Variations in ribosomal DNA and mitochondrial DNA among chromosomal species of subterranean mole rats

Restriction site variations in nuclear ribosomal DNA (rDNA) spacers and mitochondrial DNA (mtDNA) were examined in several populations of mole rats with variable numbers of chromosomes, which represented the two superspecies Spalax leucodon (2n = 38, 54, or 62) and Spalax ehrenbergi (2n = 52, 54, 58, or 60). Sequence divergence of rDNA spacers between the members of the superspecies was approximately 8%, while the variation within each superspecies was 4% on average. The intrasuperspecies differentiation of rDNA spacers was generally associated with changes in the diploid number of chromosomes. However, substantial divergence (approximately 1.5%) was also detected among populations with the same diploid number of chromosomes. The sequence divergence of mtDNA among haplotypes of S. ehrenbergi was 10% or higher and among haplotypes of S. leucodon it was approximately 12%. By contrast, the range of sequence divergence between superspecies was 7.4%-12%. The large divergence of mtDNAs within each superspecies of mole rats may be explained by the preservation of ancient mtDNA polymorphisms.      

Variations of molar morphology in the Spalax ehrenbergi superspecies: adaptive and phylogenetic significance

This study compares and contrasts variations of enamel fold pattern on the crowns of the molars of subterranean mole rats ( Spalax ) belonging to two superspecies, S. ehrenbergi and S. leucodon , and involving about 20 chromosomal species, distributed parapatrically and ranging on different soil types. The sample studied involved 397 skulls, subdivided as follows: 280 of S. ehrenbergi from Israel, representing the four chromosomal species 2n = 52, 54, 58 and 60; 59 of S. ehrenbergi from Egypt, Lebanon, Syria, Jordan and Turkey; 58 of S. leucodon from Turkey.
In making the comparisons it was necessary to take into account the marked changes of molar pattern that take place as a result of wear.
There is a reasonably good agreement between differences of molar pattern and genetic and immunological distances. The species in Israel and Egypt form a compact group, clearly differentiated from Turkish leucodon; ehrenbergi from Turkey and Syria are intermediate, in accordance with their geographical distribution.
In Israel, differences associated with soil type are at about the same level as differences between species. Local populations show a higher level of diversity, and in a number of cases neighbouring populations differed significantly, sometimes over very short distances. Population differences within species are generally associated with differences of soil and vegetation, indicating the influence of natural selection. There are also differences across species boundaries, implying a phylogenetic (heritage) effect. The nature of the adaptive link between molar pattern and soil is unknown: possible factors are contamination of the food by soil, and toughness of the food which necessitates changes of enamel fold length to improve grinding efficiency.     

MtDNA polymorphisms: evolutionary significance in adaptation and speciation of subterranean mole rats

Patterns of mtDNA diversity in subterranean mole rats of the Spalax ehrenbergi superspecies (2n = 52, 54, 58 and 60) were previously studied in the main ranges of the four chromosomal species, and specifically in the 2n = 60 species and its peripheral steppe semi-isolates and desert isolates. In the present study we correlated mtDNA diversity indices, nucleon diversity, h , and nucleotide divergence, π, with physical (climatic), biotic (parasites) and biological (genetical, morphological, physiological and behavioural) factors, showing that mtDNA diversity is structured ecogeographically and biologically. The following significant correlations of mtDNA diversity were indicated with: (i) climatic heterogeneity and unpredictability; (ii) levels of ecto- and endoparasites; and (iii) biological diversities, primarily with physiological diversity associated with the energy budget. Small steppe semi-isolates and desert isolates harbour high levels of mtDNA haplotype diversity, some novel, which may be a prerequisite for future speciation events. We conclude that the ecogeographical and biological correlates, as well as the maintenance of mtDNA polymorphisms in small isolated populations, strongly suggest that mtDNA diversity is not neutral. Diversifying natural selection appears to be an important differentiating factor of mtDNA diversity in the twin evolutionary processes of adaptive radiation and active speciation. We suggest critical experiments to substantiate our conclusions and highlight the contribution of mtDNA diversity to fitness, i.e. to the biological function of mtDNA diversity in the evolutionary process.     

