New polymorphic microsatellite loci for different camel species

New microsatellite loci were screened and sequenced from the genomic DNA of male Camelus bactrianus. Among 32 loci, 23 were amplified in bactrian and dromedary species, 19 in llama and 20 in alpaca. The different species had similar fragment lengths per locus, with more striking similarities between bactrian and dromedary and between llama and alpaca, respectively. Seven loci had more than 10 alleles each, nine were monomorphic in all species, and one was monomorphic in Old World and polymorphic in New World camels. The results show that the informative microsatellite loci can be widely applied to several species.     

High mitochondrial differentiation levels between wild and domestic Bactrian camels: a basis for rapid detection of maternal hybridization

Hybridization between wild species and their domestic congeners often threatens the gene pool of the wild species. The last wild Bactrian camel (Camelus ferus) populations in Mongolia and China are examples of populations facing such a hybridization threat. To address this key issue in the conservation of wild camels, we analysed wild, hybrid and domestic Bactrian camels (Camelus bactrianus) originating from Mongolia, China and Austria. Through screening of an 804‐base‐pair mitochondrial fragment, we identified eight mitochondrial haplotypes and found high sequence divergence (1.9%) between C. ferus and C. bactrianus. On the basis of a mitochondrial DNA sequence fixed difference, we developed a diagnostic PCR restriction fragment length polymorphism (PCR‐RFLP) assay to differentiate between wild and domestic camel samples. We applied the assay to 81 individuals and confirmed the origin of all samples including five hybrids with known maternal ancestry. The PCR‐RFLP system was effective for both traditional (blood, skin) and non‐invasive samples (faeces, hair), as well as for museum specimens. Our results demonstrate high levels of mitochondrial differentiation between wild and domestic Bactrian camels and that maternal hybridization can be detected by a rapid and reliable PCR‐RFLP system.     

Genetic diversity and phylogeographic structure of Bactrian camels shown by mitochondrial sequence variations

The Bactrian camel includes various domestic (Camelus bactrianus) and wild (Camelus ferus) breeds that are important for transportation and for their nutritional value. However, there is a lack of extensive information on their genetic diversity and phylogeographic structure. Here, we studied these parameters by examining an 809‐bp mtDNA fragment from 113 individuals, representing 11 domestic breeds, one wild breed and two hybrid individuals. We found 15 different haplotypes, and the phylogenetic analysis suggests that domestic and wild Bactrian camels have two distinct lineages. The analysis of molecular variance placed most of the genetic variance (90.14%, P < 0.01) between wild and domestic camel lineages, suggesting that domestic and wild Bactrian camel do not have the same maternal origin. The analysis of domestic Bactrian camels from different geographical locations found there was no significant genetic divergence in China, Russia and Mongolia. This suggests a strong gene flow due to wide movement of domestic Bactrian camels.     

Foraging ecologies of giraffe (Giraffa camelopardalis reticulata) and camels (Camelus dromedarius) in northern Kenya: effects of habitat structure and possibilities for competition?

The foraging ecologies of reticulated giraffe (Giraffa camelopardalis reticulata) and domestic camels (Camelus dromedarius) were examined in the Laikipia District of Kenya, where these species have recently become sympatric. Camels increased popularity in the region has lead to concerns about their environmental impacts and possible competition with wild giraffe for resources. We gathered foraging data on both species using 2‐min group scans that recorded feeding heights and plant food preferences. Transects sampled the vegetation in areas where foraging observations were recorded. Giraffe females feed at lower elevations than males, while female camels feed below both sexes of giraffe. There was very little observed overlap in food preferences between the species. However, habitat type has an effect on foraging ecologies of both giraffe sexes, but habitat did not influence camel foraging. Camel herder husbandry techniques also influence camel foraging dynamics. These findings have important implications in achieving the twin objectives of wildlife conservation and pastoralist livestock production in northern Kenya.     

