Abhandlungen der Senckenberg Gesellschaft für Naturforschung

Abhandlungen der Senckenberg Gesellschaft für Naturforschung (SGN) publishes monographs with a special focus on palaeontology, zoology, biology, systematics and taxonomy. 

Abhandlungen der SGN, published by the Senckenberg Nature Research Society (SGN), Frankfurt am Main, is an international peer-reviewed journal. 

There is not schedule for publication and you may submit a monograph for publication at any time. We expect two volumes of  Abhandlungen  to be published annually. 

Before 2009: ISSN 0365-7000 
Since 2009: ISSN 1868-0356

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The most recent issue of Abhandlungen der SGN is Volume 575, published on 30th April 2021. (See below)

Zbyněk Roček, Jean-Claude Rage (†) & Márton Venczel

Fossil frogs of the genus Palaeobatrachus (Amphibia: Anura)

Palaeobatrachidae are an extinct family of frogs whose earliest members are recorded from the Middle Eocene (Geiseltal, Messel), almost 50 million years ago, although there are indications that they appeared as early as before Cretaceous mass extinction and survived almost to our times. Their last survivors are recorded from the Middle Pleistocene, less than 0.5 million years ago, such that they were contemporaries of Neanderthal man. Their last fossil evidence is from the regions that were adjacent to the Pleistocene continental glacier. Like today’s pipid frogs (e.g. Xenopus), they were obligate water dwellers, not able to make long excursions onto dry land. Without exaggerating, they literally died out frozen in their ponds.

Permanent life in water has a uniforming effect. This is why they all looked similar to one another, which is a source of difficulty for taxonomists. On the other hand, one cannot overlook features they have in common with pipids, restricted to the southern hemisphere. Palaeobatrachus and Xenopus are undoubtedly related, but when and why ancestors of Palaeobatrachus immigrated to the western part of northern Eurasia but not to its eastern part and not to North America remains a puzzling question. Thus, being acquainted with Palaeobatrachus means to be invited on an excursion not only into comparative anatomy, but also into paleogeography and many other fields of palaeontology.

Last but not least, Palaeobatrachus was the first fossil frog scientifically described, as early as in 1831. Since that time, many excavation sites from which original material came have ceased to exist; thus there is no possibility that further material will be recovered. What is now deposited in scientific collections is the unique and limited source of data for future comparative studies. This is one of the main reasons why this rich illustrated review has appeared.

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Introduction       2

Material and Methods    3
  Localities          3
  Material            3
  Character modules        5
  Determination of the relative individual age         7
  Morphometric analysis   7

Results             8
  Development of the frontoparietal during metamorphosis             8
  Frontoparietal in postmetamorphic development             9
  Notes on frontoparietal structure and its variation in adults           10
    Variation of frontoparietals in adults from Bechlejovice     10
    Interspecific variation of frontoparietals   10
  Development of the transverse processes (including ribs)            11
  Development of the synsacrum              12
  Variations of the synsacrum in adults      12
  Clustering character modules     13
  Morphometric evaluation            17

Systematic palaeontology          17
Palaeobatrachus Tschudi, 1838             17
  Palaeobatrachus tobieni (Wuttke in Sanchiz, 1998)         19
  Palaeobatrachus sp.      20
  Palaeobatrachus gracilis Meyer, 1857    21
  Palaeobatrachus novotnyi Špinar, 1972              22
  Palaeobatrachus minutus sp. nov.          23
  Palaeobatrachus bohemicus Meyer, 1860           24
  Palaeobatrachus laubei Bieber, 1881     26
  Palaeobatrachus luedeckei Wolterstorff, 1886     28
  Palaeobatrachus diluvianus (Goldfuss, 1831a)    29
  Palaeobatrachus grandipes (Giebel, 1851)         30
  Palaeobatrachus gigas Meyer, 1852       31
  Palaeobatrachus sp.      33
  Palaeobatrachus robustus Hossini & Rage, 2000            34
  Palaeobatrachus hauffianus (Fraas, 1909)          36
  Palaeobatrachus hiri Venczel, 2004        37
  Palaeobatrachus codreavladi sp. nov.     39
  Palaeobatrachus eurydices Villa, Roček, Tschopp, van den Hoek Ostende & Delfino, 2016     40
  Palaeobatrachus langhae (Fejervary, 1917)        42

