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11 pages, 3 figures, 1 table, supplementary material https://doi.org/10.1126/sciadv.abm7452.-- Data and materials availability: All data needed to evaluate the conclusions in the paper are present in the paper, the Supplementary Materials, Dryad, and Zenodo, Phylogenetic relationships and the timing of evolutionary events are essential for understanding evolution on longer time scales. Cheilostome bryozoans are a group of ubiquitous, species-rich, marine colonial organisms with an excellent fossil record but lack phylogenetic relationships inferred from molecular data. We present genome-skimmed data for 395 cheilostomes and combine these with 315 published sequences to infer relationships and the timing of key events among c. 500 cheilostome species. We find that named cheilostome genera and species are phylogenetically coherent, rendering fossil or contemporary specimens readily delimited using only skeletal morphology. Our phylogeny shows that parental care in the form of brooding evolved several times independently but was never lost in cheilostomes. Our fossil calibration, robust to varied assumptions, indicates that the cheilostome lineage and parental care therein could have Paleozoic origins, much older than the first known fossil record of cheilostomes in the Late Jurassic, This project is funded by the European Research Council (ERC) under the European Union’s Horizon 2020 research and innovation programme (grant agreement no. 724324 to L.H.L.). Extra funding for specific sampling expeditions were provided by the National Science Centre of Poland (grant PANIC/2016/23/B/ST10/01936 to P.K.), the ASSEMBLE Plus (Horizon 2020), the Leverhulme Trust (Research Project Award RPG-2016-429 to A.W.), the Russian Science Foundation (grant 18-14-00086 to A.N.O.), Conselho Nacional de Desenvolvimento Científico e Tecnológico, Brazil [PQ-CNPq 308768/2018-3], Fundação de Amparo à Pesquisa do Estado de São Paulo, Brazil [FAPESP 19/17721-9] and the DISTANTCOM project (CTM2013-42667/ANT to C. Avila). B.F. was supported by Beatriu de Pinós (2019-BP-00183), funded by the AGAUR (Government of Catalonia) and by the Horizon 2020 programme under the Marie Skłodowska-Curie grant (no. 801370). The CAML-CEAMARC cruise of RV Aurora Australis (IPY project no. 53) were supported by the Australian Antarctic Division, the Japanese Science Foundation, the French Polar Institute IPEV, and the Muséum national d’Histoire naturelle, led by voyage leader, M. Riddle, With the institutional support of the ‘Severo Ochoa Centre of Excellence’ accreditation (CEX2019-000928-S)

Located on the northern coast of Santa Maria Island (Azores Archipelago, central North Atlantic), the Lagoinhas section preserves a carbonate buildup correlated with Marine Isotope Substage (MIS) 5e, the warmest interval of the Last Interglacial. The buildup is formed mainly by crustose coralline algae (CCA) identified as Spongites sp., and some subordinate crusts of Lithophyllum sp. and Neogoniolithon sp., as well as cf. Titanoderma sp. Extant CCA buildups are not recorded in the archipelago. Herein, we describe in detail the morphological and taphonomical features of the Lagoinhas CCA buildup and interpret the environment in which it grew. Additionally, this buildup is compared with another of similar age, exposed in the Prainha-Praia do Calhau section on the island's opposite southern coast. The hydrodynamic regime appears to play a crucial role in the development of Azorean CCA buildups during the MIS 5e. We thank Direccao Regional da Ciência e Tecnologia (Regional Government of the Azores), FCT (Fundação para a Ciência e a Tecnologia) of the Portuguese Government, and Camara Municipal de Vila do Porto for financial support. We also acknowledge the field assistance of Camara Municipal de Vila do Porto. We are grateful to the organizers and participants of the 16 editions of the International Workshop Palaeontology in Atlantic Islands who helped with fieldwork (2002 2020). A.C.R. was supported by a grant SFRH/BPD/117810/2016 from FCT (Fundação para Ciência e Tecnologia), Portugal. S.P.A. and R.S.R. acknowledge their research contracts (IF/00465/2015 and IF/01641/2015, respectively) funded by Fundação para a Ciência e a Tecnologia (FCT). C.S.M. is benefiting from a PhD grant M3.1.a/F/100/2015 from Fundo Regional para a Ciência e Tecnologia (FRCT). This work was supported by FEDER funds through the Operational Programme for Competitiveness Factors -COMPETE, by Portuguese National Funds through FCT under the projects UID/BIA/50027/2013, UID/BIA/00329/2013-2023, POCI-01-0145-FEDER-006821, PLATMAR (PTDC/GEO-GEO/0051/2014) and under DRCT-M1.1.a/005/Funcionamento-C-/2016 (CIBIO-A) project from FRCT, and by FEDER funds (in 85%) and by funds of the Regional Government of the Azores (15%) trough Programa Operacional Acores 2020, in the scope of the projects "AZORESBIOPORTAL -PORBIOTA": ACORES-01-0145FEDER-000072 and VRPROTO -Virtual Reality PROTOtype: the geological history of "Pedra-que-pica": ACORES-01-0145-FEDER000078. A.U. benefited from additional support by the Jagiellonian University. info:eu-repo/semantics/publishedVersion

The Turonian–Coniacian continental fossil record in Europe is scarce. Here we present a new fossil assemblage of early Coniacian age that was systematically collected from the coal-bearing Gosau Group of the Tiefengraben locality near St. Wolfgang, Austria. The diverse assemblage is composed of at least 60 taxa including sporomorphs and Normapolles-related pollen, seeds and leaves of angiosperms and gymnosperms, charophytes, gastropods, bivalves, ostracods, termites, fishes, crocodiles and dinosaurs. Concerning charophytes, ostracods, gastropods, crocodiles and dinosaurs, the discovered specimens either extend the temporal and spatial range of specific groups (in some cases as possible relict forms) or suggest the occurrence of new taxa. The discovered remains of algae, molluscs, ostracods, calcareous nannofossils and lepisosteid fish represent a mixed faunal assemblage from different palaeohabitats, from marginal marine to low salinity and freshwater or terrestrial environments. As Normapolles-related angiosperm plants dominate the flora with a relatively high number of dentate leaves, a slightly cooler microenvironment compared to other Turonian–Coniacian Central European localities is indicated. The characteristically grooved crocodylian teeth of Tiefengraben differ from the previously known Upper Cretaceous European crocodyliform teeth and suggest a more diverse crocodyliform fauna in the region. Dinosaurs are represented by teeth of at least three different theropods, the largest of which is referred here to as basal tetanurans. The fossil assemblage of this early Gosau Group occurrence is of great importance for our understanding of the continental floristic and faunistic composition of the western Tethyan archipelago during the Cenomanian–Campanian gap.

