[Entomology • 2024] Phylogeographic Analyses of western Palearctic Scaurus (Coleoptera: Tenebrionidae) reveal undetected taxonomic substructure along the pre-Saharian Atlantic Coast of western Africa

[a–c] Scaurus gigas Waltl, 1835  and  [d–f] S. ferreri  Español, 1943.
Specimens photographed are from (a) Meia Praia, Lagos (Portugal), (b) Chipiona, Cádiz (Spain), (d) Aglou Plage (Morocco) and (e) Deghaimis (Morocco).
Landscape images represent (c) the northernmost range of S. gigas (near Sines, Portugal)
and (f) one of the northern locations of S. ferreri at Aglou Plage (Morocco).

in Duque-Amado, García-París et Sánchez-Vialas, 2024. 

DOI: doi.org/10.3897/asp.82.e132546

Abstract

Phylogeography stands as a key tool to explore evolutionary patterns and processes, playing a crucial role in delimiting evolutionary units. Identifying these units is essential for providing robust taxonomic decisions. In this study, we present a comprehensive phylogeographical framework of Scaurus uncinus (Forster, 1771) and Scaurus gigas Waltl, 1835 across the Iberian Peninsula and North-western Africa, where they are widely co-distributed, coexisting in several localities on both sides of Strait of Gibraltar. Our results show that the Strait of Gibraltar did not act as a geographical barrier for these species, revealing shared mitochondrial haplotypes and nuclear alleles between populations on both sides. However, the Souss Valley in Morocco appears to have historically served as a significant geographical barrier within the S. gigas lineage, leading to the divergence of two morphologically distinct sublineages, one to the north (S. gigas) and the other to the south (S. ferreri stat. nov.). In addition, we point out a case of cytonuclear discordance between S. uncinus and S. gigas in the southwest corner of the Iberian Peninsula, suggesting the occurrence of an ancient event of genetic introgression between the two species.

Keywords: Allele networks, cyto-nuclear discordances gene flow, genetic isolation, haplotype networks, introgression, mitochondrial DNA

Live specimens and typical habitat of Scaurus gigas (a–c) and S. ferreri (d–f).
Specimens photographed are from (a) Meia Praia, Lagos (Portugal), (b) Chipiona, Cádiz (Spain), (d) Aglou Plage (Morocco) and (e) Deghaimis (Morocco).
Landscape images represent (c) the northernmost range of S. gigas (near Sines, Portugal) and (f) one of the northern locations of S. ferreri at Aglou Plage (Morocco).  

Carmen Duque-Amado, Mario García-París, Alberto Sánchez-Vialas. 2024. Phylogeographic Analyses of western Palearctic Scaurus (Coleoptera: Tenebrionidae) reveal undetected taxonomic substructure along the pre-Saharian Atlantic Coast of western Africa. Arthropod Systematics & Phylogeny. 82: 707-717. DOI: doi.org/10.3897/asp.82.e132546

[PaleoIchthyology • 2024] Afrocascudo saharaensis • A Saharan Fossil and the Dawn of Neotropical Armoured Catfishes in Gondwana

Afrocascudo saharaensis

Brito, Dutheil, Gueriau, Keith, Carnevale, Britto, Meunier, Khalloufi, King, de Amorim & Costa, 2024

 DOI: 10.1016/j.gr.2024.04.008

  

Highlights: 

• We describe a Cretaceous endemic Neotropical freshwater catfish from Morocco.

• Overlooked evolutionary events took place at the outset of the break-up of Gondwana.

• Iconic Neotropical freshwater lineages diversified after surviving the K/Pg extinction.

Abstract

Siluriformes are considered as primarily freshwater and have frequently been a model for the study of historical biogeography. Among catfishes, the most diverse clade is the Loricarioidei, a Neotropical group for which the fossil record extends back to the Palaeocene of Argentina. Here we describe a fossil from the early Late Cretaceous of Morocco, exhibiting typical morphological traits of the Loricariidae. A phylogenetic analysis integrating morphological characters with a multigene database for the main loricarioid lineages and outgroups highly supports inclusion of the fossil within the Loricariidae. A time-calibrated analysis corroborates the origin of loricarioids at about 112 MYA. The presence of this loricariid in Africa provides evidence that loricarioids have diversified before the separation of Africa and South America. The Moroccan loricariid shows an ancient evolutionary history that, in Africa, ended in the Late Cretaceous but persisted in South America, later surviving the K/Pg extinction.

 

Afrocascudo saharaensis gen. et sp. nov.

 

Paulo M. Brito, Didier B. Dutheil, Pierre Gueriau, Philippe Keith, Giorgio Carnevale, Marcelo Britto, François J. Meunier, Bouziane Khalloufi, Andrew King, Pedro F. de Amorim and Wilson J.E.M. Costa. 2024. A Saharan Fossil and the Dawn of Neotropical Armoured Catfishes in Gondwana. Gondwana Research. In Press. DOI: 10.1016/j.gr.2024.04.008

 twitter.com/justin_an74/status/1783944379209855058

[Arachnida • 2020] On Sahastata (Araneae: Filistatidae): Complementary Description of the Generotype and Two New Species from Oman and Morocco

[upper] Sahastata wesolowskae from Oman, &

 [lower] S. wunderlichi from Morocco.