Discussion        45
  Species diversity and anatomical evolution         45
  Unresolved taxonomic problems            48
  Palaeogeographic context          50

Taxa removed from the list of valid species of Palaeobatrachus   52

Acknowledgements       53

References       53

Table 1             58

Plates 1–35      63

Appendices       142

Peter Müller & Gerhard Hahn

Die Trilobiten der Erdbach-Kalke von Erdbach (Hessen) und die der „Phillipsien-Bank“ im Raum Warstein (Nordrhein-Westfalen), sowie eine Revision der Cystispininae (mittleres Mississippium)

Die Erdbach-Kalke nehmen zur Zeit des mittleren Mississippiums eine Sonderstellung innerhalb der Kulm-Fazies ein. Während in den Becken überwiegend lebensfeindliche Bedingungen herrschten, entwickelten sich auf Tiefschwellen über devonischen Riffruinen und vulkanischen Ergüssen überaus reiche Faunen. Diese lokal eng begrenzten Vorkommen sind durch hochdiverse und überwiegend endemisch geprägte Trilobiten-Faunen charakterisiert. Nach fast 130-jähriger Erforschungsgeschichte werden hier die Trilobiten der klassischen Lokalitäten im Raum Erdbach (Hessen) revidiert bzw. neu beschrieben. Zusätzlich werden die Trilobiten-Fauna der „Phillipsien-Bank“ im Raum Warstein (Nordrhein-Westfalen) sowie die Unterfamilie Cystispininae neu interpretiert.

Mit Erdbachaspis n. gen., Amphiroaspis n. gen., Liebsteinella n. gen. und Xylomelon n. gen. werden vier neue Gattungen vorgeschlagen. Zusätzlich werden 26 neue Arten bzw. Unterarten von den unterschiedlichen Lokalitäten beschrieben: Liobole (Liobole) meyeri n. sp., Liobole (Loibole) io n. sp., Liobole (Panibole) subaequalis hschmidti n. ssp., Liobole (Panibole) emrichterae n. sp., Liobole (Panibole) paraglabroides n. sp., Liobole (Panibole) isis n. sp., Liobole (Panibole) groeningae n. sp., Cyrtoproetus pegasus n. sp., Cyrtoproetus galapagos n. sp., Cyrtoproetus eremus n. sp., Crassibole ate n. sp., Crassibole? walliseri n. sp., Latibole pleon n. sp., Latibole gamma n. sp., Latibole parapaprothae n. sp., Brevibole archinalae n. sp., Brevibole halszkae n. sp., Chlupacula (Chiides) pandora n. sp., Diacoryphe (Diacoryphe) phoenix n. sp., Belgibole kalypso n. sp., Spatulata (Spatulata) tilsleyi n. sp., Spatulata (Spatulata?) pachycraspedata n. sp., Tawstockia longispina germanica n. ssp., Liebsteinella sauerlandica n. sp., Liebsteinella weberi n. sp. und Xylomelon xenios n. sp.

Die „Diagnosen“ sind zusätzlich in englischer Sprache verfasst und erleichtern dem internationalen Leser somit den Zugang zu dieser Abhandlung.

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Problematik und Zielsetzung
Die Fundpunkte und ihre jeweiligen Trilobiten-Assoziationen
  Liebstein (Erdbach)
  Kramberg (Erdbach)
  Neumühle (Erdbach)
  Homberg (Kulm-Steinbruch, Erdbach)
  Massenkalk-Bruch (Erdbach)
  „Phillipsien-Bank“ (Kattensiepen-Member) im Raum Warstein
  Faunen aus karbonatischer Kulm-Fazies
  Faunen aus schiefriger Kulm-Fazies
Systematische Paläontologie
Stamm Arthropoda von Siebold,
  Unterfamilie Archegoninae G. Hahn & Brauckmann,
  Unterfamilie Weaniinae Owens,
  Unterfamilie Cystispininae Hahn & Hahn, sensu novo
  Unterfamilie Cummingellinae Hahn & Hahn,
  Unterfamilie Bollandiinae G. Hahn & Brauckmann,
Tafeln 1–21