The Mediterranean bryozoan fauna is considered to be well studied compared to other marine areas of the world. However, in the Levantine Basin, bryozoan diversity has not yet been adequately documented. This report presents the first systematic and most comprehensive study of bryozoans sampled in Cyprus during census surveys from 2011 to 2018. The specimens were collected between 9 and ~620 m depth from several habitat types (mainly soft-bottom environments but also hard natural/artificial substrata, ancient shipwrecks, a marine cave, and deep-water coral habitats) around the island by means of bottom trawls, remotely operated vehicles, and scuba diving. The surveys produced a total of 91 species, 26 of which (=28%) are new records for the Levantine Basin, and 10 (=11%) are probably new to science. Our results thus show that the diversity of bryozoans in the eastern Mediterranean Sea is still significantly underestimated.

The bowl-shaped trace fossil Piscichnus waitemata Gregory 1991 appears in Pliocene sandstones from Santa Maria Island (Azores Archipelago), extensively excavated during a stage of island evolution when the volcanic edifice was a guyot (flat-topped seamount) isolated in the NE Atlantic. The host sediments were deposited at depths from the intertidal zone to fair-weather wave base in a tropical climate under the influence of periodic storms and hurricanes. The traces were produced by ray fishes hunting for polychaetes, crustaceans and bivalves living in the sediment, similar to present-day nearshore, warm waters in the Azores, Baja California Sur (Mexico), and New Zealand, from which examples of feeding depressions are drawn (incipient Piscichnus). While P. waitemata is abundantly present in planar sediments on top of the guyot, far fewer trace fossils occur in sandstone deposited on the guyot's margins. Presumably, the different densities of ray holes in the two sedimentary bodies were a response to lesser availability of prey, lower seawater temperatures (due to greater depths), and a more dynamic environment in which life conditions were less favorable. Moreover, the potential preservation of bowl-shaped depressions was lower in this setting, given the steepness of the seafloor, stronger currents, and constant sediment mobility. The top of the guyot was a more favorable habitat, refuge and/or nursery ground for many ray fishes. Measurement of the diameters of the ray holes indicate three distinct size classes, which may suggest that several species were responsible for their formation.

The Azores, Madeira, Selvagens, Canary Islands and Cabo Verde are commonly united under the term “Macaronesia”. This study investigates the coherency and validity of Macaronesia as a biogeographic unit using six marine groups with very different dispersal abilities: coastal fishes, echinoderms, gastropod molluscs, brachyuran decapod crustaceans, polychaete annelids, and macroalgae. We found no support for the current concept of Macaronesia as a coherent marine biogeographic unit. All marine groups studied suggest the exclusion of Cabo Verde from the remaining Macaronesian archipelagos and thus, Cabo Verde should be given the status of a biogeographic subprovince within the West African Transition province. We propose to redefine the Lusitanian biogeographical province, in which we include four ecoregions: the South European Atlantic Shelf, the Saharan Upwelling, the Azores, and a new ecoregion herein named Webbnesia, which comprises the archipelagos of Madeira, Selvagens and the Canary Islands., SCOPUS: ar.j, info:eu-repo/semantics/published

Hazards and disasters have occurred throughout Earth's History and thus the geological record is an important resource for understanding future hazards and disasters. The Earth Science Group (ESG) of the Consortium of European Taxonomic Facilities (CETAF) carried out a “Hazard and Disaster Event Survey” to identify Earth Science collections in European museums that represent hazards and disasters throughout the geological record, and recent times. The aim is to use the collections within the survey as an educational and research resource that promotes the importance of museum collections for understanding past and future hazard and disaster events. The survey pinpointed a wide variety of hazards (e.g. earthquakes, volcanism, floods, impact events, etc.), representing a vast time span in Earth’s history (Proterozoic to Holocene), that are documented in the collections of the participating museums. Each hazard and disaster event has been described in terms of how they are preserved (e.g. fossil record or rock record), spatial scale, impact on life, and geological age. Here we showcase seven examples in detail which include well-known and less-known events from the survey that have contributed to our understanding of hazard and disaster processes and their impact on life. Also we present general conclusions and lessons learnt from the “Hazard and Disaster Event Survey”.

Pliocene body fossils from Santa Maria Island, Azores, have been studied for decades, but only more recently have ichnofossils received their due attention. Calcareous Pliocene deposits from the Baia de Nossa Senhora section contain numerous, diverse, well-preserved natural casts of invertebrate borings. The study of this type of fossils adds to knowledge on the dispersal of benthic faunas across oceans to volcanic oceanic islands. The borings belong to seven ichnogenera and twenty-two ichnotaxa at the ichnospecies level with more than half pertaining to Entobia, which is produced by clionaid sponges. Other borings found were produced by bivalves (Gastrochaenolites), polychaete worms (Caulostrepsis and Maeandropolydora), sipunculid worms (Trypanites), phoronid worms (Talpina) and ctenostome bryozoans (Iramena). The occurrence, ichnogeny, distribution and preservational state of the borings suggest that the bearing bioclasts have been exposed for several years on the sea floor. The borings derive from different bathymetric zones on the shelf, and their formation took place during several bioerosional phases. The association of borings belongs to the Entobia ichnofacies, which is typical of carbonate rocky shores, and shows close similarity to those described from the Paratethys, Mediterranean and partly the eastern Atlantic regions. This fits the idea that most of the Neogene shallow-water marine fauna in the Azores is biogeographically related to the eastern Atlantic shores.

Hazards and disasters have occurred throughout Earth's History and thus the geological record is an important resource for understanding future hazards and disasters and its Impact. The Earth Science Group (ESG) of the Consortium of European Taxonomic Facilities (CETAF) carried out a “Hazard and Disaster Event Survey” to identify Earth Science collections in European museums that represent hazards and disasters throughout the geological record, and recent times. The aim is to use the collections within the survey as an educational and research resource that promotes the importance of museum collections for understanding past and future hazard and disaster events. The survey pinpointed a wide variety of hazards (e.g. earthquakes, volcanism, floods, impact events, etc.), representing a vast time span in Earth’s history (Proterozoic to Holocene), that are documented in the collections of the participating museums. Each hazard and disaster event has been described in terms of how each is preserved (e.g. fossil record or rock record), spatial scale, impact on life and biodiversity, and geological age. The results were published in Research ideas and outcomes journal RIO (https://riojournal.com/article/34087/) as seven examples, which include well-known and less-known events from the survey that have contributed to our understanding of hazard and disaster processes and their impact on life and biodiversity. For better visualisation and education purposes we also created and specialised website, which will be launched during summer 2019. In the presentation we will talk about general conclusions and lessons learnt from the “Hazard and Disaster Event Survey”.