Magalhaes, Stockmann, Marusik & Zonstein, 2020

DOI:  10.11646/zootaxa.4899.1.12 

twitter.com/6eyes8legs 

Abstract

The Filistatinae genus Sahastata Benoit, 1968 is distributed in arid and semi-arid areas, from westernmost Sahara to India, and includes seven known species. Four of these are only known from one sex, including Sahastata nigra (Simon, 1897), the type species. Here we present the first description of a male of this species collected near the type locality in Muscat, Oman. Additionally, two new species are described: S. wunderlichi sp. nov. (♂♀, Morocco) and S. wesolowskae sp. nov. (♂♀, Oman). Sahastata infuscata (Kulczyński, 1901) is newly recorded from Kenya and Yemen and S. nigra is newly recorded from the United Arab Emirates. DNA barcodes are given for S. nigra and the two new species. We observed some details of the life cycle of three Sahastata species, including clutch size, time to maturation, and a biased sex ratio for individuals raised from egg sacs, indicating that only 20–25% of specimens reaching adulthood are males. We provide SEM images of spiders of this genus, some observations on the morphology of spinnerets and male palps, and a distribution map of the species included in the genus.

Keywords: Araneae, Arabia, barcode, desert, Filistatinae, Maghreb, Sahara, spider, taxonomy

FIGURE 1. Sahastata spp., living specimens. 
A–B  Sahastata nigra (Simon, 1897) from Jebel Shams, Oman, male (A) and female (B); 
C–D Sahastata wesolowskae sp. nov. from Salalah, Oman, male holotype (C) and female (D); 
E-F Sahastata wunderlichi sp. nov. from Zag, Morocco, male (E) and female (F). 

Scale bars: 10 mm (except E, 5 mm). 

FIGURE 2. Sahastata habitats and webs. 
A–B, G Sahastata nigra (Simon, 1897) from Jebel Shams mountains, Oman; 
C, H Sahastata wesolowskae sp. nov. from Al Wusta, Oman; D S. wesolowskae sp. nov. from Salalah, Oman (type locality); 
E Sahastata sp. from Sudh, Oman; 
F Sahastata wunderlichi sp. nov. from Assa-Zag, Morocco.

FIGURE 24. 
A–B Sahastata nigra (Simon, 1897): A Male from Jebel Shams mountains, Oman (MACN-Ar 39464), left bulb, prolateral; B Female from Jebel Shams mountains, Oman (MACN-Ar 38368), spermathecae, dorsal. 
C–D Sahastata wunderlichi sp. nov.: C Male paratype from Zag, Morocco (MACN-Ar 39465), left bulb, prolateral; D Female paratype from the same locality (MACN-Ar 38366), spermathecae, dorsal;
 E–F. Sahastata wesolowskae sp. nov., E Male holotype from Salalah, Oman (SMNH), right bulb, prolateral, mirrored; F Female paratype from the same locality (MACN-Ar 39466), spermathecae, ventral. Scale bars: 100 μm, all figures to scale. 

Abbreviations: BBS—basal bulb sclerite; CIC—cymbium internal crest; F—fundus; K—embolus keel; tM29—tendon of the claw flexor muscle. 

Ivan L. F. Magalhaes, Mark Stockmann, Yuri M. Marusik and Sergei L. Zonstein. 2020. On Sahastata (Araneae: Filistatidae): Complementary Description of the Generotype and Two New Species from Oman and Morocco. Zootaxa. 4899(1); 215–246. DOI:  10.11646/zootaxa.4899.1.12

 twitter.com/6eyes8legs/status/1344317808814010368/

Researchgate.net/publication/348013410_On_Sahastata

      

[Botany • 2019] Zahora ait-atta • A New Monotypic Genus from tribe Brassiceae (Brassicaceae) endemic to the Moroccan Sahara

Zahora ait-atta Lemmel & M.Koch

in Koch & Lemmel, 2019. 
Zizaou n’oudad  ||  DOI: 10.3897/phytokeys.135.46946

Abstract
Zahora ait-atta Lemmel & M.Koch, a new species from the Moroccan Sahara, is described and documented here and constitutes a monotypic new genus. The new taxon belongs to the tribe Brassiceae (Brassicaceae), and cytogenetic and phylogenetic analyses reveal that this diploid species has a remote status of Miocene origin in the northwestern Sahara Desert. We examined the morphological differences between morphologically related genera and provide photographs of the new species. The new genus may play a key role in future Brassica-Raphanus crop research since it is placed phylogenetically at the base of a generically highly diverse clade including Raphanus sativus, and it shows affinities to various Brassica species.

Keywords: Brassiceae, Brassicaceae, flora of the Sahara, Morocco, new genus, Zahora ait-atta

Figure 1. Zahora ait-atta in its natural environment. Border region with Algeria. Near Errachidia.
Oued Bou-Ibourine – type locality a sandy habitat b flowering plant c rosette during winter d lyrate leaf from lower part of the plant e rosette starts building the inflorescence f ripening heteroarthrocarpic fruits g flowers and detailed view on sepals h siliques releasing seeds from dehiscent distal part of fruit.

