RESUMO

In insect taxa with homogeneous external morphology, genital structures often emerge as essential traits for interspecific differentiation. In the tribe Ptomaphagini (Coleoptera, Leiodidae, Cholevinae), precise identification often depends on analyzing the male genital morphology, even at the genus level. Here, we present a new character for diagnosing the genera Paulipalpina Gnaspini & Peck, 1996 and Parapaulipalpina Gnaspini, 1996. This feature, which we dub 'paralobe', is a projection arising from the internal surface of the right lobe of the aedeagal apex. Based on its absence in other beetles, including other Ptomaphagini, we recognize it as a putative synapomorphy for those genera. The recognition of this previously overlooked structure adds important information for understanding the sequence of changes that occurred in the male genitalia among the genera of Ptomaphagini.

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Although the recent advances on the relationship of its major groups, the systematics of the rich fauna of Neotropical snakes is far from being a consensus. In this sense, derived groups presenting continental distributions have represented a main challenge. The taxonomy of the snake tribe Echinantherini is one of the most contentious among the diverse family known as Dipsadidae. The tribe is poorly sampled in phylogenetic studies, resulting in conflicting hypotheses of relationships among its taxa. Moreover, several rare and micro endemic species of Echinantherini have never been evaluated within a comprehensive phylogenetic framework. Here, we assess for the first time the phylogenetic position of the rare Echinanthera amoena within Echinantherini. We based our analyses on a comprehensive multilocus dataset including 14 of the 16 species described for the tribe. Our results support the monophyly of Echinantherini and strongly indicate E. amoena as a unique lineage, phylogenetically positioned apart from all other congeners. From the three current genera (Echinanthera, Taeniophallus, and Sordellina) our results indicate that Echinanthera and Taeniophallus are paraphyletic, since the T. affinis species group is positioned as sister to Echinanthera (except E. amoena) clustering apart from the clade formed by the T. brevirostris and T. occipitalis groups. We describe new genera for the T. affinis and T. occipitalis species groups and an additional monospecific genus for E. amoena. Although we did not evaluate the phylogenetic position of T. nebularis, we described a new genus and removed it from Echinantherini since its morphology strikingly departs from all species now included in the tribe. Finally, we redefine the genera Echinanthera and Taeniophallus and we provide comments about further directions to study the biogeography and the evolution of morphological traits in Echinantherini.

RESUMO

Although the recent advances on the relationship of its major groups, the systematics of the rich fauna of Neotropical snakes is far from being a consensus. In this sense, derived groups presenting continental distributions have represented a main challenge. The taxonomy of the snake tribe Echinantherini is one of the most contentious among the diverse family known as Dipsadidae. The tribe is poorly sampled in phylogenetic studies, resulting in conflicting hypotheses of relationships among its taxa. Moreover, several rare and micro endemic species of Echinantherini have never been evaluated within a comprehensive phylogenetic framework. Here, we assess for the first time the phylogenetic position of the rare Echinanthera amoena within Echinantherini. We based our analyses on a comprehensive multilocus dataset including 14 of the 16 species described for the tribe. Our results support the monophyly of Echinantherini and strongly indicate E. amoena as a unique lineage, phylogenetically positioned apart from all other congeners. From the three current genera (Echinanthera, Taeniophallus, and Sordellina) our results indicate that Echinanthera and Taeniophallus are paraphyletic, since the T. affinis species group is positioned as sister to Echinanthera (except E. amoena) clustering apart from the clade formed by the T. brevirostris and T. occipitalis groups. We describe new genera for the T. affinis and T. occipitalis species groups and an additional monospecific genus for E. amoena. Although we did not evaluate the phylogenetic position of T. nebularis, we described a new genus and removed it from Echinantherini since its morphology strikingly departs from all species now included in the tribe. Finally, we redefine the genera Echinanthera and Taeniophallus and we provide comments about further directions to study the biogeography and the evolution of morphological traits in Echinantherini.