The family Strombidae is one of the twenty-three families and subfamilies of gastropods associated with tropical environmental conditions, and therefore useful as a biogeographical and paleoclimatic proxy. Today, the strombid genus Persististrombus is represented in the NE Atlantic by a single species restricted to the tropical Mauritanian–Senegalese Province. This work reports the occurrence of Persististrombus coronatus from the lower Pliocene of Santa Maria Island in the Azores Archipelago. Based on this occurrence, and on the Mio-Pliocene fossil record of the NE Atlantic oceanic islands, paleoclimatological considerations are discussed, which allow, for the first time, to include the Azores and the other Macaronesian islands in a wider context of the NE Atlantic paleobiogeographical molluscan provinces. Late Miocene to present day molluscan biogeographic units, ranging in latitude from 60°N down to 17°S, are here defined and the boundaries of the proposed climatic zones are outlined. We suggest that during the upper Miocene–lower Pliocene, the paleoclimate at Santa Maria Island was drastically different from that seen at those latitudes today, with mean annual sea surface temperatures (SSTs) about 3.7 °C to 6.3 °C higher than the present-day 20.6 °C, and with mean monthly SSTs ranging from 20 °C to 28 °C, with six months with mean SSTs over 24 °C, conditions typical of a tropical setting.

Massive fossil shell accumulations require particular conditions to be formed and may provide valuable insights into the sedimentary environments favouring such concentrations. Shallow-water shell beds appear to be particularly rare on reefless volcanic oceanic islands on account of narrow, steep and highly-energetic insular shelves where the potential for preservation is limited. The occurrence of an exceptional coquina (Pedra-que-pica) within the Miocene–Pliocene deposits of Santa Maria Island (Azores), therefore provides a rare opportunity to understand the conditions that led to the formation and preservation of a massive shell bed at mid-ocean insular setting. This study provides a detailed analysis regarding a 10–11-m-thick bivalve-dominated fossil assemblage exposed at Pedra-que-pica on Santa Maria Island in the Azores. Integration of taphonomical, palaeoecological and sedimentological observations are used to reconstruct the genesis of the coquina bed and related events, and to discuss why such exceptional sedimentary bodies are so rare on shelves around reefless volcanic oceanic islands. The sequence at Pedra-que-pica demonstrates a complex succession of sedimentary environments in response to the drowning of an existing coastline during a period of rapid sea-level rise. The Pedra-que-pica shell bed incorporates storm-related materials and possible debris falls that originated nearby in a shallow and highly productive carbonate factory. Deposition took place below fair-weather wave base, at around 50 m depth, as inferred from the overlying volcanic succession. The preservation of this coquina was favoured by deposition on a platform laterally protected by a rocky spur, combined with rapid burial by water-settled volcanic tuffs and subsequent volcanic effusive sequences. The recent exhumation of the deposit is the result of island uplift and subsequent erosion.

Ten new species belonging to three new genera ( Atlantisina gen. nov., Bathycyclopora gen. nov., Calvetopora gen. nov.) of umbonulomorph bryozoans from northeastern Atlantic seamounts, islands, and the continental slope are introduced. We furthermore erect the new family Atlantisinidae fam. nov. for these genera. Eight new species belong to the new genus Atlantisina : Atlantisina atlantis gen. et sp. nov. (type species), A. acantha gen. et sp. nov., A. gorringensis gen. et sp. nov., A. inarmata gen. et sp. nov., A. lionensis gen. et sp. nov., A. meteor gen. et sp. nov., A. seinensis gen. et sp. nov., and A. tricornis gen. et sp. nov. The genus Bathycyclopora gen. nov. is introduced for ? Phylactella vibraculata Calvet from the Azores, and also includes Bathycyclopora suroiti gen. et sp. nov. The type species of Calvetopora gen. nov. is Lepralia inflata Calvet from the Gulf of Cadiz; this genus also includes Calvetopora otapostasis gen. et sp. nov. and another species left in open nomenclature. Of the 13 species described herein, 11 occur on seamounts and islands, and nine species are endemic to a single seamount, island or station. The present results show that bryozoans provide striking examples of the function of seamounts as areas of endemism, most likely intrinsically linked to the low dispersal abilities of bryozoan larvae.

Pywackia baileyi Landing in Landing et al., 2010, from the upper Cambrian Yudachica Member of Oaxaca State, southern Mexico, consists of small, phosphatic, proximally tapering cylindrical rods covered by shallow polygonal calices. The bryozoan-like morphology of this fossil prompted its interpretation as the first bryozoan known from the Cambrian. However, restudy of some of the original material, employing scanning electron microscopy for the first time, questions the assignment of Pywackia to the Bryozoa. Striking similarities between Pywackia and the modern pennatulacean octocoral Lituaria lead to an alternative hypothesis interpreting Pywackia an early fossil octocoral. While Pywackia is probably not a true pennatulacean, a group with a definitive fossil record stretching back only to the Late Cretaceous, it can be envisaged as having had a similar skeletal structure and ecology to Lituaria, the rods representing mineralized axes of tiny colonies that lived with their proximal ends buried in the sediment and distal ends covered by feeding polyps. Landing et al. (2010) considered the phosphatic composition of Pywackia specimens to be the result of diagenetic replacement, but the evidence is equivocal. If Pywackia had a primary phosphatic skeleton, this would support the hypothesized existence of phosphatic biomineralization early in the evolutionary history of Cnidaria, as well as providing further evidence that Pywackia is not a bryozoan.

