm maturation seems to be a synchronized development, but reality is far from that. The fact that spermatozoa can survive for days up to years in the female genital tract makes an exact synchronisation of spermatogenesis and oogenesis unnecessary. As a rule mating preceeds ovulation several days or weeks in lizards, 2-3 months in snakes, and spermatozoa await ovulation in the female genital tract. In species with sperm storage (Chamaeleo chamaeleon, Natrix maura) one copulation is sufficient for several clutches. Female reproductive cycles One of the simplest types of reproductive cycles observed in temperate latitudes is the production of one clutch per year during a short and fixed reproductive period. Female reptiles following this schedule are called monestrous. The opposite type with a year-round reproductive phase and many clutches is limited to the tropics. In males of these species spermatogenesis is continuous. This type of reproductive cyle is lacking in NAF, but two or several clutches (diestrous and polyestrous species) are common. The following list of female reproductive strategies is far from being complete. Most data are based on single fortuities rather than continuous observations, and the influences of age, climate and weather are badly known. We can but repeat that observations on the biology of quite a number of speciesare completely lacking. For simplicity we also add data concerning amphibians. As one indispensable condition for vitellogenesis is a good nutritive state of the females, no clutches may be formed in unfavourable years. Obviously the start of vitellogenesis depends on nutrients stored in the two abdominal fat bodies. While in most species ovule growth in the ovaries is a matter of weeks, the Amphisbaenians (Blanus and Trogonophis) do not ovulate yearly, but show a biennial or even longer cycle. This slowing down of the cycle is explained by the fact that Amphisbaenians are a tropical group and the two NAF genera live at ity northern distribution margin (Bons & St. Gikons, 1963). One reproductive phase per year: Pelobates varaldii, Bufo bufo, Bufo viridis, Hyla meridionalis, Rana saharica, most snakes, Mauremys leprosa, Chamaeleo chamaeleon, Chalcides chalcides, Chalcides ocellatus, Eumeces schneider Varanus niloticus. i, Mabuya vittata, Ophisaurus koellikeri, Varanus griseus, ‘Two clutches possible: Bufo mauritanicus, Emys orbicularis, Testudo graeca, Agama impalearis, Uromastyx acanthinura, Acanthodactylus erythrurus, Acanthodactylus spinicauda (at least 2), Mesalina guttulata, Psammodromus algirus.72 ‘Seyeral clutches possible (clutch numbers in brackets): Pleurodeles walt! (several), Alytes obstetricans (2-4), Discoglossus pictus (3-6), Geckonia chazaliae (4-5), Hemidactylus flaviviridis (3), Hemidactylus turcicus (3- 5), Pryodactylus ragazzii (3-6), Saurodactylus (m.) brosseti (up to 8), T. annularis (2-8), T. deserti (4-6), T. mauritanica (up to 15!), Tropiocolotes tripolitanus (up to 6). Acanthodactylus pardalis (2-3), Lacerta andreanskyi (up to 3), Ophisops elegans (up to 3), Podarcis hispanica (2-3), Psammo- dromus blanci (up to 5). Male reproductive cycles (Boys & Saint Girons, 1982) A simple seasonal spermatogenesis shows the following phases: 1. A pause afier mating, 2. Start of spermatogenesis 3. Maturation (meiotic) di formation, 4. Transformation of spermatids into ripe spermatozoa, 5. Sperm storage, 6. Mating and expulsion of stored spermatozoa. mitotic multiplications of spermatogonia, sions of spermatocytes resulting in spermatid ‘As spermatogenesis proceeds (apparently with few exceptions) only above 20°C, most, if not all, North African reptiles insert another resting period during winter. The situation of this hibernal diapause (after phase 5, amidst phase 3 or after phase 2 of the above list) allows to distinguish three main types of spermatogenesis: a. Postnuptial or estival ty and is finished before winter: Emys orbicularis, Mauremys leprosa, Testudo graeca, Macroprotodon cucullatus, Natrix maura. Chamacleo chamaeleon is unique with estival spermatogenesis and ovulation in October. perm production starts after the mating period spring summer autumn winter mating multiplication maturation storage pause b. Mixed type: Winter pause interrupts maturation divisions: Lizards and vipers from temperate regions: Quedenfeldtia trachyblepharus, Hemidactylus turcicus, Lacerta, Podarcis, Psammodromus with several ovulations; Chalcides, Eumeces, Ophisaurus koellikeri with a single ovulation. spring summer autumn winter matur.end — mating multiplication _matur. start pause . Prenuptial type: Winter pause afier the multiplication phase: Agama impalearis, Uromastyx acanthinura, Acanthodactylus erythrurus with two ovulations; Scincus scincus, Varanus, Coluber, Malpolon, Cerastes73 with one ovulation; Blanus, Trogonophis wiegmanni, apparently do not ovulate every year. spring summer autumn winter maturation mating multiplication pause ‘There is no case of a continuous spermatogenesis known from Northern Africa but Tarentola mauritanica (and other Tareniola species?). Mesalina, Acanthodactylus pardalis, come close to it, with a regression in winter and a second, but weaker one in summer. Other examples of more complicated rythms occur in Acanthodactylus erythrurus with two successive phases of sperm production and Acanthodactylus pardalis with a prolonged one. Reproductive modes of North African reptiles In addition to the different number of ovulations there are some further alternative strategies which in various combinations result in a wide spectrum of different reproductive modes. Asa rule snakes mate 2-3 months before early summer ovulation or even as early as in autumn, In oviparous species eggs can be retained by the female for several weeks before oviposition. In ovoviviparous species eggshells are reduced to a thin membrane, but no placenta develops. The offspring leave their eggs shortly after oviposition. In (cu)viviparous species egg membranes and an oviducal wall form a specialized contact zone, the placenta. In our region ovoviviparous and viviparous species reproduce once a year and at a still slower rate (biennial or slower) in Trogonophis wiegmanni. In chamacleons the embryonal development is exceptionally long because of a long diapause during the cool season. Reproductive modes are visualized on the next page for the following species: 1. Varanus griseus: Monestrous, oviparous 2. Chalcides chaicides: Monestrous, viviparous Agama impalearis: Diestrous, oviparous 4. Psammodromus blanci: Polyestrous, oviparous 5. Chamaeleo chamaeleon: Monestrous, prolonged incubation period 6. Natrix maura: Prolonged egg retention, double mating period (data from France, in Norther Africa about one month earlier) 7. Trogonophis wiegmanni: biennial estrus cycle, ovoviviparous; (most data are from Saint Gikoxs, 1985)74 Signs: vevwy vitellogenesis; mmmmm: mating: O: ovulation; Ja: egg laying; ...... egg retention in oviduct; egg incubation; ******* gestation. months JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC 1. Varanus griseus vwyw0...la—hatching ‘one annual clutch mmm 2.Chalcides chalcides vwwwwwO* ‘one annual litter mm 3.Agama impalearis ww0.....vwvO.le——hatehing, ‘two anual clutches m om | la hatching 4.Psammodromus blanci ~ several annual clutches vww0 vO wO wO la-—hatching, mm ni mim Ja—-hatching Ja-—hatching la—hatching AUG SEP OCT NOV DEC JAN FEBMAR APR MAY JUN JUL AUG ‘5. Chamaeleo chamaeleon - winter diapause wowy0.. la —$ 6.Natrix maura ‘two mating periods vwww0..a—hatehing ‘mm ‘mmm JNJL AU SE OC NO DEJA FEMR APMY JNJL AU SE, 7. Trogonophis wiegmanni - biennial cycle v vv Vv VV Most North African lizards apparently follow the modes 3 and 4, snakes mostly follow mode 6, 53 Population density and biomass The great technical difficulties of solid estimates make data very scarce. They are only available for a few intensely studied species, and mostly for one locality during one yr. Densities per surface unit make only sense for evenly distributed species on rather homogeneous ground, and are