RESUMO

Shearwaters are among the most diverse groups of Procellariiformes, and the systematics of genera Calonectris, Ardenna and Puffinus (sensu stricto) is particularly complex. Here we present the first morphology-based phylogenetic hypothesis for the group, using 104 osteological characters (23 continuous and 81 discrete) scored for 20 extant species of shearwaters, representing 62.5% of species currently recognized. Both equal and implied weighting were used for phylogenetic analyses, and the final hypothesis presented herein is the strict consensus of the most stable trees produced by the latter. Calonectris was recovered as monophyletic (Relative Bremer Support = 75), while all other members of shearwaters were included in a single, monophyletic grouping: Puffinus (sensu lato) (RBS = 76). The monophyly of Ardenna, as suggested by all molecular-based hypotheses, was not recovered, whereas a monophyletic Puffinus (sensu stricto) is present within the larger clade. The final hypothesis, wherein shearwaters fall into the sister taxa Calonectris and Puffinus (sensu lato), is consistent with earlier classifications based on osteological evidence.

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Despite descending from heavily calcified ancestors, the holothuroid skeleton is fully internal and composed of microscopic ossicles and a ring of plates bound by connective tissue, the calcareous ring. The calcareous ring exhibits a complex and poorly understood morphology; as a result, establishing unambiguous homology statements about its macrostructure has been challenging and phylogenetic studies have had to simplify this important structure. Here, we provide the first broad comparative study of Dendrochirotida calcareous rings using micro-computed tomography (µCT). A detailed description of the three-dimensional macrostructure of the calcareous ring of 10 sea cucumber species, including rare and type specimens, is presented. The structures observed were highly variable at the subfamily level, especially at the point of tissue attachment. The relationship between the calcareous ring and its associated organs, and their functional morphology are discussed. To aid future phylogenetic studies, we listed 22 characters and performed a preliminary cladistic analysis. The topology obtained supports the idea that the simple, cucumariid ring is ancestral to the mosaic-like phyllophorid ring; however, it did not support the monophyly of the cucumariids. It also did not support the family Sclerodactylidae, which was described based on the ring morphology. Differently from the dermal ossicles, which are highly homoplastic, the general homoplasy index of the calcareous ring characters was relatively low. This result highlights the importance of this structure for phylogenetic inference. Unfortunately, time since collection, rough collection methods and fixation can damage the skeleton, and the calcareous ring is often overlooked in taxonomic descriptions. The data presented here will improve our understanding of holothuroid relationships and facilitate studies on holothuroid functional morphology and biomechanics.

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Tantilla is one of the most diverse genera among colubrids, with 67 species arranged in six phenotypically recognized species groups. Tantilla boipiranga is the most recently described species within the T. melanocephala group, and it was described based on a small type series, collected in the rupestrian grasslands of south-eastern Brazil. The morphological diversity and the phylogenetic affinity of this species remain poorly known. Here, based on the assessment of recently collected specimens in combination with results of a molecular phylogenetic analysis, we evaluate the morphological variation within T. boipiranga and its phylogenetic position. Our analyses confirm T. boipiranga as genetically distinct from its generic counterparts, and diagnosable based on a combination of colour pattern, meristic counts, and hemipenial morphology. However, contrary to its original description, the hemipenial ornamentation does not differen-tiate T. boipiranga from the highly variable T. melanocephala. In our phylogenetic analysis, T. boipiranga is retrieved as a monophyletic group, nested within the diversity of T. melanocephala and sister to a clade composed by specimens from south-eastern Brazil. Tantilla melanocephala is recovered as a highly diverse lineage, indicating the possible presence of undescribed species. Additionally, our analysis indicates that the T. coronata and T. planiceps species group are mono-phyletic, while the T. taeniata species group is paraphyletic and the only sampled species for the T. calamarina group is nested within the T. melanocephala group. Our results suggest that the phenotypic evolution within the genus is probably more complex than previously recognized, and some colouration patterns used to define the groups can in fact represent phenotypical convergences. Moreover, our phylogenetic analysis suggests a strong signal of geographical structure in the tree topology. Three main geographical lineages were found, a North American, a South American and a Central Ameri-can. The first two lineages were recovered as monophyletic, while the latter is paraphyletic, with Central American species positioned as sister groups of both, the North American and the South American clades. Although a more comprehensive phylogenetic analysis is needed, our study strongly indicates the existence of hidden diversity within the T. melanocephala group and that Tantilla represents an ideal model to evaluate the validity of phenotypical groups in snake systematics and to study the driven mechanisms of morphological evolution.

