Testing Bergmann's rule in the presence of potentially confounding factors: a case study with three species of Galerida larks in Morocco

Aim  To test Bergmann's rule (which predicts a larger body size in colder areas within warm-blooded vertebrate species) in three partially sympatric species of larks ( Galerida theklae , Galerida cristata and Galerida randonii ) that occur in Morocco.
Location  Morocco.
Methods  Restriction fragment length polymorphism techniques applied on cytochrome b haplotypes were used to discriminate G. cristata and G. randonii , and to investigate the effects of interspecific hybridization in their contact zone. A comprehensive statistical framework was then designed to test Bergmann's rule in our three Galerida species (using altitude as a proxy for cold temperatures), while controlling for the possible influence of interspecific hybridization and competition and accounting for spatial autocorrelation. The method we propose is conservative in the sense that potentially confounding factors are adjusted so as to maximize their influence on the variable of interest.
Results  Bergmann's rule was strongly supported in G. theklae and G. randonii . However, body size did not respond to altitude in G. cristata , a result that was not simply explained by species-specific differences in geographical ranges and altitudinal span. In G. cristata , we detected a tendency for body size to increase with aridity, in agreement with an alternative definition of Bergmann's rule. However, since G. cristata also hybridizes with G. randonii in a contact zone located in the most arid part of the range of G. cristata , we could not tease apart the relative contribution of selection and hybridization in driving this pattern.
Main conclusions  This study highlights the need for careful statistical designs that allow meaningful variables to be picked out from large sets of potential factors. When taking these factors into account, we found that Bergmann's rule was still strongly supported in two out of the three species examined.     

Phenotypic variation in Galerida larks in Morocco: the role of history and natural selection

The relative influences of history, natural selection and hybridization in shaping phenotypic variation in closely related taxa is a crucial issue in current evolutionary biology. In this study, we used as a model two sibling but paradoxically highly variable species of larks (Galerida theklae and Galerida cristata) of Morocco to separate the impacts of these evolutionary forces. In the former species, variation is manifested mainly in colouration, while in the latter, variation also encompasses bill size and shape. Mitochondrial and nuclear DNA sequencing were used to identify the historical relationships among the subspecies and species. According to our analyses, G. cristata and G. theklae diverged about 3.7 million years ago (Ma), and we found no evidence for a role of hybridization in maintaining their similarity. In G. theklae, there was no further subdivision, while in G. cristata two major mtDNA groups were identified (divergence approximately 1.1 Ma). These two lineages are parapatric and regroup, respectively, the three short-billed subspecies [G. (cristata) cristata] and the two long-billed subspecies [G. (cristata) randonii]. Patterns of morphological variation were then contrasted to this pattern of neutral relationships: we found that G. (c.) cristata was morphologically more similar to G. theklae than to G. (c.) randonii. Overall, these results point towards the prominent role of (i) natural selection and/or phenotypic plasticity in adapting the plumage to local conditions and (ii) natural selection in combination with historical isolation in driving the divergence in size and bill morphology in the crested larks.     

Global warming and body mass decline in Israeli passerine birds

Global warming may affect the physiology, distributions, phenology and adaptations of plants and animals. In Israel, minimum summer temperatures increased by an average of 0.26 degrees C per decade during the second half of the 20th century. Bergmann's rule predicts that, in warm-blooded animals, races from warm regions are smaller than races from cold regions. Numerous studies have reported general correlations between body mass in fossil animals and independently established palaeoclimatic changes from various parts of the world in accordance with this rule. Using museum specimens, I tested the prediction that the body mass and tarsus length of five resident passerine species in Israel declined between 1950 and 1999. The body mass of four species (the graceful warbler Prinia gracilis, the house sparrow Passer domesticus, the yellow-vented bulbul Pycnonotus xanthopygos and the Sardinian warbler Sylvia melanocephala, but not of the crested lark Galerida cristata) declined significantly during this period. Tarsus length also declined significantly during this period for two species (the graceful warbler and the house sparrow). Body condition (body mass-to-tarsus length ratio) decreased in the Sardinian warbler, the yellow-vented bulbul and the crested lark. It is suggested that the above declines in body mass and tarsus length are due to global warming and also in accordance with Bergmann's rule. The above explanation does not exclude the possibility that other factors, such as a decrease in food availability, contributed to the decline in body mass. These declines may have serious implications for community structure and competition among bird species and may affect the survival of small passerines.     