Haytham El Atay, Rainer Brocke & Dieter Uhl 

Palynology of the Miocene Rudeis and Kareem formations (Gharandal Group), GH 404-2A Well, Gulf of Suez, Egypt

The Gulf of Suez is the main oil province in Egypt, with oil being produced from Paleozoic, Mesozoic and Cenozoic rocks. The Miocene sediments are the most prolific hydrocarbon-bearing sequences both in onshore and offshore fields. Major research activities have been so far concentrated on structural geology, geophysics, organic geochemistry, stratigraphy, and micropaleontology. In contrast, very little has been published on the Miocene palynology from this region.

A comprehensive palynological investigation on the GH 404-2A Well (southern part of the Gulf of Suez), comprising the Miocene Rudeis and Kareem formations of the Gharandal Group has produced diverse palynomorph assemblages. They contain dinocysts, spores, pollen, algae, fungal palynomorphs, scolecodonts and microforaminiferal linings. For the first time in Egypt, the present study introduces a detailed systematic overview of the palynomorphs with respect to their biostratigraphy and paleoenvironment within this time interval. Biostratigraphically, three dinocyst and one sporomorph biozones are established and correlated with other Miocene schemes from Egypt and other areas worldwide.

In addition, the palynomorphs are used to get further insights both into the adjacent land vegetation and climate of the Gulf of Suez area, during the Miocene, and relate these paleoenvironmental conditions to depositional processes in the Gulf of Suez basin.

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Geologic setting   
   Rudeis Formation   
   Kareem Formation
Previous palynological studies on the Miocene of Egypt – an overview   
Material and methods   
   Samples and overview    
   Extraction methods    
   Presentation of palynological results   
Systematic palynology   
   Dinoflagellate cysts   
   Sporomorphs (spores and pollen)
   Green and blue-green algae   
   Fungal palynomorphs   
Biostratigraphy and age assignment    
   Dinoflagellate cyst biozones  
   Sporomorph biostratigraphy and age assignment    
Paleoenvironmental interpretations   

Martin Basse & Peter Müller 

Trilobiten aus dem Ober-Emsium und frühen Eifelium der südlichen Lahnmulde (Rupbach-Schiefer, Leun-Schiefer und Ballersbach-Kalk)

Der als autochthon gewertete Rupbach-Schiefer der südwestlichen Lahnmulde (Rhenoherzynikum) ist ein Sediment des Epikontinentalschelfs des nördlichen Rheia-Ozeans vor der Küste Ostavalonias. Er umfasst mittleres Ober-Emsium bis jüngeres Givetium (Unter- und Mitteldevon). Abschnitte bis zum frühen Eifelium liefern eine sukzessiv verarmende Trilobitenfauna. Der tiefe Rupbach-Schiefer führt die geologisch älteste biofaziell rheinisch-herzynische Mischfauna im rechtsrheinischen Schiefergebirge mit sechs Ordnungen, 14 Unterfamilien und 16 Gattungen (mit Scabriscutellum (Rheiscutellum) und Barrandeops (Lahnops) als neuen Untergattungen) mit bis auf eine Ausnahme je einer Art. Acht davon neu: Cyphaspis kweberin. sp., Diademaproetus habenichti n. sp., Rhenocynproetus vanvierseni n. sp., Tropidocoryphe werneri n. sp., Acastoides (?n. subgen.) poschmanni n. sp., Destombesina schumacherorum n. sp., Barrandeops (Lahnops) steinmeyeri n. subgen. n. sp., Koneprusia martini n. sp. 

