Morphological integration and pleiotropy in the adaptive body shape of the snail‐feeding carabid beetle Damaster blaptoides

The snail‐feeding carabid beetle Damaster blaptoides exhibits diverse head and thorax morphologies, and these morphotypes are linked with two alternative feeding behaviours. Stout‐shaped beetles feed on snails by crushing the shells, whereas slender‐shaped beetles consume snails by inserting their heads into the shells. A trade‐off exists between these feeding strategies. Because intermediate‐shaped beetles are less proficient in these two behaviours, stout–slender morphological divergence occurs between related species feeding on land snails. To examine the genetic basis of these morphotypes, we conducted morphological analyses and quantitative trait locus (QTL) mapping using backcross offspring between the stout and slender subspecies. The morphological analyses showed that the width and length of the beetle body parts were correlated with each other; in particular, the head width (HW) and thorax length (TL) were strongly negatively correlated. QTL mapping showed that QTLs for HW and TL are located in close proximity to one another on the longest linkage group and that they have positive and negative additive genetic effects. Our results suggest that the adaptive phenotypic sets of a wide head and short thorax and a narrow head and long thorax are based on the closeness of these QTLs. Morphological integration between the head and thorax may play an important role in the adaptive divergence of these beetles.     

Molecular genetic relationships and some issues of systematics of rock lizards of the genus Darevskia (Squamata: Lacertidae) based on locus analysis of SINE-type repeats (Squam1)

To study the molecular genetic relationships and correlate them with the taxonomy within the complex of lacertid lizards of the genus Darevskia, the locus analysis of the copies of the SINE-type repeat (Squam1) specific for the order Squamata was used. It was demonstrated that one of the loci (No. 34) contained the Squam1 copy insert in all species and subspecies of the examined genus. SINE allelic copies in some of the loci contained large indels and specific sets of mutations. The allelic variant M (medium, about 340 bp) was found most frequently; it was detected in all subspecies of D. saxicola (saxicola, darevskii, szczerbaki, lindholmi) and in most of the other species of the genus. Two species, D. derjugini and D. praticola, differed from the other species in the presence of long (L) and short (S) alleles. The longest allele was characteristic of the D. derjugini population from the Northern Caucasus (L, 379 bp, ssp. silvatica), while the shortest allele (97 bp) united the derjugini and barani subspecies. The second allele S (279 bp) characterizes the subspecies D. praticola praticola, some individuals of which also carry allele M. The second subspecies, D. p. pontica, contains allele L2, which differs from all other medium alleles in the presence of strictly specific short indel. In addition to apomorphic indels, the specificity and mutation distribution patterns among the Squam1 alleles were also examined. An analysis of the NJ tree indicated the concordance between morphological and molecular genetic characters of the species derjugini, praticola, and saxicola. Furthermore, four subspecies of D. saxicola were much closer to each other than the subspecies within the first two species; D. d. silvatica and the group of D. d. derjugini + barani were clearly separated. It cannot be excluded that populations from Azerbaijan and Serbia can be treated as the independent subspecies of D. praticola.     

Differentiation of subspecies of the polymorphic species Hyalomma marginatum (Acari: Ixodidae) based on immature stages

Study of morphological characters to identify the subspecies of Hyalomma marginatum immature stages is based on material collected throughout all the geographical range of the species. As it was found, the discrimination of subspecies of H. marginatum immature stages needs a complex use of structural and morphometrical characters. Only H. m. marginatum larvae and H. m. isaaci nymphs may be easily differentiated from other subspecies based on the structural (qualitative) characters. Besides, morphometric (quantitative) characters should be used only in combinations for identification of the subspecies. In regard to structural characters of larvae, it was found that the shape and size of coxal spurs of H. marginatum allow differentiating this subspecies from the others (Fig. 3, 5, 6). Based on morphometrical characters of larvae, the following characteristic features of subspecies have been found: in H. m. marginatum, the capitulum is wide, the palpi, hypostome and genua I are short and narrow; in H. m. turanicum, the capitulum is narrow, the palpi are short and narrow, the hypostome is short and wide, the genua I are long and narrow; in H. m. rufipes, the scutum is large, the capitulum is wide, the palpi are long and narrow, the hypostome and genua I are long and wide; in H. m. isaaci, the scutum is small, the capitulum is narrow, the palpi and hypostome are long and narrow, the genua I are short and narrow. Among structural characters of nymph, several discriminative features of subspecies have been found. The shape of the scutum in H. m. isaaci (Fig. 2, 2) clearly differentiates this subspecies from the others (Fig. 2, 1). In H. m. marginatum, the setae of alloscutum as a rule have bluntly rounded apices (Fig. 1, 2), while in the other subspecies these setae are more tapering apically (Fig. 1, 1). The shape of spiracular plates is rather variable within the species, but in H. m. marginatum and H. m. isaaci (Fig. 1, 3) the plates are larger and more perforated than in H. m. rufipes and H. m. turanicum (Fig. 1, 4). In H. m. isaaci, the spurs of coxae I are narrower than in other subspecies (Fig. 1, 8). Based on morphometrical characters of nymphs, the following discriminative features of subspecies have been found: in H. m. marginatum, the capitulum is wide, the palpi are short and wide; in H. m. turanicum, the capitulum is narrow, the palpi are long and narrow; in H. m. rufipes, the scutum is wide, the capitulum is narrow, the palpi are long and narrow, the hypostome is long; in H. m. isaaci, the scutum is long and narrow, the capitulum is wide, the palpi are short and wide, the hypostome is short.     