A karyological study on subterranean mole rats of the Spalax leucodon Nordmann, 1840 superspecies in Turkey

The karyotypes of 179 specimens of the subterranean mole rats of the Spalax leucodon Nordmann, 1840 superspecies across 40 localities in Turkey were analysed. It was determined that S. leucodon has 2n = 56, NF = 76 in the European part of Turkey (Eceabat population) but 2n = 38, NF = 74; 2n = 50, NF = 70; 2n = 52, NF = 70; 2n = 54, NF = 72; 2n = 56, NF = 74; 2n = 58, NF = 78; 2n = 60, NF = 78; 2n = 60, NF = 80 in populations analysed from the Asian part of Turkey. According to these karyological findings the diploid chromosome numbers of 2n = 50, 2n = 52, 2n = 56 and 2n = 58 determined from the Asian part of Turkey are new fo Spalax leucodon in Turkey. Because diploid numbers of these populations were formerly found from geographically distant localities in Turkey, they were designated as 2n = 50 N, 52 N, 56 N and 58 N, to differentiate from the other forms having the same diploid chromosome numbers but different chromosome morphology.

The occurrence of so many chromosomal forms in such small areas suggested that many other new forms may be found by new studies in Turkey. The presence of 2n = 50, 56 and 58 populations in very small areas is strongly supportive of the opinion of a peripatric chromosomal evolution in Spalax.     

Genetic polymorphism of urine deoxyribonuclease I isomerases of subterranean mole rats,Spalax ehrenbergi superspecies,in Israel: Ecogeographical patterns and correlates

Genetic polymorphism of urine deoxyribonuclease I (DNase I) of mole rats was analyzed by isoelectric focusing in a thin-layer polyacrylamide gel (IEF-PAGE). One hundred and three subterranean mole rats, comprising 13 populations belonging to the four chromosomal species (2n = 52, 54, 58, 60) of the actively speciating Spalax ehrenbergi superspecies in Israel, were tested. The following results were indicated. (i) Spalax DNase I consisted of 6-12 major isozymes. (ii) Four phenotypes (numbers in parentheses) were 1 (92), 1-2 (5), 1-3 (4), and 2 (1). The decreasing order of genetic diversity, He, in the four species was 0.37, 0.13, 0.10, and 0.0 for 2n = 58, 52, 54, and 60, respectively. (iii) Spearman rank correlations and multiple regression analyses indicated associations of allele frequencies and genetic diversity with climatic and vegetation factors. We concluded that (a) climatic selection, either directly or indirectly through plant (i.e., food resources) diversity, plays an important role in DNase genetic differentiation and (b) no gene flow and introgression occur between the recent derivative of speciation (2n = 60) and its ancestor (2n = 58), suggesting the operation of reproductive isolation between both species despite natural hybridization.     

DNA sequence variation in the mitochondrial control region of subterranean mole rats,Spalax ehrenbergi superspecies,in Israel

The complete mitochondrial control region was sequenced for 60 individuals representing different populations for each of the four species of the subterranean mole rat Spalax ehrenbergi superspecies in Israel: Spalax galili (2n = 52), S. golani (2n = 54), S. carmeli (2n = 58), and S. judaei (2n = 60). The control region of all species and populations is very similar both in length (979 to 983 bp) and in base composition. As in agreement with previous surveys on mitochondrial control regions on mammals, the mole rat control region can be divided into a central domain and two flanking domains, ETAS (extended termination associated sequences) and CSB (conserved sequence blocks). Along with the common conserved blocks found in these domains (ETAS1, ETAS2, CSB1, CSB2, and CSB3), we have also detected in all individuals an ETAS1-like and a CSB1-like element, both in the ETAS domain. The most conserved region was the central domain, followed by the CSB and ETAS domains, showing important differences in the four species analyzed. Phylogenetic analysis supported the existence of two clades. One clade contained individuals belonging to Spalax galili (2n = 52) and S. golani (2n = 54), separated in two different branches depending on the species. The other clade contained individuals belonging to S. carmeli (2n = 58) and S. judaei (2n = 60) mixed together, suggesting a more recent event of speciation. Within species we have observed a southward trend of increasing variability. These results have been explained as a consequence of the adaptation of the species to ecological factors such as aridity and temperature stresses.     

