Patterns and causes of morphological population differentiation in the Tenerife skink, Chalcides viridanus

Recent studies indicate that differential selection between mesic and xeric habitats is the most plausible explanation of within-island geographic variation in the Gran Ganarian skink, Chalcides sexlineatus. The island of Tenerife shows mesic/xeric heterogeneity similar to that on Gran Canaria, so we tested the prediction of parallel geographic patterns of morphological population differentiation in the Tenerife skink, Chalcides viridanus. Geographic variation was found to be complex; patterns of differentiation in body dimensions and scalation show evidence of both mosaic and latitudinal facets. Using randomization and regression methods, significant relationships were found between a hypothesized xeric/mesic model of population differentiation and geographic variation in female body dimensions, as well as with substantial proportions of the geographic variation in male and female scalation and, to a lesser extent, body dimensions. Matrix comparisons based on the entire morphological distance matrices show that the patterns of variation are not concordant with an alternative hypothesis of historical secondary contact, or with an isolation-by-distance model. It is argued that a substantial component of the geographic variation in body dimensions and scalation of C. viridanus is due to differential selection along an aridity gradient.     

Highly polymorphic microsatellite loci for the Gran Canarian skink (Chalcides sexlineatus) and their applicability in other Canarian Chalcides

The scincid lizard Chalcides sexlineatus is endemic to the island of Gran Canaria, within which it shows a pronounced phylogeographical pattern. Understanding the maintenance of this pattern requires knowledge of nuclear DNA structuring. Eleven polymorphic tetranucleotide microsatellite loci were isolated from a library constructed from genomic DNA enriched for AAAG repeat sequence. The number of alleles per locus ranged from 12 to 19, and expected heterozygosities spanned from 0.738 to 0.933 (n = 30). The majority of these could be cross‐amplified in other endemic Canarian Chalcides. The loci will be used to investigate gene flow between the principal mitochondrial clades within C. sexlineatus.     

Within-island geographic variation of the dangerous Taiwanese snake, Trimeresurus stejnegeri, in relation to ecology

Multivariate analysis shows that geographic variation occurs in the scalation and body proportions of the dangerously venomous snake Tritneresurus stejnegeri , within the island of Taiwan. Canonical correlation analysis reveals a high correlation between constellations of morphological characters and ecological factors. Numerical hypothesis testing, using partial Mantel tests, provides evidence of a causal relationship with ecology. Head shape was found to be primarily associated with mean annual temperature and altitude, and head scalation with the annual range of temperature. Body scalation was found to be influenced by altitude, mean annual temperature and mean annual rainfall. Tail length is primarily associated with the range of temperature and rainfall and secondarily with the mean annual temperature and altitude. Geographic proximity was found to be less important in the explanation of the observed geographic pattern than some ecological factors, supporting the hypothesis of an ecogenetic origin of morphological variation.     

Phenotypic variation in colour pattern and seasonal plasticity in Eristalis hoverflies (Diptera: Syrphidae)

Abstract.

• 1 An examination of phenotypic variation in colour pattern was carried out on four Eristalis hoverfly species using museum material.
• 2 The amount of phenotypic variation varied substantially among the species with E.arbustorum being the most variable. The other species showed a wide colour pattern range but less variation within that range (E.abusivus and E.nemorum), or a narrow range of colour variation (E.horticola).
• 3 Sexual colour dimorphism was apparent in all four species, but most pronounced in E.abusivus and E.nemorum.
• 4 There were good phenotype-season relationships shown by both sexes in all species, except for female E.abusivus and E.nemorum, with paler insects being more abundant during the warmer summer months.
• 5 Female, but not male, E.arbustorum collected at inland sites were on average paler than those collected at coastal sites. This observation is considered with respect to temperature during the developmental stages.
• 6 The function of colour plasticity in hoverflies is discussed with reference to the need to maintain optimal thermal conditions for activity.

     

Growth and carapacial colour variation of the leopard tortoise, Geochelone pardalis babcocki, in northern Tanzania

Observations were made of growth rate and variation in colour pattern of the leopard tortoise ( Geochelone pardalis babcocki ) in northern Tanzania between October 1993 and June 1996. Growth rate differed significantly between tortoises, with immature animals (6.9 ± 1.4 mm month⁻¹) and females (5.7 ± 1.3 mm month⁻¹) exhibiting higher rates of growth than adult animals (2.9 ± 0.8 mm month⁻¹) and males (3.2 ± 1.1 mm month⁻¹), respectively. The faster growth rate in immature animals suggested that growth rate declined with age. The difference in growth rates between the sexes was ascribed to sexual dimorphism, adult females being larger and therefore growing faster than males.
The carapacial colour pattern observed in the northern Tanzanian tortoises was generally similar to that described elsewhere in the species' range. The colour pattern varied between Arusha and Serengeti District tortoises. Animals at the Arusha sites tended to be sparsely coloured (55.6%) compared to the Serengeti tortoises, which had a buff colouration with radiating spots (89.3%).; The sparse colouration provides effective camouflage in the drier Arusha habitats, as does the buff colouration with black spots in the wetter, darker habitats in the Serengeti.     