Images taken by C. Lemmel and Z. Attioui.

Zahora ait-atta Lemmel & M.Koch, gen. et, sp. nov.

Type: Morocco. Meknés-Tafilalet/Drâa-Tafilalet: Border region with Algeria. Near Errachidia. Oued Bou-Ibourine, « Zizaou n´oudad », gps 31.4114, -3.7220, 900 m a.s.l., 11th March 2019, C. Lemmel s.n. (Holotype, HEID 505689; Isotype, G00394714, Conservatoire et jardin botanique de Genève; Paratype, HEID 505749, 505750, ex. cult. Botanical Garden Heidelberg 2019). 

Description: Herbs, woody at base, monocarpic, simple trichomes; rhizome fleshy, 2–3 cm in diam. Stems 80–140(-180) cm tall, robust, up to 1.4 cm in diam, erect, simple at base, often alternately branched in lower part. Basal leaves rosulate, fleshy; leaves lyrate, distal lobecordate, (10-)15–25(-40) cm, margin entire to distantly dentate, numerous simple trichomes on lower surface mostly along veins, upper side loosely covered with simple trichomes; cauline leaves similar but apex obtuse to weakly subacute, 10–15 × 5–7 cm. Raceme ebracteate, elongating in fruit, 40–100 cm; often branched. Sepals erect, saccate ca. 8 mm long, with few simple trichomes; petals pale-yellow,1.5–1.7 cm long, 6–7 mm wide, petal claw 8 mm long, obtuse at apex, glabrous. Filaments tetradynamous, ca. 9 mm long; nectar glands 4, rounded, elateral pair larger. Stigma entire. Infructescence with up to 100(-200) siliques, (30-)40–45(-48) mm, petiolate (9–11 mm). Fruits heteroarthrocarpic with a distal indehiscent balloon-like structure with two viable seeds (3.5–5 × 6–8 mm); proximal part dehiscent, terete (30–45 mm); 20–40 ovules; septum complete. Seeds biseriate, mucilaginous, 1.3–1.4 × 1.4–1.5 mm.

Etymology: Zahora means “flower” in Arabic, indicating the attractive and peculiar appearance of the plant. “Aït-atta” are a Berber tribal confederation of south eastern Morocco who locally know the plant under the name «Zizaou n’oudad» (Barbary-sheep’s cabbage).

Habitat: All places are in sandy beds of oueds flowing from the base of the kreb (cliff) of the Hamada du Guir or the Bin el Korbine.

Ecology: Greenhouse and pollination experiments showed that the species is largely self-compatible. At its natural stands the plant is annual and monocarpic. However, in cultivation the plant species can be kept growing when cutting frutescence. There are two different options of seed release, either directly into a local soil seed bank from the dehiscent part of fruit or via the distal indehiscent part carrying two seeds, which may allow distributing effectively with water in the wadi systems at rare and occasional events.

Figure 2. Distribution of known localities (red dots) of Zahora ait-atta documented from 2015 to 2019 (satellite map was taken from image metadata Copernicus/Landsat).

Figure 3. BEAST analysis of tribe Brassiceae based on ITS DNA sequence data (Suppl. material 2). The new genus Zahora is highlighted, and its respective stem group node is indicated (red dot). Divergence times are given as Mya (million years ago). Agronomically important species, Brassica oleracea and Raphanus sativus, are indicated and shown with their respective clades. Brassica nigra and B. carinata are also indicated with an asterisk (orange).

Conclusion: 

Zahora ait-atta is described as a new species of a new monotypic genus. Zahora shows a peculiar fruit feature, namely heteroarthrocarpic fruits, and the species might mediate evolutionary between Core Oleracea clade (e.g. Brassica oleracea, Brassica napus) and Raphanus sativus and related genera. Both represent important crop plant groups with seeds playing an enormous agronomical role. The diploid new species might, therefore, serve as important germplasm reservoir to study traits and characters in a number of Brassiceae crop plants.

   

 Marcus A. Koch and Claude Lemmel. 2019. Zahora, A New Monotypic Genus from tribe Brassiceae (Brassicaceae) endemic to the Moroccan Sahara. PhytoKeys. 135: 119-131. DOI: 10.3897/phytokeys.135.46946

[Mammalogy • 2019] Parahypsugo happoldorum • A New Genus and Species of Vesper Bat (Chiroptera: Vespertilionidae) from West Africa, with Notes on Hypsugo, Neoromicia, and Pipistrellus

Parahypsugo happoldorum

Hutterer, Decher, Monadjem & Astrin, 2019

 DOI: 10.3161/15081109ACC2019.21.1.001 

 twitter.com/MuseumKoenig ZFMK.de

Abstract
We describe a new species of vespertilionid bat from Guinea and Liberia, West Africa. In this context we evaluate previously described taxa from West Africa assigned to Pipistrellus, Neoromicia, and Hypsugo. Based on genetics, morphology and ecology we conclude that the taxon Pipistrellus eisentrauti bellieri should be elevated to species level, and that the taxa bellieri, crassulus, eisentrauti plus the new species form a monophyletic clade for which a new genus name is proposed. The new genus occurs in forested regions south of the Sahara from Senegal to Ethiopia and Somalia, from where further taxa remain to be described.