Nest predation and nestling growth rate of two lark species in the Negev Desert, Israel

Clutch-size, nestling growth and predation rates on eggs and nestlings in two sympatric alaudids, the Crested Lark Galerida cristata and the Desert Lark Ammomanes deserti, were studied in the Negev desert of Israel. Logistic growth constants were similar for both species, and experimental brood reduction did not accelerate the growth rate of their nestlings. It is suggested that the predation rate on nests of both species has been sufficiently high to select for maximal growth rate of nestlings. Overall probability of predation was 0.85 on a Desert Lark nest and 0.76 on a Crested Lark nest. The smaller clutch-size of the Desert Lark (3.7) may be an adaptation to a higher predation risk, as compared to that of the Crested Lark (4.6) which is more vigilant and nests in more concealed sites.     

Archeology, population genetics and studies of human racial ancestry.

Using traditional classification, many people have tried to determine the place and time of origin of the major human races. Two basically conflicting views have prevailed, one asserting that races developed as local variations in a species evolving phyletically over much of the Old World for up to one million years, and the other view holding that present races are local variations on populations recently expanding into most areas and replacing the previous hominid inhabitants, this occurring since the time of the Neanderthals. Both views are based on the same archeological data. Gene frequency data and time-calibrated genetic distance measures have recently been applied to this problem. Here we show that the probable demographic nature of Pleistocene populations has obscured genetic distances to such an extent that they cannot be used to discriminate between the two viewpoints of racial origins. The racial classifications themselves are probably not useful in this context and obscure the question, so that we presently do not have a scientifically valid understanding of racial origins.     

Mitochondrial DNA analysis confirms the existence of two glacial races of rainbow smelt Osmerus mordax and their reproductive isolation in the St Lawrence River estuary (Quebec, Canada)

A phylogeographic analysis of mitochondrial DNA variation was performed in order to test the hypothesis that north-eastern North America has been postglacially recolonized by two races of rainbow smelt Osmerus mordax . This was accomplished by documenting the geographical distribution of two major mtDNA phylogenetic clades among 1290 smelt from 49 lacustrine and anadromous populations covering most of the species' native range. The data set was built by combining previously published results with those generated in this study. The two mtDNA clades showed a geographical dichotomy, independent of life-history types, whereby the more eastern populations were either fixed or largely dominated by one clade and western populations for the other. Such geographical pattern implying a phylogenetic discontinuity provided strong evidence for the persistence of smelt in two distinct glacial refugia as well as their differential postglacial dispersal. The most likely refugium for the so-called Atlantic race was the Atlantic coastal plains, whereas that of the so-called Acadian race was the exundated Grand Banks area. Patterns of postglacial dispersal interpreted from palaeogeographic events suggested that the Atlantic race recolonized northern regions about 5000 years prior the Acadian race. Both races came into contact in the St Lawrence River estuary. While gene flow has been possible, the sympatric occurrence in the estuary of anadromous populations alternatively dominated by one mtDNA clade or the other indicated that reproductive isolation mechanisms between the two races developed within this contact zone. This represents the first evidence of secondary intergradation among distinct races of aquatic organisms in an estuarine environment.     