Longidorus biformis n. sp. and L. glycines n. sp. (Nematoda: Longidoridae): Two Amphimictic Species from Arkansas

Two new amphimictic species of Longidorus were found in Arkansas. Longidorus biformis n. sp., found in the rhizosphere of hardwood trees along streams in sandy soil in 14 Arkansas locations, is characterized by its long body (5.42-9.50 mm), wide expanded flattened head end, head width 20.0 to 26.0 µm, odontostyle 96 to 125 µm, guide ring 29 to 38 µm posterior to the anterior end, elongate conoid tail, and c'' = 0.9-2.1. Females with 2 to 11 vetromedian supplement-like structures were found in 2 of 14 populations of this new species. Longidorus biformis n. sp. is closest to L. seinhorsti Peneva, Loof &Brown, 1998 and L. closelongatus Stoyanov, 1964. Among North American species it is closest to L. glycines n. sp. A distinguishing feature of L. biformis n. sp. is the presence of supplement-like organs in some females. Longidorus glycines n. sp., found in soybean microplots at the Main Research Station, Fayetteville, Arkansas, is characterized by its long body (6.14-8.31 mm), wide offset flattened head end, head width 20.3 to 23.3 µm, odontostyle 87.3 to 99.5 µm, guide ring 22.3 to 26.4 µm posterior to the anterior end, short conoid tail with rounded terminus, and c'' = 0.9-1.4. Longidorus glycines n. sp. is closest to L. lusitanicus Macara, 1985. Among North American species it is close to L. biformis n. sp., L. breviannulatus Norton and Hoffman, 1975, and L. crassus Thorne, 1964. Both new species are believed to have four juvenile stages; the first stage was not found for L. biformis n. sp.     

Evolutionary diversification and geographical isolation in Dubautia laxa (Asteraceae), a widespread member of the Hawaiian silversword alliance

Background and Aims

The Hawaiian silversword alliance (Asteraceae) is one the best examples of a plant adaptive radiation, exhibiting extensive morphological and ecological diversity. No research within this group has addressed the role of geographical isolation, independent of ecological adaptation, in contributing to taxonomic diversity. The aims of this study were to examine genetic differentiation among subspecies of Dubautia laxa (Asteraceae) to determine if allopatric or sympatric populations and subspecies form distinct genetic clusters to understand better the role of geography in diversification within the alliance.

Methods

Dubautia laxa is a widespread member of the Hawaiian silversword alliance, occurring on four of the five major islands of the Hawaiian archipelago, with four subspecies recognized on the basis of morphological, ecological and geographical variation. Nuclear microsatellites and plastid DNA sequence data were examined. Data were analysed using maximum-likelihood and Bayesian phylogenetic methodologies to identify unique evolutionary lineages.

Key Results

Plastid DNA sequence data resolved two highly divergent lineages, recognized as the Laxa and Hirsuta groups, that are more similar to other members of the Hawaiian silversword alliance than they are to each other. The Laxa group is basal to the young island species of Dubautia, whereas the Hirsuta group forms a clade with the old island lineages of Dubautia and with Argyroxiphium. The divergence between the plastid groups is supported by Bayesian microsatellite clustering analyses, but the degree of nuclear differentiation is not as great. Clear genetic differentiation is only observed between allopatric populations, both within and among islands.