Diese Fauna wirkt stärker gondwanisch als jede andere bekannte rheinisch-herzynische Fauna aus dem autochthonen Unterdevon des deutschen Rhenoherzynikums. Noch im Ober-Emsium wird sie durch eine herzynische mit Cornuproetinae abgelöst. Die jüngste, herzynische Fauna (frühes Eifelium) ist mit vier Ordnungen, fünf Unterfamilien und sechs Gattungen mit je einer Art formenarm. Aulacopleura (Paraaulacopleura) lemkei n. sp. und Cyphaspides (Cyphaspides) malbertii n. sp. sind neu. Cyphaspides (Cyphaspides) weugi n. sp. (Deutschland, frühes Eifelium), Pelitlina? corbachoi n. sp. (Spanien, Unter-Emsium) und Psychopyge angeles n. sp. (Marokko, Ober-Emsium) liefern ergänzende Daten. Faunistisch vergleichbar sind vor allem folgende Gebiete mit Sedimenten des Rheia-Ozeans: Die nordwestliche Türkei, Tschechien, andere Teile Deutschlands, Nordwestafrika. Jedoch bestehen für den Rheia-Ozean noch zu große Kenntnislücken, um dies paläo(bio)geografisch auszuwerten.

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Content (in parts)

Allgemeiner Teil    
   Rupbach-Schiefer: Geologische und biostratigrafische Übersicht    
   Paläogeografie und Fazies     
   Status quo der Trilobitenforschung     
   Potenziale der Trilobitenfaunen    
   Potenziale einzelner Taxa    
   Zielsetzung und Methodik    
   Profile und Stratigrafie    
   Stratigrafie und Fauna von einigen verglichenen Vorkommen     
Systematische Paläontologie   
   Stamm Arthropoda von Siebold 1845     
   Unterstamm Trilobitomorpha Størmer 1944    
   Klasse Trilobita Walch 1771      
   Ordnung Corynexochida Kobayashi 1935     
   Unterordnung Illaenina Jaanusson 1959    
   Ordnung ?Aulacopleurida Adrain 2011       
   Ordnung Proetida Fortey & Owens 1975     
   Ordnung Harpetida Whittington 1959b    
   Ordnung Phacopida Salter 1864     
   Unterordnung Phacopina Struve 1959a    
   Ordnung Lichida Moore 1959     
   Ordnung Odontopleurida Whittington 1959a    
Tafeln 1–33     
   A. Tabellen 1–94 des Untersuchungsmaterials und seiner Maße     
   B. Trilobiten aus „exotic limestones“ des Lahn-Dill-Gebiets im Vergleich zu denen des Rupbach-   Schiefers im weiteren Grenzbereich Emsium/Eifelium

Josef Gandl, Enric Ferrer, Josep Magrans & Javier Sanz López 

Trilobiten aus dem Unter-Karbon des Katalonischen Küstengebirges (NE-Spanien)

Aus 4 Lokalitäten des Katalonischen Küstengebirges (NE-Spanien), nämlich Aiguafreda, Cànoves, Papiol und Scala Deï werden insgesamt 18 Trilobiten- (Unter-) Arten der Proetidae-Unterfamilien Drevermanniinae, Cyrtosymbolinae und Mirabolinae beschrieben. Eine Gattung, 4 Untergattungen und 16 (Unter-) Arten, davon 15 voll benannt, sind neu beschrieben.

Als Folge zeichnen sich zunächst zwei nahe beieinander liegende Zeithorizonte ab. In den ersten drei der vier Lokalitäten entstammen die Trilobiten der „Formación El Papiol“, deren Fazies und Fauna auch an der N-Küste von Menorca vorkommt. Durch den Faunenvergleich mit dieser Insel und einigen Conodonten der bilineatus-Zone wird schließlich ein tieferes Ober-Visé Alter, etwa Go α4/?beta;str, wahrscheinlich. So gut wie alle Trilobiten sind blind oder fast blind und zeigen einen überaus flachen Körperbau. Sie werden zudem von einer benthonischen „Kümmerfauna“ begleitet, was auf ein Leben am Boden eines tieferen Schelfs schließen lässt und auf die Fähigkeit, sich im schlammigen Sediment einzugraben.