Parahypsugo happoldorum


Rainer Hutterer, Jan Decher, Ara Monadjem and Jonas Astrin. 2019. A New Genus and Species of Vesper Bat from West Africa, with Notes on Hypsugo, Neoromicia, and Pipistrellus (Chiroptera: Vespertilionidae). Acta Chiropterologica. 21(1); 1-22. DOI: 10.3161/15081109ACC2019.21.1.001  

twitter.com/MuseumKoenig/status/1184014396475035648

Neue Fledermausgattung aus Westafrika

zfmk.de/de/zfmk/presse/neue-fledermausgattung-aus-westafrika

[Paleontology • 2019] Stratigraphy and Paleobiology of the Upper Cretaceous-Lower Paleogene Sediments from the Trans-Saharan Seaway in Mali

 Reconstruction of the Trans-Saharan Seaway waters from the Late Cretaceous–early Paleogene. Sunlight of the tropics illuminated seawater filled with mangrove roots near shore where freshwater influence was highest. Mollusks lined the floor of the shallow sea, and tube-shaped burrows opened onto the seafloor.

in O'Leary, Bouaré, Claeson, et al., 2019.  

 digitallibrary.AMNH.org

 Illustration: Carl Buell   AMNH.org

Abstract 

An epicontinental sea bisected West Africa periodically from the Late Cretaceous to the early Eocene, in dramatic contrast to the current Sahara Desert that dominates the same region today. Known as the Trans-Saharan Seaway, this warm and shallow ocean was a manifestation of globally elevated sea level associated with the rapid break-up of the supercontinent Gondwana in the late Mesozoic. Although it varied in size through time, the Trans-Saharan Seaway is estimated to have covered as much as 3000 km2 of the African continent and was approximately 50 m deep. The edges of the sea were defined in part by the high topography of the Precambrian cratons and mobile belts of West Africa. Over its approximately 50 million year episodic existence, through six major periods of transgression and regression, the Trans-Saharan Seaway left behind extensive nearshore marine sedimentary strata with abundant fossils. The waters that yielded these deposits supported and preserved the remains of numerous vertebrate, invertebrate, plant, and microbial species that are now extinct. These species document a regional picture of ancient tropical life that spanned two major Earth events: the Cretaceous-Paleogene (K-Pg) boundary and the Paleocene-Eocene Thermal Maximum (PETM). Whereas extensive epeiric seas flooded the interior portions of most continents during these intervals, the emerging multicontinental narrative has often overlooked the Trans-Saharan Seaway, in part because fundamental research, including the naming of geological formations and the primary description and analysis of fossil species, remained to be done. We provide such synthesis here based on two decades of fieldwork and analyses of sedimentary deposits in the Republic of Mali. Northern parts of the Republic of Mali today include some of the farthest inland reaches of the ancient sea. 

We bring together and expand on our prior geological and paleontological publications and provide new information on ancient sedimentary rocks and fossils that document paleoequatorial life of the past. Ours is the first formal description of and nomenclature for the Upper Cretaceous and Lower Paleogene geological formations of this region and we tie these names to regional correlations over multiple modern territorial boundaries. The ancient seaway left intriguing and previously unclassified phosphate deposits that, quite possibly, represent the most extensive vertebrate macrofossil bone beds known from anywhere on Earth. These bone beds, and the paper shales and carbonates associated with them, have preserved a diverse assemblage of fossils, including a variety of new species of invertebrates and vertebrates, rare mammals, and trace fossils. The shallow marine waters included a wide range of paleoenvironments from delta systems, to hypersaline embayments, protected lagoons, and carbonate shoals. 

Our overarching goal has been to collect vertebrate fossils tied to a K-Pg stratigraphic section in Africa. We provide such a section and, consistent with prior ideas, indicate that there is a gap in sedimentation in Malian rocks in the earliest Paleocene, an unconformity also proposed elsewhere in West Africa. Our phylogenetic analyses of several vertebrate clades across the K-Pg boundary have clarified clade-by-clade species-level survivorship and range extensions for multiple taxa. Few macrofossil species from the Trans-Saharan Seaway show conspicuous change at either the K-Pg boundary or the PETM based on current evidence, although results are very preliminary. Building on our earlier report of the first record of rock-boring bivalves from the Paleocene of West Africa, we further describe here a Cretaceous and Paleogene mollusk fauna dominated by taxa characteristic of the modern tropics. Among the newly discovered fossil osteichthyans, large body size characterizes both the pycnodonts and a new freshwater Eocene catfish species, one of the largest fossil catfishes found in Africa. Our new paleoecological and faunal reconstructions show an evergreen, broadleaf forest that included some of the oldest mangroves known. The ancient Malian ecosystem had numerous apex predators including Crocodyliformes, Serpentes, and Amiidae, some of which were among the largest species in their clades. The Trans-Saharan Seaway exhibited intermittent isolation from major seas. This environmental variable may have created aquatic centers of endemism, stimulating selection for gigantism as previously observed for species on terrestrial islands.