Conclusions

These results indicate that geographical separation has aided diversification in D. laxa, whereas ecologically associated morphological differences are not associated with neutral genetic differentiation. This suggests that, despite the stunning ecological adaptation observed, geography has also played an important role in the Hawaiian silversword alliance plant adaptive radiation.     

Meloidogyne petuniae n. sp. (Nemata: Meloidogynidae), a Root-knot Nematode Parasitic on Petunia in Brazil

Meloidogyne petuniae n. sp. is described and illustrated from specimens parasitic on petunia (Petunia hybrida L.) in Brasilia, Brazil. The perineal pattern of the female is elongate to ovoid with a high, squarish arch and widely spaced, coarse striae. The stylet of the female is 12.9-16.5 µm long and has three small, rounded knobs that are distinctly set off from the shaft. Each knob is marked by a deep longitudinal indentation posteriorly and anteriorly. In SEM the base of the shaft appears to be divided into six distinct ridges. The excretory pore opens about 15.4-53.6 µm from the head end. Males are approximately 0.8-2.2 mm long. Most specimens have a high and narrow head cap, but in some the head cap is narrow and low. The stylet of the male is 21.1-26.0 µm long and has small, rounded knobs, set off from the shaft, but not indented as in the female. Second-stage juveniles are 353-464 µm long; the labial disc is fused with the medial lips to form a dumbbell-shaped head cap; the medial lips are indented posteriorly; and the head region is marked with one to two irregular annulations. The stylet is 9.2-10.8 µm long and has rounded, posteriorly sloping knobs. The tail is slender, approximately 46.4-57.2 µm long, and has a short hyaline terminus, 10.3-13.5 µm long. The somatic chromosome number is 2n = 41 and the esterase phenotype is VS1-S1, with S1 being a weak band. The malate dehydrogenase phenotype is N1, which is unique for this species. Petunia, tomato, tobacco, pea, and bean are good hosts; pepper, watermelon, and sweet corn are poor hosts; and peanut, cotton, and soybean are non-hosts. Galls produced by this species are smaller on petunia than on tomato.     

拟平鳅头背部斑纹和下唇结构变异及其分类学意义

针对拟平鳅(Liniparhomaloptera disparis)的两个亚种(L.disparis disparis和L.disparis qiongzhongensis)现有形态分类特征的含糊,通过观察标本室保存的15条水系441尾标本,首次提出利用下唇结构类型作为拟平鳅亚种的补充形态鉴别依据,结合头背部斑纹特征,能有效地鉴别拟平鳅各地理种群和亚种,为拟平鳅的分类和鉴定提供参考依据。依据头背部斑纹差异归纳出4种类型,分别为小斑块型(云开山脉以东的7条水系及海南岛万泉河),虫蚀纹型(云开山脉以西的5条水系),混合斑纹型(海南岛的万泉河、南渡江、昌化江共3条水系),大斑块型(海南岛的昌化江)。依据下唇结构变化情况总结出3种下唇类型,分别是半圆型(云开山脉以西的5条水系),几字型(云开山脉以东的7条水系),马蹄型(海南岛的3条水系)。依据头背部斑纹特征结合下唇结构特征,对照模式标本的描述,重新定义了拟平鳅原有的两个亚种,将分布于云开山脉以东水系的种群鉴定为指名亚种,拟平鳅L.disparis disparis,其鉴别特征为:头背部斑纹为小斑块型并且下唇类型为几字型;将海南岛种群鉴定为琼中拟平鳅L.disparis qiongzhongensis,它们都具有独特的马蹄型下唇类结构,且头背部斑纹以混合斑纹型为主。重要的发现是,分布于云开山脉以西的种群与已命名的2个亚种在形态上有明显的差异,其头背部斑纹为独特的虫蚀纹型且下唇类型为半圆型。这可能是一个新亚种。分子生物学方法也许能证明云开山脉以西的种群在遗传上的独特性以及形成新亚种的可能性。云开山脉东、西两侧以及海南岛不同水系的拟平鳅的种群分化问题值得进一步探究。     

Genetic and morphological divergence within the Sebastes pachycephalus complex (Scorpaeniformes: Scorpaenidae)

