Courtship in Two Morphotypes of the <Emphasis Type="Italic">Anastrepha fraterculus</Emphasis> (Diptera: Tephritidae) Cryptic Species Complex and their Implications for Understanding Mate Recognition

Ritualized courtship behaviors are used to recognize potential mates and behavioral patterns are inevitably different among populations that demonstrate reproductive incompatibility. We characterized and compared the courtship behaviors of two morphotypes of the cryptic species complex Anastrepha fraterculus: Brazil-1 morphotype and Brazil-3 morphotype. Courtship behaviors were filmed to analyze the behavioral sequences of these two morphotypes during homotypic crossings. The behavioral units Alignment (AL) and Abdominal movements (AB and AB-call) were newly recognized in the courtship ethogram of Anastrepha fraterculus males. The two morphotypes show distinct behavioral sequences leading up to copulation. Some behaviors were repeated frequently during the courtship process, while others were more restricted to the final moments of courtship. The three behavioral units that contributed most to copulation success were Contact, Alignment, and Arrowhead 1 in the Brazil-1 morphotype and Alignment, Arrowhead 1, and Fanning in the Brazil-3 morphotype. Some behavioral routines differed across the two morphotypes. Significant differences were also noted between the frequencies of the behavioral units displayed during courtship in the two morphotypes. The relationships between the pre-zygotic incompatibilities of the Brazil-1 and Brazil-3 morphotypes and the differences between the courtship behaviors of their males are discussed. Our results indicate that behavioral isolation is involved in the process of pre-zygotic reproductive isolation of Brazil-1 and Brazil-3 morphotypes.     

Variability of Acartia clausi in the Black Sea

There are two morphotypes of A. clausi in the Black Sea. The diversity in the spine ornamentation of the last prosomal and urosomal segments generally has been decreasing in both morphotypes during the period of this study (1976–1990). Seasonal data showed increased variability in April.     

The intraspecific differentiation of the burrow tick Ixodes crenulatus (Ixodidae)

In four extensive disjunct areas of the distribution range of Ixodes crenulatus Koch, 1844 complexes of samples in 8 locations, and separate samples in two locations have been studied (fig.). Morphological characters (sizes of organs of idiosome, gnathostoma, legs, as well as some proportions of organs), which show statistically significant differences between complexes of samples on all corresponding stages of ontogenesis were revealed (tabl. 1, 2). Statistically significant differences were determined by Student's criterion (table. 3, 4). We use the term "transit" characters to denote these characters if they have to addition a similar tendency at all stages of ontogenesis. Complexes of samples showing statistically significant differences of transit characters are considered by us as morphotypes. Apparently these differences of morphotypes were formed evolutionally. In European disjunct area morphotypes divided into two groups on the basis of the degree of differences: western (A, B), and eastern (C, D, E). Degree of differences of morphotypes within each group is low, whereas one between of these two groups is high and corresponds to differences of morphotypes A, B from all Asian morphotypes (F, G, H, I, J), and of morphotypes C, D, E from Asian ones in mountain disjunct areas I. crenulatus (G, H, I, J). At the same time Eastern-European complex of morphotypes (C, D, E) is morphologically similar to the morphotype from north Kazakhstan disjunct area (F). All Asian morphotypes have high degree of differences one from the other. Species of the genus Marmota are initial hosts of I. crenulatus, they retain the main role as hosts in Eastern-European and Asian morphotypes: C, D, E, F--M. bobac bobac (Muller, 1776), G--M. baibacina centralis (Thomas, 1909), I, J--M. sibirica sibirica (Radde, 1862). The most aberrant in all characters morphotypes are the ones inhabiting European areas (A, B), where marmots are exterminated. These morphotypes parasitize on hibernating carnivores.     