Brain size diversity in adaptation and speciation of subterranean mole rats

We have tested brain size diversity and encephalization in the actively speciating subterranean mole rats of the Spalax ehrenbergi superspecies in Israel. Our sample involved 171 individuals comprising 12 populations and 4 chromosomal species (2n = 52, 54, 58 and 60) distributed parapatrically from the northern Mediterranean region southward (2n = 52, 54→+58→60) into increasingly more arid and unpredictable climatic regimes, approaching the Negev Desert. Our results indicate that relative brain size and encephalization are highest in 2n = 60 as compared with 2n = 52, 54 and 58. We hypothesize that this pattern is adaptive and molded by natural selection. Brain evolution and higher encephalization in the S. ehrenbergi complex appears to be associated with increasing ecological stresses of aridity and climatic unpredictability.     

Morphometrics of speciating mole rats: Adaptive differentiation in ecological speciation

The morphometrics of subterranean mole rats, Spalax ehrenbergi superspecies in Israel, were studied. A total of 42 skull and body variables of 327 adults from 44 populations across the ranges of the 4 chromosomal species (2n = 52, 54, 58 and 60) were measured. The results showed: a. in general, significantly higher morphological values in males than in females; b. significant interspecific quantitative diffferences in 15 skull variables of males, and in 3 of females; but no qualitative diagnosis of any of the species; c. an intercorrelation between most skull variables at different orgamsmal levels, and a uniform and complete correlation with weight; d. a statistical discrimination in both metric and nonmetric multivariate analyses between most species pairs; e. an indication of high morphological similarity based on the relatively small multivariate Mahalanobis distances of dissimilarity between species; f. a possible explanation for a significant part of the variance in most skull and body variables resulting from a combination of temperature variables and water availability; g. a declination in size in skull and body variables between northern and southern species; h. the order of speciation events to be 2n = (54, 52)→58→60. Based on these results we conclude: 1. no morphological breaks are associated with the speciation of the S. ehrenbergi complex, and morphological differentiation is quantitative rather than qualitative between the species: 2. morphological diversity is significantly explained by climatic selection, and is therefore adaptive and explicable on even very low selective pressures over evolutionary time: 3. the morphospecies seem unable to reliably reflect the biological species in the S. ehrenbergi complex, and morphological differentiation, at least in this superspecies, evolves in a gradual rather than a punctual way: this would conform with Neo-Darwinian evolutionary theory.     

Banded karyotypes of mole rats (Spalax, Spalacidae, Rodentia) from Turkey: a comparative analysis

Chromosome banding (G-, C- and Ag-NOR) analysis was carried out on 27 specimens of Sphalax ehrenbergi from seven localities and two specimens of S. leucodon from one locality, all from Turkey. No chromosomal variation was detected in S. ehrenbergi populations from Elazig, Siverek, Diyarbakir and Birecik having the same diploid numbers (2n = 52) and morphology of chromosomes (NFa = 72). The karyotypes of mole rats from Tarsus and Gaziantep possessed the identical diploid number (2n = 56) but different numbers of autosomal arms: NFa = 68 in the Tarsus and NFa = 78 in the Gaziantep populations. Chromosomes of S. leucodon from Malaty (2n = 60, NFa = 74) differed distinctly in the C-banding pattern from all S. ehrenbergi cytotypes by the almost entire absence of heterochromatin in acrocentric autosomes and the presence of heterochromatin arms iin subtelocentric autosomes. Nucleolar organizing regions were found mainly on three pairs of chromosomes, but some differences in their localization were revealed. Comparison of G-banded chromosomes showed, that most chromosomes have a similar pattern. The types of chromosomal rearrangemetns were revealed due to the banding methods.     

Total body water and adaptive water turnover rate in four chromosomal species of subterranean mole rats of the Spalax ehrenbergi superspecies in Israel