Repeated trans-Atlantic dispersal catalysed a global songbird radiation

Aim Turdus thrushes are one of the most speciose and widespread songbird genera, comprising nearly 70 species that combined have a near‐global distribution. Herein, we use molecular phylogenetic, molecular clock and behavioural evidence to examine the historical biogeography of the genus. Ancestral area reconstructions in conjunction with divergence estimates and palaeoclimatogical data are used to test whether the long‐standing paradigm of Beringian colonization or trans‐Atlantic dispersal best explains modern distributions in the New and Old Worlds. Location Worldwide, with emphasis on New World–Old World biotic interchange. Methods Using a molecular phylogenetic hypothesis of Turdus thrushes, we reconstructed ancestral area relationships utilizing the five major continental or regional areas occupied by species in the genus. We also examined the evolution of behaviours on the phylogeny, and estimated the timing of major lineage divergences via a molecular clock. Results Turdus originated in Eurasia, and following the colonization of Africa underwent a series of five trans‐Atlantic sweepstake dispersals. The data reject the alternative hypothesis that connections between Old and New World Turdus species can be attributed to movement through Beringia with subsequent extinction. Divergence estimates indicate that these dispersals all occurred near the Miocene–Pliocene boundary, 5 Ma. A significant phylogenetic correlation between migratory and flocking behaviour is evident in the genus. Main conclusions The initial divergence of Turdus in the Old World was followed by a series of trans‐Atlantic sweepstake dispersal events. These dispersals are temporally correlated with a specific palaeoclimatic system, which would have facilitated transport of Turdus from the Caribbean to the Old World across the Atlantic. Uplift of the Central American Seaway 4.7 Ma effectively shut down the palaeoclimatic system, and no additional trans‐Atlantic dispersals are evident in Turdus after this time. Migratory movements by ancestral lineages in flocks, rather than as single individuals, suggest an increased likelihood of successfully colonizing new areas, post‐dispersal.     

New Pliocene remains of Camelus grattardi (Mammalia,Camelidae) from the Shungura Formation,Lower Omo Valley,Ethiopia, and the evolution of African camels

Abstract

Camels are exceptionally rare in the Plio-Pleistocene fossil record of Africa, hindering attempts to understand the evolution of this family on the continent. Here we describe recently collected camel specimens from the Shungura Formation, Lower Omo Valley, Ethiopia, and attribute these remains to Camelus grattardi. The new specimens date to the late Pliocene (~3 to 2.6 Ma) and consist of three upper molars, one upper premolar, and two proximal metatarsals. The dental specimens confirm this species’ small P4 relative to its molars, a trait that differs significantly from all extant and fossil Old World camels. The metatarsals indicate that C. grattardi was similar in size to the living Bactrian camel, C. bactrianus. Phylogenetically, we find no suitable ancestor, sister, or descendant of the eastern African fossil camel, which suggests greater lineage diversity in Plio-Pleistocene Camelus than previously recognised. Microwear analyses suggest that C. grattardi was likely a mixed-feeder preferring browse, which is consistent with carbon isotopes of enamel from the Turkana Basin. A review of the fossil record of African camels suggests no clear paleoenvironmental association, as fossil camels occur in a range of environments from dry savannas with no permanent water bodies to closed woodlands along the paleo-Omo River.     

Identification and management of a single large population of wild dromedary camels

The dromedary camel (Camelus dromedarius) is a significant invasive species in Australia. It is an unusual pest species that is of large body size with relatively low fecundity compared with other pest species. Camels are highly adapted to the arid regions that characterize a large proportion of Australia and occupy an almost completely undisturbed area of ≥3 million km². They have no history of invasion elsewhere in the world. Despite this, their population has been expanding at approximately 80,000 camels per annum, with the most recent estimate of population size around 1,000,000 individuals. We employed a landscape-genetic approach to evaluate the population structure and molecular ecology of Australian camels. We combined mitochondrial control region sequence (n = 209 animals) with 18 microsatellite markers to profile over 800 adult camels to identify the presence of a single panmictic population. We showed that demographically defined neighborhoods for wild camels are about 200 km; this value was supported by home range estimates. Distances greater than this display no pattern of isolation by distance across the Australian continent. The result is the largest single geographical population so far recorded for an invasive species in Australia. This pattern may be explained by the impressive and near-nomadic dispersal pattern of camels, in combination with an unpredictable environment virtually devoid of barriers to movement and predatory suppression. Although it is technically feasible, the reality is that it would not be economically or politically viable to have continental eradication of wild camels in Australia because of the vast size and movement dynamics of the camel population. As such, we advocate a change away from an expensive solution to an intractable reduction program (that is almost entirely focused on protection of biological refugia) and moves to include cultural, economic, and biodiversity asset protection for the management of this most unorthodox of invasive species. © 2012 The Wildlife Society.     