RESUMO

The monotypic genus Metasiphonella, originally described from Costa Rica, is now formally recorded from South America with a new species from Brazil-M. amorimi sp. nov. The taxonomic revision includes a detailed description of adult external morphology, female and male terminalia, and the egg stage for all species. A lectotype of M. magnifica Duda is designated and new country records are given.

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Snakes have high morphological variation, including between sexes. Most of these variations are evidenced only by linear measurements, which are generally restricted to size traits. Using traditional and geometric morphometrics methods we analyze how body size attributes and head size and shape varies between sexes and species of two sympatric Tachymenini snakes, Thamnodynastes strigatus and Tomodon dorsatum. We took body size measurements from 87 specimens of T. strigatus and 83 specimens of T. dorsatum. We also extracted head shape and size of 146 specimens of T. strigatus and 156 specimens of T. dorsatum. Our results showed sexual differences in some body size attributes and in head shape and size. Females of both species had larger SVL, BM, HW, BW and heads than in males. But, T. strigatus is not sexually dimorphic in head size. Females of the T. strigatus also exhibited an enlargement of the parietal region compared to the males, while T. dorsatum females exhibited a wider snout than males. These sexual differences could be a result of fecundity selection and may also indicate that females consume larger preys than conspecific males. At the species level, T. strigatus is larger in body and head size than T. dorsatum. The head shape of T. strigatus is characterized by a more elongated head with larger frontal and prefrontal scales and slender snout compared to T. dorsatum. Shape differences between these two snakes probably evolved in association with divergences in habitat use e.g. terrestrial and arboreal and diet specialization e.g. piscivory, and malacophagy.

RESUMO

Metachromadora (Bradylaimoides) benepapillata, was first discovered from the coastal mangroves of Bangladesh by Timm (1961). This species was later synonymised with Pseudochromadora cazca, which was described from the Brazilian coast by Gerlach (1956). The present paper proposes to update the taxonomic status of the Timm's species as Pseudochromadora benepapillata comb. n. along with its distribution and comparative morphology. The major differences between P. cazca and P. benepapillata are found in the characteristics of the head capsule; length of cephalic sensilla; length and position of somatic setae; shape of amphideal fovea; shape of spicules and gubernaculum; number and morphology of copulatory and postcloacal thorns. The present report on the occurrence of this species along the eastern most coast of India extends its distribution from the eastern part of Bay of Bengal to its western part.

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Callichthyidae comprises the subfamilies Callichthyinae and Corydoradinae, both of which are morphologically distinct and monophyletic. Although there is consensus regarding the monophyly of the family, the relationships of about 80% of its species, currently included in the genus Corydoras, remain poorly known. Despite the vast amount of osteological information for Teleostei, knowledge regarding the phylogenetic implications of encephalon anatomy is sparse and represents a poorly explored source of potential characters. The present study aims to describe the encephalon morphology in members of the Callichthyidae in order to propose new characters that may help address phylogenetic questions regarding this group. In addition to representatives of Callichthyidae, specimens belonging to the Nematogenyidae, Trichomycteridae, Scoloplacidae, Astroblepidae and Loricariidae were dissected for comparative purposes. Head dissection revealed information on the structure of the medulla spinalis, rhombencephalon, mesencephalon, diencephalon and telencephalon. The conditions observed on the encephalons examined suggest that representatives of Callichthyidae have great taste perception and processing, while Corydoradinae stand out for visual acuity and Callichthyinae for mechanoreception processing subunits. Our results also indicate that the encephalon has important features for systematic studies of the family bringing greater resolution to current phylogenetic hypotheses.(AU)

Callichthyidae é composto por Callichthyinae e Corydoradinae, ambos morfologicamente distintos e monofiléticos. Apesar do consenso em relação ao mofiletismo da família, as relações de cerca de 80% de suas espécies, atualmente incluídas no gênero Corydoras, permanecem pouco conhecidas. Apesar da grande quantidade de informação osteológica sobre Teleostei, o conhecimento sobre as implicações filogenéticas da anatomia do encéfalo é escasso e, por isso, considerado uma fonte inexplorada de caracteres. O objetivo do presente estudo é a descrição morfológica dos encéfalos de Callichthyidae, fornecendo novos caracteres que podem elucidar questões filogenéticas para o grupo. Além dos representantes de Callichthyidae, espécimes pertencentes a Nematogenyidae, Trichomycteridae, Scoloplacidae, Astroblepidae e Loricariidae foram dissecados para fins comparativos. A dissecção do crânio revelou informações sobre a estrutura da medulla spinalis, rhombencephalon, mesencephalon, diencephalon e telencephalon. As condições observadas nos encéfalos sugerem que representantes de Callichthyidae possuem grande capacidade de percepção e processamento químico, enquanto os Corydoradinae se destacam pela acuidade visual e os Callichthyinae pelas unidades de processamento mecanoreceptoras. Nossos resultados indicam que os encéfalos detêm características importantes para contribuir com estudos sobre a sistemática da família, trazendo maior resolução para as hipóteses atuais de reconstrução filogenética.(AU)

