The holotypes of the upper Pleistocene Crocuta crocuta spelaea (Goldfuss, 1823: Hyaenidae) and Panthera leo spelaea (Goldfuss, 1810: Felidae) of the Zoolithen Cave hyena den (South Germany) and their palaeo‐ecological interpretation

The rediscovered holotype skulls of Late Pleistocene Panthera leo spelaea ( Goldfuss, 1810 ) (Felidae) and Crocuta crocuta spelaea ( Goldfuss, 1823 ) (Hyaenidae) from the Zoolithen Cave at Burggeilenreuth, southern Germany, are discussed. The cave became famous mainly due to its rich cave bear bone remains from the late Saalian (OIS 6–8) to Eemian/Weichselian (OIS 3–6) including additionally a third holotype of Ursus spelaeus Rosenmüller, 1794 (Ursidae). The ‘Felis spelaea’ holotype represents an adult male with a strong bite mark on the saggital crest, which was in an early stage of healing. Compared with other European Late Pleistocene lion skulls and skeletons, and with modern African lions, it provides evidence of intraspecific conflict between male Ice Age lions. The holotype of ‘Hyaena spelaea’ is one of several hundred hyena remains from a well‐frequented hyena den cave. The cave was used intensively by Late Pleistocene hyena clans, for collecting lion carcasses in addition to their accustomed prey, as happened in many caves throughout Europe. Ice Age spotted hyena clans might have killed Ice Age steppe lions for many reasons, such as fights over prey and territory, and the protection of cubs, but they did not always scavenge on their carcasses. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 822–831.     

Is the morphological disparity of lemur skulls (Primates) controlled by phylogeny and/or environmental constraints?

The cranial morphology of Lemur catta and of the five species of the genus Eulemur is investigated here by landmark identification and Procrustes superimpositions. This geometrical morphometric method makes it possible to describe pure shape differences independent of size effects, and to quantify differences between specimens. The aim of this study is to determine whether the morphological disparity of lemur skulls is constrained by the environment and/or by the group's phylogenetic history. First, Procrustes residuals are analysed by principal component analysis and the scatter-plots interpreted against the geographical distributions of taxa to determine whether morphology is correlated with geography. Then, a morphological distance tree is computed and compared with various cladograms reported in the literature to test for any correlation between morphology and phylogeny. Morphological disparity is found to be closely correlated with geographical distribution but independent of phylogeny. This confirms that the morphological disparity of lemur skulls is associated with a high degree of homoplasy, probably as a result of ecological constraints.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 577–590.     

Geometric morphometrics, neural networks and diagnosis of sibling Taterillus species (Rodentia, Gerbillinae)

The lack of diagnostic traits in sibling species means that great quantities of biogeographical and ecological data held in museum collections cannot be utilized effectively, leading to underestimates of biodiversity. In this study we applied neural networks (NN) and canonical variates analyses (CVA) to landmark measurements on the skulls of the West African species Taterillus arenarius , T. petteri , T. gracilis and T. pygargus in an attempt to discriminate species previously identified unambiguously from their karyotypes. Among suggested differences, the relationship between inflation of the tympanic bullae and lower population density is discussed. Cross-validated classification rates did not exceed 73%. Two hypotheses are proposed to explain such high phenotypic similarity: morphological plasticity limited by environmental constraints and the possibility that speciation has been too recent to allow significant morphological divergence.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 319−327.     

A new species of Encephalartos (Zamiaceae) in Sudan

Encephalartos mackenziei Newton is described as a new cycad species occurring in south-east Sudan. It is characterized by its branching at the base to produce up to ten trunks per plant and by its broad leaflets, as well as some details of the cone scales. The population is large and widespread, and the presence of small seedlings indicates successful regeneration.  © The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 187–192.     

An analysis and revision of Cyclamen L. with emphasis on subgenus Gyrophoebe O. Schwarz

Studies in the areas of comparative morphology and palynology of the genus Cyclamen L. (Primulaceae) are outlined, and used in combination with observations and analyses of past works and publications and of field data (gathered by the Cyclamen Society and others) to generate new data sets. These are subject to cladistic parsimony analysis for the entire genus, and to phenetic ordinal analysis for subgenus Gyrophoebe O.Schwarz, to reach a conclusion on the validity of the latter classification, and the separation of taxa within it. A phylogeny and subgeneric reclassification is proposed with the reinstatement of Cyclamen elegans Boiss. & Buhse at species level based on the results gathered. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 138 , 473–481.     

