Cytochrome b sequences reveal Acomys minous (Rodentia, Muridae) paraphyly and answer the question about the ancestral karyotype of Acomys dimidiatus

Sequences of the cytochrome b (cyt b) mitochondrial gene show that the spiny mouse Acomys from Crete, known as the endemic species A. minous, is composed of two distinct maternal lineages ("A" and "B"). Group "A" sequences cluster with A. nesiotes (Cyprus) and group "B" sequences cluster with A. cilicicus (Turkey), which is evidence of paraphyly of A. minous in regard to these two species. From cyt b sequences, the three taxa are very closely related to A. cahirinus (Egypt): the maximum divergence found among these sequences is 1.6%, which is equivalent to the intraspecific diversity observed in other Acomys species. Paleozoology evidenced that man unintentionally introduced Acomys into Crete and Cyprus during antiquity. The divergence time between Acomys cyt b sequences found in Crete was estimated at 0.4 Myr, which means that the diversity observed did not appear after the introduction but reflects a much more ancient polymorphism. Cytochrome b phylogeny and cytogenetic data together comprise evidence that, within the species A. dimidiatus (Saudi Arabia, Israel, Egypt), it is the karyotypic form with 36 chromosomes that derives from the form with 38 chromosomes, due to a single acrocentric fusion.     

Taxonomic relationships between Pachycerus and Rhabdorrhynchus (Coleoptera: Curculionidae: Lixinae), with descriptions of two new species of Rhabdorrhynchus from the Arabian Peninsula

Taxonomic relationships between Rhabdorrhynchus and Pachycerus are discussed, and two new species of Rhabdorrhynchus are described: Rhabdorrhynchus sauditus (type locality: Saudi Arabia, As Shāqqah al Yamānīyah, 19°42'N, 40°48'E) and Rhabdorrhynchus emir (type locality: United Arab Emirates, Jebel Ali, 24°59'N, 55°43'E). Some hypotheses on the distribution of the new species in the Arabian Peninsula, based on a biogeographical analysis, are proposed.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 152 , 25–37.     

A morphological, karyological and chemical study of the Apteranthes (Caralluma) europaea complex

The Apteranthes europaea complex (Apocynaceae–Asclepiadoideae–Ceropegieae–Stapeliinae) distributed from Morocco/southern Spain along the North African coast of the Mediterranean Sea up to Sinai, Negev and southern Jordan is studied with emphasis on stem and flower morphology by SEM, chromosome sizes and flavonoid compounds. Apteranthes europaea was found to be rich in rather uncommon flavone glycosides. Of these, luteolin 4'-neohesperidoside represents the major flavonoid of all samples; luteolin-3'- O -(6'- O -sinapoylglucoside)-4'- O -neohesperidoside and luteolin-3'- O -(6'- O -feruloylglucoside)-4'- O -neohesperidoside are reported here for the first time. Flavonol glycosides also occur, but in much smaller quantities. The different flower morphs, variation in stem and corolla epidermal structures, slight variations in the length of the 2 n  = 22 chromosomes and in quantitative flavonoid composition are taxonomically best reflected by subdividing the complex into a western var. europaea (Europe and Africa) and an eastern var. judaica (Sinai, Arabia).  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 419–432.     

New data on the evolution of the Cretan spiny mouse,Acomys minous (Rodentia: Murinae), shed light on the phylogenetic relationships in the cahirinus group

