Comparative morphology of mandibular structures in lycid larvae and its phylogenetic implications (Polyphaga, Hexapoda)

The morphology of the larval mandibular structures of the family Lycidae (Polyphaga) is characterized by facultative dicondyly, posterior articulation with long mandibular rods, lateral location of the anterior condyle and its articulation with a paired non‐cranial structure. It is compared to that of Eucrustacea, Chilopoda, Entognatha, Microcoryphia and Zygentoma and found to be more reminiscent of the Entognatha. The phylogenetic implications of this conclusion are discussed, with the Pterygota and Dicondylia hypothesized to be non‐monophyletic.     

The controversial origin of the abdominal appendage‐like processes in immature insects: Are they true segmental appendages or secondary outgrowths? (Arthropoda hexapoda)

In this article, I review the major characteristics of different types of appendage‐like processes that develop at the abdominal segments of many immature insects, and I discuss their controversial morphological value. The main question is whether the abdominal processes are derived from segmental appendages serially homologous to thoracic legs, or whether they are “secondary” outgrowths not homologous with true appendages. Morphological and embryological data, in particular, a comparison with the structure and development of the abdominal appendages in primitive apterygote hexapods, and data from developmental genetics, support the hypothesis of appendicular origin of many of the abdominal processes present in the juvenile stages of various pterygote orders. For example, the lateral processes, such as the tracheal gills in aquatic nymphs of exopterygote insects, are regarded as derived from lateral portions of appendage primordia, homologous with the abdominal styli of apterygotan insects; these processes correspond either to rudimentary telopodites or to coxal exites. The ventrolateral processes, such as the prolegs of different endopterygote insect larvae, appear to be derived from medial portions of the appendicular primordia; they correspond to coxal endites. These views lead to the rejection of Hinton's hypothesis (Hinton [1955] Trans R Entomol Soc Lond 106:455–545) according to which all the abdominal processes of insect larvae are secondary outgrowths not derived from true appendage anlagen. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

Deep phylogeographic divisions and long‐term persistence of forest invertebrates (Hexapoda: Collembola) in the North‐Western Mediterranean basin

The North‐Western Mediterranean basin is well known for its high number of relictual endemic taxa, and has been indicated as one of the world’s major biodiversity hotspots at the species level. A possible contributing factor may be long‐term persistence of populations and their prolonged stability. This study was designed to investigate the phylogeographic structure of three common species of the genus Lepidocyrtus (Hexapoda: Collembola), soil‐dwelling arthropods characterized by limited dispersal capabilities and generally associated with forest habitats. We observed a remarkable geographic structure, with numerous deeply divergent genetic lineages occupying islands as well as mainland sites with no apparent gene flow among most sites, even across distances of only tens of kilometres. The reconstructed time frame for the evolution of these lineages suggests divergence between 5 and 15 Ma. This indicates a remarkably ancient origin and long‐term persistence of individual lineages over a fine geographic scale despite the occurrence of abrupt sea level and climatic fluctuations in the area. This further suggests that currently recognized morphological species might be a serious underestimation of the true springtail biodiversity within this region.     

Diversity and distribution of Collembola (Arthropoda: Hexapoda) of Brazil

Objectives of this study are to summarize the current state of knowledge of the diversity and distribution of Collembola in Brazil; to identify areas of most importance, interest, and need of additional research; and to enable development of experimental hypotheses for future research on Collembola in this region. The total number of collembolan species currently known from Brazil is 199, distributed among 19 families and 80 genera. The greatest numbers of species have been recorded from the states of Rio de Janeiro and Amazonas (with 69 and 56 species, respectively). Few or no species are known from most Brazilian states. Most of the species for which specific Brazilian environmental habitat information is available (93, 66%) are known only from forest environments. Most of the species (127, 64%) are known only from Brazil (most likely being endemic); 33 species (17%) are known only from Brazil and other neotropical areas; and 39 species (20%) have a distribution beyond the neotropical region. Results of this study indicate that much remains to be learned about the Brazilian collembolan fauna. This is especially true for areas of the northeastern, central-western, and southern regions. Studies to determine the species composition of collembolan communities in specific environments in Brazil are needed. The great diversity of Collembola species in Brazil is largely unknown and there are many opportunities for additional research on these environmentally important organisms in this area. Such additional research on the Collembola in Brazil is also essential for a better understanding of the neotropical (and world) collembolan fauna.     

Systematics, cladistics and biogeography of the American genus Farrodes (Ephemeroptera: Leptophlebiidae: Atalophlebiinae)

Cladistic and biogeographic analyses of the genus Farrodes are presented. Two species groups are delineated within Farrodes: F. caribbianus and F. bimaculatus. Three species formerly placed in other genera– Thraulus caribbianus Traver, Thraulus roundsi Traver and Homothraulus maculatus (Needham & Murphy)–are transferred to Farrodes. The species of the F. caribbianus species group are revised. Three new species are described: F. savagei from Venezuela, F. maya and F. mexicanus from Mexico. Keys to separate the two species groups of Farrodes and the species of the F caribbianus species group are provided. Successive cladistic analyses were carried out on both adult and nymphal characters using Hennig86 and CLADOS. The matrix was composed of all available data (nymphal characters were missing for some species), from nymphal and adult stages separately and on taxa represented by both adult and nymphal characters. Species of the genera Simothraulopsis and Homothraulus (components of the Farrodes lineage) were included in the analyses, and Ecuaphkbia was used as the outgroup. Results of the four analyses are compatible. The historical biogeography of Farrodes , with a distribution from nordiern Argentina to southern Texas, is analysed using the program COMPONENT. Areas of endemism are established, and some of their relationships compared with those of other groups available in the literature.     