Interspecific interactions drive cultural co-evolution and acoustic convergence in syntopic species

1.?Antagonistic interactions have been favourite subjects of studies on species co-evolution, because coexistence among competing species often results in quantifiable character displacement. A common output for competitive interactions is trait divergence, although the opposite phenomenon, convergence, has been proposed to evolve in some instances, for example in the communication behaviour of species that maintain mutually exclusive territories. 2.?I use here experimental and observational evidence to study how species interactions drive heterospecific signal convergence and analyse how convergence feeds back to the interaction itself, in the form of aggressive behaviour. I recorded the learned territorial signals of two non-hybridizing larks, Galerida cristata and G.?theklae, and used allopatric populations as controls for evaluating acoustic convergence in syntopy. Acoustic variation was analysed with respect to social conditions controlling for other potential agents of natural selection, habitat and climate. 3.?Interspecific convergence of Galerida calls peaked in syntopy. Although call acoustic structure was affected by climate and habitat, it matched gradients of density and proximity to congeners even at small local scales. The process of cultural transmission, in which individuals may acquire components of behaviour by copying neighbours, enhances the correlation between call acoustics and the local social milieu. 4.?Territories were defended against both species, but playback stimuli of convergent congener calls elicited a stronger aggressive reaction than congener calls from allopatric locations. 5.?This study shows that learned behaviours may co-evolve as a consequence of antagonistic interactions, determining reciprocal cultural evolution or cultural co-evolution. As for (biological) co-evolution, the distribution of competing species influences whether a particular area becomes a syntopic environment in which convergence is occurring, or an allopatric environment lacking interactions and reciprocal change. Because of their plastic nature, cultural coadaptations may rapidly shift in response to fluctuating social selection, thus propelling dynamic interactions and fine adjustments to the local environment.     

The tobacco mouse and its relatives: a "tail" of coat colors, chromosomes, hybridization and speciation

The article reviews over 30 years' study of the chromosomal variation of the western house mice (Mus musculus domesticus) from the neighboring valleys of Poschiavo and Valtellina on the Swiss-Italian border. This is done in the context of the social and political history of this area, on the grounds that mice, as commensals, are influenced by human history. The chromosomal study of mice in this area was initiated because their unusual black coat color led a 19th century naturalist to describe the "tobacco mice" from Val Poschiavo as a separate species (Mus poschiavinus). The special coloration of the Val Poschiavo mice is matched by their chromosomes: they have 26 chromosomes instead of the usual 40. The Val Poschiavo mice are not a separate species according to the Biological Species Concept; instead they constitute a chromosome race (the "Poschiavo", POS) that is related to other races with reduced chromosome numbers that occur in N Italy (of which only those races in Val Poschiavo and Upper Valtellina have black coats). A phylogenetic analysis of mitochondrial DNA sequences suggests that the lineage of chromosome races found in N Italy was not formed during an extreme population bottleneck, although such bottlenecks have apparently occurred during the origin of individual races and certainly have influenced single populations. In one small, isolated population in Valtellina (Migiondo), two chromosome races (the POS and the "Upper Valtellina", UV, 2n = 24) became reproductively isolated from each other. In another small population (Sernio) bottlenecking led to fixation of a hybrid form with the UV karyotype and coat color, but with allozyme and microsatellite alleles characteristic of mice with the standard 40-chromosome karyotype. Two of the chromosome races in Valtellina (the UV and the "Mid Valtellina", MV, 2n = 24) also appear to be the product of hybridization. The dynamic history and patchy distribution of the house mouse chromosome races in Val Poschiavo and Valtellina in part reflects extinction-recolonization events; the formation of the UV and MV races and the introduction of the pale brown Standard race mice are believed to reflect such events. Dynamism in the chromosomal constitution of single populations is also evident from 25 years of data on the population in Migiondo. Due to change in agricultural practices, house mice in Valtellina and Val Poschiavo are becoming rarer, which is likely to have further impacts on the distribution and characteristics of the chromosome races in this area.     

Races of Ammomanes deserti in Arabia.

T he Arabian patchwork of differently-coloured races of the "Desert Lark" (see Ibis, 1936, pp. 332–4) can now be exhibited in a more definite shape than three years ago, since we have much additional material to work on.     