The karyotype of the Cretan spiny mouse Acomys minous was examined with chromosome banding techniques in 53 individuals from 12 localities of Crete, aiming to gain a more detailed knowledge on the chromosomal constitution and variability of its natural populations. We found that it consists of three Robertsonian (Rb) populations with 2n = 38, 2n = 40 and 2n = 42, respectively, the last one being reported for the first time, and with stable fundamental number (FNa = 66, FN = 68). The G‐banding pattern proves that the Rb populations are closely linked phylogenetically by the many common Rb fusions and the lack of monobrachial homologies. In addition, they appear to freely mate at their contact areas, producing viable and fertile hybrids. No other type of chromosomal rearrangement appears to have played part in the chromosomal evolution of this species, at least in the recent past, as indicated also by the study of the telomeric sequences. Heterochromatin appears to be restricted to the pericentromeric position of all acrocentric and most biarmed autosomes, as well as of the X chromosome, whereas the Y chromosome is uniformly, yet faintly heterochromatic. Chromosome banding comparison of the karyotypes in A. minous with those of the other species in the cahirinus group (i.e. Acomys cahirinus, Acomys cilicicus, and Acomys nesiotes) proves their very close phylogenetic relationship, further reinforced by the study of the cytochrome b sequences, and that A. minous possesses the ancestral karyotype of the group. It is suggested that at least two of the karyotypes that characterize A. minous today, pre‐existed in North Africa before it colonized Crete and that the specific status of the four members in the cahirinus group may need to be revisited. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 498–509.     

Karyotype differentiation in Chromaphyosemion killifishes (Cyprinodontiformes, Nothobranchiidae): patterns, mechanisms, and evolutionary implications

Chromaphyosemion killifishes are a karyotypically highly diverse group of small, sexually dimorphic fishes living in rainforest rivulets in tropical West and Central Africa. In the present study, we used various chromosome banding and staining techniques to analyse the karyotypes of 13 populations representing seven described species ( Chromaphyosemion loennbergii , Chromaphyosemion punctulatum , Chromaphyosemion splendopleure , Chromaphyosemion volcanum , Chromaphyosemion malumbresi , Chromaphyosemion melanogaster , Chromaphyosemion bitaeniatum ) and two undescribed forms ( Chromaphyosemion cf. lugens , Chromaphyosemion sp. Rio Muni GEMHS00/41). Diploid chromosome numbers (2 n ) and the number of chromosome arms (NF) ranged from 2 n  = 24 in C. malumbresi to 2 n  = 40 in C. bitaeniatum and from NF = 40 in C. volcanum and C. cf. lugens to NF = 54 in one population of C. loennbergii . A tentative XX/XY sex chromosome system was revealed in C. loennbergii , C. melanogaster , C. malumbresi , and Chromaphyosemion sp. Rio Muni GEMHS00/41. Mapping cytogenetic data for all described Chromaphyosemion species onto a recently published mitochondrial DNA phylogeny revealed a complex pattern of chromosomal evolution with several independent reductions of 2 n and independent modifications of NF and nucleolus organizer region phenotypes. Together with the results of preliminary crossing and mate choice experiments, the cytogenetic and molecular phylogenetic data suggest that, contrary to previous hypotheses, chromosomal rearrangements are probably not the most important and certainly not the only factor driving speciation in Chromaphyosemion killifishes.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 143–153.     

Chromosomal variability in the genus Primula (Primulaceae)

Chromosome numbers and measurements were recorded in 47 individuals of ten taxa of Primula , representing eight species and two subspecies from ten populations. The basic numbers of chromosomes were x  = 8, 9 or 11, and they were mostly metacentric, medium-long to medium-small, ranging in length from c . 3.6 µm to 1.6 µm. Diploid chromosome numbers of 2 n  = 2 x  = 24 and 22 were scored for the first time in P. loeseneri Kitag. and P. prenantha Balf. & Sm., respectively. A new ploidy level of 2 n  = 4 x  = 44 was found in P. burmanica Balf. & King. The recorded diploid numbers of the remaining species confirmed earlier reports. Interspecific variability in chromosome numbers was correlated with heterogeneity in their mean length. Comparison of the data with those in the literature revealed that the observed variability of chromosomal characters was compatible with the other taxonomic criteria and supports the current taxonomic delimitation. Chromosomal variation at the diploid level is the predominant feature in Primula evolution, and allopatric speciation has played a major role in its specific diversity. Subgen. Aleuritia could represent the main clade from which the other subgenera have evolved. The origin of the chromosome numbers, geographical distribution and evolution of the species were assessed, and the parallel polyphyletic mode of evolution in the genus was confirmed.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 211–219.     