Total body water (TBW) and water turnover rate (WTR) were measured in 24 subterranean mole rats comprising four populations, each belonging to a different chromosomal species of the Spalax ehrenbergi superspecies in Israel. The four species range in different climates: humid-cool (2n = 52); semiarid-cool (2n = 54); humid-warm (2n = 58) and arid-warm (2n = 60). TBW, as a percentage of body weight, measured by tritiated water (HTO), was 72.4%±4–7 in 2n = 52, significantly ( P < 005) higher than the similar estimates 61.7%± 7.2, and 59.4%± 5–3, for 2n = 60 and 58, respectively. A comparison of HTO space, as a percentage of TBW, closely approximated TBW, ranging from 97% to 108%. WTR was high, 218.1 and 230.9 ml/kg^0-75/day in the mesic populations of 2n = 58 and 52, respectively. By contrast, WTR estimates were significantly lower , ( P < 0.001), 150.2 and 148.9 ml/kg^0-75/day in the xeric populations of 2n = 54 and 60, respectively. The biological half-life time, T_1/2, was similar and faster, 32.7 and 27.9 hours in the mesic populations of 2n = 52 and 58, as compared with slower, 47.9 and 40.8 hours in the xeric populations of 2n = 54 and 60, respectively. Urine osmolarity in the most xeric northern Negev steppic population of 2n = 60 (737 ± 45 mmol/kg) was significantly ( P < 0.001) higher than in the other species.
We conclude that adaptive radiation in the Spalax ehrenbergi superspecies involves speciation in semiarid (2n = 54) and arid (2n = 60) climates by physiological adaptations of kidney water conservation, along with multiple morphological, physiological and behavioural syndromes of climatic adaptations to increasing aridity (Nevo, 1986).     

Speciation and adaptive radiation of subterranean mole rats, Spalax ehrenbergi superspecies, in Jordan

The major initial mechanism of speciation in subterranean blind mole rats, Spalacidae, is chromosomal, primarily through Robertsonian rearrangements. Here we highlight another scenario of chromosomal rearrangement leading to ecological speciation and adaptive radiation apparently initiated by pericentric inversions and genie divergence to different ecologies in mole rats in Jordan. We analysed karyotype, allozyme, size and ecological diversity across the range of mole rats in Jordan from mesic Irbid in the north to xeric Wadi Musa (Petra region) in the south, a transect of 250 km. We examined mole rats for chromosome ( N =71), size ( N =76), and allozyme ( N =67) diversities, encoded by 32 loci, in 12 populations of the Spalax ehrenbetgi superspecies in Jordan. By a combination of chromosome morphology, genetic distance, body size and ecogeography, we identified four new putative biological species. All species (except two animals in Madaba) share 2 n = 60 but vary in chromosome morphology, caused by pericentric inversions and/or centromeric shifts. The 'north Moav' species is karyotypically polymorphic for 2 n (2 n = 60; including locally also two animals with 2 n = 62). The distribution of the four species is associated with ecogeographical different domains and climatic diversity. Genetic diversity indices were low, but like chromosome arms (NFa) were positively correlated with aridity stress. Discriminant analysis correctly classified 91% of the individuals into the four species utilizing combinatorially chromosome, allozyme and size diversities. It is hypothesized that mole rat evolution underground is intimately associated with climatic diversity stress above ground.     

Mitochondrial DNA polymorphisms in subterranean mole-rats of the Spalax ehrenbergi superspecies in Israel, and its peripheral isolates

Patterns of mitochondrial DNA (mtDNA) variation were examined in 133 mole-rats constituting all four chromosomal species (2n = 52, 2n = 54, 2n = 58, and 2n = 60) of the Spalax ehrenbergi superspecies in Israel, as well as the peripheral isolates of 2n = 60. In the main range of the complex, a total of 28 mtDNA haplotypes were found in 64 mole-rats, with most haplotypes being unique to either a single chromosomal species or population. mtDNA divergence increased from low to high diploid number in a north-to-south direction in Israel. Overall levels of mtDNA diversity were unexpectedly the highest in the 2n = 60, the youngest species of the complex. The mtDNA haplotypes can be separated into two major groups, 2n = 52-54 and 2n = 58-60, and a phylogenetic analysis for each group revealed evidence of a few haplotypes not sorted by diploid number. The overall patterns of mtDNA divergence seen within and among the four chromosomal species are consistent with the parapatric mode of speciation as suggested from previous studies of allozyme and DNA hybridization. In a separate data set the patterns of mtDNA variation were examined across the main geographic range and across peripheral semi-isolates and isolates of the 2n = 60 chromosomal species. Fifteen haplotypes were found in 69 mole-rats. High levels of mtDNA diversity characterized the main range, semi-isolated, and even some desert isolated populations. The peripheral isolates contain much mtDNA diversity, including novel haplotypes.      