easiest to assess for diumal forms. The concept cannot be applied for species with75 linear distribution, e.g. along a watercourse, or local concentrations, e.g. geckos on rock outcrops. Biomass estimates (g/ha) must take into account the considerable weight differences between juveniles and adults, Species population biomass remarks Jha Hyla meridionalis 3.800 Canarian Isl, banana plantation > 10.000 shores of artificial lake, Oukaimedene Lacerta andreanskyi 2.950 6000 g Oukaimedene, near brook Acanthodactylus 379 2154g Tunisia, dunes maculatus Ophisaurus koellikeri 20 Moyen Atlas, stone walls near brook Chalcides chaleides 0,0077-0,218 S-France, grassland Uromastyx 01-1 general value acanthinura 1,465 268,98 g Dj. Rhenouma 05-2 Monts d’Ougarta 0.1504 19,622 g Grand Erg Occidental Varanus griseus 0,02 NW-SAH, open dunes 0,06 depression between dunes 5.4 Trophic complexes 5.4.1 Role in food chains The food chain is a simple linear model for trophic interrelations, starting ‘with producers and showing the way of matter and energy through consumers of different orders, ending with large carnivores as the final consumers. ‘This model disregards the use and decomposition of waste material. While a caterpillar and a sand wasp predating on it are easy to locate in this model, other correlations are often more complicated. Many carnivores feed on herbivores and small carnivores, ranging as consumers 2nd order aswell as 3rd order. So do lizards preying on grasshoppers and spiders. For this reason the chain model is rather simplistic although it may help to show some points of interest. Another well-known model is the food pyramid where producers figure at the base and final consumers at the top. In contrast to the food chain this model visualizes biomass differences.76 Ina stable ecosystem each floor of the food pyramid subsists on the surplus production of the inferior one. Only a fraction of food intake is converted into biomass. Ideal producers (as some algae are) double their biomass daily, ideal consumers save energy and substance, converting biomass economically. While mammals and birds spendcrudely 90% of energy intake for heat production, ectothermal animals avoid this expenditure and convert biomass more economically. For this reason the same food base can support a much higher biomass of reptiles than of mammals, provided that temperature is high enough. For this reason reptile biomass is relatively high in warm deserts where plant production is at times insufficient for small mammals, In ,,true“ North African deserts as the Tanezroutt lizards of the genus Mesalina are therefore the only ground-dwelling vertebrates over hundreds of kilometers. Birds and large mammals move in and out, profiting from ,desert blooms* after rains, but cannot maintain stationary populations. Amphibians and reptiles in the trophic chain: Producers_| Consumers | 1. order 2. order 3. order| 4. order plants herbivores | camivores... important amphibian and reptile food: algae | zooplankton leaves worms, insects, flowers snails seeds | FISNES 0.000000 Urodela | larvae & adult tadpoles i adult frogs and toads Mauremys leprosa | Emys orbicularis Testudo spec. most lizards Scincus scincus Agama impalearis Uromastyx juv. Uromastyx ad. Varanus griseus. | | Amphisbaenians [Dpblops & ‘Leptotyphlops most snakes.7 In North Africa amphibians and reptiles range in the same places of the food web as in other regions, but some species merit special attention. Among turtles the common Mauremys leprosa uses a wide spectrum of plant food while the rare and local Emys orbicularis is a rather strict camivore. Uromastyx acanthinura (precise information on other species of Uroma- ‘tye is missing) juveniles are insectivores while adults almost exclusively feed on plants. One advantage of herbivores is the easier accessability to food, but they need a relatively voluminous body (in Uromastyx most plant material is stored in the caecum) and high temperatures to process vegetable matter. Another Agamid, Agama impalearis, eats the yellow flowers of the composite Anvillea radiata as a supplement to its animal diet. The flowers do not only serve as an important source of water, but are also ingested when the lizards have sufficient opportunities to drink. A less pronounced turn to herbivorous nutrition takes place in Chalcides ocellatus (and possibly its close relatives). The Common skink (Scincus scincus), besides its insect prey, eats a substantial amount of seeds, mostly from grasses. Their percentage rise: summer when insects withdraw to moist and cool depths and ripe seeds of dune plants fall down, Additionally preying on scorpions, the species also enters as a consumer 3rd order. ‘The Desert monitor Varanus griseus shows the broadest prey spectrum from small insects to adult snakes. With increasing growth the monitors steadily move up the food chain and finally reach the highest rank taken by a desert reptile, It is remarkable that three of the mentioned effective consumers, namely Uromastyx acanthinura, Scincus scincus and Varanus griseus are hunted by human desert dwellers as a staple food. Chased by children they are more important than big game today. 5.4.2 Role in food webs ‘Trophic interrelations, symbolized by arrows for material and energy flux, become very complex in rich biocoenoses. For this reason desert systems offer themselves for model cases, and even these are massively simplified in the following figures with only vetrebrates represented as species. ‘The increase in the number of ecological niches from sandy to rock biotopes becomes very evident in the comparison between the erg and djebel formations (pp. 78,79). Mind that the trophic arrow between scorpions and ,,Wind spiders" (Galeodes, p.78) on one side and small reptiles on the other may point in opposite directions with the reptiles being predator or prey. The vernacular names for bird and mammal predators can be retrieved in the appendix (8.3).78 Trophic web - Grand Erg Occidental (based on Verner & Grenor, 1972) For names of bird and mammal predators see appendix 8.3 HERBIVORES —INSECTIVORES CARNIVORES caterpillars Lytorhynchus diademe | | Vulpes ruepelli Subterraneous insect larvae plants and dead organic matter Poecilictis libyca79 ‘Trophic web - Djebel formation, Monts d’Ougarta, W of Beni Abbés (based on Grenot, 1976) For names of bird and mammal predators see appendix 8.3, HERBIVORES INSECTIVORES CARNIVORES small rodents. Nephron percnopterus large herbivorous ‘mammals Accipiter nisus plants dung Ke LiF Hilvus migrans, rT SWS ° x sfatco meee WY Falco biaraicus ea A! rropi WAS A WY \ N RN TP cor icolli: 5 Vr scanaas A Vi eppcorves ruticottis, : yp sintcus NEG : w\\\ 8 i [FerentoTa sree }\oy i i : A\\ne ; Sa) 3 ZS Alaisle Uy 7 > LX van tenebrionid beetles [Hal polon J ON Stel) =>80 5.5 Herpetological communities (herpetocoenoses) One characteristic for a highly evolved biocoenosis is species diversity. Though the majority of amphibians and reptiles predates on small animals, specialisation allows their coexistence by resource partitioning and diffe- rent ambiental preferences. Niche overlapping is avoided by a number of diversifications, e.g. diffe annual and die! activity periods, activity temperature ranges, substrate choice, food spectrum, water dependence, presence of cover, shelter and basking sites. Theoretically the number of ecological niches depends on the complexity of biotope structure which is greatest in dense forests and poorest in uni- form deserts. Both of them harbour poor herpetocoenoses: Shade tolerant species in the woodlands (Bufo bufo, Salamandra algira) and Desert racers (Mesalina) in extreme deserts close to ,abiotic* areas. As most North African reptiles are heliophilous the most complex communities are encountered in biocoenoses with open vegetation. The columns in the biotope list (5.1) give merely an idea of communities because species of different geographic distribution are listed together. More precise conclusions can be drawn from the transsects (p. 36ff). The following surveys are based on the field researches of several authors. We tried to present a wide range of regions and biotopes. Their general arrangement is from west to east and from north to south (fig. 81,2). Two very different aspects are covered: 1. The herpetofauna of a limited areal (several hectares at most) which was studied for a relatively short time, possibly only several hours (lizard communities of northern Morocco after Metabo (1985, p.81), ruins of Lixus, Ifrane, Col du Zd, Oukaimedene, pp.82,83). 