The indigenous honey bees of Saudi Arabia (Hymenoptera, Apidae, Apis mellifera jemenitica Ruttner): Their natural history and role in beekeeping

Apis mellifera jemenitica Ruttner (= yemenitica auctorum: videEngel 1999) has been used in apiculture throughout the Arabian Peninsula since at least 2000 BC. Existing literature demonstrates that these populations are well adapted for the harsh extremes of the region. Populations of Apis mellifera jemenitica native to Saudi Arabia are far more heat tolerant than the standard races often imported from Europe. Central Saudi Arabia has the highest summer temperatures for the Arabian Peninsula, and it is in this region where only Apis mellifera jemenitica survives, while other subspecies fail to persist. The indigenous race of Saudi Arabia differs from other subspecies in the region in some morphological, biological, and behavioral characteristics. Further taxonomic investigation, as well as molecular studies, is needed in order to confirm whether the Saudi indigenous bee populations represent a race distinct from Apis mellifera jemenitica, or merely an ecotype of this subspecies.     

Revision of the Micromeria (Labiatae) in tropical to southern Africa and on the Arabian Peninsula

The material of Micromeria s.s. ( M. sect. Micromeria ) from tropical to southern Africa and the Arabian Peninsula is revised. The endemic Ethiopian M. unguentaria Schweinf. is retained as a species, while the other African and Arabian material of Micromeria is included in M. imbricata (Forssk.) C. Chr. M. imbricata is divided into the following three varieties: var. imbricata , which is very polymorphic, widespread in the African highlands, and also occurs in Arabia; var. rhodesiaca (Elly Walther & K.H. Walther) Ryding, which occurs in the highlands of Malawi, Mozambique and Zimbabwe; and var. villosa (Elly Walther & K.H. Walther) Ryding, which occurs in montane to alpine areas of Ethiopia and East Africa. Discontinuities in the variation within var. imbricata , observed at some localities, are suggested to be a result of partial autogamy. Problems with delimitation between M. imbricata , the East Asiatic M. biflora (Buch.-Ham. ex D.Don) Benth. and some Mediterranean species are discussed.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 427–446.     

Heterosis in Cepaea

A test has been made of the association of heterozygosity with shell breadth in the polymorphic snail Cepaea nemoralis. The material was collected by C. B. Goodhart from a series of paired sites at which individuals reached different adult breadths. Dominant phenotypes, in which a large fraction was heterozygous, had a greater breadth and lower variance than recessive phenotypes regardless of whether the measurement was of shell ground colour, banding or the double recessive vs. the rest. Most of the difference was contributed by samples from the habitat where animals reached the largest size. The result is consistent with existence of heterotic sections of chromosome that include the colour and banding loci, and may help to explain the persistence of the polymorphism.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 49–53.     

Cytogenetic studies of four South American species of Paullinia L. (Sapindaceae)

Four South American species of Paullinia ( P. elegans , P. meliaefolia , P. pinnata , and P. rhomboidea ) were compared using conventional chromosome staining, C-Giemsa and C-chromomycin A₃/4',6-diamidino-2-phenylindole (C-CMA₃/DAPI) banding, and fluorescence in situ hybridization (FISH) with a 45S ribosomal DNA (rDNA) probe. All species showed a somatic complement of 2 n  = 24 chromosomes, agreeing with earlier records in some cases, and showing a tendency for the chromosome number to be conserved in this genus. The chromosome number of P. rhomboidea is a new report. The karyotypes differed in chromosome size and degree of karyotype asymmetry. The chromosomal band patterns and location of the 45S rDNA sites are reported for the first time in the genus. Terminal C-CMA₃ bands were associated with the 45S rDNA sites, but varied in number and size between the species. The occurrence of other C-Giemsa bands that were not revealed by CMA₃ suggests that more than one family of repetitive DNA may be involved in karyotype differentiation. The systematic implications of these results on the infrageneric relationships are discussed.   © 2007 The Linnean Society of London . Botanical Journal of the Linnean Society , 2007, 154 , 313–320.     