Genetic and morphological divergence among the four subspecies in the Sebastes pachycephalus complex (S. pachycephalus pachycephalus, S. p. nigricans, S. p. nudus and S. p. chalcogrammus) was clarified. Principal coordinate analysis (PCoA) based on AFLP clearly divided 55 specimens of the complex into two groups, the S. p. pachycephalus?CS. p. nigricans group (P-Ni group) and the S. p. nudus?CS. p. chalcogrammus group (Nu-C group), although three specimens occupied intermediate positions. The minimum spanning network (MSN) based on partial sequences of the mitochondrial control region (mtCR) failed to separate either the P-Ni and Nu-C groups or the four subspecies into distinct clades, although restricted gene flow and genetic differentiation between the former were indicated by the F _ST estimation. Differences in morphological characters, including counts of pectoral fin rays and counts of dorsal fin spines lacking basal scales, were also evident between the two groups. However, little or no genetic or morphological difference was found between the two subspecies within each group. It was concluded that the P-Ni and Nu-C groups of the S. pachycephalus complex actually represent two different species, which is further supported by their sympatric distribution. Differences in dorsal body coloration and the presence or absence of brown spots on the ventral surface, which were formerly used to discriminate between four ??subspecies,?? may simply represent intraspecific variation. The three specimens occupying intermediate positions in the AFLP PCoA also occupied equivocal positions between the two species in the principal component analysis (PCA) based on morphometric characters, suggesting that they were hybrids between the two species. The star-shaped MSN of mtCR, which lacks distinct clades representing the two species, may be due to not only interspecific hybridization but also the sharing of ancestral haplotypes.     

Interspecific gene flow in a multispecies oak hybrid zone in the Sierra Tarahumara of Mexico

Background and Aims

Interspecific gene flow can occur in many combinations among species within the genus Quercus, but simultaneous hybridization among more than two species has been rarely analysed. The present study addresses the genetic structure and morphological variation in a triple hybrid zone formed by Q. hypoleucoides, Q. scytophylla and Q. sideroxyla in north-western Mexico.

Methods

A total of 247 trees from ten reference and 13 presumed intermediate populations were characterized using leaf shape variation and geometric morphometrics, and seven nuclear microsatellites as genetic markers. Discriminant function analysis was performed for leaf shape variation, and estimates of genetic diversity and structure, and individual Bayesian genetic assignments were obtained.

Key Results

Reference populations formed three completely distinct groups according to discriminant function analysis based on the morphological data, and showed low, but significant, genetic differentiation. Populations from the zone of contact contained individuals morphologically intermediate between pairs of species in different combinations, or even among the three species. The Bayesian admixture analysis found that three main genetic clusters best fitted the data, with good correspondence of reference populations of each species to one of the genetic clusters, but various degrees of admixture evidenced in populations from the contact area.

Conclusions

The three oak species have formed a complex hybrid zone that is geographically structured as a mosaic, and comprising a wide range of genotypes, including hybrids between different species pairs, backcrosses and probable triple hybrids.     

Aphelenchoides microstylus n. sp. and Seinura onondagensis n. sp. (Nemata: Aphelenchina) from New York

Aphelenchoides microstylus n. sp. and Seinura onondagensis n. sp., a nematode predator, are described from dead Scots pine (Pinus sylvestris L.) in Onondaga County, New York. Females of A. microstylus are 370 to 485 µm long. The body is slender and tapers posteriorly to an amucronate, pointed terminus. The head is continuous with the body, and lips bear a stylet guide. Diagnostic characters of females are three incisures in the lateral field, a short stylet (6-7.5 µm) with small basal knobs, a single row of oocytes, and a long postuterine sac (25-50 µm). Males are characterized by small spicules (10-11µm); two pairs of post-anal, subventral papillae; and a single row of spermatocytes. A bursa and gubernaculum are absent. Seinura onondagensis females are characterized by a body of moderate length (475-595 µm), finely annulated cuticle, and a slightly set-off head. Diagnostic characters are four incisures in the lateral field, long stylet without basal knobs (17-22 µm), single row of oocytes, and presence of a postuterine sac (14-38 µm). Males are unknown. The monospecific genus Indaphelenchus is proposed as a synonym of Seinura, and S. siddiqii n. comb. is proposed for the only species, I. siddiqii.     