The influence of environmental factors on the morphology of red‐backed voles Myodes gapperi (Rodentia,Arvicolinae) in Québec and western Labrador

Geographical patterns of morphological variation in small mammals are often associated with environmental factors. The southern red‐backed vole Myodes gapperi is a widespread and abundant small mammal in Canada, occurring in environments as diverse as mixed‐wood forests and taiga. First upper molars and skulls from nine populations of southern red‐backed voles distributed across three ecozones and approximately 10° of latitude were analysed by means of geometric morphometric techniques, and their relationships with environmental variables were examined. A weak, non‐linear trend of size increase towards higher latitudes was observed in voles' skulls. Environmental variables appeared to be important drivers of shape differentiation among populations from the three distinct ecozones analysed. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 204–218.     

The morphology and systematics of Mammalodon colliveri (Cetacea: Mysticeti), a toothed mysticete from the Oligocene of Australia

Mammalodon colliveri is an unusual toothed archaic mysticete (Cetacea) from the Upper Oligocene Jan Juc Formation of south‐east Australia. The morphology of the holotype skull and postcrania are described in detail. Superimposed on the generally plesiomorphic archaeocete‐like morphology of Mammalodon are subtle mysticete synapomorphies. Derived features of Mammalodon include a short and bluntly rounded rostrum, reduced premaxillae, and anterodorsally directed orbits. Within Mysticeti, this suite of features is unique. The aberrant rostral morphology of Mammalodon suggests specialization for suction feeding. Janjucetus hunderi is placed in an expanded family Mammalodontidae. Phylogenetic analysis corroborates the monophyly of Basilosauridae, Neoceti, Odontoceti, and Mysticeti, and yields a novel hypothesis of toothed mysticete relationships: a basal clade of undescribed toothed mysticetes from South Carolina, a Llanocetidae + Mammalodontidae clade, and monophyletic Aetiocetidae are posited as successive sister taxa to edentulous baleen whales (Chaeomysticeti). Toothed archaic mysticetes clearly employed diverse prey capture strategies, with exaptations for filter feeding evolving sequentially in stem group Mysticeti. A stratigraphically calibrated phylogeny implies that the initial diversification of Mysticeti occurred during the Late Eocene. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 367–476.     

The most basal beaked whale Ninoziphius platyrostris Muizon, 1983: clues on the evolutionary history of the family Ziphiidae (Cetacea: Odontoceti)

Ninoziphius platyrostris, from the late Neogene of Peru, is one of the best‐known fossil beaked whales (Odontoceti: Ziphiidae), with a holotype including the skull with ear bones, mandibles, teeth, and postcranial elements. Furthermore, based on several characters, including a complete functional upper and lower dentition, it is usually considered as one of the most archaic ziphiids. However, the poorly preserved dorsal portion of the holotype skull has led to unresolved phylogenetic relationships. With the addition of two newly prepared skulls from the same Peruvian locality we redescribed N. platyrostris. In the light of recent ziphiid discoveries, an emended diagnosis of the species is proposed here. In our cladistic analysis Ninoziphius is the most basal stem ziphiid. Newly observed or reassessed morphological traits allow functional and ecological considerations. The morphology of the oral apparatus suggests that Ninoziphius was less specialized for suction feeding than most extant ziphiids. Tooth wear in the holotype may indicate benthic feeding. Although the vertebral column of Ninoziphius corresponds to less developed locomotor abilities for deep dives, its cranial morphology does not provide definitive arguments for an echolocation system less efficient than in deep diving extant ziphiids. Finally, the phylogenetic tree produced was used to detail the evolutionary history of several major ziphiid features (dental reduction, development of mandibular tusks, and increased body size). © 2013 The Linnean Society of London     

Reassessment of cf. Halticosaurus orbitoangulatus from the Upper Triassic (Norian) of Germany – a pseudosuchian,not a dinosaur