Assuntos

RESUMO

Callichthyidae comprises the subfamilies Callichthyinae and Corydoradinae, both of which are morphologically distinct and monophyletic. Although there is consensus regarding the monophyly of the family, the relationships of about 80% of its species, currently included in the genus Corydoras, remain poorly known. Despite the vast amount of osteological information for Teleostei, knowledge regarding the phylogenetic implications of encephalon anatomy is sparse and represents a poorly explored source of potential characters. The present study aims to describe the encephalon morphology in members of the Callichthyidae in order to propose new characters that may help address phylogenetic questions regarding this group. In addition to representatives of Callichthyidae, specimens belonging to the Nematogenyidae, Trichomycteridae, Scoloplacidae, Astroblepidae and Loricariidae were dissected for comparative purposes. Head dissection revealed information on the structure of the medulla spinalis, rhombencephalon, mesencephalon, diencephalon and telencephalon. The conditions observed on the encephalons examined suggest that representatives of Callichthyidae have great taste perception and processing, while Corydoradinae stand out for visual acuity and Callichthyinae for mechanoreception processing subunits. Our results also indicate that the encephalon has important features for systematic studies of the family bringing greater resolution to current phylogenetic hypotheses.(AU)

Callichthyidae é composto por Callichthyinae e Corydoradinae, ambos morfologicamente distintos e monofiléticos. Apesar do consenso em relação ao mofiletismo da família, as relações de cerca de 80% de suas espécies, atualmente incluídas no gênero Corydoras, permanecem pouco conhecidas. Apesar da grande quantidade de informação osteológica sobre Teleostei, o conhecimento sobre as implicações filogenéticas da anatomia do encéfalo é escasso e, por isso, considerado uma fonte inexplorada de caracteres. O objetivo do presente estudo é a descrição morfológica dos encéfalos de Callichthyidae, fornecendo novos caracteres que podem elucidar questões filogenéticas para o grupo. Além dos representantes de Callichthyidae, espécimes pertencentes a Nematogenyidae, Trichomycteridae, Scoloplacidae, Astroblepidae e Loricariidae foram dissecados para fins comparativos. A dissecção do crânio revelou informações sobre a estrutura da medulla spinalis, rhombencephalon, mesencephalon, diencephalon e telencephalon. As condições observadas nos encéfalos sugerem que representantes de Callichthyidae possuem grande capacidade de percepção e processamento químico, enquanto os Corydoradinae se destacam pela acuidade visual e os Callichthyinae pelas unidades de processamento mecanoreceptoras. Nossos resultados indicam que os encéfalos detêm características importantes para contribuir com estudos sobre a sistemática da família, trazendo maior resolução para as hipóteses atuais de reconstrução filogenética.(AU)

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Heteroptera have diverse feeding habits with phytophagous, zoophagous, and haematophagous species. This dietary diversity associated with the monophyly of Heteroptera makes these insects a good object for comparative studies of the digestive tract. This work compares the ultrastructure of the middle midgut region in the phytophagous Coptosoma scutellatum (Plataspidae), Graphosoma lineatum (Pentatomidae), Kleidocerys resedae (Lygaeidae), and zoophagous Rhynocoris iracundus (Reduviidae), Nabis rugosus (Nabidae), and Himacerus apterus (Nabidae), to verify if diet affects midgut cells in phylogenetically related insects. The middle region of the midgut was used for comparison because it is the main site for digestion and absorption of the midgut. The digestive cell ultrastructure was similar in the six species, with features of secretory, absorptive, transport, storage, and excretory cells, suggesting a stronger correlation of middle digestive cell ultrastructure with the phylogeny of these species than with the different heteropteran feeding habits.