Many ways to make a novel structure: a new mode of sex comb development in Drosophilidae

On macroevolutionary time scales, the same genes can regulate the development of homologous structures through strikingly different cellular processes. Comparing the development of similar morphological traits in closely related species may help elucidate the evolutionary dissociation between pattern formation and morphogenesis. We address this question by focusing on the interspecific differences in sex comb development in Drosophilids. The sex comb is a recently evolved, male‐specific structure composed of modified bristles. Previous work in the obscura and melanogaster species groups (Old World Sophophora) has identified two distinct cellular mechanisms that give rise to nearly identical adult morphologies. Here, we describe sex comb development in a species from a more distantly related lineage, the genus Lordiphosa. Although the expression of key regulatory genes is largely conserved in both clades, the cell behaviors responsible for sex comb formation show major differences between Old World Sophophora and Lordiphosa. We suggest that the many‐to‐one mapping between development and adult phenotype increases the potential for evolutionary innovations.     

Sorting out the Snakebirds: The species status,phylogeny, and biogeography of the Darters (Aves: Anhingidae)

Although the avian family Anhingidae is unequivocally monophyletic, the number and relationships of the component species within the single genus (Anhinga) have long remained unclear. Here, we use extensive mitochondrial and nuclear DNA sequence data (8,878 bp) to show that four species should be recognized. Our fully resolved and well‐supported tree shows that the American Anhinga (Anhinga anhinga) is sister to the three Old World species, with the Oriental (A. melanogaster) and African (A. rufa) Darters sister within the Old World clade, which also includes the Australian Darter (A. novaehollandiae). We estimate that the divergence between the New World and Old World branches occurred 19–22 mya, with the Australian Darter separating from its Old World congeners 14–16 mya and the Oriental and African species splitting ~10 mya. The genus is yet another example of osteological conservatism in the Suliformes, which is comparable to that shown by the cormorants and shags. Nevertheless, the relationships we infer are congruent with recent plumage studies and are biogeographically plausible. We suggest that further investigation of the variation within the African and Australian Darters would be of interest.     

Dynamic colonization exchanges between continents and islands drive diversification in paradise‐flycatchers (Terpsiphone,Monarchidae)

Aim We use parametric biogeographical reconstruction based on an extensive DNA sequence dataset to characterize the spatio‐temporal pattern of colonization of the Old World monarch flycatchers (Monarchidae). We then use this framework to examine the role of dispersal and colonization in their evolutionary diversification and to compare plumages between island and continental Terpsiphone species. Location Africa, Asia and the Indian Ocean. Methods We generate a DNA sequence dataset of 2300 bp comprising one nuclear and three mitochondrial markers for 89% (17/19) of the Old World Monarchidae species and 70% of the Terpsiphone subspecies. By applying maximum likelihood and Bayesian phylogenetic methods and implementing a Bayesian molecular clock to provide a temporal framework, we reveal the evolutionary history of the group. Furthermore, we employ both Lagrange and Bayes‐ Lagrange analyses to assess ancestral areas at each node of the phylogeny. By combining the ancestral area reconstruction with information on plumage traits we are able to compare patterns of plumage evolution on islands and continents. Results We provide the first comprehensive molecular phylogenetic reconstruction for the Old World Monarchidae. Our phylogenetic results reveal a relatively recent diversification associated with several dispersal events within this group. Moreover, ancestral area analyses reveal an Asian origin of the Indian Ocean and African clades. Ancestral state reconstruction analyses of plumage characters provide an interpretation of the plumage differentiation on islands and continents. Ancestral plumage traits are inferred to be close to those of the Asian paradise‐flycatcher (Terpsiphone paradisi), and island species display a high degree of plumage autapomorphy compared with continental species. Main conclusions Terpsiphone paradisi is polyphyletic and comprises populations that have retained the ancestral plumage of the widespread Terpsiphone genus. The genus appears to have colonized south‐west Asia, the Indian Ocean and Africa from eastern Asia. The phylogeny and divergence time estimates indicate multiple simultaneous colonizations of the western Old World by Terpsiphone. These results reinforce a hypothesis of range expansions of a Terpsiphone paradisi‐like ancestor into eastern Asia and the western Old World.     