Reconstruction of the Trans-Saharan Seaway waters from the Late Cretaceous–early Paleogene. Sunlight of the tropics illuminated seawater filled with mangrove roots near shore where freshwater influence was highest. Mollusks lined the floor of the shallow sea, and tube-shaped burrows opened onto the seafloor.

During the Late Cretaceous–early Paleogene, the shallow waters of the Trans-Saharan Seaway waters were teeming with aquatic species which ranged from small mollusks to giant sea snakes and catfish.

 Illustration: Carl Buell 

Maliamia gigas, Cretalamna (Serratolamna) maroccana, Palaeophis colossaeus,
 Pycnodus jonesae, Plesielephantiform mammal, Lavocatodus giganteus,
Acleistochelys maliensis, Rhabdognathus aslerensis, Myliobatis wurnoensis 
and Adrar des Iforas massif. 

Maureen A. O'Leary, Mamadou L. Bouaré, Kerin M. Claeson, Kelly Heilbronn, Robert V. Hill, Jacob A. McCartney, Jocelyn A. Sessa, Famory Sissoko, Leif Tapanila, Elisabeth Wheeler and Eric M. Roberts. 2019. Stratigraphy and Paleobiology of the Upper Cretaceous-Lower Paleogene Sediments from the Trans-Saharan Seaway in Mali. Bulletin of the American Museum of Natural History. 436; 1–177.   digitallibrary.AMNH.org/handle/2246/6950

Large Sea Snakes, Giant Catfish Once Swam in the Sahara AMNH.org/explore/news-blogs/research-posts/sea-snakes-catfish-sahara

Ancient Saharan seaway shows how Earth's climate and creatures can undergo extreme change phys.org/news/2019-07-ancient-saharan-seaway-earth-climate.html via @physorg_com

[Herpetology • 2017] A Phylogeny of Open-habitat Lizards (Squamata: Lacertidae: Ophisops) Supports the Antiquity of Indian Grassy Biomes

scroll.in
DOI: 10.1111/jbi.12999 

Abstract

Aim: India is dominated by tropical grassy biomes (TGBs), traditionally considered seres or degraded forest, with low diversity relative to the restricted, ancestral wet zone. It is unclear if Indian grasslands and other open habitats are anthropogenically derived or native, old-growth habitats; without a clear timescale of grassland evolution. One way to understand grassland evolution is to study the diversification in taxa restricted to open habitats. We use a dated phylogeny of Ophisops to address questions related to the origin, diversification and inter-relationships of Indian and Saharo-Arabian Ophisops, and ultimately the origin of Indian grasslands and open habitats.

Location: The Indian subcontinent; the Saharo-Arabian Realm.

Methods: We generated up to 2736 base pairs of aligned sequence data (one mitochondrial, two nuclear genes) for Indian lacertids and reconstructed phylogenetic relationships using maximum likelihood and Bayesian inference. We use a fossil-calibrated timetree, diversification analyses and ancestral area reconstructions to test the hypotheses of origin and relationships with Saharo-Arabian Ophisops.

Results: Ophisops is strongly supported as monophyletic, with a basal split into a large-bodied (LBC) and small-bodied clade (SBC). The Saharo-Arabian species are nested within the Indian species in the LBC. Species diversity in Indian Ophisops is grossly underestimated, with 26–47 candidate species. Ophisops began diversifying in the late Oligocene with significant rate shifts in the late Miocene-Pliocene and Pleistocene within the SBC.

Main conclusions: Our results are consistent with an ancient origin of grassland taxa and TGBs in India. Ophisops is a dramatic example of overlooked cryptic diversity outside forests, with ~30 species where five were known. Ophisops dispersed into India from the Saharo-Arabian Realm in the Oligocene with a back dispersal in the Middle Miocene, a novel biogeographical pattern. Diversification in the SBC of Ophisops increased 8-fold during the global C4 grassland expansion. Indian TGBs are old-growth ecosystems that need urgent conservation attention.

Habitats of Ophisops leschenaultii species complex, Tumkur District, Karnataka. 

  

Ishan Agarwal and Uma Ramakrishnan. 2017. A Phylogeny of Open-habitat Lizards (Squamata: Lacertidae: Ophisops) Supports the Antiquity of Indian Grassy Biomes.  Journal of Biogeography. DOI: 10.1111/jbi.12999

How lizards revealed the millennia-old evolution story of India’s grasslands

scroll.in/magazine/836758/how-lizards-revealed-the-ancient-evolution-story-of-indias-grasslands via @scroll_in

  

  

[Mammalogy • 2017] Neoromicia stanleyi • A New Species of Neoromicia (Chiroptera: Vespertilionidae) from southern Africa: A name for “N. cf. melckorum”