鸡冠花植株内无机元素的分析研究

用电感耦合等离子体发射光谱法（ＩＣＰ－ＡＥＳ）测定了鸡冠花３个栽培品种的２３种无机元素,即Ａｌ、Ｂａ、Ｂｅ、Ｃａ、Ｃｄ、Ｃｏ、Ｃｒ、Ｃｕ、Ｆｅ、Ｋ、Ｌｉ、Ｍｇ、Ｍｎ、Ｍｏ、Ｎａ、Ｎｉ、Ｐ、Ｐｂ、Ｓｉ、Ｓｒ、Ｔｉ、Ｖ、Ｚｎ。结果表明,不同品种及不同器官,其无机元素含量存在显著差异。总体上看,该种植物无机元素含量丰富,不同器官中以种子尤为突出,３个品种中以园绒鸡冠突出。     

Competition between birds and mammals: A comparison of giving-up densities between crested larks and gerbils

We combined the concept of mechanisms of co-existence with the approach of giving-up densities to study inter-taxon competition between seed-eating birds and mammals. We measured feeding behaviour in food patches to define and study the guild of seed-eating vertebrates occupying sandy habitats at Bir Asluj, Negev Desert, Israel. Despite a large number of putatively granivorous rodents and birds at the site, two gerbil species (Allenbys gerbil, Gerbillus allenbyi, and the greater Egyptian gerbil, G. pyramidum) dominated nocturnal foraging, and a single bird species (crested lark, Galerida cristata) contributed all of the daytime foraging. We used giving-up densities to quantify foraging behaviour and foraging efficiencies. A low giving-up density demonstrates the ability of a forager to profitably harvest food at low abundances and to profitably utilize the foraging opportunities left behind by the less efficient forager. Gerbils had lower giving-up densities in the bush than open microhabitat, and lower giving-up densities in the semi-stabilized than stabilized sand habitats. Crested larks showed the opposite: lower giving-up densities in the open than bush, and on the stabilized than semi-stabilized sand habitats. Despite these patterns, gerbils had substantially lower giving-up densities than crested larks in both microhabitats, all sand habitats, and during each month. Several mechanisms may permit the crested lark to co-exist with the gerbils. Larks may be cream skimmers on the high spatial and temporal variability in seed abundances. Larks may rely on insects, fruit or smaller seeds. Or, larks may rely on adjacent rocky habitats.     

THE SPECIFIC CHARACTERS OF THE CRESTED PORCUPINES, SUBGENUS HYSTRIX

The characters used for specific diagnosis in the subgenus Hystrix have been studied to determine their constancy as specific characters and their geographical variation within the species. As a result three species are recognized, namely H. indica, H. cristata and H. africaeaustralis. H. galeata is considered a synonym of H. cristata , and H. stegmanni a synonym of H. africaeaustralis . The morphological differences between cristata and africaeaustralis in East Africa are not continued clinally within these species. Between cristata and indica some of the diagnostic characters vary clinally, others change abruptly.     

鸡冠花叶蛋白质营养价值的评价研究

应用模糊识别法和氨基酸比值系数法,分别以鸡蛋蛋白南为标准蛋白,以ＷＨＯ／ＦＡＯ氨基酸参考模式为评价标准,对３种鸡冠花叶蛋白质营养价值进行了全面评价,并与１０种常见叶菜蛋白进行对照比较。结果表明,３种鸡冠花叶（干品）蛋白质含量为２３．７％￣２７．４％,蛋白质中氨基酸种类齐全,其含量为８３．４７％￣８６．９４％,必需氨基酸（ＥＡＡ）占总氨基酸量的４０．２％￣４１．７％,第一限制性氨基酸为含硫氨基酸（Ｍ     

The status of Anostraca,Notostraca and Conchostraca (Crustacea: Branchiopoda) in Yugoslavia