On the genetic diversity of spiny mice (genus Acomys) and gerbils (genus Gerbillus) in the Arabian Peninsula

Using non-destructive sampling we provide further genetic characterisations for spiny mice (Acomys dimidiatus/cahirinus) and gerbils (Gerbillus sp.) in three regions in Saudi Arabia. All individuals were sequenced for a fragment of the cytochrome b gene, and compared against available conspecifics and closely related taxa. We confirm the existence of a second Acomys dimidiatus/cahirinus lineage specific to the Arabian Peninsula as seen previously. The Arabian Gerbillus nanus is shown to group with Middle Eastern rather than African conspecifics. A second cryptic Gerbillus lineage was also sampled across multiple locations, which may be an uncharacterised G. dasyurus.     

New perspectives on the mechanisms of chromosome evolution in parasitic flowering plants

In an attempt to find any association between chromosomal characters and parasitism, a cytological study of parasitic flowering plants in Israel has been carried out. While no such association was found, evidence for three levels of chromosome evolution could be discerned: intra-chromosomal modifications, polyploidy per se , and genome restructuring in polyploids. Our conclusions may serve as a paradigm of the multiple pathways of chromosomal evolution in plants in general. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 138 , 117–122.     

Contribution to the karyological knowledge of Echinops (Asteraceae, Cardueae) and related genera

Twenty-six chromosome counts were made of several genera of the tribe Cardueae from various European and Asian provenances: Acantholepis (one species studied), Amphoricarpus (1), Chardinia (1), Echinops (14 species, 15 populations), Siebera (1), Staehelina (3) and Xeranthemum (4). Eleven of the reports are made for the first time, ten confirm previous counts, while the remainder report disparities with earlier records. The existence of different basic chromosome numbers and ploidy levels suggests dysploidy and polyploidy as the main mechanisms of chromosomal evolution in the taxa considered.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 337−344.     

Systematics and phylogeny of the cane rats (Rodentia: Thryonomyidae)

The family Thryonomyidae is known from the Eocene up to the present. Today, this group comprises just two closely related species, which are restricted to sub-Saharan Africa. However, various thryonomyids have been recorded in strata of Miocene age, when the group spread out of Africa eastward to southern Asia (Pakistan). A systematic revision and a cladistic analysis shows that 20 species can be referred to this family: Thryonomys swinderianus (Temninck), T. gregorianus (Thomas), Paraphiomys pigotti Andrews, P. occidentalis Lavocat, P. simonsi Wood, P. hopwoodi Lavocat, P. shipmani Denys et Jaeger, P. australis Mein, Pickford et Senut, P. roessneri Mein, Pickford et Senut, P. afarensis Geraads, Paraphiomys sp. nov. from Saudi Arabia López-Antoñanzas et Sen , P. renelavocati sp. nov. , Neosciuromys africanus Stromer, Apodecter stromeri Hopwood, Paraulacodus indicus Hinton, Paraulacodus johanesi Jaeger, Michaux et Sabatier, Gaudeamus aegyptius Wood, Epiphiomys coryndoni Lavocat, Kochalia geespei (de Bruijn et Hussain), Paraphiomys sp. nov. from Saudi Arabia, and Paraphiomys orangeus Mein et Pickford. The unresolved basal position of Sacaresia moyaeponsi with respect to Metaphiomys schaubi and the clade comprising the above-cited species, suggest that this taxon should not be allocated to the family Thryonomyidae. A phylogenetic definition of the family Thryonomyidae is proposed as an outcome of the phylogenetic analysis: Epiphiomys corindoni , Thryonomys swinderianus , their most recent common ancestor and all its descendants (node-based taxon).  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 142 , 423–444.     

Genetic diversity among different species of the genus Leiurus (Scorpiones: Buthidae) in Saudi Arabia and the Middle East

The molecular phylogenetic relationship among two species of genus Leiurus, from Saudi Arabia with additional comparative sequence data available from Egypt, Oman and Turkey is presented. The molecular phylogeny was performed using maximum parsimony, neighbor joining and bayesian inference. Our results indicate a clear deep splitting between the Western clade, which represented by L. quinuestriatus sequences from Egypt and those from the Eastern clade which encompassing different Leiurus species from Saudi Arabia, Oman and Turkey was shown. Also, the phylogenetic relationship represents additional support for the taxonomic status of Arabian Leiurus species.     