 Neoromicia stanleyi 

 Goodman, Kearney, Michèle, Ratsimbazafy & Hassanin, 2017 

 DOI:  10.11646/zootaxa.4236.2.10 

Abstract

The taxonomy of sub-Saharan small insectivore bats of the family Vespertilionidae is unresolved and currently five named species of the genus Neoromicia are recognized from southern Africa, with N. melckorum considered a synonym of N. capensis. Since several years, the name “N. cf. melckorum” has been used in the literature to designate an apparently undescribed and moderately large bodied vespertilionid bat known from different localities in southern and southeastern Africa. Using new data from molecular genetics, bacular morphology, and cranio-dental characters, we conclude that N. melckorum sensu stricto is indeed nested within N. capensis and obtain the needed evidence to formally describe “N. cf. melckorum”, named herein as Neoromicia stanleyi sp. nov. On the basis of molecular and bacular evidence, N. stanleyi is found in Botswana, Zimbabwe, and Zambia, and using a combination of other characters is presumed to occur in northern South Africa and Malawi. Bayesian and maximum likelihood analyses based on 12S rRNA sequences indicate that it belongs to a clade containing four species of Neoromicia (N. capensis, N. malagasyensis, N. matroka, and N. robertsi) and Laephotis. Neoromicia stanleyi shows at least 3.2% nucleotide divergence from its closest relatives. It is larger in cranial characters than other members of the capensis group occurring in the southern portion of Africa, and a number of bacular characters distinguish N. stanleyi from N. capensis.

Keywords: Mammalia, taxonomy, morphology, molecular genetics, Neoromicia, new species, southern Africa

 Steven M. Goodman, Teresa Kearney, Malalatiana Michèle, Ratsimbazafy and Alexandre Hassanin. 2017. Description of A New Species of Neoromicia (Chiroptera: Vespertilionidae) from southern Africa: A name for “N. cf. melckorum”.
Zootaxa. 4236(2); 351–374. DOI:  10.11646/zootaxa.4236.2.10

[Mammalogy • 2016] Petrosaltator gen. nov., A New Genus Replacement for the North African Sengi Elephantulus rozeti (Macroscelidea; Macroscelididae)

FIGURE 3. Images of the three genera of the tribe Macroscelidini.
A. Basking North African sengi (Petrosaltator rozeti). Adult male near Salas village, Jhilet Mountains, Marrakesh, Morocco on 18 July 2005, specimen number CAS MAM 27982. Photo G. Rathbun.
B. The four-toed sengi (Petrodromus tetradactylus), Mareja Community Reserve, Pemba, northern Mozambique, 17 June 2011, specimen number CAS MAM 29347, Photo G. Rathbun.
C. The Namib round-eared sengi (Macroscelides flavicaudatus), south of the Micberg formation, Kunene Region, Khorixas District, 7 May 2010, specimen number CAS MAM 29700, photo J.P. Dumbacher.

DOI: 10.11646/zootaxa.4136.3.8

Abstract

In 2003, a molecular phylogeny was published that examined the role of the Sahara Desert as a vicariant event in the evolution of sengis (also known as elephant-shrews.) The phylogeny included a single sample from the North African sengi, Elephantulus rozeti (Duvernoy, 1833), which was found to be more closely related to the sengi genus Petrodromus Peters, 1846 than to other Elephantulus. Here we independently test the monophyly of Elephantulus using an additional specimen of E. rozeti and similar phylogenetic analyses, and discuss additional morphological and behavioral data that support the phylogeny. We propose a revised taxonomy that reflects the current paraphyly of Elephantulus and the sister relationship of E. rozeti and Petrodromus, including a new genus name for the North African sengi, Petrosaltator rozeti gen. nov., nov. comb. We additionally define two tribes within the subfamily Macroscelidinae, the Macroscelidini (including Macroscelides, Petrodromus, and Petrosaltator), and the Elephantulini (including all other members of Elephantulus).

Keywords: Mammalia, Sengi, Elephant-shrew, Petrosaltator, North African Sengi

Basking North African sengi Petrosaltator rozeti. Adult male near Salas village, Jhilet Mountains, Marrakesh, Morocco on 18 July 2005, specimen number CAS MAM 27982.

Photo G. Rathbun.   DOI: 10.11646/zootaxa.4136.3.8

Petrosaltator Rathbun and Dumbacher, new genus

Type species: Petrosaltator rozeti (Duvernoy, 1833)

Holotype. The type specimen is located at Musée Zoologique de la ville de Strasbourg, France; specimen number MZSMAM03685.

Geographic distribution. Petrosaltator has a unique range, and is currently the only species of the family Macroscelididae that occurs north of the Sahara Desert. It is known from the Maghreb Region of northern Africa, in Mediterranean, sub-desert, and montane zones from near sea level to 2725m elevation (Fig. 1) (Corbet & Hanks 1968; Cuzin & Séguignes 1990).

Diagnosis and description. The genus Petrosaltator is monotypic (P. rozeti) (Corbet & Hanks 1968; Duvernoy 1833; Perrin & Rathbun 2013). Features that distinguish the genus from all other Macroscelidinae are rare (see below), thus explaining why it was included in Elephantulus by earlier workers. Genetic data are among the most useful diagnostic characters, and have been used here and elsewhere to identify P. rozeti and clearly align it with Petrodromus and Macroscelides (Douady et al. 2003; Kuntner et al. 2011; Smit et al. 2011).