Confirmation of the species status of the blackfly Simulium galeratum in Britain using molecular taxonomy

Since 1920 Simulium reptans (Linnaeus) (Diptera: Simuliidae) has been reported as exhibiting two different larval morphotypes, a typical S. reptans and an atypical S. reptans var. galeratum, which differ in the markings of the larval head capsule. Inconsistent variation in adults and no apparent variation in the pupae have led taxonomists to conclude that these types in Britain are a single species. We investigated populations in Britain where either the typical form or var. galeratum is found, and one population where the two exist sympatrically. A phylogenetic study based upon a region of the mitochondrial cytochrome c oxidase 1 gene (DNA barcoding) produced a tree that delineated the morphotypes into two distinct monophyletic clades. The average Kimura-2-parameter distances within each clade (i.e. within each morphotype) were very low (0.67% and 0.78%), with the distances between morphotypes being 9-10-fold greater (mean 7.06%). This is concordant with differences within and between species in other taxa; based upon the strict correlation between the molecular variation and the morphotypes, we propose the re-instatement of S. galeratum to species status.     

Variation in foraging behavior facilitates resource partitioning in a polymorphic cichlid, Herichthys minckleyi

We examined foraging behavior (microhabitat use and feeding behavior) in a trophically polymorphic cichlid fish, Herichthys minckleyi, to address several questions regarding resource partitioning in this threatened species. These include: (1) do morphotypes demonstrate different foraging behaviors? (2) do individuals within a morphotype vary in their foraging behavior (e.g. are some individuals specialists, only using a subset of available resources, while other are generalists)? (3) do foraging behaviors vary between isolated pools? (4) do foraging behaviors vary across seasons? We quantified microhabitat use and feeding behavior for over 100 individuals (of two morphotypes) feeding freely in two isolated pools (populations) and across two seasons (winter and summer). We found differences in foraging behavior between morphotypes and individual specializations within morphotypes; i.e. some individuals specialize on certain food resources by using a few feeding behaviors within a subset of microhabitats, whereas others employ a range feeding behaviors across many microhabitats. Foraging behavior also varied between pools and across seasons. This spatial and temporal variation in foraging behavior and resource use may serve to maintain this polymorphism, as the relative fitness of the each morph may vary over space and time.     

Successive changes in the epimural bacterial community of young lambs as revealed by scanning electron microscopy.

Scanning electron microscopy was used to determine the time of initial colonization of the rumen epithelium of young lambs and successive changes with time in the morphological composition of the epimural community. Tissue samples were obtained from two groups of lambs at 1, 2, 4, 6, 8, and 10 weeks of age. Comparisons were made with the epimural communities observed at 12 well-distributed sites in the rumen of a mature wether. Epimural bacteria were already present on the epithelium at 1 week of age. The morphological composition of the epimural community changed with age, with the pattern of succession being similar in both groups of lambs. A total of 24 morphotypes were distinguished by scanning electron microscopy; 17 were rod shaped, 4 were cocci, 2 were spiral, and 1 was filamentous. These morphotypes were further subdivided into: (i) those persisting after their initial colonization in young lambs and present in the adult (7 morphotypes), (ii) those seen only in the adult (2 morphotypes), and (iii) those present only in young lambs (15 morphotypes). The seven morphotypes present in both the lamb and the adult could be considered indigenous members of the epimural community. Several morphotypes appeared restricted in their colonization to certain regions of the papillae, suggesting the presence of microhabitats within the epithelial habitat. Two rod-shaped bacteria were repeatedly seen specifically attached to one another, suggesting an interspecific association.     

Evidence for cryptic hybridization between different evolutionary lineages of the invasive clam genus Corbicula (Veneroida,Bivalvia)

We studied two Corbicula morphotypes in a syntopic population in the Rhine River in order to reveal their taxonomic, reproductive and phylogenetic relationship, using morphometrics, DAF‐fingerprinting, mitochondrial COI and nuclear ITS1 sequence variation. Morphometric analysis showed that two statistically distinguishable morphotypes with few intermediates were present.Mitochondrial sequence analysis detected two divergent clades. DAF‐fingerprinting revealed three highly distinctive multilocus genotypes. Two of the multilocus genotypes were significantly associated with different morphotypes and mitochondrial lineages. The third genotype B, however, was found in both morphotypes, intermediates and mitochondrial lineages. Conclusive evidence for hybridization came from RFLP analysis of the nuclear ITS1 locus. We interpret the hybrids as F1 hybrids between different evolutionary lineages. Integration of Corbicula sequences from all over the world into Maximum Parsimony analysis suggested a simultaneous radiation resulting in several evolutionary lineages whose species status remained doubtful. An unequivocal taxonomic assignment of the two evolutionary lineages in the Rhine population was therefore not possible.     