2. Summaries of extensive research covering months or even years in a large area are contracted. They often contain numerous species and show their distribution over different biotopes, which may be miles apart (Rheraia valley, erg near Beni Abbés, Monts d’Ougarta, mouth of Qued Massa, Presahara of SW Morocco). On the illustrations data are arranged in the following order: landscape forms; typical plants; sketch of the biotopes; distribution of amphibians and reptiles. Continuous lines indicate a regular occurrence, interrupted ones indicate marginal biotopes. Where data were available frequency and need for water are noted: f frequent, ff = very frequent, r = rare, rr = very rare; w = water necessary; (w) = vicinity of water preferred. A general list of biotopes is presented in chapter 5.1 while the spatial distribution over different landscape types is shown in the transsects of chapter 4.5.81 9, Fig. 1. Localities for table 5.5.1 5.5.1 Lizard communities in northern Morocco (after MeLLADo, 1985, rearranged; nr. 14 after Boxs & Saner Ginons, 1963). Locality (fig. 1) elevation inm vegetation 1 Oued Moulouya 2 Dj. Kebdana 3 Guercif 4 Driouch 5 Safsafte 6 Exsaft 7 Tafrata SAL Ateuf 9 Bab el Arba 10 Souk es Sebt 11 Tafrant 12 Bab Bou Idir 13 Dj. Tazekka 14 Oulmes Taxon Saurodact. m. mauritan. Saurodact. fasciatus Stenodact. sthenodact. Tarentola mauritanica Agama impalearis Trapelus mutabilis Uromastyx acanthinura Chalcides chalcides Chalcides mauritanicus Chalcides ocellatus Chatcides ghiarai Chalcides polylepis Eumeces algeriensis Acanthod. boskianus Acanthod. erythrurus Acanthod. maculatus Lacerta pater Mesalina olivieri Podarcis hispanica Podarcis perspicillata Psammodromus algirus Psammodromus blanci Ophisaurus koellikeri Blanus tingitanus Trogonophis wiegmanni Pistacia shrubs Chamaerops humilis (Dwarf fan palm) Zizyphus lous (Sujubethorn shrubs) Haloxylon articulatum, Mugwort (Artemisia) steppe Zizyphus lotus Stipa tenacissima (Halia steppe) Quercus ilex (Holm oak) evergreen forest Cedrus atlantica Quercus ilex wood Localities 1-2-3. 4-5-6- 7- 8-9-10- 11-12-1314 xXx x X XXX xXx xx x XX KXXX x x x x x x XX XXXKXXXK x x x XX XX KX x Xx x Xx x ox x x x x XXX xX Xx x x XXX xX Xx xXx x xx ox XXX xX XX x xx XxX x x x xx ox x x82 5.5.2 Landscape and biotope profiles (survey fig. 81/2) 1 alluvial plain cascle rock reine of Lines mecca sparse plane growth Meaid. turcieus tyla weridioaalis Pan comm! Bufo viridis Bufo meuritanicus £ Macrix moura ff aureays leprosa ff faye orbicularis © ‘ Morecco, Moyen Atlas Valley near Col du Zad, 22000. ‘2 fron Steaaler, 1965a) streas sorthers slope vith debris At antic Cedar forest Sedua (Stonecrop) Codrus atlanti HWeltantheam (Rock Rose) Quercus iex (Im Oak) Wieractua (lavkweed) Verbascur (Hullein) Dianehus (Pink) Cencaurea Canpansia (Warebell) Ononis (Rest Harrow) Myla seridionalis Rana compl. Bifo viridis The Vipera "latastei" specimen has been lost and might have been Vipera monticola,83 Morocco, Moyen Aas Oued Iicane Kovards FL fajeb, 16400 asia of Helge vith ate evgatin ditch being Testo grec Calserhipocepis 2 Aslpioe wnsprsulous Taectate reer Teese fF onrtt, Bays ecicalris 2 Main marin? ita tf Lacerta pater ‘The “Chalcides potylepis” was a misidentified Chalcides lanzai. Norocco, Haut Atlas Mateau of Oskainedene, 24008 alkeus (1981), Bisact (1487) boulders brook debris with ‘snp rock outcrops stone Assit A Legoy spur vegetacion (cme 2") alt Leceres dresses ff Quedeafelitia track. ff Ayla erid. Dis pictus ¢ Podacis perspcill. £f Uacerta aio aurit. r Pedals bsp. £ pater f8 5. 6. ‘The "Saurodactylus mauritani- cus" is brosseti; Acanthodacty- lus pardalis is maculatus sensu Satvavor (1982) ued Heraia valley, 1300-17008 foot of Djebel Toubkal (Bate fron Saint Cirons, 1953) rock face allwial soils, dense shrubs single shrubs open ground elders ‘srigatd caltres lire, walt Govdefelaia tach. (o)ff Padcis persplcill. wf Tarencale saritania ¥ Podsrcishisp.vauctert (eof ho inp! ¢ ——Uphisuras hell. (wre ‘fo azsritanices Chaacleo chameleon iscghsss pict.» iacerta pater ocrovipera auuritacica ¥ ic coast south of tgadir 0-5 al., 1988; Mellado & Olaelo, 1981) Aclmic costal stepe ——aportia steppe Arjanierstramub ‘an dels ‘thik Layer of dead Leaves ges sonspernn——fportin echines Agua spinosa (hon aati Bpbaris droite eye Tate [dings sims Scie snteprhion es ari Swe Petes laevigata 3B. pa Besa Testo greca sr —_—_‘Sesalia liviet f Toremala muritanicn f ‘Agana inpaleris. ££ Gmclen shenclon er at (aleides sioncton £ ar aiesées yiepis £ ‘Blaos settetal er fy mecid.r—Srodctyles eritanices ff aw conn, Maren Lp. tt Ieris sore F ‘anthotectylas sures chon cazaize £ (aides einen { ‘Spbempe epenjsif.t85 . Sourtvostora Moraceo Fresahara between Oued Massa and Ovet Oras (dace from Bons, 1959) Acanthodactylus pardalis is sand story oust rock reg earth debris) rock face ‘maculatus sensu Satvavor (1982) steps slope ArganiaNeriuw oleaader Haloxylon scoparive Argan Tasarix Artenisia berba-alba Vitex deabasie apiylla ‘Alzoon canariense ‘sptodelus tenuifolius Tecentele seare Quedent. crach. Cv) Varanus griseus Wipers naar S-- = = Tropioclotes tripol. Stenodectylus then sos Pryotactylus axis Scincas scincus ‘Sphenops toulenger Mesalina olivieri (+) ‘Aeanth, boskianss- ‘canth. pacdalis Gerastes cerastes Uromastyx scanthinars (0) Agana ipalearis--— ato brongersaat Bufo aauritanicus » Bale viridis » Ghalcides ocellatus Coluber algicus ¥ Psanaphis schokari Aaja haje ¥ 8. Algeria, Monts d'Ougarta W of Beni Abbés, . Fock formations (Jebel), 500-700e. (data frow Grenot, 1976) ree sandstone formations dry riverbed debris slope vith eroded groore ‘Acacia raddiana — Louna arborescens eLianthemua Lipii Uinoniactrun foot Panicun rurgidoe Linaria aegyptia Ephedre alata ptt Acanthodactylus boskiams Pryodactylus oudrit Tarentola spec Tropiocolotes tripolitanus “Haipoten moileasis -Psanaophis schokari “Cerastes cerastes86 9 10. Algeria, Grand Erg Occidental. Dunes near Beni Abbés, ca. 500n (data from Gauthier, 1967; Grenot, 1970; Vernet & Grenot, 1972; draving after Vernet & Grenot) shrubs herbs Ephedra alata ‘Stipagrostis pungens (Drinn grass) Genista sabarae Cyperus congloseratus Calligonus comsus firm sand loose sand dead Drinn grass Trapelus tourvevillei Scincus scincus Sphenops boulengeri Tarentala neglecta ‘Acanthodact ylus Jongipes Cerastes vipera Varanus griseus (den) -Mesalina pasteuri ‘Stenodactylus petri Lytorbynchus diademe -Malpolon soi lensis Exposed plateau betveen dunes (taiert) near Beni Abbés, ca, Soom (same reference) 103 cued decaying crust calcareous crust (50 ca) sandy and salty ground Traganun nudatue Salsola foot ida -Acenthodact ylus scutellatus ‘Scenodact ylus sthenodact ylus Trapelus sutabil Nesalina rubropunctet Cerastes cerastes87 u. Southern Tunesia Border of the Grand Erg Oriental, 200-3008 (after ¢. Blane, 1966) wmbile dones rom dunes hillock owed stony grad. boulder of solid sand (cebia) (res) Seinew se. 7 ‘eanthod. Jon, fe. scutell. Ac. boskiamas Ae, pardelis ‘Sphesops boul. Aeselina gurtulata, ¢ Trapeins mtabilis Tarentole nauit. {f-- Tropiocolotes tripolitans-~ 12, Litya, Cyrenaice oat fat (after Schlei srk sear HL Baydab, 0-308 1987) Moor ‘wach coastal plain rocky plates ‘sa Pistia lestiscas drbetas neds saline denidactylustarcieas ff ry in suner Mesa ne gutalata Gorets cares nests f Chaleidesocllats-- Trapolus sabilis Acarchosctys Dostamus { Colwer rpersi aa comp, in sources ad cisterns Chameleo chameleot Tarotola asuritania ff For climatological data see fig. 12/1,2; 15/1,2; 18/1.5.6 Impact of man ‘The urge to feed a rapidly increasing population