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Etymology. The roots of Petrosaltator (masculine gender) are Greek (petro) and Latin (saltator), together meaning “rockdancer”. This genus name reflects the habitats occupied by this species, which are dominated by rocks and boulders (Séguignes 1988). Petrosaltator also alludes to the phylogenetic relationship with Petrodromus (meaning rockrunner with Greek roots), although oddly Petrodromus tetradactylus is not specifically a petrophile (Jennings & Rathbun 2001). We suggest that the common name of Petrosaltator rozeti continue to be the North African Sengi or Elephant-shrew

The taxonomic hierarchy for the subfamily Macroscelidinae follows:

Class: Mammalia Linneus, 1758

 Supercohort: Afrotheria Stanhope et al., 1998

 Order: Macroscelidea Butler, 1956

 Family: Macroscelididae Bonaparte, 1838

 Subfamily: Macroscelidinae Bonaparte, 1838

 Tribe: Macroscelidini, new tribe

 Genus: Macroscelides A. Smith 1829

 Genus: Petrodromus Peters 1846

 Genus: Petrosaltator, new genus

 Tribe: Elephantulini, new tribe

 Genus: Elephantulus Thomas & Schwann 1906

Macroscelidini, New Tribe

Type genus: Macroscelides A. Smith 1829

Description: The tribe is defined primarily by the genera included—Macroscelides, Petrodromus, and Petrosaltator. Synapomorphies recovered from genetic analyses include portions of vWF, IRBP, and mtDNA 12s–16s ribosomal RNA loci, which are all consistent with the monophyly of the tribe. Morphological synapomorphies for the group are not easy to discern, although these may include a penis with two lateral lobes and a narrowing end (Woodall 1995b) and the presence of a fully ossified stapediofacial tube (Benoit et al. 2013). Analyses of basal skull morphometrics also appear to support the monophyly of Macroscelidini (Scalici & Panchetti 2011).

Elephantulini, New Tribe

Type Genus: Elephantulus Thomas and Schwann, 1906, by monotypy

Description: The tribe provisionally includes only members of the genus Elephantulus (not including Petrosaltator rozeti) and is supported primarily by genetic synapomorphies. All members have three pairs of mammae, hallux present, auditory bullae not grossly inflated (Corbet & Hanks 1968), and penis morphology in which the urethra does not extend beyond the lateral lobes (Woodall 1995b). Not all members of the genus Elephantulus have been included in phylogenetic studies, so we include all members provisionally. 

In addition to the placement of P. rozeti with Petrodromus and Macroscelides into the tribe Macroscelidini, our genetic data suggest a possible phylogenetic split within Petrodromus tetradactylus, with one population in the Udzungwa Mountains of Tanzania (CASMAM28170 and CASMAM28171) and another occurring from at least southeastern Tanzania (Douady et al. 2003) to KwaZulu Natal in South Africa (GenBank numbers EU136156, EU136145, and EU136138), at the southern end of the current distribution of the genus (Rathbun 2015). Divergence within Petrodromus tetradactylus suggests that more research is needed to understand subspecies diversity in this monotypic genus. 

With our renaming of Petrosaltator, the subfamily Macroscelidinae now contains four genera (Elephantulus, Macroscelides, Petrosaltator, and Petrodromus). The number of extant species in the order, however, remains the same at 19 (Dumbacher et al. 2014). The diversity of extant taxa within the order continues to slowly increase with a better understanding of the underlying phylogenetics. Although this trend may continue as we learn more, extant species diversity in the order Macroscelidea remains remarkably low compared to other non-Afrotherian mammalian radiations in Africa (Kingdon et al. 2013; Rathbun 2009).

John P. Dumbacher, Elizabeth J. Carlen and Galen B. Rathbun. 2016. Petrosaltator gen. nov., A New Genus Replacement for the North African Sengi Elephantulus rozeti (Macroscelidea; Macroscelididae). Zootaxa. 4136(3); DOI: 10.11646/zootaxa.4136.3.8

[Paleontology • 2016] Morphofunctional Analysis of the Quadrate of Spinosauridae (Dinosauria: Theropoda) and the Presence of Spinosaurus and a Second Spinosaurine Taxon in the Cenomanian of North Africa

two species of Spinosaurinae, and ascribed to Spinosaurus aegyptiacus and ?Sigilmassasaurus brevicollis from the Cenomanian of North Africa

Illustration: S. Krasovskiy atrox1.deviantart.com DOI:  10.1371/journal.pone.0144695

Abstract

Six quadrate bones, of which two almost certainly come from the Kem Kem beds (Cenomanian, Upper Cretaceous) of south-eastern Morocco, are determined to be from juvenile and adult individuals of Spinosaurinae based on phylogenetic, geometric morphometric, and phylogenetic morphometric analyses. Their morphology indicates two morphotypes evidencing the presence of two spinosaurine taxa ascribed to Spinosaurus aegyptiacus and ?Sigilmassasaurus brevicollis in the Cenomanian of North Africa, casting doubt on the accuracy of some recent skeletal reconstructions which may be based on elements from several distinct species. Morphofunctional analysis of the mandibular articulation of the quadrate has shown that the jaw mechanics was peculiar in Spinosauridae. In mature spinosaurids, the posterior parts of the two mandibular rami displaced laterally when the jaw was depressed due to a lateromedially oriented intercondylar sulcus of the quadrate. Such lateral movement of the mandibular ramus was possible due to a movable mandibular symphysis in spinosaurids, allowing the pharynx to be widened. Similar jaw mechanics also occur in some pterosaurs and living pelecanids which are both adapted to capture and swallow large prey items. Spinosauridae, which were engaged, at least partially, in a piscivorous lifestyle, were able to consume large fish and may have occasionally fed on other prey such as pterosaurs and juvenile dinosaurs.