The holotype of cf. Halticosaurus orbitoangulatus Huene, 1932, comprises an incomplete and macerated but associated skull of an archosaurian reptile from the middle (second) Stubensandstein (middle Löwenstein Formation; Upper Triassic: Norian) of Baden‐Württemberg, Germany. It was originally interpreted as a theropod dinosaur but more recently it has been suggested that this taxon has crocodylomorph affinities. Detailed preparation of the holotype of cf. H. orbitoangulatus has revealed much new anatomical information and permitted reassessment of its affinities. The maxilla lacks both a distinct antorbital fossa and a medial bony lamina bordering the antorbital fenestra. The lateral surface of the dentary bears a pronounced horizontal ridge. The squamosal differs from that of basal crocodylomorphs in being L‐shaped rather than arcuate in dorsal view, lacking a dorsolateral overhang, and lacking an interlocking contact with the paroccipital process as, for example, in the basal crocodylomorph Saltoposuchus connectens from the same horizon and locality. Phylogenetic analysis placed cf. H. orbitoangulatus amongst loricatan pseudosuchians (but not amongst Crocodylomorpha) rather than amongst theropod dinosaurs. The holotype of cf. H. orbitoangulatus represents a previously unrecognized taxon of loricatan pseudosuchian, which is here named Apatosuchus orbitoangulatus and set apart from other known Norian‐age non‐crocodylomorph loricatans by its apparently much smaller size. © 2013 The Linnean Society of London     

Reproductive effort and sex allocation strategy in Commelina benghalensis L., a common monsoon weed

Commelina benghalensis L. exhibits variability in both foliar and floral features; every plant bears three types of branches and four types of flowers. The branches are negatively geotropic, positively geotropic and diageotropic. The flowers are uni- or bisexual, chasmogamous and cleistogamous. This variability influences the breeding system as well as resource allocation to male and female functions. The plants allocate c. 15% of their total resources to reproduction, the major part of which (68.9%) is devoted to production of aerial branches. The proportion of reproductive effort (RE) allocated to various branch systems is correlated with the availability of resources at the time of their differentiation. The pollen/ovule (P/O) ratio, female : male biomass ratio and reproductive output vary between different flower and branch types; variation is more pronounced in the latter. These variations notwithstanding, the results are in line with Charnov's sex allocation theory. The cleistogamous flowers of aerial branches are, however, an exception, being male- rather than female-biased. The reason behind the deviation is, in all probability, their recent evolution from chasmogamous flowers.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 403−413.     

Chromosomal criteria and their phylogenetic implications in the genus Onobrychis Mill. sect. Lophobrychis (Leguminosae), with special reference to Egyptian species

Chromosome numbers and measurements were recorded in 22 individuals from six populations of six species of Onobrychis , including the Egyptian species and most representatives of section Lophobrychis . The basic number of chromosomes was either x = 7 or x = 8 and the chromosomes were medium to medium-small, ranging in length, from c . 1.6 μm to 2.6 μm. Two new ploidy levels were found, 2 n = 4 x = 28 in O. bobrovii Grossh. and 2 n = 4 x = 32 in O. pulchella Schrenk. The origin of the chromosome numbers, geographical distribution and evolution of the species were assessed. Comparison of the data with those in the literature revealed that the observed interspecific variability among section Lophobrychis can be useful in taxonomic delimitation and demonstrates a complexity of evolution between the diploid and polyploid species. Section Lophobrychis has a comparatively highly derived organization and can be considered as a heterogeneous unit in the genus Onobrychis .  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 409–414.     

Two new basal crocodylomorph archosaurs from the Lower Jurassic and the monophyly of the Sphenosuchia

We report on and name two new taxa of basal crocodylomorph archosaurs from the Lower Jurassic, Litargosuchus leptorhynchus gen. et sp. nov. , from the upper Elliot Formation (Stormberg Group) of South Africa, and Kayentasuchus walkeri gen. et sp. nov. , from the Kayenta Formation (Glen Canyon Group) of Arizona, USA. Examination of this material led to a reconsideration of basal crocodylomorph interrelationships. A phylogenetic analysis found no support for the monophyly of Sphenosuchia.  © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society , 2002, 136 , 77–95.     