Sexual selection and natural selection in body size differentiation of subterranean mole rats

We exlored indirectly, the operation of sexual selection in subterranean mole rats of the Spalax ebrenbergi superspecies in Israel comprising four chromosomal species, 2n = 52, 54, 58 and 60. We reanalzed two previously available data sets of 1. body size differentiation (Nevo et al. 1986a) and 2. the intensity of “Total Aggression” in mole rats (Nevo et al. 1986b). We correlated the mean size difference between the two sexes, in each of the 12 populations of the chromosomal species, with the mean level of agression, and with climatic factors, both displaying significant correlations. The results indicated that for 2n = 52, 54, 58 and 60, the population averae difference in body weight between the sexes decreases southward as follows: 37.7g (30.8 % of females body weight), 39.3g(29.0%) 26.3g(22.8%) and 20.3g (19.3%), respectively. We interpret the higher body size diherential ketween the sexes in the north as due to sexual selection.     

Microsatellite diversity in populations of blind subterranean mole rats (Spalax ehrenbergi superspecies) in Israel: speciation and adaptation

The genetic diversity, structure, and relationships of 12 populations of blind subterranean mole rats belonging to four species of the Spalax ehrenbergi superspecies were examined. Diversity levels and divergence of 19 nuclear microsatellite markers were analysed. Results indicate that the 12 populations fall into two major clusters representing the species pairs Spalax galili (2 n  = 52) and S. golani (2 n  = 54) vs. S. carmeli (2 n  = 58) and S. judaei (2 n  = 60). There is very little gene flow between the species pairs, except for one population (Afiq) of S. carmeli (2 n  = 58), which is located near the hybrid zone between S. golani (2 n  = 54) and S. carmeli (2 n  = 58). Higher values of the mean for alleles per locus, gene diversity, and number of unique alleles were observed for the southern species pair, S. carmeli and S. judaei, than for the northern species pair, S. galili and S. golani . Microsatellite diversity increases southward toward the Negev Desert and is positively correlated with aridity stress. Climatic factors of water availability coupled with temperature account for 76.7% of microsatellite variance. Natural selection appears to adaptively determine microsatellite evolution in Spalax regionally across Israel.  © 2004 The Linnean Society of London , Biological Journal of the Linnean Society , 2004, 83 , 229–241.     

The effect of protein and salt loading on urinary concentrating ability in four chromosomal species of Spalax ehrenbergi

Urinary concentrating ability was tested under protein and salt load in the four chromosomal species of subterranean mole-rats ( Spalax ehrenbergi superspecies) found in Israel. Protein stress induced by a diet of soyabeans supplemented by agar gel, demonstrated a significant increase in urinary osmolarity (UO). In the species living in the driest and warmest region (2n = 60). UO (1423·101 mOsmol/kg) was significantly (P<0·05) higher than in the other three species (2n = 52. 1172·31 mOsmol/kg; 2n = 54, 1160·116 mOsmol kg; and 2n = 58, 1216·145 mOsmol/kg). Upon salt loading this diet with 0·3 mol NaCl, UO increased significantly. However, when the salt load was increased to 0·45 mol NaCl, UO decreased significantly in all but one species (2n = 60) which maintained UO at 1522·65 mOsmol/kg. A decline in UO was attributed to diuresis resulting from a significant increase in urine and sodium excretion. The kidney, of only the xeric ranging species (2n = 60), demonstrated the ability to produce a hyperosmotic urine, in spite of the high salt load. These results might explain the restricted distribution of S. ehrenbergi: the only species (2n = 60) found in an environment rich in succulents and halophyte plants (steppe). This species appears to push speciation and adaptive radiation to the southern limit of its superspecies.     

Chromosomal polymorphisms in subterranean mole rats: origins and evolutionary significance

Two distinct chromosomal polymorphisms, among others, characterize the short (p) and long (qh) arms of chromosome number 1 of the Spalax ehrenbergi complex in Israel. We have studied the geographic distributions of these two polymorphisms in 60 animals belonging to four chromosomal species (2 n = 52, 54, 58 and 60). These comprised 15 populations: 12 from continuously distributed populations, two from semi-isolates, and one isolate in the northern Negev Desert.
Our results indicate that: (i) the two polymorphisms are widespread not only within populations and species but most strikingly between different bone marrow cells within an individual; this may reflect a diversity generating mechanism; (ii) the two polymorphisms of p and qh, primarily of p, are correlated with climatic factors of water availability and temperature, as well as geographical distances; these correlations are significantly above those expected by chance, (iii) The semi-isolates and the isolate display high levels of polymorphism in both p and qh, but particularly in qh, despite the small size population effects.
We suggest that at least the polymorphism in p is involved in the adaptive radiation of mole rats into diverse climatic regimes. Furthermore, we speculate that the forces generating chromosomal polymorphisms relevant to speciation (i.e. Robertsonian mutations), coupled with the evolutionary forces operating in small peripherally isolated populations, may be appropriate for the origin of new chromosomal species through peripatric speciation.     