Systematic Paleontology

Dinosauria Owen, 1842 

Saurischia Seeley, 1887 

Theropoda Marsh, 1881 

Tetanurae Gauthier, 1986 

Megalosauroidea (Fitzinger, 1843) Walker 1964 

Spinosauridae Stromer, 1915 

Spinosaurinae (Stromer, 1915) Sereno et al., 1998 

Description: The six isolated quadrates from the Kem Kem beds of Morocco clearly belong to two morphotypes (Figs 2–4) based on the size and outline of the quadrate foramen, shape of the mandibular articulation, and outline, surface, and orientation of the quadratojugal contacts. Measurements taken on each quadrate (Fig 5A–5D) are provided in Table 1.

Spinosaurus Stromer, 1915 

Spinosaurus aegyptiacus Stromer, 1915 

?Sigilmassasaurus Russel, 1996 

?Sigilmassasaurus brevicollis Russel, 1996 

Spinosaurus aegyptiacus and ?Sigilmassasaurus brevicollis

by S. Krasovskiy atrox1.deviantart.com

Conclusion

The description and identification of six isolated quadrates, among which two most probably come from the Kem Kem beds of Morocco, provide additional information on the Cenomanian dinosaur fauna of North Africa. Based on cladistic, geometric morphometric, and phylogenetic morphometric analyses, two morphotypes have been successfully identified as belonging to two species of Spinosaurinae, and ascribed to Spinosaurus aegyptiacus and ?Sigilmassasaurus brevicollis. This study provides the first convincing evidence of two spinosaurine taxa in the Cenomanian of North Africa based on cranial material, casting doubt on the recent reconstruction of a quadrupedal Spinosaurus which may be based on individuals belonging to two different species of Spinosaurinae.

Ontogenetic changes occurring in the spinosaurid quadrates include the suture of the quadrate and quadratojugal, delimitation of the mandibular condyles and squamosal capitulum, and development of a ventral projection of the dorsal quadratojugal contact and a second quadrate ridge ventral to the quadrate head. Based on the quadrate proportions and estimated skull length of Baryonyx and Spinosaurus, quadrates of mature individuals from Morocco belong to animals with a skull length of no more than 120 cm. This suggests that very large forms of Spinosaurus may have been rare in the Kem Kem assemblages.

Morphofunctional analysis of the spinosaurid quadrates has revealed peculiar jaw mechanics in these specialized theropods. An helicoidal and strongly lateromedially oriented joint of the jaw articulation allowed the lateral displacement of the mandibular ramus when the lower jaw was depressed. This lateral movement of the ramus was possible due to a movable mandibular symphysis as the dentaries were joined by connective tissues, and allowed the pharynx to be widened. A similar jaw articulation was convergently present in pterosaurs and particularly pelecanids which also have a mandibular symphysis restricted to the anterior extremity of the mandible. Spinosauridae, which are considered to be semi-aquatic and partially piscivorous animals, were able to swallow large prey such as fish in the same way as pelecanids.

  

Fig 15. Jaw mechanics in the spinosaurid Spinosaurus.
A–D, Mandibular articulation; and F, G, skull in A, C, F–G, lateral; and B, D, anterior views; when A–B, F, the mouth is closed; and C–D, G, fully open, illustrating the lateral movement (in red) of the mandibular ramus for a 45° rotation of the lower jaw (courtesy of Jaime A. Headden); E, skeletal reconstruction of Spinosaurus aegyptiacus by Ibrahim et al. [22]) in swimming position in lateral view with a human (1.8 m) as a scale (modified from Ibrahim et al. [22]). This model is based on spinosaurid cranial and postcranial remains (colored in red) from the Albian-Cenomanian of Northern Africa which possibly belong to two spinosaurine taxa (see also Evers et al. [27]); H, reconstruction of a semi-aquatic Spinosaurus in fishing position (i.e., jaws wide open) in anterolateral view (courtesy of Jason Poole).

Abbreviations: an, angular; ar, articular; d, dentary; ecc, ectocondyle; enc, entocondyle; j, jugal; m, maxilla; n, nasal; p, parietal; pm, premaxilla; po, postorbital; pt, pterygoid; ptf, pterygoid flange; q, quadrate; qf, quadrate foramen; qj, quadratojugal; retp, retroarticular process of the articular; sa, surangular; sq, squamosal.
DOI: 10.1371/journal.pone.0144695

Christophe Hendrickx , Octávio Mateus and Eric Buffetaut. 2016. Morphofunctional Analysis of the Quadrate of Spinosauridae (Dinosauria: Theropoda) and the Presence of Spinosaurus and a Second Spinosaurine Taxon in the Cenomanian of North Africa.
 PLoS ONE. 11(1): e0144695.  DOI: 10.1371/journal.pone.0144695