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Lage und Alter der Trilobiten-Fundpunkte
   El Papiol
   Altersstellung der Fundpunkte Papiol 1-3 sowie Aiguafreda und Cànoves
   Scala Deï
Beschreibung der Trilobiten (Josef Gandl)
Superfamilie Proetacea Hawle & Corda 1847
   Familie Proetidae Hawle & Corda 1847
   Unterfamilie Drevermanniinae Maximowa 1960
   Drevermannia Rud. Richter 1909 
   Drevermannia (Pseudodrevermannia) P. Müller & Brauckmann 2010 
   Drevermannia (Paradrevermannia) n. sg.
   Menorcaspis G. & R. Hahn & Brauckmann 1994 
   Brachymetaspis n. g.
   Unterfamilie Cyrtosymbolinae Hupé 1953
   Semiproetus Reed 1943 
   Weyeraspis G. & R. Hahn & Gawlick 1995 
   Weyeraspis (Canovesia) n. sg.
   Chlupacula G. Hahn & Wunn-Petry 1983 
   Chlupacula (Chlupacula) G. Hahn & Wunn-Petry 1983
   Chlupacula (Avenconia) n. sg.
   Unterfamilie Mirabolinae Yuan & Xiang 1998
   Liobole Rud. & E. Richter 1949
   Liobole (Quadratibole) n. sg.
Ergebnisse, Folgerungen und Vergleich
Tafel 1 – 10

Margaret E. Collinson, Steven R. Manchester &Volker Wilde 

Fossil Fruits and Seeds of the Middle Eocene Messel biota, Germany

The oil shale of the Middle Eocene Messel Formation as exposed in the Messel Pit near Darmstadt, Hessen, Germany, is a famous and widely known source for extremely well preserved fossils documenting a wealth of terrestrial biota. For this reason the Messel Pit Fossil Site was awarded the status of a UNESCO World Heritage Site documenting the ecosystem on the European mainland which existed under a paratropical climate during Paleogene greenhouse conditions.

A survey of the extensive fruit and seed collections from the Middle Eocene oil shale of the Messel Formation now reveals at least 140 genera, representing more than 34 families of seed plants. The flora includes occasional conifer and numerous angiosperm remains. There are 34 extant angiosperm families represented of which ten are new records for Messel, plus 65 morphotypes of unknown familial affinity. Three extant genera are recorded for the first time from the Paleogene. The assemblage indicates a wide range of dispersal strategies including pods, capsules, explosive dehiscence, a single arillate seed, two seed-types with dispersal hairs and most modern categories of winged disseminules. In terms of mammalian frugivory the flora contains examples of all potential dietary categories. Tough and hard materials are abundant and soft material is common. Gut contents preserved in many birds and mammals prove that fruits and seeds played an integral part in vertebrate diets and borings in one seed type indicate seed predation by weevils. Previous quantitative studies suggesting an equable warm and humid palaeoclimate with some seasonality for Messel are supported by the newly recognised taxa. Judging from the habit of related living taxa, the vegetation appears to have been a multilevel canopy forest, including a high proportion of lianas in addition to shrubby to arborescent taxa. Herbaceous components are also present but relatively underrepresented. Among other large and well-studied Eocene macrofloras, the Messel assemblage shows overlap with the genera known from the London Clay flora of England and the Clarno Nut Beds Flora of Oregon, but relatively little similarity with floras known from eastern Asia. Compared with extant floras, the Messel flora includes a temperate component with mostly Asian endemics, and some genera that are now disjunctly distributed in the Northern Hemisphere. A large tropical-paratropical component includes genera now confined to the Old World tropics, particularly southeastern Asia and Malaysia, but there are also a few exclusively Neotropical elements.

Altogether, the Messel flora can be regarded as one of the most diverse Paleogene floras worldwide.