entails a steady destruction of natural habitats which are partially replaced by artificial ones. The ecological damage caused by modern mechanized agriculture is of course much greater than the effect of traditional extensive pasturing, or farming and gardening on small plots. Especially affected by this development are biocenoses with special demands and of such restricted areas as the Argan forests (Argania spinosa) of SW-MOR (Metabo, 1989). ‘The balance inclines more and more toward a few common species while those inhabiting special ecological niches disappear. Of course the pressure ‘on snakes which are, seen as a whole, rater, larger and more suspect than other amphibians and reptiles, is especially hard. The large, conspicious and feared species of Macrovipera have become very rare. 5.6.1 Direct impact ‘Use as food: Uromastyx acanthinura, Scincus scincus, Varanus. griseus Use as popular remedies or talismans: Uromastyx acanthinura, Chamaeleo chamaeleon Persecution by snake hunters: Naja haje, Bitis arietans, Cerastes cerastes, Coluber hippocrepis, Spalerosophis diadema Persecution of large venomous snakes: Naja haie, Macrovipera species, Bitis arietans Souvenir trade: Turtle shells (Testudo graeca), skins of Coluber hippocrepis for drums etc. Pet trade: Venomous snakes. Illegal export threatens Uromastyx acanthinura, Testudo graeca, 5.6.2 Habitat change Intense use of pastures: Dayas used to water the herds are heavily polluted and make breeding of Pleurodeles walt! and Pelobates varaldii impossible in many places. Cultivated ground is inhabited by a few common ubiquists. Their number decreases with the change towards modem agriculture. Fields are inhabited by Chalcides chalcides, Chalcides ocellatus, Acanthodactylus erythrurus, Podarcis hispanica vaucheri, Trogonophis wiegmanni, Eryx jaculus, Coronella girondica, Natrix maura ‘Stone piles and walls around fields serve as refuges for Hyla meridionalis, Bufo mauritanicus, Hemidactylus turcicus, Tarentola mauri- tanica, Chalcides mionecton, Chalcides ovellatus, Chaleides polylepis,Lacerta pater, Macroprotodon cucullatus, Malpolon monspessulanus, Naje haje, Macrovipera species Anundo donax hedges inhabited by: Hyla meridionalis, Chamaeleo chamaeleon Opuntia hedges: serve as refuges for Coluber hippocrepis, Malpolon ‘monspessulanusFig. 1. Original (hatched) and recent (black) woodland Gardens and groves :Bufo mauritanicus, Hyla meridionalis, Psammodromus algirus, Trogonophis wiegmanni Parks : Coluber hippocrepis Refuse dumps: Chalcides ocellatus Deforestation (fig. 1) destroys habitats of forest dwellers such as Bufo bufo and Salamandra algira and of forms digging in humid soil like Ophisaurus koellikeri. The disturbed water balance above all hits forms which prefer cool and clear water, e.g. Emys orbicularis and Natrix natrix. Eucalyptus plantations are avoided by most species. Only Testudo graeca, Psammodromus algirus and. Macroprotodon cucullatus are met here. Irrigation projects Cisterns : Discoglossus pictus, Mauremys leprosa Reservoirs: Hyla meridionalis: A demographic explosion of this species was observed near Oukaimeden after the creation of an artificial lake. Rana saharica, Natrix maura Ditches: Pleurodeles wali! also in dry ones, Rana saharica, Concrete tubes: Discoglossus pictus, Hyla meridionalis, Pryodactylus oudrii, Tarentola spec., Lacerta pater. Subterraneous channels (Foggaras): Pleurodeles waltl larvae. Stone gutters along roads: Tarentola mauritanica, Lacerta pater, Podarcis hispanica Buildings: Hemidactylus turcicus, Tarentola bochmei, T. desert, T. mauritanica, Coluber algirus, C. hippocrepis. Ruins: Discoglossus pictus, Hyla meridionalis, Bufo mauritanicus, Chalcides ocellatus, Chalcides colosii, Chalcides polylepis, Trogonophis wiegmanni, Macroprotodon cucullatus, Natrix maura, Naja haje, Macrovipera mauritanica and the species mentioned for buildings. Road embankments: Bufo viridis, Agama impalearis, Chalcides ocellatus, Acanthodactylus erythrurus, Lacerta pater, Podarcis hispanica vaucheri, Psammodromus algirus, Coluber hippocre~ pis, Malpolon monspessulanus. Road signs (stone piles): Trapelus mutabilis. 5.6.3 Protective measures Nature reserves have been established or proposed in all countries. However the enforcement of protective measures is mostly rather weak with overgrazing, poaching and felling of trees going on. Often the demands of tourism seem to prevail over nature conservation in these projects. Export limits: MOR: CITES permits are needed for the export of turtles, Chameleo and Uromastyx species. ‘The export of wild animals is generally not allowed. Tourist souvenirs made from parts of protected animals, e.g. turtle shells, can be exported. References: Dupots (1982), Mettapo (1989), Scuweicer (1992a), STEMMLER & Horz (1969).6. Special part: Amphibians and reptiles of the Maghreb and Libya Scheme for the presentation of data on the species: In most species information is available for only a fraction of the following topics. Scientific denomination (author in brackets if the first description used another generic name). ‘Text figures in other chapters and plates referring to the species. ‘These references consist of the page resp. plate/ and figure resp. foto number. (eg. fig. 141/2; pl.28/79). Etymology: Derivation of the generic and specific names, mostly from Greek © or Latin (I). synonyms: Obsolete or alternative scientific denominations. fg: a, b: English, French, German, Arabic and Berber denominations. If several names are applied, the more usual or appropriate one is cited first. Regional names are followed by the region in brackets. ‘To uniformalize the different varieties of spelling of vernacular names in English or French we follow the proposal of Srumpet-Rienks (1982, pp32/ 33): Their first part is written with a capital. In composed names the subsequent words begin with a lower case, except when they are derived from geographic or personal names. In German all parts of names are written with a capital. Please take into account that the names are often artificial constructions, ied for a larger public and often derived from the scientific form. The French prefer to take the easiest way, e.g. calling Psammodromus algirus the ,,.Psammodrome algire“ or Tropiocolotes iripolitanus ,,Tropiocolote de Tripolitanie“. We did not hesitate to propose German names where none were available, generally preferring forms which correspond to the scientific denomination. Arabic and Berber denominations are of course not in conformance with scientific terms. In many cases collective names, e.g. for all frogs toads or small lizards, are used and listed for a family or genus. On the other hand many local, iruly vernacular, terms exisi. The respective region is then indicated in brackets. ‘The most important sources for Arabic and Berber names are: Le Berke (1989) who also tenders the denominations in written Arabic and Tamachek, Pasteur & Bons (1959), and Vatveroe (1957). Identification: Characters concerning size and shape. General characteristies: Typical exterior characterisation. Measurements: All measurements are given in mm. SVL: Snout-vent-length; max.: maximum, maximal. Pholidosis: Scalation (in reptiles): External Morphology (in amphibians): Pileus: Upper head plates; head sides, gular, dorsal, ventral. Scale counts follow certain rules and often require the use of a binocular microscope. A needle is very useful for pointing. Limbs, pores, tail. Coloration: Pattern elements and their variability. Colour change: Physiological (caused by rapid neural and/or hormonal processes). Developmental: Changing with age. In most lizards the hatchlings bear a juvenile pattern which during the following weeks/months gradually changes into the often different patterns of adult males and females.91 Sexual dimorphism: Differences in colour and external morphology. Similar species: Possibilities of confounding with other species living in the same region. Osteology/odontology: Details on peculiarities of the skeleton and dentition. Ecology and general behayiour ‘The