Coccidia in Camels (Camelus dromedarius) in Saudi Arabia1

Saudi Arabian camels {Camelus dromedarius) are infected with three species of Eimeria: E. dromedarii (28.4%), E. rajasthani < 22.2%), and E. cameli (19.2%); 41.6% of the animals examined were positive. The highest prevalence of infection was reported in the western region of the country. Mixed infection with two Eimeria species is most common; E. dromedarii was most frequently and generally the most predominant species. Eimeria dromedarii and E. rajasthani are described for the first time from Saudi Arabian camels.     

Phylogeny,host association and biogeographical patterns in the diverse millipede‐parasitoid genus Myriophora Brown (Diptera: Phoridae)

Myriophora is the most species‐rich group of parasitoids that attack toxic, chemically defended millipedes in the superorder Juliformia and order Polydesmida—a resource that few insect predators and parasitoids are able to exploit. Worldwide, there are an estimated 200 species of Myriophora, with the majority of the diversity centred in the Neotropical region. The phylogeny of Myriophora is unknown, biogeographical patterns are not documented, and known host associations have not been assessed in a phylogenetic context. We provide the first phylogenetic study of the genus from a data set composed of 52 taxa primarily from the Neotropical region including 10 outgroups, 40 morphological characters, and molecular data from three mitochondrial (16S, COI and ND1) and one nuclear marker (AK). We find that Myriophora dispersed from the New World to the Old World in a single event before subsequently spreading to the Afrotropical region. The ancestral hosts reconstructed for Myriophora are the benzoquinone‐producing Juliformia, and this association has been retained in the Old World clade. In the Neotropical region, Myriophora that are associated with cyanide‐producing polydesmidan millipedes are confined to a single clade that shows remarkably little genetic variation between clearly morphologically diagnosable species.     

Ticks (Acari: Ixodidae) infesting the Arabian Camel (Camelus dromedarius) in the Sinai,Egypt with a note on the acaricidal efficacy of Ivermectin

In this study, tick burdens on camels (Camelus dromedarius) were determined in the vicinity of the St. Catherine monastery, Sinai, Egypt. In total 2,545 ticks (1,491 adults and 1,054 nymphs) were collected and identified. Mean tick burdens were relatively heavy and the range in number of ticks per camel was very broad (6–173). Hyalomma dromedarii was the predominant tick species and accounted for 95.6% of the adult ticks. Other ticks found were H. marginatum subspp. and H. anatolicum excavatum. All nymphs collected were Hyalomma spp.In addition, the effect of ivermectin (Ivomec MSD AGVET) on tick burdens, when given subcutaneously at 0.2 mg/kg¹, was evaluated in nine camels in the date-palm plantation of Kibbutz Yahel in the Arava valley, Israel. Initial tick burdens on these camels (half-body tick counts) ranged from 20–105 ticks per camel. Seven camels from the same herd with half-body tick counts ranging from 40–107 ticks per camel were not treated and served as a control group. Ivermectin was not effective against Hyalomma tick infestation in camels under these conditions.     

Tropical Asian species show that the Old World clade of ‘spiny solanums’ (Solanum subgenus Leptostemonum pro parte: Solanaceae) is not monophyletic