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Content (in parts)

Geologic setting and age
Modes of preservation
Material and methods
Floristic composition
Comparison with diversity known from leaves and pollen
Fruit and seed biology, dispersal and animal diets
Taphonomic considerations 
Growth habits and vegetation reconstruction
Climatic interpretations
Biogeographic considerations
Future directions for Messel palaeobotanical research
Family Doliostrobaceae KVAÈEK, Family Alangiaceae DC., Family Altingiaceae LINDL., Family Anacardiaceae R. BR., Family Apocynaceae JUSS., Family Arecaceae BERCHT. & J. PRESL,Family Bignoniaceae JUSS., Family Burseraceae KUNTH 1824, Family Cannabaceae MARTINOV, Family Cyclanthaceae POIT. ex A. RICH., Family Cyperaceae JUSS., Family Elaeocarpaceae JUSS., Family Euphorbiaceae JUSS., Family Hamamelidaceae R. BR., Family Icacinaceae MIERS, Family Juglandaceae DC., Family Lauraceae JUSS., Family Leguminosae JUSS., Family Lythraceae J. ST.-HIL., Family Magnoliaceae JUSS., Family Mastixiaceae CALEST., Family Menispermaceae JUSS., Family Myristicaceae R. BR., Order ?Nymphaeales, Family Nyssaceae JUSS. ex DUMORT., Family Pentaphylacaceae ENGL., Family Rhamnaceae JUSS., Family Rutaceae JUSS., Family Sabiaceae BLUME, Family Sapotaceae JUSS., Family Simaroubaceae DC., Family Tapisciaceae TAKHT., Family Theaceae MIRB., Family Toricelliaceae HU, Family Ulmaceae MIRB., Family Vitaceae JUSS. 
Incertae Sedis

Josef Gandl

Die Karbon-Trilobiten des Kantabrischen Gebirges (NW-Spanien), 5: Trilobiten des höheren Westfal

This paper is the fifth part of a comprehensive study, dealing with the Carboniferous trilobites of the Cantabrian Mountains. Trilobites are described from more than 90 different localities. With the exception of two localities of Westphalian A (Langsettian) age and a few ones corresponding to the lower Cantabrian substage of the basal Stephanian, they all belong to the Westphalian C (Bolsovian) or D (Asturian), thus following closely upon the late Namurian and early Westphalian trilobites described in 1987. The localities involved mainly occur in 4 principal areas, viz (1) the region of the “Manto del Ponga”, east of the Central Asturian coalfield (province of Oviedo); (2) the area around the “Puerto del Pando” (province of León); (3) the Casavegas Syncline (province of Palencia); and (4) the southern flank of the Castillería Syncline (province of Palencia). Additionally, an upper Westphalian trilobite locality is recorded from the Sierra de la Demanda (at Valmala, province of Burgos) for the first time.

Most of the localities are dated quite precisely, with reference to brachiopods, foraminifera (Fusulinacea) and/or land plants, and are situated in well-studied sections. On the other hand, it has been possible to propose certain correlations by means of trilobites with a higher degree of precision than it had been feasible previously.

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Content (in parts)

Vorwort und Dank 
Lage und Alter der wichtigsten Trilobiten-Fundpunkte 
   I. Die Fundpunkte östlich des Zentralasturischen Kohlebeckens im Bereich der „Manto del Ponga“ 
   II. Die Fundpunkte PR 1 – 14 in der Pando-Region (Provinz León)
   III. Die Fundpunkte CS 1 – 10 in der Casavegas-Synkline (Provinz Palencia)
   IV. Die Fundpunkte RS 1a – c in der Redondo-Synkline (Provinz Palencia) 
   V. Die Fundpunkte NCS 1 und NCS 2a – b an der Nordflanke der Castillería-Synkline (Provinz Palencia)
   VI. Die Fundpunkte SCS 1 – 4 an der Südfl anke der Castillería-Synkline (Provinz Palencia) 
   VII. Ergänzung: Der Fundpunkt Valmala am N-Rand der Sierra de la Demanda, (Provinz Burgos)
Beschreibung der Trilobiten 
Superfamilie Proetacea HAWLE & CORDA 1847 
   Familie Phillipsiidae OEHLERT 1886 
   Unterfamilie Archegoninae G. HAHN & BRAUCKMANN 1984 
   Unterfamilie Weaniinae OWENS 1983 
   Unterfamilie Cummingellinae G. & R. HAHN 1967  
   Unterfamilie Bollandiinae G. HAHN & BRAUCKMANN 1988  
   Unterfamilie Ditomopyginae HUPÉ 1953 
   Familie Brachymetopidae PRANTL & PRIBYL 1950  
   Unterfamilie Brachymetopinae PRANTL & PRIBYL 1950
Tafel 1 – 15 