terminology used by Carpenter & FERGUSON (1977) to describe act systems is applied as far as possible. Habitat: Characteristic living space; data on refuges; altitudinal distribution, Activity range: Space used during foraging excursions e.g, Data are very rare. Activity pattern: Verbal description, often graphs, which are rarely quan- titative, but depict general observations. Diel: Activity changes during the day: Diurnal, crepuscular and nocturnal, Annu: During the year with data on hibernation and estivation. ‘Thermal behaviour: Methods to regulate body temperature; temperature measurements; all data in centigrades. Locomotion: Different types. Population density: Specimens per surface unit; exact data are extremely rare (see 5.3). Population structure: Representation of age groups and sexes within a population. Data are extremely rare. Population dynamics: Fluctuation within a population caused by birth and death of specimens. Data are extremely rare. Longevity: Records are mostly from captives. Social behaviour: Behaviour released by and directed towards conspecifics. Herpetological community: Other reptiles and amphibians inhabiting the same habitat. Feeding strategy: Methods applied to acquire food (and water). Food spectrum: Type (and possibly percentage) of different food items, in some cases from siomach content or fecal pellet analyses. Predators, ext: extralimital data, often from Spai predator names is compiled in chapter 8.3. A list of vernacular Antipredator behaviour: Includes evasive or agressive behaviour toward humans. Effect of poison on man: Data for some venomous snakes, Reproduction: In some species type of spermatogenesis (see chapter 5.2.5); data on sexual maturity; mating; clutches: Number, season, size; eggs: Dimensions. oviposition: incubation and hatching. Altematively gestation and birth; hatchlings/neonates: Description, develop- ment; data on the change of colour patierns with development are also cited under colour change, developmental.92 re “Zoogeographic region(s) ot province(s) in which the species fives alo dat on eeemism and telictary areals (see also chapter 4.2). Northern Africa: Comprising the Maghreb states (Morocco, Algeria, Tunisia, Libya (LIB) and Egypt (EGY). Other regions: If the range of a species extends south into one of the adjacent states (Mauritania, Mali, Niger, Chad), a remark at the lower border of the map indicates the southernmost vegetation zone with records. PAPENFUSS (1969) characterizes them as follows: Subdesert steppe: Low grass in widely separated tufts, the bare ground between them with sand or stony flats; local low thorny trees, especially along wadis. ‘Wooded steppe: Herbaceous plants and grass do not form acontinuous cover; trees occur in thickets, but are mostly scattered. Typical are deciduous, small-leaved and thorny acacias. Dry savanna: Undisturbed grass grows 1-1.5 mhigh, trees are mostly single. ‘There are permanent rivers; pools are filled during summer rains. During the dry season fires often burn down the old grass cover. The Distribution maps show the distribution by two principal methods: A more or less continuous range is limited by a broken line, isolated records are marked with dots. Where the limits meet the shore arrows point towards the area. Limits of subspecies are indicated with a thin broken line. If maps contain ranges of several species, additional signs had to be adopted. An arrow crossing the Strait of Gibraltar indicates that the species also occurs in southern Spain, one along the eastern Mediteranean coast indicates a distribution into SW-Asi Distribution maps are ext sly susceptible to errors. Area limits are never documented by a continuous line of records in our region, and the wide spans between known localities have to be connected somehow. The unreliability of most maps is caused by quite a number of defects, some of whichare: Obsoleteness of records, some of which are more than one century old; shifts of distribution limits caused by agricultural enterprises or desertifi rarity of records from extreme desert regions, especially from Algeria and Libya; wrong records by faulty identification or subsequent division of a species into several ones; impossibility to localize old records: Old sites have since been abandoned and are no longer listed in modern ‘maps, nomads use the same name for different localities and different names for the same locality; even towns changed their names several times. Another impediment in the identification of localities is the different spelling, of Arabic or Berber names. One famous village near which the Atlas dwarf lizard (Lacerta andreanskyi) was found spells Around, Around, Arroumd, Agound, or Aremd. Any new locality record should therefore be accompanied by exact data, ¢.g. distance from a town, geographic coordinates or a map etch. A disastrous effect is caused by the import of pet animals which are bought by tourists, but also by the local population. The inclusion of Egypt into area of Testudo graeca is probably based on a single imported specimen (LA. Buskirk in litt. 16.10.1985), Systematics: Remarks on relationships. Comments: Remarks concerning systematic problems. Subspecies: Geographic ,races“ belonging to the same species with intergrades at the zones of contact. Notes: Additional general informations. References: Literature sources for informations, mostly articles in journals.93 Key to amphibians and reptiles: 1 Skin without scales: 1’ Skin with scales (they may lack on large parts of the body; e.g. Soft-shelled turtles): 2 Aquatic forms with gill respiration, tail with avery thin translucent fin : amphibian larvae 2 Aquatic or terrestrial forms with lung respiration: 3 Body slender, with tail: Caudata , Fam. Salamandridae (p. 94) 3° Tailless: ‘Anura, frogs and toads (p. 102) 4. With shell: Testudines, turtles etc. (p. M1) 4 Without shell: . 5 Four legs and/or movable eyelids: Sauria, lizards ete. (p. aii 5 Legless: 6 6 7 Dorsal scales arranged in oblique rows: _Serpentes, snakes (p. i, Dorsal scales arranged in straight longitudinal and transverse rows: ..7 Eyes covered by skin: Sauria, Amphisbaenidae (p. 464) 7 Eyes not covered; a lateral fold: Sauria, Ophisaurus koellikeri (p. 453) Amphibia Key to amphibian eggs based on Gruturrscx & al, (1989) and Pasteur & Boxs (1959), (not applicabie to the Central Sahara). 1 Eggs in strands: 5 1’ Eggs single or in lumps: 2 2 Ens onee or in Toor and small lumps: 3 2 Eggs in larger and agglutinated masses: 4 3 Eggs attached to plants: Pleurodeles 3° Eggs on bottom: Discoglossus pictus 4 Small compact clusters of yellowish eggs; egg diameter less than 5 mm: Hyla meridionalis 4 Large clusters of eggs; egg diameter over 5mm: Fam Ranidae, Rana saharica 5. Short thick eggstrands under 1 m lon and over 1 em thick, mostly around plants: Pelobates varaldii 5S’ Eggstrands long and thin, over 1 m long, and under 1 cm thick: Bufo Exact determination very risky. Key to amphibian larvae 1 Body clongate, not set off from ti is visible until metamorphosis; front limbs visible fi ‘Caudata larvae (p. 94) 1’ Body shor, distinctly set off from tail; gills are free at hatching and covered with a skin fold early during larval development, with a spiracle left n for the expulsion of respiratory water, hindlimbs visible first; mouth with external keratinized feeding organs: ‘Tadpoles, Anura larvae (p. 102). 1. Salamandra algira Fig. 2. Pleurodeles Fig. 3. Larvae: a) Pleurodeles b) Salamandra ORDER: CAUDATA Key to adults 1 Skin smooth, large parotoids (fig, 1), black with yellow spots: Salamandra algira (p. 100) 1' Skin rough, no parotoids (fig. 2): Pleurodeles......2 2 Arow of yellowish lateral spots: Pleurodeles wall! (p. 96) 2° No yellowish lateral spots: Pleurodeles poireti (p. 94) Key to Caudata larvae 1 In stagnant water; gills large, tail fin high and pointed on end (fig. 3a): Pleurodeles ‘The larvae of the two species cannot be identified specifically, but the separated areas make confounding impossible. 