Hypoxic survival differs between two mole rat species (Spalax ehrenbergi) of humid and arid habitats.

1. Two chromosomal species, 2n = 52 and 2n = 60 of the mole rat superspecies (Spalax ehrenbergi), occupy humid (2n = 52) and arid (2n = 60) habitats in Israel. 2. Gas conductivity of the soil of the 2n = 52 mole rat is lower than that of the 2n = 60 mole rat, and the 2n = 52 mole rat is better adapted to hypoxia. 3. The hypothesis that the 2n = 52 mole rat can survive to a lower pO2 than the 2n = 60 mole rat was tested. 4. Terminal pO2 (Torr) of four females 2n = 52 was lower, 18.0 +/- 2.9 (SD), than the terminal pO2 of five females 2n = 60, 28.2 +/- 5.1 (SD). 5. The hypoxic survival of the 2n = 52 mole rat as compared to that of the 2n = 60 mole rat correlates with other physiological traits: breathing and heart frequencies, blood hemoglobin and tissue gas tensions.     

Mhc Class II DNA Polymorphisms within and between Chromosomal Species of the Spalax Ehrenbergi Superspecies in Israel

Restriction fragment length polymorphisms (RFLPs) of two major histocompatibility class II genes (P alpha 1 and Q beta) were studied in 13 populations of four chromosomal species (2n = 52, 54, 58 and 60) of the mole rat, Spalax ehrenbergi superspecies in Israel. A substantial frequency of allelic fragments was found in both genes for all populations, including a desert isolate. In the P alpha 1 gene, one allelic fragment is a result of a deletion mutation which is diagnostic of the 2n = 52 chromosomal species. All other ten allelic variants are the result of point mutations. All mutations are located in a short region flanking the 3' end of the gene. Based on Mhc polymorphisms we confirm earlier evidence that gene flow does not occur between the older chromosomal species (2n = 52, 54, 58), and that reproductive isolation decreases, progressively from the oldest to the youngest species (2n = 60).     

Penial differentiation in speciation of subterranean mole rats Spalax ehrenbergi in Israel

Geographic and ontogenetic variation was examined in the glans penes and bacula of 92 specimens representing four biological species with the karyotypes 2n=52, 54, 58 and 60 of the Spalax ehrenbergi superspecies in 13 populations from Israel and one disjunct population from northern Egypt. Five measured and three calculated penial characters were analysed and employed in multivariate statistics. Comparison was made between penial morphological characters and craniometric biochemical and chromosomal data. The baculum is strongly species-specific and its length bears no significant relationships with head and body length. The results of these analyses suggest that different bacular morphologies may provide a basis for species-specific prezygotic isolating mechanisms, thus, together with other factors, contributing to reproductive isolation and speciation.     

Aggression patterns in adaptation and speciation of subterranean mole rats

The patterns and correlates of aggressionwithin andbetween actively speciating subterranean mole rats of theSpalax ehrenbergi superspecies in Israel were tested in an attempt to approach an evolutionary theory of aggression related to both adaptation and speciation. Laboratory experiments were conducted on 314 adult mole rats (188 males and 126 females) caught in nature, representing 12 populations and comprising four chromosomal species (2n = 52, 54, 58, 60). The present analysis concentrates on “total aggression” comprising four aggressive variables: attacks, bitings, head-ons and sniffing with open mouth, out of 23 recorded variables. The results indicate that (a) aggression is distributed as a multipeak, discontinuous phenotypic parameter, displaying polymorphism within all 12 populations and 4 species tested; (b) the level of aggression was higher in males than in females, in the breeding than in the nonbreeding season, and displayed clinal geographic variation in both sexes across the superspecies range, where “militancy” increased northwards; (c) aggression was significantly correlated with ecological, physiological, genetical and ethological factors. These results support an evolutionary theory of aggression related to the intertwined processes of adaptation and speciation.