萼花臂尾轮虫多态性及形态型转化

在实验室条件下研究了萼花臂尾轮虫3种形态型(双短侧棘刺、单短侧棘刺和无侧棘刺)个体之间的关系。从轮虫的培养水体中随机挑出若干3种形态型雌体的幼轮虫(龄长<24h),分别培养在正常培养液和晶囊轮虫培养液过滤液中。将实验轮虫的后代(F1代)置于200倍显微镜下检查其个体的形态型。实验结果表明,在正常培养液中,每种形态型轮虫个体都可以产生3种形态型的后代。当培养在晶囊轮虫培养液过滤液中时,这3种形态型轮虫的后代都具有2个显著增长的侧棘刺。此外,这3种形态型个体的轮虫能够共存于同一培养环境中,而且每种形态型个体的轮虫都具有3种生殖类型的雌体。由此推测萼花臂尾轮虫是一个由3种基础形态型个体组成的形态型轮虫复合体。     

Morphological and genetic differences in ecologically distinct populations of Petrosia (Porifera, Demospongiae)

The taxonomic status of two sponge tnorphotypes living sympatrically in Mediterranean caves and usually ascribed to Petrosia ficiformis was elucidated on the basis of morphological, morphometric and genetic features. The two morphotypes, spherical and cylindrical, showed differences in the shape and size of spicules and in the morphology of the aquiferous system. Electrophoretic analyses demonstrated that the two morphotypes are reproductively independent and so they should be considered as distinct biological species. The spicular features allow attribution of the cylindrical morphotype to P.ficiformis and the spherical one to P. clavata. For these two species parasympatric speciation is proposed.     

Morphological and ecological differences in two electrophoretically detected species of Cliona (Porifera, Demospongiae)

The taxonomie status of different morphotypes generally ascribed to the species of boring sponge Cliona viridis is examined. Morphometric, ecological and genetic differences between two morphotypes living sympatrically on rocky shores of the Ligurian Sea are considered. Morphotype 1 is characterized by small, well separated papillae (α stage), exclusively boring into the crust of coralline algae on the exposed rocky bottoms of the upper sublittoral zone. Morphotype 2 shows bigger papillae, often connected by portions of encrusting tissue ( β stage), and is predominantly found boring into organogenic concretions in slightly lighted habitats. Measurements of tylostyles indicate that morphotype 1 has spicules significantly smaller than morphotype 2. Electrophoretic analysis shows that the two morphotypes are fixed for different alleles at all four loci scored, indicating absence of gene flow between the two populations, i.e. they appear to be distinct biological species.     

Difference in antifungal activity of morphotypes of clavicipitaceous endophytes within and between species

In a previous study, a total of 484 endophytic fungi were isolated and purified from seven populations of Achnatherum sibiricum (L.) Keng collected at six geographical locations in Inner Mongolia, China. Based on growth rates as well as morphological characteristics, the isolates were classified into five morphotypes. Among them, morphotypes A, B and C were ascribed to the same species, Neotyphodium chisosum, based on ITS sequences. Morphotype E was identified as Epichlo? amarillans. In the present study, four morphotypes, A, B, C and E, belonging to two species, were chosen for an in vitro pathogen trial. The results showed that both endophyte colonies and endophyte filtrate of all morphotypes could inhibit the mycelia growth and spore germination of the pathogen fungi tested. The magnitude of inhibition varied not only between species, but also among morphotypes of the same species. Overall, the antifungal ability of E. amarillans (morphotype E) was higher than that of N. chisosum. Within N. chisosum, the antifungal ability was highest in morphotype C, followed by morphotype A, and lowest in morphotype B. This variability suggests that different morphotypes might represent different genotypes of endophyte. The effect of endophyte infection on the host grass should be examined not only on the species level but also on the morphotype level to determine the possible interactions.     