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The objective of this study focused on the comparative morphological and histochemical analysis of the duodenum of fetuses, juveniles and adult coypu (Myocastor coypus bonariensis), the major socioeconomic wildlife resource of Argentina. Histological and histochemical procedures for in situ characterization of glycoconjugates (GCs) were used. This study evidenced that fetal mucins differ histochemically in many respects from their adult counterparts. Only in fetuses from 90 days-post coitus (dpc) glycogen-rich sites were observed throughout the duodenal epithelium. The goblet cells appeared from 105 dpc and their secretory content varied considerably after birth. Duodenal glands presented scanty neutral and sulphated GCs in the 30-day juveniles; in adults the proportion of these GCs increased, and carboxylated and sialylated GCs were also observed. The results obtained in this work may be used in future studies to evaluate the effects of diet and intestinal pathologies in the glycosylation pattern of GCs. Also, knowledge of the normal glycoprofile of the duodenum of M. coypus bonariensis during its ontogenetic development may constitute a basis for the study of this organ in other Hystricognathi rodents.

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RESUMO

Among arthropods, ticks of the genus Amblyomma are of great medical and veterinary importance and present phylogenetic and taxonomic divergences given polymorphisms and phenotypic plasticity between subpopulations. Generally, the male reproductive system and spermatozoon exhibit diversified morphology and ultrastructure species-specific, bringing new possibilities for phylogenetic and taxonomic issues. Therefore, the present study aimed to describe and compare the morphology of the male reproductive system and its germ cells of Amblyomma aureolatum, A. sculptum, and A. triste, intending to identify possible diagnostic features. Couples of the three tick's species were kept in colony, infested on rabbits and collected over 12 days of feeding. The males had their reproductive systems dissected, fixed and processed for histology and scanning electron microscopy. The results obtained here allowed the description of spermiogenesis stages and the comparison of spermatids morphology in the last stage of development. Furthermore, the testis of A. triste present an isthmus connecting the distal region of both, while in the other two species this structure could not be observed. Some anatomical features were identified which can be used for taxonomic and phylogenetic studies, like the presence or absence of the isthmus connecting testis, spV cell shape, the shape of the operculum and the presence or absence of the rim on its base.

RESUMO

The brain of Brycon orbignyanus is described as a model for future studies of the gross morphology of the central nervous system in Characiformes. The study of brain gross morphology of 48 distinct taxa of Characiformes, one of Cypriniformes, two of Siluriformes and two of Gymnotiformes, allowed us to propose, for the first time, six putative brain synapomorphies for the Characiformes and also two possibly unique gross brain morphology characters for the Siluriformes. A detailed protocol for the extraction of the brain in Characiformes is also provided.(AU)

O encéfalo de Brycon orbignyanus é descrito como um modelo para futuros estudos da anatomia externa do Sistema Nervoso Central de Characiformes. O estudo da morfologia externa de 48 táxons distintos de Characiformes, um de Cypriniformes, dois de Siluriformes e dois de Gymnotiformes, permitiu-nos propor, pela primeira vez, seis prováveis sinapomorfias encefálicas e também duas possíveis características encefálicas para Siluriformes. Um protocolo detalhado para a dissecção e extração do encéfalo de Characiformes é também apresentado.(AU)

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RESUMO

The brain of Brycon orbignyanus is described as a model for future studies of the gross morphology of the central nervous system in Characiformes. The study of brain gross morphology of 48 distinct taxa of Characiformes, one of Cypriniformes, two of Siluriformes and two of Gymnotiformes, allowed us to propose, for the first time, six putative brain synapomorphies for the Characiformes and also two possibly unique gross brain morphology characters for the Siluriformes. A detailed protocol for the extraction of the brain in Characiformes is also provided.(AU)

O encéfalo de Brycon orbignyanus é descrito como um modelo para futuros estudos da anatomia externa do Sistema Nervoso Central de Characiformes. O estudo da morfologia externa de 48 táxons distintos de Characiformes, um de Cypriniformes, dois de Siluriformes e dois de Gymnotiformes, permitiu-nos propor, pela primeira vez, seis prováveis sinapomorfias encefálicas e também duas possíveis características encefálicas para Siluriformes. Um protocolo detalhado para a dissecção e extração do encéfalo de Characiformes é também apresentado.(AU)