Genus Lentiarenium Voss gen. nov. urn:lsid:zoobank.org:act:A67C71FB-CBC3-460E-953C-5774A3EA9939 Type species Halitherium cristolii Fitzinger, 1842 Diagnosis Dugongid based on the following combination of synapomorphies: absence of alisphenoid canal, open foramen ovale, loss of permanent ffth premolar, and squamosal reaching temporal crest. Differs from the Dugonginae (Crenatosiren Domning, 1991, Nanosiren Domning & Aguilera, 2008, Dugong Lacépède, 1799, Rytiodus Lartet, 1866, Corystosiren Domning, 1990, Callistosiren Vélez-Juarbe & Domning, 2015, Bharatisiren Bajpai & Domning, 1997, Domningia Thewissen & Bajpai, 2009, Kutchisiren Bajpai, Domning, Das, Vélez-Juarbe & Mishra, 2010, Dioplotherium Cope, 1883 and Xenosiren Domning, 1989), the Hydrodamalinae Simpson, 1932 (Dusisiren Domning, 1978 and Hydrodamalis Retzius, 1794), and other dugongid genera (Eotheroides Palmer, 1899, Prototherium De Zigno, 1887, Eosiren Andrews, 1902, Caribosiren Reinhart, 1959, Priscosiren Vélez-Juarbe & Domning, 2014b, Metaxytherium De Christol, 1840, and other species formerly lumped under “ Halitherium ”) by displaying the following unique combination of plesiomorphies: frontal roof fat; supraorbital process of the frontal dorsoventrally fattened with well-developed, prominent posterolateral corner; supraoccipital wider dorsally than ventrally; exoccipitals meet in a suture dorsal to the foramen magnum; posttympanic process with a prominent anteroventral process for attachment of m. sternomastoideus; paroccipital process of exoccipital long, reaching as far ventrally as occipital condyles; accessory mental foramina present; horizontal ramus of mandible slender dorsoventrally; permanent premolars (P2/p2 – P4/p4) present. Synapomorphies: nasals reduced and not meeting in midline (shared with Eosiren imenti Domning et al., 1994, Caribosiren, Priscosiren, and all later dugongid taxa), nasal incisure at posterior end of the mesorostral fossa deep, extending posterior to the supraorbital processes of the frontals (shared with some Metaxytherium spp., dugongines, and Hydrodamalis); strongly concave ventral border of the horizontal mandibular ramus (shared with Priscosiren, Metaxytherium spp., and dugongines), masticating surface of mandible broad (shared with Metaxytherium ssp., dugongines, and hydrodamalines). In contrast to the similar species Priscosiren atlantica it has a supraorbital process with the posterolateral corner projecting posteriorly; prominent temporal crests on frontal and parietal; supraoccipital distinct in height, only slightly wider than high. Etymology A combination of the Latin terms for the Linz Sands (Lentia and arenium), the informal name for the upper Oligocene sediments in which the species was found.