Comparison of ITS RFLP patterns of Gracilaria (Rhodophyceae, Gracilariales) populations from Chile and New Zealand and an examination of interfertility of Chilean morphotypes

Restriction fragment length polymorphism (RFLP) patterns of the internal transcribed spacer (ITS) of the nuclear ribosomal cistron and crossability trials were used to characterize four morphotypes of Gracilaria from Lenga, Isla Santa María and Maullín, Chile, and two morphotypes from sites in New Zealand. PCR products from all Chilean morphotypes resulted in a major single band of ca. 1198 bp. ITS-RFLP profiles generated with the restriction enzymes Cla I, Hae III, Pst I, Hha I, Rsa I and Taq I, were identical in all cases. All crosses within, as well as between, morphotypes resulted in cystocarp differentiation, with the production of viable carpospores. Based upon these data, it is concluded that the four morphotypes from Chile correspond to a single species, G. chilensis, and that the ITS-RFLP pattern is a useful marker to predict genetic relatedness at the specific level in Gracilaria. A comparison of the ITS-RFLP patterns of the Chilean morphotypes with the patterns of two samples of G. chilensis from New Zealand revealed that the sample from Scorching Bay, Wellington, fits the Chilean ITS-RFLP patterns. The population from Blockhouse Bay, Auckland, appears to correspond to another species. This revised version was published online in July 2006 with corrections to the Cover Date.     

Biochemical comparison of two Hypostomus populations (Siluriformes, Loricariidae) from the Atlântico Stream of the upper Paraná River basin, Brazil

Two syntopic morphotypes of the genus Hypostomus - H. nigromaculatus and H. cf. nigromaculatus (Atlantico Stream, Paraná State) - were compared through the allozyme electrophoresis technique. Twelve enzymatic systems (AAT, ADH, EST, GCDH, G3PDH, GPI, IDH, LDH, MDH, ME, PGM and SOD) were analyzed, attributing the score of 20 loci, with a total of 30 alleles. Six loci were diagnostic (Aat-2, Gcdh-1, Gpi-A, Idh-1, Ldh-A and Mdh-A), indicating the presence of interjacent reproductive isolation. The occurrence of few polymorphic loci acknowledge two morphotypes, with heterozygosity values He = 0.0291 for H. nigromaculatus and He = 0.0346 for H. cf. nigromaculatus. F(IS) statistics demonstrated fixation of the alleles in the two morphotypes. Genetic identity (I) and distance (D) of Nei (1978) values were I = 0.6515 and D = 0.4285. The data indicate that these two morphotypes from the Atlantico Stream belong to different species.     

Difference in antifungal activity of morphotypes of clavicipitaceous endophytes within and between species

In a previous study, a total of 484 endophytic fungi were isolated and purified from seven populations of Achnatherum sibiricum (L.) Keng collected at six geographical locations in Inner Mongolia, China. Based on growth rates as well as morphological characteristics, the isolates were classified into five morphotypes. Among them, morphotypes A, B and C were ascribed to the same species, Neotyphodium chisosum, based on ITS sequences. Morphotype E was identified as Epichloë amarillans. In the present study, four morphotypes, A, B, C and E, belonging to two species, were chosen for an in vitro pathogen trial. The results showed that both endophyte colonies and endophyte filtrate of all morphotypes could inhibit the mycelia growth and spore germination of the pathogen fungi tested. The magnitude of inhibition varied not only between species, but also among morphotypes of the same species. Overall, the antifungal ability of E. amarillans (morphotype E) was higher than that of N. chisosum. Within N. chisosum, the antifungal ability was highest in morphotype C, followed by morphotype A, and lowest in morphotype B. This variability suggests that different morphotypes might represent different genotypes of endophyte. The effect of endophyte infection on the host grass should be examined not only on the species level but also on the morphotype level to determine the possible interactions.     