1" Predominantly in flowing and relatively cool water; short gills, tail fin low and rounded on end (fig. 3b); with yellow dots on the bases of the legs: Salamandra algira Family Salamandridae Genus Pleurodeles Etymology: gr. pleuron: rib; delos: visible, apparent. Large salamanders with a granular skin and flat head which ist almost triangular seen from above. Pleurodeles poireti (Gervais, 1835) Etymology: Dedicated to the botanist Abbé Point who landed in Al in 1785, visited the northern parts and wrote a ,Voyage en Barbarii Animaux amphibies de I’Algérie.* which appeared 1789. Synonyms: Lacerta palustris Linnaeus, 1758 Triton poireti Gervais, 1835 Triton nebulosus Guichenot, 1850 Glossoliga hagenmuelleri Lataste, 1881 Molge poireti Boulenger, 1891 Pleurodeles poireti Alluaud, 1923 €: Poiret’s newt, Algerian newt f: Triton de Poiret 1 Algerischer Rippenmolch At the beginning of the century this form was divided into two species: jireti* and ,Hagenmuelleri*, and at the beginning of the last century there had even been a third one, e.g. .nebulosus*. They were described on the basis of few specimens of different provenience which differed in the95 Fig. 1. Spermatophore transfer: In Pleurodeles walt! (left) the female proceeds to contact the spermatophore (x), in P. poireti (right) the couple rotates with their interlaced arms forming, the pivot. Fig, 2. Temperature induced sex determination in the two Pleurodeles species position of palatine teeth and body proportions, but later proved to be forms of a single species, Identification General characteristics: A greenish newt with very granular skin, flat head and short snout. No yellow spots on body sides. Measurements: Total length max. not much over 185 mm, tail 95 mm; a specimen which was 124 mm long had a tail of 66 mm; head length (to gular fold) 15 mm, frontleg 17 mm, hindleg 19 mm. External morphology: Skin warty, above all on the upper side of the body and the tail base. Head strongly flattened, about as broad as long, with rounded snout, broadest at the parotoids or the occipital region. Eyes rather small. Body somewhat flattened. Ribs do not penetrate the skin. Hindlegs somewhat stronger than frontlegs. During the stay in water dorsal and ventral tail fins develop. Their borders run parallel over most of the tail length. Coloration: Brown with greenish or yellowish touch; irregular dark patches without distinct borders. Head. back and tail with very small yellow or brown dots, Flanks yellowish, but in contrast to Pleurodeles walil, without yellowish spots at the rib sites. Dorsal tail crest and upper side of digits can ibe yellow to orange or pale, lower sides of limbs and body are pale grey, the latter often with round dark brown spots. Sexual dimorphism: Male with swollen cloacal region during mating period - In some males the colours are more contrasting and the dark ventral spots are much more pronounced. Reddish brown nuptial pads during mating period Tail longer than SVL in adult males, shorter in females and juveniles. Similar species: Pleurodeles waltl, which is not sympatric. Ecology and general behaviour: Almost no field records. Logevity record: 15 yr in captivity, a hybrid (x Pleurodeles walt!) 20 yr. Habitat: Mostly encountered in coo! cisterns. Antipredator behaviour: No observations, but it would be of greatest interest to know if this species, in which the ribs do not penetrate the skin, shows similar defensive acts as the closely related Pleurodeles waltl. Reprodu Mating: Act systems similar to Pleurodeles waltl, but with a different sper- matophore transfer (fig. 1): After ventral captation (fig. 1 above; fig. 98/4) the male releases the grip on one side, then, with the fixed arm as rotation axis, shifts position, creeping under the female’s venter to the side of the fixed arm until both animals are parallel, with heads showing into opposite directions. Then the spermatophore is deposited. A rotation of 180° = with the fixed arms as an axis again - brings the female into the position former taken by the male, and she can collect the spermatozoans from the spermatophore. Sex determination: This species, together with its close relative P. waltl, has shown to be an excellent laboratory object to study sex-reversal by keeping the larvae at abnormal temperatures (fig. 2). While normal 50:50, sex ratios result between 16 and 24°C, high or low temperatures can shift this relation, and small numbers of intersexes develop. The same temperatures induce different effects in the two Pleurodeles specics.Fig. 1. Lateral line system Hatchlings: SVL longer than tail length. Geographic range, Northern Africa: M & E-ALG (probably from the Algiers region eastwards), coastal region of N-TUN. ‘Zoogeography: Mediterranean, MAG, endemite. Systematics: Pleurodeles poireti and P. waltl are apparent sibling species. They crossbreed, but offspring is not very viable. Few survive meta ‘morphosis, and these then die mostly as young adults, References: BouLencer (1891), Doumerave (1901), Dournon (1990), Kuncetnorrer (1956), Le Berre (1989), Pasteur (1958). Pleurodeles walil Michahelles, 1830 (Pl WA) Etymology: Dedicated to Wat. who discovered the species inan Andalusian cistern. Synonyms: Pleurodeles walt! Michahelles, 1830 Triton costatus Wagler, 1830 Molge waltlii Boulenger, 1891 Pleurodeles walil Mertens & L. Miiller, 1940 harp-ribbed salamander, Ribbed newt, Spanish newt : Pleurodéle de Waltl, Triton ’Espagne ‘g: Spanischer Rippenmolch a: Bobreis diel ma b: Tiquajdarte oua mane, Zroleul oua mane Literature data do not always distinguish between animals of Iberian and Moroccan provenience. If possible only data on Moroccan specimens are presented. Identification General characteristics: Similar to Pleurodeles poireti, but with yellowish lateral spots. Measurements: Average SVL 58.8 mm; max. total length 252 mm, tail 112.5 mm, frontleg 39 mm, hindleg 40 mm (male from the forest of Mamora, Pasteur & Boxs, 1959). In NAF specimens aver 20 cm are very rare: the record for Iberian specimens is 40 cm/>450g. External morphology: Skin very rough with exception of the midventral region, Head very at, almost as broad as long and without distin! neck. Eyelids fused at both ends, leaving only a small part of the eyeball visible. Gular fold mostly distinct. Body depressed. The long ribs end in a sharp point and penetrate the skin during antipredator reaction. Rib penetration is released easiest in undernourished and half i specimens. Hindlegs stronger than frontlegs. The tail is si than SVL in recently metamorphosed specimens, but grows faster, attaining SVL in semiadults, surpassing it in elder specimens. Males attain longer mean tail lengths. Coloration: Basic colour dirty ochre, grey or olive green with dark brown or blackish spots; reddish in males, blackish in females; each flank with 8-1097 Fig. 1. Sex differences in arm and humerus (male at left) Fig. 2. Skull in visceral (top) and dorsal view Fig. 3. Body-arched defensive posture spots, whitish, greyish yellow to vivid orange, at the rib penetration sites, attaining a diameter of 4-5 mm in large adults; of the same colour are the gular fold, inner sides of arms, ventral side of feet, cloacal lips, tail crests and tai Sexual dimorphism; Male with swollen cloaca during mating period, frequently with dorsal and ventral tail crests, The latter can grow much larger than in Iberian specimens and appear and disappear in a very short time. Nuptial pads develop on arms (upper and lower) and hands. They are dark brown and strongly keratinized, Reddish spots appear on rib penetration sites, vertebral line, tail, finger and toe tips. The male’s humerus is thicker and carries a ventral process (fig. 1). Similar species: Pleurodeles poireti, which is allopatric. Ecology and general behaviour Habitat: Stagnant waters, in favourable conditions totally aquatic. If waters dry up or overheat they leave and seek shelter on land or under stones, logs, straw heaps, even in rather dry sites. In drying mud or clay they withdraw into fissures to a depth of 50 cm, where the soil is rather moist. One specimen was found near Azrou far from any water