Lentiarenium cristolii (Fitzinger, 1842) comb. nov. Figs 2–10, Tables 1–5 Halitherium cristolii Fitzinger, 1842: 71, pl. 1. Metaxytherium (?) pergense Toula, 1899: 459, pl. 12. Halitherium abeli Spillmann, 1959: 36, pls. 1–4, fgs 19–20, 22, 24–27, 29b, 31–32, 33b, 34. Manatus christolii – De Blainville 1844: 122. Metaxytherium christolii – Laurillard 1846: 172. Halianassa collinii – Meyer 1847: 189, 578 (partim). — Ehrlich 1850: 14–15, fgs a–c. — Ehrlich 1855: 3, pls. 1–2, corresponding fgs on pages 14–17. Halitherium schinzii – Peters 1867: 310. Halitherium schinzi – Lepsius 1882: 164 (partim). Halitherium christoli – Abel 1904: 25, pl. 1, fgs 12–13, pl. 2, fgs 4, 11, 17, pl. 5, fg. 8, fg. 1. — Spillmann 1959: 17, fgs 8–18, 21, 23, 28, 29a, 30, 33a; 1969: 62, pl. 8, fg. 2; 1973: 198, pl. 40, fg. 4. Halitherium pergense – Spillmann 1959: 11, fgs 6–7; 1969: 61, fg. 1; 1973: 197, pl. 39, fgs 1–2. Halitherium abeli – Spillmann 1969: 62, pl. 9, fg. 3; 1973: 205. Diagnosis As for the genus. Type Material Lectotype (present designation) A mandible (OLL 2012/1) of an adult individual (based on the presence of a large retromolar space and erupted m3 in wear) with left dp5–m2 and right m1–3, “Sicherbauer” sandpit, Linz. Paralectotypes Fragment of right maxilla with worn M1 crown and remnants of DP5 roots (OLL 2012/2), and an isolated crown of M3 (OLL 2012/3) from the right side, both from the “Sicherbauer” sandpit, Linz. Holotype (by monotypy) of Metaxytherium pergense Cast of a parietal-supraoccipital skullcap (OLL 1899/11), sandstone quarry near the town of Perg. Holotype (by original designation) of Halitherium abeli Mandible of an adult individual (based on the presence of a large retromolar space and erupted m3 in wear), fragments of basicranium, cervical and other vertebrae (OLL 1939/257), “Limoni” sandpit, Linz. Referred material OLL 1854/327, 1863/184, 1917/7, 1921/71, 1926/394, 1926/395, 1927/200, 1928/82, 1931/21, 1931/263, 1948/33, 2012/4, 2012/5, 2012/6, 2012/7, 2013/1. For a detailed listing of the preserved skeletal parts see Table 1. Type horizon and locality “Sicherbauer-Sandgstätte”, a former sandpit within the city limits of Linz (Upper Austria). Linz-Melk Formation, Linzer Sande (= Linz Sands), upper Oligocene, lower Egerian (Chattian). Range and distribution Known only from the upper Oligocene of Linz and the surrounding area. Description The following description is based on the available sirenian material housed in OLL, regardless of the different species that were proposed in the past. It will be outlined in detail in the Discussion why the respective skeletal material represents a single species. Lentiarenium cristolii is known from cranial and postcranial elements. However, the premaxilla and lacrimal are unknown as are the pelvis, zeugopod and autopod. The following description is mainly based on a partial skull (OLL 1926/394) and two mandibles (OLL 1939/257 and OLL 2012/1). PREMAXILLA. The premaxilla is not preserved in any specimen but some morphological characters can be indirectly determined. The external nares are retracted and enlarged (Fig. 2), as in all sirenians, and reach beyond the anterior margin of the orbit. Judging from the recesses in the anterior margin of the frontal, the nasal processes are not broadened and bulbous at their posterior ends but taper, having lengthy overlap with the frontals. The angle of the rostrum most likely exceeds 50° considering the defection of the mandibular symphysis of about 60° (Fig. 7). NASAL. The exact status of the nasals cannot be determined without doubt due to the poor preservation of the nasal area in specimen OLL 1926/394 (Fig. 2). Judging from the right anteromedial margin of the frontals, which is not damaged, but smooth without an attachment area for the nasals, these elements are at least considered to have been small without meeting in the midline. A nasal incisure is present at the posterior end of the mesorostral fossa, but unlike in Metaxytherium albifontanum (Velez-Juarbe & Domning, 2014a) and Priscosiren atlantica (Velez-Juarbe & Domning, 2014b) it is deep, extending posterior to the supraorbital processes of the frontals, though not to the extent seen in dugongines like Crenatosiren olseni (Domning 1997). It is uncertain whether the nasals were fused or coalesced with the frontals. ETHMOIDAL REGION. The ethmoid is incompletely preserved and not easily observable in all aspects. A prominent perpendicular plate of the mesethmoid is visible in dorsal and anterior views of the skull OLL 1926/394 (Fig. 2). This vertical wall measures 10 mm to 15 mm in width, is narrower dorsally and ventrally, and apparently also becomes thinner posteriorly. Ventrally, the perpendicular plate is fused with the likewise distinctly developed vomer. On the right side of the skull, and medial to the frontal (Fig. 2), the almost complete large ethmoturbinal (concha maxima ethmoidalis (Kaiser 1974)) extends nearly parallel to the mesethmoid. Its left counterpart is only fragmentarily preserved. The crista galli and lamina papyracea are either not preserved or not visible. VOMER. The vomer is exposed on the ventral side of the skull OLL 1926/394 but broken anteriorly (Fig. 3). It is fused with the presphenoid posteriorly and forms the cranial extension of its median crest. The vomer is triangular in cross section, frmly fused with the ethmoid via its fat dorsal surface, and contacts the maxilla laterally. In lateral view (Fig. 4B), the vomer is also visible through the orbit due to the incomplete preservation of the skull. FRONTAL. In dorsal view (Fig. 2), the frontal roof is fat between the temporal crests, and bears no knoblike bosses as present in some dugongines (e.g., Crenatosiren olseni (Domning 1997)). The straight to slightly concave anterior margin of the frontal has no internasal process. The temporal crests form distinct keels (morphotype B; Domning 1988) and are as prominent on the frontal as on the parietal. In lateral view (Fig. 4B), the supraorbital process is dorsoventrally fattened with its dorsal surface inclined gently ventrolaterad. Its lateral margin is not divided, and it has a prominent posterolateral corner that projects posteriorly. An orbitotemporal crest is distinct relative to what is observed in Metaxytherium albifontanum (Velez-Juarbe & Domning 2014a), and forms a craniocaudally-extending ridge. The lamina orbitalis is signifcantly less than 10 mm thick. PARIETAL. The parietal roof (Fig. 2) is fat between the temporal crests and characterised by a strong intertemporal constriction that reaches its maximum behind the centre of the skull roof with the parietal bulging laterally. An external sagittal crest is not developed. The frontal processes of the parietal are short and do not exceed half the length of the frontal in the midline, a condition similar to that in Priscosiren atlantica (Velez-Juarbe & Domning 2014b). The parietal is longer than the frontal, and the overall proportions of the cranial roof indicate a slight brachycephaly when the value of the frontal length is divided by the width of the supraoccipital, i.e., the ratio l FP/w SO, which is below 2. In endocranial view (Fig. 3), the bony falx cerebri extends from a prominent tentoric process and fattens out anteriorly before reaching the frontoparietal suture. The tentorium osseum is well developed and an internal parietal spine (or tentorial process as illustrated in Voss (2008: Fig. 4B)) is missing. On both sides of the bony falx, the fat internal parietal surface shows the depressions for the superior parts of the brain hemispheres. SUPRAOCCIPITAL. This element is widest dorsally rather than ventrally (Fig. 4A), relative to what is observed in Metaxytherium spp. (e.g., M. albifontanum (Velez-Juarbe & Domning 2014a)). The supraoccipital is only slightly wider than high (width to height ratio Crenatosiren olseni (Domning 1997). Dorsally, the nuchal crest is constantly narrow along its transversal length and relatively sharp-edged. It makes up the dorsolateral margin of the supraoccipital. The external occipital protuberance rises above the parietal roof and is also prominent posteriorly. Ventrad, the protuberance continues as the external occipital crest that forms a distinct ridge, which slightly fattens out after one third of the supraoccipital’s height, almost reaching its ventral margin. Dorsolateral to this median ridge the deep and large insertions for the semispinalis capitis muscle occupy about the upper third of the external lamina. The area of the muscle insertion is triangular and defned by distinct ridges medioventrally and the nuchal crest dorsolaterally. The ventral margin of the supraoccipital forms an angle of approximately 135° in specimen OLL 1926/394. Specimen OLL 1899/11 (Fig. 5), formerly designated as the holotype of “ Halitherium ” pergense but now referred to Lentiarenium cristolii, also shows a distinct nuchal crest. However, the protuberance, median ridge and defnitions of the muscle insertions are less prominently developed than in OLL 1926/394, indicating its juvenile status. In interior view (Fig. 3), the supraoccipital is fat with the exception of longitudinal bulges dorsolaterally that are merged in the median plane and separated by a deep transverse sulcus from the tentorium osseum. Posterolaterally, the parietals extend between