Admixture mapping of quantitative traits in Populus hybrid zones: power and limitations

Uncovering the genetic architecture of species differences is of central importance for understanding the origin and maintenance of biological diversity. Admixture mapping can be used to identify the number and effect sizes of genes that contribute to the divergence of ecologically important traits, even in taxa that are not amenable to laboratory crosses because of their long generation time or other limitations. Here, we apply admixture mapping to naturally occurring hybrids between two ecologically divergent Populus species. We map quantitative trait loci for eight leaf morphological traits using 77 mapped microsatellite markers from all 19 chromosomes of Populus. We apply multivariate linear regression analysis allowing the modeling of additive and non-additive gene action and identify several candidate genomic regions associated with leaf morphology using an information-theoretic approach. We perform simulation studies to assess the power and limitations of admixture mapping of quantitative traits in natural hybrid populations for a variety of genetic architectures and modes of gene action. Our results indicate that (1) admixture mapping has considerable power to identify the genetic architecture of species differences if sample sizes and marker densities are sufficiently high, (2) modeling of non-additive gene action can help to elucidate the discrepancy between genotype and phenotype sometimes seen in interspecific hybrids, and (3) the genetic architecture of leaf morphological traits in the studied Populus species involves complementary and overdominant gene action, providing the basis for rapid adaptation of these ecologically important forest trees.     

Recombinational and mutational analyses of subspecific differentiation in Oryza sativa L.

Recombinational and mutational analyses were performed in an attempt to elucidate the nature of the genic differences and the ancestral relationship between the indica and japonica subspecies of Oryza sativa L. by taking representative varieties from each group. The absence of cytological aberrations at meiosis in indica×japonica hybrids suggest that genetic differences are predominantly genic rather than chromosomal. The pattern of variation induced in the key characters, and the frequency and spectra of mutations obtained in each group, failed to support the hypothesis that indica has given rise to japonica, and suggested that the two subspecies have originated from a common ancestor.     

Longidorus paravineacola n. sp. (Nematoda: Longidoridae), a New Species from Arkansas

Longidorus paravineacola n. sp., described herein, was found in a survey of longidorids of Arkansas. It is a parthenogeneticspecies characterized by its long body (6.68-9.85 mm); slightly expanded and rounded head, head width 21-27 µm; odontostyle length 95-114 µm; guide ring 28-37 µm posterior to the head end; short rounded tail, and c'' = 0.6-1.0. Longidorus paravineacola n. sp. is similar to the amphimictic species L. vineacola Sturhan &Weischer, 1964; L. balticus Brzeski, Peneva &Brown, 2000; L. kuiperi Brinkman, Loof &Barbez, 1987; and parthenogenetic species L. crassus Thorne, 1974, which also occurred in the type locality.     

Flow cytometry confirms reticulate evolution and reveals triploidy in Central European Diphasiastrum taxa (Lycopodiaceae, Lycophyta)

Background and Aims

Interspecific Diphasiastrum hybrids have been assumed to be homoploid and to produce well-formed spores serving sexual reproduction. If this were the case, forms intermediate between hybrids and parents or hybrid swarms should be expected. The purpose of this study was: (1) to check whether homoploidy consistently applies to the three hybrids throughout their Central European range; (2) to examine whether their genome sizes confirm their parentage as assumed by morphology; and (3) to perform a screening for detection of ploidy levels other than diploid and variation in DNA content due to backcrossing.

Methods

Flow cytometry was used first to measure the relative DNA values [with 4′,6-diamidino-2-phenylindole (DAPI) staining] and ploidy level as a general screening, and secondly to determine the absolute DNA 2C values [with propidium iodide (PI) staining] in a number of selected samples with the main focus on the hybrids.

Key Results

A considerable variation of DNA 2C values (5·26–7·52 pg) was detected between the three European Diphasiastrum species. The values of the diploid hybrids are highly constant without significant variation between regions. They are also intermediate between their assumed parents and agree closely with those calculated from their putative parents. This confirms their hybrid origin, assumed parentage and homoploid status. Considerably higher DNA amounts (9·48–10·30 pg) were obtained for three populations, suggesting that these represent triploid hybrids, an interpretation that is strongly supported by their morphology.

Conclusions

Diploid hybrids have retained their genetic and morphological identites throughout their Central European range, and thus no indications for diploid backcrossing were found. The triploid hybrids have probably originated from backcrossing between a diploid gametophyte of a hybrid (derived from a diplospore) and a haploid gametophyte of a diploid parental species. By repeated crossing events, reticulate evolution patterns arise that are similar to those known for a number of ferns.     