under a stone on rather dry soil. They have also be found 60-70 m deep in absolutely dark caves (near Ben ‘Slimane = Boulhaut, MOR). ‘Altitudinal distribution up to 1100 m (Oulmés, MOR). Activity range: Hidden specimens are often found very far from water. Activity patter, diel: Land activities are exclusively nocturnal, and in Spain ‘mass migrations occur during warm spring rains. Courtship” and feeding parently not bound to a diel cycle. Captives perform extended daylight Annual activity: Estivations during dry periods, may be crowded in great numbers. Locomotion: Small specimens are capable to ascend almost vertically on smooth surfaces. Longevity record: 20 years in captivity (kept continually in water). Feeding strategy: During their aquatic phase they are fed much easier, being atttracied by any small moving object. Kept dry they often deteriorate. Food spectrum: Any prey that can be overwhelmed; own eggs and especially larvae. In captivity fish pellets. chopped raw meat, earthworms, slugs, even baby mice. Predation: predators: ext. Nycticorax nycticorax, Bulbucus ibis, Ardea purpurea, Ciconia ciconia, Milvus milvus, Milvus migrans, Neophron percnopterus, Himantopus himantopus, Athene noctua; Rattus norvegicus. Antipredator behaviour: The exceptionally long and sharp ribs penetrate the lateral humps if the newt assumes antipredator postures; in emaciated juveniles they do so spontaneously. Toxic skin secretions from poison glands just beneath the epidermis of the dorsal body and tail are squeezed out. If the animal is harrassed, e.g. head or trunk are tapped, it assumes the body-arched-posture (fig. 3). In this posture the skin is contracted towards the middorsum and the ribs in anterior direction so that their tips are exposed (fig. 98/1) and may inflict wounds to the predator. Picked up the newtsFig. 2. Tail-elevated defensive posture Fig. 3. Vacuum captation in a captive (isolated?) male, SS Fig. 5. Larval head with balancer writhe from side to side, eventually pinching the predator between rib ends. A twitching of the tail triggers the tail-elevated posture (fig. 2). Other antipredator actions are vocalizing (the sound is transscibed as ,quotch”, nktcheck“ or ,kréék and can be very feeble); it also lunges and attempts to bite Reproduction Sexual maturity: Within 6 months after metamorphosis in well-fed juveniles. In sexually mature specimens there is no narrow neck as is in larvae. The smallest reproductive male was 47+50 mm long. Mating: In N-AFR mostly during the rainy season in winter, October to April, in captives throughout the year, above al if the aquarium is provided with fresh water. Couttshi male female sees female and approaches from behind touches her with snout creeps under the female’s venter adpresses head to female’s throat rubs throat tries to grip her frontlegs from below, rejecting female often holding her with his tail adpresses forelegs; receptive female spreads forelegs It can last up to one hour until ventral captation is achieved (fig. 4); sometimes the male holds the female with his tail rolled around her. Male carries female for hours, even days with or without interruptions, even when the female already starts to lay eggs. Both animals engulp air together at the water surface. Ifa rival male appears, the copulating male propulses the couple away with his tail. releases the grip of one foreleg bends body toward female’s head to deposit a spermatophore (disturbed by a rival he resumes the double grip and carries the female away) deposits spermatophore in front of female (see Pleurodeles poireti: fig. 95/1) leads female to spermatophore collects spermatozoa The gelatinous spermatophore is tetrahedric and 12 mm high. It carries the spermatozoa mass on top. A male produces 6-7 spermatophores during one mating season, Females mate with several males. Clutches: Oviposition starts 24-48 h after mating. The number of eggs small in juvenile females (150) and reaches 800 (even 2000) in larger ones. Eggs are fixed to stones or vegetation in groups of 9-20 which are laid in intervals of 10-15 min. If the egg supply is exhausted, two or three (even a single) eggs are laid at a time, Even one mth after mating a female can still lay fertilized eggs. The eggs are rather small (1.7mm) with a thick colloid measuring 7mm in diameter. ‘They are deposited on water plants or on the bottom. The inferior pole of the ovum is yellowish green, the superior one dark greyish green. Data on incubation period vary from 2 days to 2 weeks (at 18°C). Hatchlings: 10-14 mm, transparent, with small dots on the tail. Captivescan be fed with protozoans (introduced with a quantity of water plants), later with small crustaceans and enchytreas. Larvae: Easily to distinguish from Salamandra larvae by their much stronger developed caudal fin and gills (see key caudata larvae fig. 94/3). From March on with hindfeet. They react vehemently to noise, but not to moving objects outside the water. Sex determination: See Pleurodeles poireti p.95. Metamorphosis: At a water temperature of 25° in 3-5 mth, under most favourable conditions in 50 d. Kept at 15°C or lower the larvae are very difficult to rear because they are very prone to mycoses and other ectoparasite attacks. Captive larve need a space of 3 liters of water each to develop normally. In free-living newts the first ones metamorphose during the second half of May, the last ones (becoming somewhat larger) end October. The resorption of the gills starts with a SVL of 55-80 mm. At the same time full pigmentation is achieved. Recently metamorphosed specimens can be found in December - January under stones, even several hundred meters away from water, Total length 14 cm with one yr, 17.5 cm with 2 yr. Canrennier (in Pasteur & Bons, 1959) reports that in 1934 an old fish basin in a palace of Meknés (MOR) was repaired and the water pipes cleaned. Some months later suddenly a huge quantity of newt larve was washed into the basin where they died after a short time. This phenomenon lasted for two weeks and cart-loads of dead larvae were carried away. After this event no more of these animals could be found, and nobody in the region knew them. The larvae of Pleurodeles wali! probably developed in subterraneous reservoirs with cold waters and grew to a large size due to retarded development. They came to light when the whole system was. flooded afier the repair works. Geographic range, Northern Africa: Northeastern plains of MOR. Pleurodeles valt! p. poireti Crs Fig. 2. Cervical (above) and . | | trunk vertebrae. Other regions: $ of Iberian Peninsula. Zoogeography: W-Mediterranean.100 Fig. 1. Dorsal pattern Fig. 2. Toes of Salamandra salamandra (left) and S. algira Systematics: Pasteur (1958) cites marked differences between Spanish and Moroccan specimens and they probably belong to different forms. Spanish Moroccan maximum total length 300mm 218mm dorsal tail crest width=muscular tail x2 3 development and reduction of crests. slow fast release call frequent rare loud feeble cron squatcht* References: Baxrierr & CoLewan (1994), Bovs (1972, 1973), Dounnow & al, (1990), Grof r (1993), Hatter-Pronst & Scunsice (1994), Le Beare (1989), KLINGELHOFFER (1956), MAxTIN & Lopez (1990), MELLADO & MATEO (1992), Nusrzke (1969), Nonte (1931), Nowak & Bropie (1978), Pastour (1958), Pasteur & Bons (1963), Satvanor (1985b) Seurat (1930), StEMMLER (1965e), THoRN (1968), VALVERDE (1960, 1967), WERNER, F, (1929, 1931b), ZiveateRMANN (1983). Salamandra algira Bedriaga, 1883 PL 1/2 Etymology: The type locality is Dj. Edough near Bone (Annaba), ALG. Synonyms: Salamandra maculosa var. algira Bedriaga, 1883 Salamandra salamandra algira Scortecci, 1936 Salamandra algira Veith, 1994 ¢: Algerian salamander f: Salamandre algire g: Algerischer Feuersalamander a: Jeulleum (pl. Jlaleum), Tsetséh or Dseudséh (Tanger), Arous chta (bride of the rain) b: Tiquajderte Identification General characteristics: The N-AFR species is characterized by a slender habit with a relatively long and laterally flattened tail and relatively few yellow spots which are