The cranial anatomy of Chinese placodonts and the phylogeny of Placodontia (Diapsida: Sauropterygia)

Placodonts are Triassic marine reptiles that inhabited the eastern and western margins of the Tethys Ocean (modern South China and Europe/Middle East). Although the crania of European taxa are relatively well understood, those of Chinese taxa have not been extensively studied, and most of them have not been incorporated into a comprehensive phylogeny. Here we present the first reconstructions of all known Chinese placodont holotype skulls using micro‐computed tomographic (μCT) scanning and/or detailed anatomical study. We also present the first phylogenetic analyses that incorporate all placodont genera using a general diapsid matrix that includes postcranial characters, and a placodont‐only cranial matrix. Results vary between the matrices; however, both support a monophyletic Placodontia with eastern taxa interspaced throughout, indicating no major separation between the eastern and western Tethyan realms. Support is strong for a western Tethyan origin of Placodontia, although the highly nested Placochelyidae first appear in the upper Middle Triassic of the eastern Tethys. Thus, all placodont clades appear to have originated in a period of intense speciation during the Middle Triassic. © 2015 The Linnean Society of London     

Killer sperm whale: a new basal physeteroid (Mammalia, Cetacea) from the Late Miocene of Italy

Zygophyseter varolai , a new genus and species of Physeteroidea (Cetacea, Odontoceti), is based on an almost complete skeleton from the Late Miocene (Tortonian) in southern Italy. The extreme elongation of the zygomatic process of the squamosal and the circular supracranial basin (probably for housing the spermaceti organ) delimited by a peculiar anterior projection of the supraorbital process of the right maxilla are the most distinctive features of this bizarre sperm whale. Large body size, large teeth present in both lower and upper jaw, and anteroposteriorly elongated temporal fossa and zygomatic process of the squamosal indicate that this cetacean (for which we suggest the English common name killer sperm whale) was an active predator adapted to feeding on large prey, similarly to the extant killer whale ( Orcinus orca ). A phylogenetic analysis reveals that Zygophyseter belongs to a Middle–Late Miocene clade of basal physeteroids, together with Naganocetus (new genus for the type of ' Scaldicetus ' shigensis ). Moreover, the phylogenetic analysis shows evidence of a wide physeteroid radiation during the Miocene and that the extant Physeter and Kogia belong to two distinct families that form a clade representing the crown-group Physeteroidea.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 148 , 103–131.     

Ecomorphological correlates of craniodental variation in bears and paleobiological implications for extinct taxa: an approach based on geometric morphometrics

Relative warp analyses of landmarks describing cranial and mandibular shape are used for investigating patterns of morphological variation among extant bears (Mammalia, Carnivora, Ursidae) indicative of diet and feeding behavior. These patterns are used for deriving inferences about the autecology of two extinct species previously assumed to have had different dietary preferences, the North American giant, short-faced bear Arctodus simus and the Eurasian cave bear Ursus spelaeus . Results reveal a set of shared craniodental traits among the herbivorous bears, including short and vaulted skulls with well-developed zygomatic arches, lateralized orbits and small canines, concave jaws with a highly positioned condyle, large moment arms for the temporalis and masseter muscles, and long cheek teeth. In contrast, those bears that consume animal resources have long skulls with small zygomatic arches, frontalized orbits and well-developed canines, and long jaws with a deep mandibular symphysis, low muscle leverages, a condyle situated at the level of the tooth row and reduced cheek teeth. The craniodental morphology of omnivorous bears is intermediate between those of faunivores and herbivores. This is also the case of the short-faced bear and the cave bear, which suggests that previous reconstructions of the feeding ecology of these extinct species (highly carnivorous for A. simus and herbivorous for U. spelaeus ) should be revised.     

A multidisciplinary approach to discriminating different taxa in the species complex Pachycondyla villosa (Formicidae)

A multidisciplinary approach provides new evidence that strongly supports the differentiation of three taxa previously confounded in a single species, Pachycondyla villosa . All specimens studied were collected in the same location at Ilhéus, Bahia, Brazil. Bivariate plots of petiole heights vs. petiole lengths and isozyme patterns clearly differentiate the three taxa. Chain lengths and chemical structures of cuticular hydrocarbons are characteristic of each taxon. This congruent evidence shows that there are three different valid species. Ecological hypotheses are discussed to explain the benefits of these species remaining sympatric. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 249–259.