Andrej Ernst & Peter Königshof 

Bryozoan fauna and microfacies from a Middle Devonian reef complex (Western Sahara, Morocco)

Devonian bryozoans from North Africa are scarcely known. The present study is devoted to the investigation of bryozoan fauna and microfacies of a Middle Devonian reef complex in Western Sahara, Morocco. In total, 26 bryozoan species were identifi ed. Two genera with two species are new: Lenapora pulchra n. g. n. sp. and Dissotrypa sincera n. g. n. sp. Furthermore, 15 new species were described: Fistuliphragma parva n. sp., Sulcoretepora moderata sp. n., Leioclema crassiparietum n. sp., Eridotrypella minutiformis n. sp., E. modesta n. sp., Atactotoechus gaetulus n. sp., Acanthostictoporella angusta n. sp., Euspilopora spinigera n. sp., Acanthoclema triangularis n. sp., Rhombopora minutula n. sp., Prolixicella lata n. sp., Rhombocladia striata n. sp., Filites gaetulus n. sp., Anastomopora clara n. sp., and A. recta n. sp. Four species are described in open nomenclature: Spinofenestella sp., Hemitrypa sp., Ptylopora sp., and Semicscinium sp. The studied fauna shows relations to the Middle Devonian of Holy Cross Mountains, Poland and Rhenish Massif, Germany, as well as to the Middle Devonian of North America.


Geological setting 
Microfacies analysis
Taxonomic descriptions 
   Taxonomic overview and palaeobiogeographic implications
Systematic palaeontology 
Plates 1 – 27 

Heinz Hofmann 

The glass frog tadpoles of Costa Rica (Anura: Centrolenidae): A study of morphology

Larvae of the neotropical frog family Centrolenidae live hidden on the bottom of rivers between stones, gravel, and leaf litter. Their lotic fossorial behavior makes it impossible to find and study these tadpoles in their natural environment. Therefore, presently only poor reports and descriptions of them are available in the literature.

In the present study all known 13 Costa Rican centrolenid species (or taxa) were reared under laboratory conditions. Their live tadpoles are described in different stages of their development. Six of the 13 descriptions are new (Cochranella euknemos, Sachatamia albomaculata, Sachatamia ilex, Hyalinobatrachium chirripoi, Hyalinobatrachium talamancae and Hyalinobatrachium vireovittatum). A species key and detailed diagnostic descriptions with abundant illustrations allow the identification of tadpoles of the thirteen species. 


Material and Methods
Genus Cochranella
   Cochranella euknemos (Savage & Starrett 1967)                                         
   Cochranella granulosa (Taylor 1949)
Genus Espadarana
   Espadarana prosoblepon (Boettger 1892)
Genus Sachatamia
   Sachatamia albomaculata (Taylor 1949)
   Sachatamia ilex (Savage 1967)
Genus Teratohyla
   Teratohyla pulverata (Peters 1873)
   Teratohyla spinosa (Taylor 1949)
Genus Hyalinobatrachium, Species group H. fleischmanni
   Hyalinobatrachium fleischmanni (Boettger 1893)
   Hyalinobatrachium valerioi (Dunn 1931)
Genus Hyalinobatrachium, Species group H. chirripoi
   Hyalinobatrachium chirripoi (Taylor 1958) 
   Hyalinobatrachium colymbiphyllum (Taylor 1949)
   Hyalinobatrachium talamancae (Taylor 1952) 
   Hyalinobatrachium vireovittatum (Savage & Starrett 1973) 
Key for the mature centrolenid tadpoles of Costa Rica