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RESUMO

The gross morphology of the brain of the pseudopimelodid Pseudopimelodus bufonius is described and compared with congeners. Observations were made on removed brains after elimination of bones from the top of the skull and severing of the cranial nerves and the spinal cord. Nine morphometric characters associated with the major subdivisions of the brain were identified, seven of which revealed significant differences among the species examined. The corpus cerebelli in all examined species of the genus is the largest structure of the brain. The behavior of the species of Pseudopimelodus is still unknown, but in other teleosts that condition is typically correlated with a higher degree of motor coordination. Relative size proportions of the tectum opticum, eminentia granularis, lobus facialis and lobus vagi, might be related to carnivory and an enhanced capacity for food selection.

A morfologia externa do encéfalo de Pseudopimelodus bufonius é descrita e comparada com seus congêneres. As análises foram feitas no cérebro removido após a eliminação dos ossos do topo da cabeça e secção dos nervos cranianos e cordão espinhal. Nove caracteres morfométricos foram obtidos das principais subdivisões do encéfalo, dos quais em sete ocorreram diferenças significativas entre as espécies. Em todas as espécies examinadas do gênero o corpus cerebelli é a maior estrutura do encéfalo. O comportamento das espécies de Pseudopimelodus ainda é desconhecido, mas em outros teleósteos esta característica é normalmente correlacionada com uma boa coordenação motora. Além disso, as proporções relativas do tectum opticum, eminentia granularis, lobus facialis e lobus vagi podem ser relacionadas a hábitos carnívoros e boa capacidade de selecionar alimentos.

Assuntos

RESUMO

The gross morphology of the brain of the pseudopimelodid Pseudopimelodus bufonius is described and compared with congeners. Observations were made on removed brains after elimination of bones from the top of the skull and severing of the cranial nerves and the spinal cord. Nine morphometric characters associated with the major subdivisions of the brain were identified, seven of which revealed significant differences among the species examined. The corpus cerebelli in all examined species of the genus is the largest structure of the brain. The behavior of the species of Pseudopimelodus is still unknown, but in other teleosts that condition is typically correlated with a higher degree of motor coordination. Relative size proportions of the tectum opticum, eminentia granularis, lobus facialis and lobus vagi, might be related to carnivory and an enhanced capacity for food selection.(AU)

A morfologia externa do encéfalo de Pseudopimelodus bufonius é descrita e comparada com seus congêneres. As análises foram feitas no cérebro removido após a eliminação dos ossos do topo da cabeça e secção dos nervos cranianos e cordão espinhal. Nove caracteres morfométricos foram obtidos das principais subdivisões do encéfalo, dos quais em sete ocorreram diferenças significativas entre as espécies. Em todas as espécies examinadas do gênero o corpus cerebelli é a maior estrutura do encéfalo. O comportamento das espécies de Pseudopimelodus ainda é desconhecido, mas em outros teleósteos esta característica é normalmente correlacionada com uma boa coordenação motora. Além disso, as proporções relativas do tectum opticum, eminentia granularis, lobus facialis e lobus vagi podem ser relacionadas a hábitos carnívoros e boa capacidade de selecionar alimentos.(AU)

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The present study compares the morphology of the head, appendages, and cervical region of three species of the butterflies Archaeoprepona demophon demophon (Linnaeus, 1758), Archaeoprepona licomedes licomedes (Cramer, 1777), and Prepona pylene pylene Hewitson, [1854], through descriptions, illustrations, and scanning electron micrographs. The results are compared with Prepona claudina annetta (Gray, 1832), Memphis moruus stheno Hübner, [1819], and Zaretis itys itylus (Westwood, 1850), showing unique characteristics for each species and/or genus, or characteristics shared among the species analyzed. The detailed morphology of these three species was previouslyunknown.

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RESUMO

The present report, the second part of a study of the external morphology of Preponini, compares the thorax and thoracic appendages of Archaeoprepona demophon demophon (Linnaeus, 1758), Archaeoprepona licomedes licomedes (Cramer, 1777) and Prepona pylene pylene Hewitson, [1854], through descriptions and illustrations. The results are compared with three other species, Prepona claudina annetta (Gray, 1832), Memphis moruus stheno Hübner, [1819] and Zaretis itys itylus (Westwood, 1850), revealing previously unrecognized similarities among species of Charaxinae.

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