The Feeding Behavior of Adult Root-knot Nematodes (Meloidogyne incognita) in Rose Balsam and Tomato

Meloidogyne incognita is a parasitic root-knot nematode that causes considerable yield loss in a wide range of plants. In this study we documented the movement of adult female nematodes for more than 2 hr in micro-slices of infected tomato (Solanum lycopersicum) and rose balsam (Impatiens balsamina) plants using light and video microscopy. Stylet thrusting was followed by short pumping actions of the esophagus, dorsal esophageal gland ampulla, and metacorpal bulb. Regular thrusting was normally accompanied by head turning and always preceded continuous stylet thrusting aimed at a single point (for 20 to 90 sec). Females often held the stylet in a protruded position, while pulsating the metacorpus bulb, for about 30 sec. Subsequently, the stylet was paused in a retracted position for 5 to 40 sec. This sequence of behavior took 290 to 380 sec to complete. The procedure developed in this study provides a useful cytological technique to investigate the interaction between root-knot nematodes and the giant cells formed by infected plants. Scanning electron microscopy revealed that the head of the adult nematode was located in the narrow intercellular spaces among several giant cells. The anterior part of the head of the adult was folded like a concertina, whereas that of the second-stage juvenile was not. The labial disc and medial lips of second-stage juveniles seemed expanded and sturdy, whereas those of the adult were star-shaped, appeared to be contracted, and softer. These morphological differences in the heads of adult and second-stage juveniles are discussed with respect to their movement.     

Definition and review of the lancangjiang species group of the termitophilous genus Orthogonius Macleay, 1825 (Coleoptera,Carabidae, Orthogoniini)

The lancanjiang species group of the termitophilous ground beetle genus Orthogonius Macleay, 1825, is defined and reviewed. This group is characterized by the black and rather elongate body, dense punctation on head and elytra, long and slender appendages, thin fore tibiae, and elytral interval 3 without a subapical setiferous pore. To date, the lancanjiang species group is composed of four species and one subspecies, including three new species and one new subspecies which are described in the present paper: O. macrophthalmus sp. n. (northern Vietnam), O. euthyphallus sp. n. (southern Vietnam), O. euthyphallus bolavenensis ssp. n. (southern Laos) and O. carinatus sp. n. (northern Laos). A distribution map and a key to all species of this group are also provided.     

Morphological variation in relation to flower use in bumblebees

To understand resource partitioning in a bumblebee community, we analyzed various morphological characters. A total of 1269 individuals of six bumblebee species, Bombus ardens, B. hypocrita, B. diversus, B. ignitus, B. honshuensis and B. beaticola, were examined and principal component analysis showed that the bumblebee species were clearly differentiated. Glossa, prementa and head lengths were positively correlated with the second component, and a longer proboscis was associated with a narrower body, which may help bees to intrude into and access deep‐lying nectar sources. Bombus diversus, with a long proboscis and narrow body, preferred flowers with a long corolla tube, whereas B. hypocrita and B. ignitus, which have short proboscises and wide bodies, visited flowers with short corollas or dish‐shaped flowers. Two pairs of consubgeneric species that have similar morphological characteristics, B. ardens and B. beaticola, and B. hypocrita and B. ignitus, divided flower resources by habitat selection and seasonal partitioning. For resource partitioning among bumblebee species, not only morphology but also other factors, such as habitat and seasonal preference, flower use, foraging behavior, and interspecific interactions, are responsible.     

Environmental factors, microgeographic patterns of endemism and hybrid zones in Monobella grassei (Insecta: Collembola: Neanuridae)

A complex pattern of narrow parapatric distributions with hybrid zones is described among subspecies of the Collembola species Monobella grassei in the eastern Pyrenees. Though hybrid zones are usually narrow, they vary from simple contact zones without recorded hybrids to relatively wide hybrid swarms, suggesting that large differences exist locally in the level of genetic divergence between subspecies. The course and structure of contact zones are analysed on morphological grounds, in relation to environmental (rainfall, vegetation, substrate) and topographical (altitude, hydrography) parameters. The results show that, unlike most cases documented so far, the courses of the contact zones in M. grassei are not connected to major ecological boundaries nor to ecological ecotones, on either the regional or the habitat scale. It is suggested that this is due to the great ecological similarity between the different parapatric subspecies of M. grassei , and their broad ecological requirements. There is no conclusive evidence that the distribution observed is linked to recent palaeoclimatic events such as glaciation. Endogenous selection could therefore be prevalent in this model to explain parapatry and the hybrid zone pattern.