The tropical Asian taxa of the species‐rich genus Solanum (Solanaceae) have been less well studied than their highly diverse New World relatives. Most of these tropical Asian species, including the cultivated brinjal eggplant/aubergine and its wild progenitor, are part of the largest monophyletic Solanum lineage, the ‘spiny solanums’ (subgenus Leptostemonum or the Leptostemonum clade). Here we present the first phylogenetic analysis of spiny solanums that includes broad sampling of the tropical Asian species, with 42 of the 56 currently recognized species represented. Two nuclear and three plastid regions [internal transcribed spacer (ITS), waxy, ndhF‐rpL32, trnS‐trnG and trnT‐trnF] were amplified and used to reconstruct phylogenetic relationships using maximum likelihood and Bayesian methods. Our analyses show that Old World spiny solanums do not resolve in a single clade, but are part of three unrelated lineages, suggesting at least three independent introductions from the New World. We identify and describe several monophyletic groups in Old World solanums that have not been previously recognized. Some of these lineages are coherent in terms of morphology and geography, whereas others show considerable morphological variation and enigmatic distribution patterns. Tropical Asia occupies a key position in the biogeography of Old World spiny solanums, with tropical Asian taxa resolved as the closest relatives of diverse groups of species from Australia and Africa.     

Ecological niche comparison and molecular phylogeny segregate the invasive moss species Campylopus introflexus (Leucobryaceae,Bryophyta) from its closest relatives

The delimitation of the invasive moss species Campylopus introflexus from its closest relative, Campylopus pilifer, has been long debated based on morphology. Previous molecular phylogenetic reconstructions based on the nuclear ribosomal internal transcribed spacers (ITS) 1 and 2 showed that C. pilifer is split into an Old World and a New World lineage, but remained partly inconclusive concerning the relationships between these two clades and C. introflexus. Analyses of an extended ITS dataset displayed statistically supported incongruence between ITS1 and ITS2. ITS1 separates the New World clade of C. pilifer from a clade comprising C. introflexus and the Old World C. pilifer. Ancestral state reconstruction showed that this topology is morphologically supported by differences in the height of the dorsal costal lamellae in leaf cross‐section (despite some overlap). ITS2, in contrast, supports the current morphological species concept, i.e., separating C. introflexus from C. pilifer, which is morphologically supported by the orientation of the hyaline hair point at leaf apex as well as costal lamellae height. Re‐analysis of published and newly generated plastid atpB‐rbcL spacer sequences supported the three ITS lineages. Ecological niche modeling proved a useful approach and showed that all three molecular lineages occupy distinct environmental spaces that are similar, but undoubtedly not equivalent. In line with the ITS1 topology, the C. pilifer lineage from the New World occupies the most distinct environmental niche, whereas the niches of Old World C. pilifer and C. introflexus are very similar. Taking the inferences from ecological niche comparisons, phylogenetics, and morphology together, we conclude that all three molecular lineages represent different taxa that should be recognized as independent species, viz. C. introflexus, C. pilifer (Old World clade), and the reinstated C. lamellatus Mont. (New World clade).     

Molecular detection of Rickettsia aeschlimannii in Hyalomma spp. ticks from camels (Camelus dromedarius) in Nigeria,West Africa

Several species of the spotted fever group rickettsiae have been identified as emerging pathogens throughout the world, including in Africa. In this study, 197 Hyalomma ticks (Ixodida: Ixodidae) collected from 51 camels (Camelus dromedarius) in Kano, northern Nigeria, were screened by amplification and sequencing of the citrate synthase (gltA), outer membrane protein A (ompA) and 17‐kDa antigen gene fragments. Rickettsia sp. gltA fragments were detected in 43.3% (42/97) of the tick pools tested. Rickettsial ompA gene fragments (189 bp and 630 bp) were detected in 64.3% (n = 27) and 23.8% (n = 10) of the gltA‐positive tick pools by real‐time and conventional polymerase chain reaction (PCR), respectively. The amplicons were 99–100% identical to Rickettsia aeschlimannii TR/Orkun‐H and R. aeschlimannii strain EgyRickHimp‐El‐Arish in GenBank. Furthermore, 17‐kDa antigen gene fragments of 214 bp and 265 bp were detected in 59.5% (n = 25) and 38.1% (n = 16), respectively, of tick pools, and sequences were identical to one another and 99–100% identical to those of the R. aeschlimannii strain Ibadan A1 in GenBank. None of the Hyalomma impressum ticks collected were positive for Rickettsia sp. DNA. Rickettsia sp. gltA fragments (133 bp) were detected in 18.8% of camel blood samples, but all samples were negative for the other genes targeted. This is the first report to describe the molecular detection of R. aeschlimannii in Hyalomma spp. ticks from camels in Nigeria.     