the supraoccipital and squamosal, forming a short fange. EXOCCIPITAL. The dorsal parts of these paired elements are not preserved in any specimen. However, the total length of the ventral margin of the supraoccipital in OLL 1926/394 (Figs 3, 4A) reveals the articulation surface for the exoccipitals, indicating that these bones evidently meet in a suture dorsal to the foramen magnum. Additionally, specimen OLL 1939/257 preserves the ventralmost parts of the exoccipitals, which Spillmann (1959: fg. 19) illustrated in ventral view. During the personal examinations, the paroccipital processes were detected to be long, projecting as far ventrally as the occipital condyles, which is similar to what can be observed in some dugongines (e.g., Nanosiren garciae Domning & Aguilera, 2008 and N. sanchezi Domning & Aguilera, 2008 (Domning & Aguilera 2008: fg. 3, 11)). Medioventral to the paroccipital processes, the hypoglossal foramen is not opened to form a notch or incisure but is well surrounded by bone. The supracondylar fossae are distinct and defne the occipital condyles across their entire width. BASIOCCIPITAL. An isolated basioccipital fused with the lower parts of the exoccipitals is preserved in specimen OLL 1939/257 (Spillmann 1959: fg. 19). It contributes to the occipital condyles ventrolaterally and extends craniad as a short, columnar bone. On its ventral side, the sphenooccipital eminences for the longus capitis muscles are concave and separated by a short but distinct ridge. Specimens OLL 1939/257 and 1926/394 (Fig. 3), in which either the basioccipital or basisphenoid is preserved, show smoothed attachment areas for the adjacent bone, indicating that the basioccipital and basisphenoid were not fused. On that basis, both specimens can be determined as subadults. BASISPHENOID, PRESPHENOID, ORBITOSPHENOID. In specimen OLL 1926/394, the sphenoidal region is clearly observable (Fig. 3). The basisphenoid has a fat ventral surface that is defned laterally by antero posteriorly-broad pterygoid processes. Cranially, the basisphenoid continues with a slight anterodorsal slope into the presphenoid. Both bones are frmly fused with each other and with the orbitosphenoid anterolaterally, the alisphenoid dorsolaterally, and the pterygoid posterolaterally. The median crest of the presphenoid is not prominently developed, which might be related to the state of preservation of the skull in this area, and only becomes distinct at the level of the adjacent vomer. On the lateral side of the skull (Fig. 4B), the orbitosphenoid is exposed and contributes to the anteromedial wall of the temporal fossa. The orbitosphenoid is defned by the frontal dorsally, the alisphenoid posterodorsally, and apparently by the maxilla ventrally. Its sutures with the palatine are not detectable. ALISPHENOID. The alisphenoid is visible in the lateral view of the skull (Fig. 4B). Its sutural contacts with the frontal, parietal and squamosal can be clearly determined. Furthermore, the alisphenoid forms the slightly uneven posterolateral side of the pterygoid process. An alisphenoid canal is absent, and the foramen ovale is opened to form a notch or incisure. PTERYGOID. As in other sirenians, the pterygoid is present on the posteromedial side of the pterygoid process but fully fused with the surrounding bones (Fig. 3). Though not well preserved in OLL 1926/394, the wing-shaped pterygoid processes each bear a dorsoventrally elongated pterygoid fossa posteriorly that extends above the level of the roof of the internal nares, relative to the condition observed in some dugongines like Dioplotherium manigaulti (Domning 1989). The distal ends of both pterygoid processes are somewhat damaged, although the distomedial angle of the right pterygoid process is still distinct and indicates a hamula process. PALATINE. Only the posteriormost parts of the palatines are preserved in OLL 1926/394, and are best observable on the right side (Fig. 3). There the palatine forms the anteromedial margin of the pterygoid process. Its sutures with the surrounding bones are only barely visible on the distal and medial sides of the pterygoid process and on the posterior side of the maxillary alveolar margin. MAXILLA. The zygomatic-orbital bridge (Fig. 3) is not completely preserved in any of the specimens. However, in the maxillary fragment of OLL 1939/257 its dimensions are 47 mm in minimum length and 17.5 mm in nearly original height (Spillmann 1959: fg. 20). On the basis of these data, the zygomaticorbital bridge can be clearly determined to be long anteroposteriorly, relative to what is observed in the genus Hydrodamalis (Domning 1978; Velez-Juarbe & Domning 2014a). Additionally, and in contrast to some hydrodamalines, e.g., Dusisiren jordani Kellogg, 1925 and Hydrodamalis cuestae Domning, 1978 (Domning 1978; Velez-Juarbe & Domning 2014a), it is only slightly elevated above the alveolar margin, and its posterior edge is thickened. Remnants of the infraorbital canal reveal no obstruction. SQUAMOSAL. The cranial part of the squamosal (Figs 2, 4B) extends up to the temporal crests but does not interrupt the course of the temporal crests, so that these reach the occipital nuchal crest. The posttympanic process is not club like distally (Fig. 4B), but concave anteroventrally for the attachment of the sternomastoid muscle. In the posterior view of the skull (Fig. 4A), a prominent sigmoid ridge is visible forming the laterocaudal margin of the squamosal. Posterolaterally, the mastoid foramen is present, flled by the periotic, and enclosed by the squamosal anteriorly, the exoccipital posteriorly, and the supraoccipital dorsally. Lateral to the braincase (Figs 2, 4B), each of the zygomatic processes projects from a zygomatic root that, though partially broken, is characterised by a distinct notch posteriorly. The zygomatic process is triangular in shape, tapering anteriorly. Its lateral and medial sides are fat to concave with the dorsal margin distinctly inclined inwards to form a sigmoid ridge. The posterodorsal end of the zygomatic process is straight to concave. The external auditory meatus is short mediolaterally and about as wide as high anteroposteriorly. Ventrally (Fig. 3), the elements of the mandibular articulation surface are elongated transversely. The mandibular fossa forms a distinct depression relative to the slightly convex tuberculum anteriorly. Posterior to the mandibular fossa, the postglenoid process forms a rounded and longitudinal knob, which rises above the level of the tuberculum. The posterior end of the zygomatic process, the processus retroversus, shows a moderate inward-directed infection in contrast to what is observed in the living dugong (e.g., Domning & Aguilera 2008; Velez-Juarbe & Domning 2014a). JUGAL. Only a fragmentarily preserved middle part of the right jugal is known from specimen OLL 1926/394 (Fig. 4B), indicating that a postorbital process is probably present. Considering the position and shape of the supraorbital processes of the frontal, the development of a postorbital bar can be excluded. The zygomatic process of the jugal is not preserved but according to its imprint on the ventral side of the zygomatic process of the squamosal it reaches the tuberculum, exceeding the diameter of the orbit. EAR REGION. In specimen OLL 1926/394, the right periotic is poorly preserved (Figs 3, 4A). It is not fused with the adjacent skull bones, and is set in a closely-ftting socket in the squamosal. The tegmen tympani is only indicated by its imprints on the dorsolateral side of the squamosal. It was most likely about as big as the mastoid or slightly smaller. The petrosal is fragmentarily preserved medioventrally, with the perilymphatic foramen apparently not separated into a fenestra rotunda and cochlea canaliculus. The processus fonticulus flls the mastoid foramen posteriorly (Fig. 4A). MANDIBLE (Table 3). The mandibular symphysis is broad, as is the masticating surface that is lacking a median furrow and houses four large and shallow alveoli for vestigial incisors and canines. This is best visible in OLL 1939/257 (Fig. 6A), which exhibits a completely preserved masticating surface, whereas in OLL 2012/1 (Figs 6B, 7A) the ventralmost end is broken. In lateral view (Fig. 7), the symphysis is higher than long and bears the mental foramen laterally, which is joined dorsoposteriorly by two accessory mental foramina on each side in specimen OLL 1939/257 (Fig. 7B). The accessory mental foramina are large relative to what is observed in more plesiomorph dugongid taxa, e.g., “ Halitherium ” taulannense (Sagne 2001a), but they are still smaller than the principal foramen. This observation is interpreted here in analogy with Trichechus (Domning 1982, 1994, and personal observations), as “true” accessory mental foramina, which are present in addition to and usually posterior to the large principal foramen. In the lectotype (OLL 2012/1; Fig. 7A), the mental foramen is broken off dorsoposteriorly, and therefore the accessory mental foramina are not identifable, but their open canals are merged with the main foramen. The overall build of the horizontal mandibular ramus appears to be broad dorsoventrally, but it is evaluated to be slender on the basis of its minimum dorsoventral height, which is smaller than 0.25 × the length of the mandible. The ventral border