An analysis of the phenotypic effects of certain colour pattern genes in Heliconius (Lepidoptera: Nymphalidae)

The colour patterns of Heliconius butterflies are built up from an array of serially homologous pattern elements known as the nymphalid groundplan. An analysis of the phenotypic effects of ten genetic loci from H. melpomene and H. cydno reveals that each alters the expression either of a single element of the groundplan or of an entire row of serially homologous elements. Five of the ten loci affect the size (or presence/absence) of specific pattern elements, two affect the colour in which a pattern element is expressed, two affect pattern-inducing activity of the wing veins, and one appears to affect an overall threshold for pattern determination. Three of the ten loci have identical effects on homologues of the fore- and hindwing. We show that most of the apparently large and qualitative phenotypic effects of these genes can be readily explained by relatively small and quantitative changes in the dimensions or positions of specific pattern elements.     

Geographic variation for colour in the sandburrowing beetle Chaerodes trachyscelides White (Goleoptera: Tenebrionidae) on New Zealand beaches analysed using CIELAB L * values

Chaerodes trachyscelides White is a highly specialized, flightless burrowing beetle confined to the narrow strip of sand at and just above high water level on sandy marine beaches in New Zealand. Although the ventral surface of the beetle is always pale, the dorsal surface varies from pale to almost black. Large samples of this beetle were taken, together with the sand, from 11 beaches on New Zealand's three main islands. The colour of the dorsal surface of each individual beetle and that of the sand samples was measured using reflectance spectroscopy and expressed as CIE L*,a*,b* (CIELAB) values. The L* values, which are objective, quantitative measures of the degree of lightness of the beetles, were subjected to statistical and frequency analysis. Although the species was very variable in colour and the variation appeared to be continuous, a highly significant correlation was obtained between the mean of the L* values for the samples on each beach and that of the sand, the correlation coefficient being 0.961. This close association between the lightness of the beetles and that of the sand suggests the variable melanism functions as cryptic colouration. On most beaches, the distribution oflightness among the beetles sampled conformed to a normal curve. For beetles from sites where the sand was relatively uniform, such as the black Taranaki beaches, the L* frequency distribution curves were narrow and the coefficient of variation of mean beetle colour was relatively small indicating low colour variability. In contrast, the greatest within-site variability occurred on the two Stewart Island beaches sampled, where in each case there was less uniformity in the colour of the sand. At one of these sites, Maori Beach, darker sand present below the high water level is often deposited on the zone occupied by the beetles after storms. On Lonneker's Beach, the distribution of L* values among the beetles sampled was actually bimodal. On this small beach, there was an area of intensely black sand in the zone occupied by the beetles, but most of the rest was covered with light golden sand. These results are interpreted as evidence that the variability of colour of Chaerodes beetles has the effect of populations being able to match the colour of the sand of their home beaches, presumably as a consequence of the differential survival of individuals.     

The importance of the fixation of colour,pattern and form in tropical Pseudocerotidae (Platyhelminthes,Polycladida)

At last a fixation method that ensures tropical pseudocerotid polyclads are fixed flat, preserved for histological preparation and which also retains their colour pattern has been developed. FCA-PGPP (Formaldehyde Calcium Acetate-Propylene Glycol, Propylene Phenoxetol) fixative is frozen and worms are coaxed onto filter paper which is then laid on the frozen fixative. As a consequence, over 230 species have been documented from the southern Great Barrier Reef and eastern Papua New Guinea (Newman & Cannon, 1994; in press). It was determined that species diagnoses need to be based on colour pattern, general morphology of living animals and serial reconstructions of the male anatomy.Christensen Research Institute Contribution No. 82     

Homologies in the colour patterns of the genus Heliconius (Lepidoptera: Nymphalidae)

The colour patterns of Heliconius butterflies are composed from a relatively simple set of pattern elements whose homologues are recognizable throughout the genus. Although Heliconius colour patterns look quite different from those of most nymphalids, these pattern elements are seen to derive from the generalized nymphalid groundplan. The differences arise primarily from the loss or positional shift of certain pattern elements, a high degree of fusion between individual pattern elements, and, in the forewing, asymmetries of the pattern elements relative to the wing-cell midline. The scheme of homologies we present is consistent with what is currently known about the comparative morphology and developmental physiology of colour pattern formation in Lepidoptera, and provides a framework for the interpretation of developmental, evolutionary and genetic studies in Heliconius.     