Allelic polymorphism in Arabian camel ribonuclease and the amino acid sequence of bactrian camel ribonuclease

Pancreatic ribonucleases from several species (whitetail deer, roe deer, guinea pig, and arabian camel) exhibit more than one amino acid at particular positions in their amino acid sequences. Since these enzymes were isolated from pooled pancreas, the origin of this heterogeneity is not clear. The pancreatic ribonucleases from 11 individual arabian camels (Camelus dromedarius) have been investigated with respect to the lysine-glutamine heterogeneity at position 103 (Welling et al., 1975). Six ribonucleases showed only one basic band and five showed two bands after polyacrylamide gel electrophoresis, suggesting a gene frequency of about 0.75 for the Lys gene and about 0.25 for the Gln gene. The amino acid sequence of bactrian camel (Camelus bactrianus) ribonuclease isolated from individual pancreatic tissue was determined and compared with that of arabian camel ribonuclease. The only difference was observed at position 103. In the ribonucleases from two unrelated bactrian camels, only glutamine was observed at that position.Part of this work has been carried out under the auspices of the Netherlands Foundation for Chemical Research (S.O.N.) and with financial aid from the Netherlands Organisation for the Advancement of Pure Research (Z.W.O.).     

啄木鸟科内谱系生物地理学新探

啄木鸟群体共包含254种,但早期对啄木鸟的分类学研究局限于形态学、行为学和鸣唱法等方向,分子研究结果又彼此存在差异,因此啄木鸟分类系统目前还有争议。分子数据的大量累积为啄木鸟系统发育的研究提供了条件。利用从GenBank获取的ND2基因序列,对啄木鸟科179种啄木鸟采用NJ法、MP法以及ML法分别进行了分子进化树构建,同时用NJ法重建了Picumnus属分子进化树。此外又利用cytb基因序列采用NJ法对Dryobates属和Picoides属进行分子进化树重建。我们的结果显示,姬啄木亚科Picumnus属新大陆物种彼此的进化关系仍需进一步研究,而该属唯一一种旧大陆种P.innominatus,种内遗传差异在种间水平依然显著,这说明其种内遗传差异一直被低估了。在啄木鸟亚科内,Hemicircus属和Nesoctites属为姐妹关系,而Geocolaptes olivaceus插入了Campethera属支系中,并非与其构成姐妹群体。此外,一些姐妹群在新旧大陆上的分布,推测这种分布可能与第四纪冰川有关。啄木鸟从旧大陆向新大陆迁移,在第四纪冰川作用下不断发生分布范围的扩张与收缩,最终形成现存的分布格局。     

A Phylogeny‐Based Comparison of Tarantula Spider Anti‐Predator Behavior Reveals Correlation of Morphology and Behavior

We studied anti‐predator strategies in nine species of tarantulas from Asia, Africa, and the Americas. Tarantulas in the New World possess urticating hairs which induce inflammation when in contact with vertebrate mucus membranes and skin. In contrast, tarantulas from the Old World lack this defense but are observed to exhibit a much greater willingness to escalate to an active defense when provoked. We had three goals: (1) describe the behaviors exhibited by each taxon in response to two levels of provocation, (2) look for the presence of alternative classes of anti‐predator strategy as predicted by the Old World–New World dichotomy in aggressive defense, and (3) examine the evolution of these behaviors in the context of the phylogeny of the group. We compared the response of nine different mygalomorph spider genera to two levels of aversive stimuli: puffs of air and prodding. We found that the overall structure of the defensive behavior was similar between the different taxa, consisting of fleeing, rearing, striking, and biting. Some genera did exhibit unique behaviors such as stridulating (Hysterocrates) or rocking (Haplopelma and Selenocosmia). We found that the genera from the New World exhibited low levels of escalation in their defense behaviors, while those from the Old World readily escalated to striking and biting. Our findings are consistent with the hypothesis that the possession of urticating hairs is associated with very low levels of active defense behaviors such as striking and biting. The phylogenetic analysis indicates that both the notable levels of aggression displayed by the African taxa tested and the relative passivity of the New World tarantulas each represents a synapomorphy.