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The two odontocete taxa Squalodon grateloupii and Patriocetus ehrlichii, both the type species of their respective genera, have been at the centre of a great deal of taxonomic confusion. Originally regarded to be conspecific, these two taxa have been the subject of a bewildering taxonomic debate lasting for more than a century, which recently led to the suggestion to abandon these widely used names and replace S. grateloupii with the similar, yet independently and later proposed name S. gratelupi as the type species of Squalodon. Here, we attempt to summarise the events leading to the current confused situation in the hope of resolving this issue once and for all and argue that the name Squalodon grateloupii, as originally proposed, should be reinstated.

Located at the interface between land and sea, marl prairies are sensitive to changes in water balance and useful recorders of climate and sea-level changes. Palustrine carbonate in marl prairies precipitates in temporary, barely flooded grasslands within microbial mats. Despite the special mode of carbonate production, descriptions of the sedimentary facies are exceptional and cursory because marl prairies are so far reported only from the recent of the Everglades (Florida, USA), where they produce an unspectacular calcite mud. We present a Pleistocene Everglades-type marl prairie from coastal Tanzania as the first fossil example. The unique preservation and high productivity (two times higher than in the Everglades) of the periphyton community in this marl prairie is due to increased calcification of coccoid and filamentous cyanobacteria. The excellent preservation allows us to characterize a marl prairie facies in great detail for the first time. Facies analyses of the sediments reveal a transition from tidal to terrestrial settings that started at ca. 44 14 C ka in response to eustatic sea-level fall and coastal tectonic uplift. The resultant drop of the groundwater table triggered the development of the marl prairie. The decline of the marl prairie was initiated at ca. 33 14 C ka due to the onset of the Last Glacial Aridity Maximum in equatorial East Africa.

In this study we revise the cheilostome bryozoan genus Buffonellaria Canu & Bassler, 1927 and its Mediterranean and north-east Atlantic species, thereby addressing several existing problems. First, a lectotype for the type species, Buffonellaria divergens (Smitt, 1873) from Florida, is chosen, which proves to be distinct from the European species. Second, the two hitherto established north-east Atlantic species [Buffonellaria nebulosa (Jullien & Calvet, 1903) and Buffonellaria porcellanum Aristegui Ruiz, 1987], are redescribed, which were poorly documented until now. Third, close inspection of material, collected from Spitsbergen to tropical West Africa, using scanning electron microscopy reveals that the actual number of species, all previously referred to either B. divergens or Stephanosella biaperta (Michelin, 1848), is distinctly greater in the north-east Atlantic than has been previously acknowledged. As a result, seven new species are introduced (Buffonellaria acorensis sp. nov., Buffonellaria antoniettae sp. nov., Buffonellaria arctica sp. nov., Buffonellaria harmelini sp. nov., Buffonellaria jensi sp. nov., Buffonellaria muriella sp. nov., and Buffonellaria ritae sp. nov.), whereas two are left in open nomenclature. With the increase in number of species, the extremely broad geographical range of distribution assumed for B. divergens breaks down to numerous restricted areas. However, although most species have only been reported from a single location, B. arctica sp. nov. seems to have a fairly wide distribution in the Arctic region. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 537–566.

Rhodoliths are a common producer of carbonates on modern and ancient shelves worldwide, and there is growing evidence that they thrive on volcanic insular shelves. However, little is still known on how rhodoliths cope with the demands of this particularly dynamic environment. In this study, the focus is placed on fossil rhodoliths from a Pliocene sequence at Santa Maria Island, Azores, in order to gain further insight into the life cycle (and death) of rhodoliths living within a mid-ocean active volcanic setting. These rhodoliths occur as a massive accumulation within a larger submarine volcano-sedimentary sequence that was studied from the macro- to the micro-scale in order to reconstruct the paleoenvironmental conditions under which the rhodolith accumulation was deposited and buried. All fossil rhodoliths from this setting are multi-specific and demonstrate robust growth forms with a lumpy morphology. Moreover, taphonomical analyses show the succession of several destructive events during rhodolith growth, suggesting life under a highly dynamic system prior to stabilization and burial. The rhodoliths therefore tell a story of an eventful life, with multiple transport and growth stages, owing to the environment in which they lived. Transport and deposition to their final resting place was storm-associated, as supported by the general sedimentary sequence. In particular, the sequence features an amalgamation of tempestites deposited under increasing water depths, sediment aggradation, and before burial by volcanic activity. This transgressive trend is also attested by the overall characteristics of the volcano-sedimentary succession, which exhibits the transition to subaerial environment in excess of 100 m above the rhodolith bed.

Eight NE Atlantic and Mediterranean species, which were originally assigned to the genus Schizoporella (Family Schizoporellidae) when introduced, are redescribed and stabilized by typification. Seven of these species are transferred to the bitectiporid genus Schizomavella: S. fischeri, S. glebula, S. neptuni, S. obsoleta, S. richardi, S. triaviculata, and S. triaviculata var. paucimandibulata, which is here raised to species rank. The eighth species, Schizoporella fayalensis, is transferred to the lanceoporid genus Stephanotheca. Schizomavella obsoleta and S. glebula are considered junior subjective synonyms of S. fischeri and S. richardi, respectively. Two new species are described: Schizomavella rectangularis n. sp. from the Strait of Gibraltar, and Schizomavella phterocopa n. sp. from the Great Meteor Bank. A new subgenus, Calvetomavella n. subgen. is established as a result of a phylogenetic analysis based on morphological characters; it includes S. neptuni, S. triaviculata, S. paucimandibulata and S. phterocopa n. sp., together with Schizomavella discoidea and Schizomavella noronhai. The rest of the species remain in the nominotypical subgenus Schizomavella. The work by ORG and JS was partially supported by the project “Fauna Ibérica: Briozoos II (Familia Cribrilinidae – Familia Watersiporidae)” (CGL2010-22267-C07-02), co-financed by the Ministerio de Economía y Competitividad (Spanish Government) and FEDER. BB acknowledges support from the SYNTHESYS Project (http://www.synthesys.info/), which is financed by the European Community Research Infrastructure Action under the FP7 “Capacities” Program, and which allowed studying type and comparative material at the MNHN (FR-TAF-1902) and the NHMUK (GB-TAF-3347). JS thanks the Austrian Science Fund (FWF, Lise Meitner Program, grant M1444-B25) for financial support (https://www.fwf.ac.at/). SI