Phylogeography of spiny mice (genus Acomys, Rodentia: Muridae) from the south-western margin of the Sahara with taxonomic implications

The present study aims to depict the overall pattern of Acomys history in south-western Sahara. We tested the specific predictions that: (1) several mitochondrial clades can be identified and that they coincide with the described species; (2) successive phases of desert expansion and contraction during the last 3 Myr have resulted in several phases of demographic expansion and population fragmentation in Acomys ; and (3) isolation-by-distance occurs. The extent of phylogeographic patterns and molecular genetic diversity (cytochrome b gene and D-loop) were addressed in a survey of 90 individuals of Acomys from 38 localities. Our phylogeographical analyses show a strong genetic structure within western Saharan Acomys , with several phylogroups displaying non-overlapping geographic distributions. Restricted gene flow with isolation-by-distance was recorded and a signal of population expansion was detected within several clades. We suggest that during arid or semi-arid paleoclimatic periods, when large sandy areas were present, Acomys was restricted to rocky massifs, whereas, in more humid periods, when savannah and/or steppe habitat prevailed, this species was able to disperse and to survive outside rocky areas because food resources were abundant. Based on a comprehensive sampling and the use of an integrative approach (i.e. combining cytogenetic, molecular and morphological data), we firmly propose that Acomys airensis should be considered as a junior synonym of Acomys chudeaui .  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 29–46.     

Morphological variation of the Oncocyclus irises (Iris: Iridaceae) in the southern Levant

Morphological traits of Iris section Oncocyclus (Siems.) Baker in the southern Levant (Israel, Jordan, The Palestinian Authority and Sinai/Egypt) were analysed in order to clarify taxonomic relationships among taxa and the validity of diagnostic characters. Floral and vegetative characters were measured in 42 populations belonging to nine species during the peak of the flowering season in 1998–2000. Pearson's Coefficient of Racial Likelihood (CRL) was used to calculate morphological distances between populations. Twelve of the measured populations, distributed along the north-south aridity gradient in Israel, were further explored for morphological changes along the gradient. Cluster analysis revealed two major clusters: the first includes most of the dark-coloured Iris populations, with populations of I. petrana Dinsmore and I. mariae W. Barbey forming a subcluster; the second consists of all the light-coloured populations but also some dark-coloured populations. Pearson's CRL and geographical distance were significantly correlated among the dark-coloured populations. Along the geographical gradient, flower, stem and leaf size traits decrease towards the south, probably as an adaptation to aridity. This suggests that natural selection promoted the differences between populations. Almost no discrete phenotypic groups exist within the Oncocyclus species of the southern Levant except for variation in the floral colours. Most of the suggested diagnostic characters proved unreliable in that they varied continuously across populations. The taxonomical difficulties encountered in this study reflect the special evolutionary state of the Oncocyclus irises as a group in the course of speciation.  © 2002 The Linnean Society of London. Botanical Journal of the Linnean Society , 2002, 139 , 369–382.     

Chromosome numbers in Brazilian species of Crotalaria (Leguminosae, Papilionoideae) and their taxonomic significance

Chromosome numbers were counted for 23 species of Crotalaria native to Brazil. Among these data there were new counts for 15 taxa, and some confirmed previous reports or represented numbers that were different from those cited previously. The chromosome numbers most frequently found were 2 n  = 16 and 2 n  = 32. Only C. incana L. had 2 n  = 14 and C. tweediana Benth. had 2 n  = 54. The counts 2 n  = 32 and 54 were found in species of section Calycinae and 2 n  = 16 and 14 in species of section Chrysocalycinae . The data revealed the importance of chromosomal parameters in the characterization of sections Calycinae and Chrysocalycinae in Brazil. We discuss the systematic significance and evolutionary aspects for the genus, comparing the results with the two sections that are native in Brazil.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 271–277.