The investigations of Anostraca, Notostraca and Conchostraca in Yugoslavia began after a great delay compared with other European countries. Intensive systematic investigations date back only to the 1970s.To date, 19 species have been recorded. All species, with the exception of Artemia salina, are faunal elements of the Pannonian region. Seven species, including Artemia salina, Branchinecta orientalis, Branchipus serbicus, Leptestheria dahalacensis, Limnadia lenticularis, Imnadia cristata and Imnadia panonica are known from single localities. Six species including Branchinecta ferox, Chirocephalus brevipalpis, Chirocephalus carnuntanus, Lepidurus apus, Eoleptestheria spinosa and Imnadia banatica have restricted distributions. Streptocephalus torvicornis and Cyzicus tetracerus are known from several localities, while Branchipus schaefferi, Chirocephalus diaphanus, Triops cancriformis and Leptestheria saetosaare common. Large branchiopods are mainly confined to anthropogenic landscapes, especially in the northern part of the country. Rare species inhabiting a single pond, or those with rather restricted distributions, are the most endangered and such species account for about 70% of the fauna. Limnadia lenticularis, Imnadia cristata and I. panonica have not been found for more than 20 years and they are probably extinct in Yugoslavia. Branchipus serbicus, Imnadia cristata and panonica have not been documented since being originally described. Eoleptestheria spinosa is currently the most endangered species since it only appears irregularly in small populations. At present, Branchinecta ferox, Chirocephalus carnuntanus and Imnadia banatica are the safest of the rare species as they regularly appear in large numbers. There are no large branchiopods on the List of protected animals in Yugoslavia. This revised version was published online in August 2006 with corrections to the Cover Date.     

Genetics and the origin of human races

In the last decades, the concept of human races was considered scientifically unfounded as it was not confirmed by genetic evidence. None of the racial classifications, which strongly differ in the number of races and their composition, reflects actual genetic similarity and genealogy of human populations inferred from variability of classical markers and DNA regions. Moreover, intercontinental ("interracial") variability was shown to be far lower than that within populations: the former constitutes 7 to 10% and the latter, about 85% of the total genetic variation. It is believed that the low level of differentiation of regional population groups contradicts their race status and suggests a recent origin of humans from one ancestral population. The results of studies of various genetic systems are in agreement with last conclusion rejecting the hypothesis of regional continuity. According to this hypothesis, the populations of continents regarded as large races have developed during long evolution from local types of archaic humans, in particular, Neanderthals. Phenotypic similarity of different, sometimes unrelated, populations united into one "race" is explained by strong selection since race-diagnostic traits characterize body surface and thus are directly subjected to the influence of environmental (primarily climatic) factors. It has been recently established that variability of the most important of these traits, body and hair pigmentation, is largely controlled by one locus (MC1R), which accounts for its high evolutionary lability. Other traits used for race identification are also likely to be labile and controlled by major genes. However, the fact that the currently existing race classifications are groundless does not mean that such classifications are impossible in principle. Commonly used argumentation (races do not exist because populations are not genetically separated) does not hold water. A polytypic species is characterized by genetic continuity of allopatric populations rather than the presence of narrow genetic boundaries between them. Borderlines between races are usually conventional and arbitrary. As to intergroup variation in humans, it is indeed low but comparable with that in some other species. There are no obstacles to the development of genetic systematics of human races.     

Reconstructing Dryopteris "semicristata" (Dryopteridaceae): Molecular profiles of tetraploids verify their undiscovered diploid ancestor

? Premise of the study: Discovering missing ancestors is essential to understanding the evolutionary history of biodiversity on Earth. Evidence from extinct species can provide links for reconstructing intricate patterns of reticulate relationships among extant descendents. When fossils are unavailable and other evidence yields competing hypotheses to explain species ancestry, data from proteins and DNA can help resolve conflicts and generate novel perspectives. The identity of a parent shared by two tetraploid species in the cosmopolitan fern genus Dryopteris has remained elusive for more than 50 years. Based on available data, four hypotheses were developed previously, each providing a different resolution to this uncertainty. ? Methods: New molecular evidence from studies of isozymes and restriction site analysis of chloroplast DNA tested the competing hypotheses about the diploid ancestors of these two extant Dryopteris polyploids. ? Key results: The results falsify two of the hypotheses, resolve the uncertainty in the third, and support the fourth. ? Conclusions: Our data validate the prior existence of Dryopteris "semicristata," which was proposed 38 years ago as a diploid progenitor of the allotetraploids D. cristata and D. carthusiana but has never been collected. After developing a phylogeny using the new molecular data, we describe a plausible morphology for D. "semicristata" by extrapolating likely character states from related extant species.     