Evidence for Two Morphotypes of Lake Charr, Salvelinus Namaycush, from Great Bear Lake, Northwest Territories, Canada

The lake charr, Salvelinus namaycush, is generally characterized as a morphologically stable specialized deepwater piscivore. Empirical observations suggested the existence of two or more morphotypes of lake charr in Great Bear Lake, NWT. I gathered morphological data over two seasons from a total of 72 individuals and used principal components analysis to summarize variation in the dataset. UPGMA classification showed that there were two groups in the dataset. I classified individuals into one of two morphotypes using discriminant function analysis. The two forms differed primarily in head proportions. I developed a function, usable in the field, that classifies 80% of all specimens correctly. This study represents one of the first documentations of sympatric morphotypes of lake charr outside the Laurentian Great Lakes and provides the basis for further investigation into origins of and genetic, morphological and ecological differences between the forms.     

Characterization and spatial distribution of ectomycorrhizas colonizing aspen clones released in an experimental field

Ectomycorrhizas (EM) from aspen clones released on an experimental field were characterized by morphotyping, restriction analysis and internal transcribed spacer (ITS) sequencing. In addition, their community structure and spatial distribution was analyzed. Among the 23 observed morphotypes, six mycobionts dominated, forming roughly 90% of all ectomycorrhizas: Cenococcum geophilum, Laccaria sp., Phialocephala fortinii, two different Thelephoraceae, and one member of the Pezizales. The three most common morphotypes had an even spatial distribution, reflecting the high degree of homogeneity of the experimental field. The distribution of three other morphotypes was correlated with the distances to the spruce forest and deciduous trees bordering the experimental field. These two patterns allowed two invasion strategies of ectomycorrhizal fungi (EMF) to be recognized, the success of which depends on adaptation of the EMF to local ecological conditions.     

Discrimination of snipefish Macroramphosus species and boarfish Capros aper morphotypes through multivariate analysis of body shape

The existence of two species of the genus Macroramphosus Lacepède 1803, has been discussed based on morphometric characters, diet composition and depth distribution. Another species, the boarfish Capros aper (Linnaeus 1758), caugth along the Portuguese coast, shows two different morphotypes, one type with smaller eyes and a deeper body than the other, occurring with intermediate forms. In both snipefish and boarfish no sexual dimorphism was found with respect to shape and length relationships. However, females in both genera were on average bigger than males. A multidimensional scaling analysis was performed using Procrustes distances, in order to check if shape geometry was effective in distinguishing the species of snipefish as well as the morphotypes of boarfish. A multivariate discriminant analysis using morphometric characters of snipefish and boarfish was carried out to validate the visual criteria for a distinction of species and morphotypes, respectively. Morphometric characters revealed a great discriminatory power to distinguish morphotypes. Both snipefish and boarfish are very abundant in Portuguese waters, showing two well-defined morphologies and intermediate forms. This study suggests that there may be two different species in each genus and that further studies on these fish should be carried out to investigate if there is reproductive isolation between the morphotypes of boarfish and to validate the species of snipefish.     

Differences in frost hardiness of two Norway spruce morphotypes growing at Mt. Brocken, Germany

Norway spruce (Picea abies (L.) Karst.) exhibits strong ecotypic variation along altitudinal gradients in morphological traits, e.g. slenderness of crowns or arrangement of second-order branches. We were interested whether montane and lowland morphotypes differ in a key trait for the survival in cold environments, i.e. frost hardiness, and asked: (i) are montane morphotypes more resistant to frost damage and (ii) do they have a lower risk of frost damage by late frosts in spring than lowland morphotypes?We used the electrolyte leakage-method to measure frost hardiness on a monthly basis from October 2006 to May 2007 in stands of the montane and lowland morphotypes at Mt. Brocken in the Harz Mountains, Germany.LT₅₀ (i.e. the temperature that results in 50% of maximum electrolyte leakage) was assessed by freezing treatments in a frost chamber and was significantly influenced by morphotype, month and minimum ambient temperatures. LT₅₀ was significantly lower in the montane than in the lowland morphotype, with −107 °C and −49 °C, respectively. However, the interactions between morphotype with minimum ambient temperature or month were not significant. Thus, as frost hardiness of the two morphotypes responded to temperature in the same way, both morphotypes can be supposed to be exposed to the same risk of frost damage during hardening in autumn and dehardening in spring.