Enzyme and colour variation in the hoplonemertean Tetrastemma nigrifrons from the Sea of Japan

Five sympatric colour varieties of the hoplonemertean Tetrastemma nigrifrons (var. pallidum, var. bicolor, var. purpureum, var. punctata, var. albino) were found in Vostok Bay (the Sea of Japan). The taxonomic status of the varieties, some of which have been known for almost a century, was uncertain. Starch gel electrophoresis was used to compare allele frequencies of these varieties at 19 isozyme loci. Neither fixed allelic differences nor statistically significant differences in allele frequencies between different colour varieties was revealed. It was concluded therefore that all the colour varieties studied are conspecific. As estimated from the electrophoretic screening of 24 isozyme loci, the species is highly variable at the molecular level. It has high mean values of observed and expected heterozygosity estimates, H _o =0.313 ± 0.049 and H _e =0.323 ± 0.050, and is one of the most variable species among invertebrates.     

The environmental control of colouration in a bushcricket, Mygalopsis marki Bailey (Orthoptera: Tettigoniidae)

The environmental factors proximately influencing variable colouration in the bushcricket, Mygalopsis marki , were studied in natural populations and under controlled laboratory conditions. Two major components of variable colouration were identified; a green/brown dimorphism in ground colour hue (dominant wave length) and continuous variation in the darkness of brown insects, representing a contrast between ground colour value (brightness) and the intensity of overlying black pattern. Ground colour hue differed between natural populations in different seasons and at different localities. The proportion of brown individuals was greater in populations developing over late summer than over spring at the same locality, and in populations from drier localities, as indicated by vegetation type, in the same season. Experimental studies showed that colour change occurred at moults, with green colouration promoted by increased water intake in developing nymphs. Ground colour value and pattern differed between natural populations at different localities, but not in different seasons. Ground colour value was less, and pattern was greater, in populations from localities with a dense cover of forbs and grasses. Experimental studies showed that ground colour value and pattern were influenced by the intensity of incident radiation received by developing nymphs, with decreased intensity promoting lower value and greater pattern.     

Sexual selection mediated by the thermoregulatory effects of male colour pattern in the ambush bug Phymata americana

Sexual dimorphism in coloration is a taxonomically widespread phenomenon often attributed to sexual selection on visual signals. However, the ambush bug Phymata americana exhibits sexual dimorphism in coloration that has no apparent signalling function. Here we provide evidence that colour pattern in this species influences male mating success indirectly through its effect on thermoregulation. We demonstrate, using experimental manipulation, that individuals with dark colour pattern achieve higher thoracic temperatures under illumination. We also show that dark colour pattern predicted mate-searching success but only under thermally challenging conditions (i.e. cool ambient temperature). As far as we are aware, this is the first study to provide evidence that sexual dimorphism can be accounted for by sexual selection on thermoregulatory performance.     

Nymphal colour/pattern polymorphism in the leafhoppers Eupteryx urticae (F.) and E. cyclops Matsumura (Hemiptera: Auchenorrhyncha): spatial and temporal variation in morph frequencies

Variation in colour/pattern morph frequencies in Eupteryx urticae and E. cyclops is described for various field populations. Eupteryx urticae populations in S Wales exhibit a steep morph-ratio cline, such that black morph frequencies are positively correlated with altitude. High melanic frequencies at high-altitude sites, and the absence of the two darker morphs in lowland populations, suggest a similar trend in E. cyclops , but the data are insufficient to confirm this statistically. No differences in morph frequencies were detected on different parts of the primary host plant or on alternative host species. Similarly, there were no consistent trends within or between the two annual generations of either species, although melanic morph frequencies in one E. urticae population were heterogeneous over 10 generations. It is suggested that the polymorphism in E. urticae is strongly influenced by climate selection, darker morphs being at an advantage in cooler environments where their coloration enhances absorption of solar radiation. The advantage gained through thermal melanism is probably balanced by visual selection against black morphs by entomophagous parasitoids.     

Homologies in the colour patterns of the genus Charaxes (Lepidoptera: Nymphalidae)

The phylogenetically and morphologically diverse patterns of Charaxes can be reduced to a simple set of pattern elements which can be homologized throughout the genus. At least five types of correspondence (homologies) exist among pattern elements: those between (1) species, (2) forewing and hindwing, (3) dorsal and ventral wing surface, (4) serial wing-cells, and (5) individual pattern elements within a single wing-cell. Differences in Charaxes colour patterns result from the distortion, elaboration, enlargement, reduction or loss of individual pattern elements. Further variation is often the result of dislocation of pattern elements from their serial homologues in neighbouring wing-cells, and fusion of individual pattern elements to create larger areas of colour. The type of analysis presented in this paper should be broadly applicable within the Lepidoptera and may prove useful in studying the systematics of colour patterns and the evolution of the developmental system that gives rise to them.