irregularly arranged as a rule. Measurements: Max. total 208 mm, tail 98 mm, foreleg 36 mm, hindleg 35 mm, External morphology: The N-AFR species differs from its European relati- ve by its slender shape, long forelegs, flatter toes (fig. 2) and a relatively Jonger tail which is laterally compressed. Coloration: No marked difference from Central European specimens. Sexual dimorphism: Reproductive male with swollen cloacal region, whiich is best seen from the side. Similar species: Error impossible. Ecology and general behaviour We do without rendering special informations on European Spotted salamanders as they belong to a different species and are treated extensively101 Fig. 2. Mating sequence in Salamandra salamandra CAS > = Fig. 3. Lateral line system of a S. salamandra \arva in the literature (KuzweN, 1988). However, observations on S. algira are extremely scarce. Habitat: Montane forsets, especially near caves. The biotope described by Doumercue (1901) near the mines of Rar-el-Maden lacked any open waters and the salamanders deposited their larvae in cisterns. Highest record from Imasinéne (Tell, MOR) at 1500 m. This relictary species is heavily threatened by deforestation, overgrazing and channelling of watercourses for irrigation (region of Taza, Moyen Atlas). It is much less tolerant against polluted water than Pleurodeles waltl. Activity pattern, diel: Normally nocturnal, but activity peaks during rains and fog. Annual activity: Winter activities at lower elevations and probably in caves; see Arabian name! Hiberates at higher altitudes. Locomotion: After Pasteur & Bons (1956) it is more aquatic than European Salamandra forms and swimming beter. Not only reproducing females are encountered in the water. Longevity record: Over 50, 43 and 33 yr for closely related captive European Spotted salamanders (KLEwEN, 1988), no records for algira. Predation: Ext. (on Salamandra salamandra). Buteo buteo, Gallus gallus, Strix aluco, Turdus torquatus. Reproduction As no data on Salamandra algira have been published we give a short survey of the mating sequence of Salamandra salamandra assuming it to be very similar in all species of the genus. Illustrations (fig. 2) are based on Kiewen (1988), the male is drawn in bold outline. a) After persecuting the female the male mounts and rubs her dorsal side with his chin. A receptive female does not try to flee. b) The male creeps under her. ) Ventral captation: The male seizes the female’s forelegs from below. ) The male rubs the female’s gular region and tail base. €) The male deposits a spermatophore, a 5-6 mm high gelatinous cone with an oval base of 5x3 mm. £) The male suddenly shifts the hind portion of his body sidewards and the female receives the sperm mass from the top of the spermatophore into her cloa on occurs internal. Data on larval birth are unknown for algira. Salanendra ae102 Fig. S. Oral region of tadpole Geographic range, Northern Africa: Very rare and relictary, MOR: Rif and Moyen Atlas: ALG: Coastal mountain ranges. ‘Zoogeography: Mediterranean, endemite, MAG; the range is fractioned into refuges with cool and moist climates. ‘Subspecies: The systematic state of the N-AFR form used to be uncertain (considered a subspecies of Salamandra salamandra by some authors) until Verr (1994) provided evidence for its specific status by isoenzyme electrophoresis. References: Bons (1972, 1973), Boutencer (1891), Doumeraue (1901), K.ewen (1988), Mettabo & Daxki (1988), MELLADO & Mateo (1992), Pasreur & Bons (1959, 1963), Seurar (1930), Verm (1994), Werner , F. (1931a,b).. ORDER: ANURA Collective names a: Dofdah, Dafdah, Dofdah tini. In N-MOR frog is Jrana, Drana (SW), Drain, Iran; toads are called Karkra, Kérkra (pl. Kraker). In S- MOR alll are Jrana, in LIB Duvda. b: Agriou, Ajerou. in MOR "frog" can be Agrou, Agerou, Ajrou; "toad" can bbe Alefsa, or Aliga, but in many regions one of these names is used for both frogs and toads. Key to adults (based on Pasteur & Bows, 1959): 1 Skin warty, parotoids (swollen glandular area behind eyes) present: aa 2 1" Skin smooth, possibly with isolated tubercles or glandular folds, no parotoids: 3 2 Pupil vertical, parotoids small (fig. 1): Alytes obstetricans (p. 104) (small area in N-MOR). 2 Pupil horizontal, parotoids large (figs. p. 117-126): Genus Bufo (toads), see _key on page: 115 3. Fingers and toes with adhesive disks (fig. 2): Hyla meridionalis (p. 131) (Tree frog; mediterranean MOR - TUN). 3° Fingers and toes without adhesive disks: 4 Tympanum indistinct (fig. 3,4): 4 Tympanum distinct (fig. 138/1): Family Ranidae (True frogs) see key on page 134. 5 Pupil vertical (fig. 3): Pelobates varaldii (p. 110) 5) Pupil almost circular (fig. 4): Discoglossus pictus (p. 107) 4 5 Key to tadpoles with interior gil Oral papillae and labial denticles (fig. 5) must be examined with a strong magnifying glass or a microscope. Abbreviations on fig. : SP: Supralabial papilla; IP: Infalabial p3 LP: Lateral p.: Denticle rows 1-n; HB: horny beak. Important measurements (fig. 103/1): BL: Body length; TL: Tail length; ‘TD: Tail depth. S: Spiracle.103 Fig. 1. BL: Body length, ‘TL: Tail length, TD: Tail depth S: Spiracle s = - Fig. 2. Aljtes obstetricans, ventral view. S: Spiracle Fig. 3. a: Discoglossus pictus b: Alytes obstetricans Fig. 4. a: Bufo bufo b: Rana saharica Pelobates varaldii G sa Fig. 6. a: Hyla meridionalis b: Ranidae Fig. 7. a: Rana oceipiualis b: Ptychadena ‘mascareniensis 1A midventral spiracle (fig. 2): Fam, Discoglossidae......2 V' Spiracle on the left side (fig. 1): 2. Spiracle at equal distance from snout and vent; fins with a fine polygonal dark reticulation; max. 33 mm (fig.3a): Discoglossus pictus 2 Spiracle in the anterior half of the venter (fig. 2,36); fin with a dark pattern of lines an angle of 90°; max. 80 mm: Alytes obstetricans (fig. 3b ) Only lateral oral papillae (fig. 4a): Genus Bufo, see key below Oral papillae not interrupted on lower lip (fig. 4b). 4 3-4 rows of denticles on lower lip (fig. 4b): 5 5 rows of denticles on lower lip (fig. 5); max. to 109 mm: Pelobates varaldii ae 5 Dorsal fin ending in the anterior third of body, above eye region (fig. 6a); max. 55 mm, tail depth up to 26 mm: Hyla meridionalis 5) Dorsal fin almost reaching midbody, ending above spiracle (fig, 6b): Fam, Ranidae Key to tadpoles of Fam. Ranidae (Rana saharica by far most abundaat!): 1A simple row of infralabial papillae (fig. 7a) record from S-LIB: Rana (Dicroglossus) occipitalis 1° A double row of infralabial papillae: 2 2 2 rows of supralabial and infralabial denticles each (fig. 7b): Prychadena mascareniensis (one dubious and unconfirmed record from the ,,Mare d’Ifédil, Tassili n’Ajjer, S-ALG). 2 rows of of supralabial and 3 of infralabial denticles (fig. 4): Rana saharica Larvae metamorphosing in summer reach 70 mm, those hibernating can measure over 100 mm next spring.104 Fig. 5. Bufo regularis Key to tadpoles of the genus Bufo (B. xeros not included): The identification within this very difficult group is somewhat facilitated taking into account the geographic range of its species. The examination of oral papillae is only possible in preserved specimens. 1 Median gap in 2nd supralabial denticle row very small, comprising less than 1/6 of the entire row (fig. 103/4a); lowest infralabial denticle row (nt. T, fig. 102/5) not more than 3/4 of nr. 2; body and muscular tail very dark, often black; max. 32 mm: Bufo bufo 1 Median gap of 2nd supralabial denticle row much wider (fig. 1): 2 2. Ist (lowest) infralabial denticle row not much shorter than 2nd_ row, the quotient 2nd/ist being less than 1.25 (fig. 2): 3 2 Ist infralabial denticle row much shorter, the quotient being more than 1.25 (fig.1,5): ose 3. Tail length over 4x tail depth (fig. 3a); lower horny beak (,,mandible“) V-shaped, its dark keratinized part less than 1/3 of its height (fig. 2); max. 30 mm: Bufo mauritanicus 3) Tail length under 4x tail depth (fig. 3b); max. somewhat over 30 mm: Bufo brongersmai 4 Tail length 3-4x tail depth (fig. 4a); lowest

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