This paper describes and illustrates a diverse parautochthonous macrofauna from a single Upper Burdigalian (lower Ottnangian) horizon at Gurlarn in Lower Bavaria (SE Germany). In total, 80 different taxa are recorded in specific or open nomenclature; some 50 % of these taxa are bryozoans, followed by bivalves (16 taxa), cirripedes (7 taxa), echinoderms, corals (5 taxa each), brachiopods, fish (4 taxa each), serpulids, and gastropods (3 taxa each). The presence of additional organisms was documented by actualistic comparison based on indirect evidence such as drillholes, bite marks, and specialized growth forms of bryozoan colonies. Analysis of autand synecological indicators suggests that the fauna thrived in a near-shore shallow marine setting at a water depth of 5-20 m. Based on particular faunal elements and overall faunal composition we hypothesize that the environment is characterized by three distinct but interfingering habitats, i.e. (1) rocky slopes and boulders, (2) seagrass meadows, and (3) bryozoan meadows. Because similar, albeit less wellpreserved, faunas occur at several localities along the northern coast of the early Ottnangian Molasse Sea, the parautochthonous assemblage from Gurlarn provides an excellent example for the structure of these typical biota.

The Madeira-Tore Rise, located ∼700 km off the NW African coast, forms a prominent ridge in the east Atlantic. The age and origin of the rise are controversial. This study presents major and trace element, Sr, Nd, Pb, Hf isotope and 40Ar/39Ar age determinations from volcanic rocks dredged from different sites along the rise. In addition, isotopic compositions of rock samples from Great Meteor Seamount in the central Atlantic are presented. The new radiometric and paleontologically constrained ages identify two major episodes of volcanism: The first is the base of the rise (circa 80 to >95 Ma) and the second is seamounts on the rise (0.5–16 Ma). It is proposed that interaction of the Canary hot spot with the Mid-Atlantic spreading center formed the deep basement of the Madeira-Tore Rise and the J-Anomaly Ridge west of the Atlantic spreading center in the Mid-Cretaceous. Age and geochemical data and plate tectonic reconstructions suggest, however, that the recovered Late Cretaceous volcanic rocks represent late stage volcanism from the time when the Madeira-Tore Rise was still close to the Canary hot spot. Long after moving away from the influence of the Canary hot spot, the Madeira-Tore Rise was overprinted by late Cenozoic volcanism. Miocene to Pleistocene volcanism at the northern end of the rise can be best explained by decompression mantle melting beneath extensional sectors of the Azores-Gibraltar Fracture Zone (African-Eurasian plate boundary). The geochemical compositions of these volcanic rocks suggest that the magmas were variably contaminated by enriched material within or derived by melting of enriched material underplated at the base of the lithosphere, possibly originating from the Cretaceous Canary plume. Alternatively, these late Cenozoic volcanic rocks may have derived from decompression melting of enriched pyroxenitic/eclogitic material in the upper mantle. Isotopically more depleted Pliocene to Pleistocene volcanism at the southern end of the Madeira-Tore Rise may be related to the nearby Madeira hot spot.

The intricate geological evolution of the Arctic Ocean is paralleled by complexities in the biogeographical and phylogenetical histories of the Arctic biota, including bryozoans. Here we present revised taxonomic descriptions for all known species of the bryozoan genus Pseudoflustra, and use the present-day distributions and phylogenetic relationships between these species to infer the historical biogeography of the genus. Nine species belonging to the genus Pseudoflustra are recognized in the Arctic and North Atlantic. One new species, previously identified as Ichthyaria aviculata, is described as Pseudoflustra radeki sp. nov. Another species, previously assigned to Smittoidea as S. perrieri, is transferred to Pseudoflustra. Biogeographical analysis of Pseudoflustra reveals that species distributions mostly match current patterns pertaining in the North Atlantic and Arctic Ocean. Distributions were probably shaped by recent geological history as present-day current directions in the Arctic Ocean are believed to have been similar for at least the last 120 000 years. Phylogenetic analysis of Pseudoflustra places the five Arctic-North Atlantic species in a clade crownward of a paraphyletic grouping of North Atlantic species. Given that the Arctic Ocean was fully glaciated until 18 000 years, the most likely explanation for this phylogeographical pattern is that species of Pseudoflustra colonized the Arctic relatively recently from North Atlantic sources. However, a fuller understanding of the origin of Pseudoflustra in the Arctic will require molecular and fossil data, neither of which are currently available.

Anopheles gambiae s.s., the most important mosquito species transmitting malaria in sub-Saharan Africa, is experiencing an incipient speciation process triggered by human-made environmental changes, which is changing malaria epidemiology. The species is subdivided into two molecular forms, named ‘M’ and ‘S’, defined on rDNA-IGS mutations and distinguished for the M form-specific insertion of a SINE transposable element (~1 Mb upstream from IGS). M and S forms represent a model for the study of speciation at its early stage since they are diversifying in West and Central Africa despite some ongoing gene flow. In fact, M and S present some genome regions which are supposed to resist introgression – especially in the low-recombinant centromeric areas of chromosomes X and 2 (‘speciation islands’) – while the rest of the genome continuously homogenise. The observed pattern rises questions on whether divergence is essentially due to physical features of the genome that reduce recombination (e.g., centromeres) despite gene flow, or to selection on adaptive genes clustered in the centromeric ‘speciation islands’ or elsewhere in the genome. Hybrids between M and S are rare in nature, but an unusually high degree of hybridization was found at the western extreme of their distribution range (i.e.The Gambia and Guinea Bissau). This secondary contact zone represents an opportunity to investigate how reduced reproductive isolation affects genetic differentiation in genomic regions known to be segregated between M and S forms elsewhere in Africa. While most studies focus on high-throughput SNPs assays on limited sampling, we genotyped a large number of A. gambiae s.s. specimens from Guinea Bissau (and from other African countries for comparison) focusing on two intronic regions known to segregate in sympatric M and S molecular forms out from the hybridization area: i) the intron-4 of a cytochrome P450, namely the Cyp4G16 gene, which maps in the low recombination region of the X-centromere, ~7 kb upstream from SINE locus, and ii) the intron-1 of the voltage-gated sodium channel (VGSC) gene mapping on chromosome 2, out from the centromere area. Our results indicate that in the secondary contact zone recombination occurs within the most divergent region of M and S genome (i.e. the X-centromere), as highlighted by the intron-4 of Cyp4G16 data and support the hypothesis of asymmetric introgression from M to S, as shown by the analysis of intron-1 of VGSC gene.