Genetics and the Origin of Human “Races”

In the last decades, the concept of human races was considered scientifically unfounded as it was not confirmed by genetic evidence. None of the racial classifications, which strongly differ in the number of races and their composition, reflects actual genetic similarity and genealogy of human populations inferred from variability of classical markers and DNA regions. Moreover, intercontinental (interracial) variability was shown to be far lower than that within populations: the former constitutes 7 to 10% of the total genetic variation and the latter about 85% of it. It is believed that the low level of differentiation of regional population groups contradicts their race status and suggests a recent origin of humans from one ancestral population. The results of studies of various genetic systems are in agreement with the latter conclusion rejecting the hypothesis of regional continuity. According to this hypothesis, the populations of continents regarded as large races have developed during long evolution from local types of archaic humans, in particular, Neanderthals. Phenotypic similarity of different, sometimes unrelated, populations united into one race is explained by strong selection since race-diagnostic traits characterize body surface and thus are directly subjected to the influence of environmental (primarily climatic) factors. It has been recently established that variability of the most important of these traits, body and hair pigmentation, is largely controlled by one locus (MC1R), which accounts for its high evolutionary lability. Other traits used for race identification are also likely to be labile and controlled by major genes. However, the fact that the currently existing race classifications are groundless does not mean that such classifications are impossible in principle. Commonly used argumentation (races do not exist because populations are not genetically separated) does not hold water. A polytypic species is characterized by genetic continuity of allopatric populations rather than the presence of narrow genetic boundaries between them. Borderlines between races are usually conventional and arbitrary. As to intergroup variation in humans, it is indeed low but comparable with that in a number of other species. There are no obstacles to the development of genetic systematics of human races.     

Cytotaxonomic study of theVicia cracca complex 1. Czechoslovak taxa

Three races ofVicia cracca L. have been found on the territory of Czechoslovakia, two with the chromosome number 2n=14 and one with 2n=28. Diploid races seem to be more primitive and are less widely distributed. They occur mostly in the primary communities while the tetraploid race is very plastic in the ecological respect and is common both in primary and secondary communities. These races are characterized by some morphological and ecological features. As far as the vertical distribution is concerned, one of the diploid races and the tetraploid occur mostly in the lowland and in the colline zone, the second diploid race has a mountain character. The related tetraploid speciesVicia oreophila ?ertová, with the chromosome number 2n=28 is distributed at similar altitudes.     

The role of the nasal passages in the water economy of crested larks and desert larks

Condensation of water vapor in the exhaled air stream as it passes over previously cooled membranes of the nasopharynx is thought to be a mechanism that reduces respiratory water loss in mammals and birds. Such a mechanism could be important in the overall water economy of these vertebrates, especially those species occupying desert habitats. However, this hypothesis was originally based on measurements of the temperature of exhaled air (Tex), which provides an estimate of water recovered from exhaled air as a proportion of water added on inhalation but does not yield a quantitative measure of the reduction in total evaporative water loss (TEWL). In this study, we experimentally occluded the nares of crested larks (Galerida cristata), a cosmopolitan species, and desert larks (Ammomanes deserti), a species restricted to arid habitats, to test the hypothesis that countercurrent heat exchange in the nasal passages reduces TEWL. Tex of crested larks increased linearly with air temperature, (Ta): Tex=8.93+0.793xTa. Following Schmidt-Nielsen and based on measurements of Tex, we predicted that crested larks would recover 69%, 49%, 23%, and -5% of the water added to the inhaled air at Ta's of 15 degrees, 25 degrees, 35 degrees, and 45 degrees C, respectively. However, with the nares occluded, crested larks increased TEWL by only 27%, 10%, and 6% at Ta's of 15 degrees, 25 degrees, and 35 degrees C, respectively. At Ta=45 degrees C, TEWL of the crested lark was not affected by blocking the nares. In contrast to our expectation, occluding the nares of desert larks did not affect their TEWL at any Ta.