Molecular biological study on speciation and phylogeny of the order Entomobryomorpha (Collembola: Hexapoda)

In order to elucidate the phylogenetic relationship among groups of the order Entomobryomorpha (Collembola), the sequences on the ITS 1 to ITS 2 fragments of the rRNA gene were analyzed in 11 species of three families. In order to avoid the potential risks and inconsistencies of a single method or data set, the phylogenetic reconstructions were based on three different approaches: methods of maximum parsimony, maximum likelihood and neighbor joining. The inferred phylogenies supported monophyly of the order Entomobryomorpha. The relationships between families were different, but the orders of branching within each family were the same. Entomobryidae and Isotomidae were paraphyletic, whereas Tomoceridae was monophyletic. Tomoceridae was subdivided into two branches; the molecular analysis provided results distinctive enough to separate the two genera by the high bootstrap value. On the other hand, two different populations of putative Homidia koreana appeared to be different species, although their chaetotaxy is identical. A wide coverage of characters, including not only morphological characters but also genetic data such as allozymes and DNA sequences, will give a more accurate picture of the classification and phylogeny of the studied group.     

Functional morphology of the dorsal vessel in the adult fly Protophormia terraenovae (diptera,calliphoridae)

The morphology and ultrastructure of the contractile tubular vessel acting as the cardiac pump in Protophormia terraenovae flies were analyzed by means of light microscopy, SEM and TEM. The results provide a novel anatomical picture of the two vessel portions, the abdominal heart and the aorta, and lay the foundations for an interpretation of the cardiocirculatory function in the fly. In the thorax, the thin and unchambered aorta is without apertures, while the abdominal heart presents a very small caudal aperture and pairs of lateral ostia, one in each of the five chambers of which it is composed. The ostia of the four more distal chambers are of the incurrent type, which is to say that they act as valves ensuring that hemolymph flows only into the heart. Conversely, the ostia in the most proximal chamber allow hemolymph to flow both into and out of the heart. The entire vessel is composed of a single layer of myofibers that are oriented circularly around the lumen in the abdominal heart and longitudinally in the thoracic aorta. The abdominal heart has a thicker wall, a far more diffused and thick distribution of tracheoles, and a far greater number of mitochondria with respect to the aorta. This arrangement ensures a greater availability of oxygen and energy in the abdominal heart compared to the aorta and leads one to suppose that the high‐ and low‐frequency contractions of the cardiac cycle (Thon, [1982] J. Insect Physiol. 28:411–416) can be attributed to the abdominal heart and the aorta, respectively. J. Morphol. 240:15–31, 1999. © 1999 Wiley‐Liss, Inc.     

Marine insects: genital morphology, phylogeny and evolution of sea skaters, genus Halobates (Hemiptera: Gerridae)

Five species of sea skaters, genus Halobates Eschscholtz, are the only insects to have successfully colonized the open ocean. In addition, 36 species are found in sheltered coastal waters throughout tropical Indo-Pacific. The taxonomy of the genus is relatively well known, but reliable hypotheses about phylogenetic relationships are required if the biogeography and evolution of sea skaters is to be discussed in a meaningful way. This work presents the results of a study of new characters from the genital segments, especially those of the male phallus and the female gynatrial complex, and a reinterpretation for several other characters. In total 64 characters were scored for 26 species of Halobates , two species ofAsclepios and one species of Metrocoris. With Asclepios and Metrocoris species as outgroups, the character state sets were analysed cladistically using the computer program Hennig86. After critical evaluations of both characters and clades, a phylogeny for Halobates is presented and its taxonomic implications are discussed. A number of monophyletic species groups are delimited. One genus-level synonymy and three species-level synonymies are suggested. The evolution of Halobates is discussed in the light of the reconstructed phylogeny and present knowledge of the ecology and behaviour of sea skaters. A hypothesis of ecological evolution in halobatine water striders is proposed and tested.     

Comparative internal structure of dorsal lips and radiolar appendages in sabellidae (Polychaeta) and phylogenetic implications

Fan worms (Sabellidae) possess paired modified prostomial structures at the base of the radiolar crown, dorso‐lateral to the mouth, called dorsal lips. The dorsal lips are involved in the sorting of particles collected by the radiolar crown. The range of variation in the morphology of dorsal lips is extensive, and probably this is not only due to adaptations to different environments and feeding preferences but also due to phylogenetic constraints. In this study, we describe and compare the morphology of dorsal lips in a range of sabellid taxa based on histological cross‐sections of these structures, and compare our data and terminology with those of previous studies. Dorsal lips are maintained erect in most taxa by a modified radiole fused to them known as dorsal radiolar appendage. We suggest that dorsal radiolar appendages with an internal supporting axis (cellular or acellular) and probably also the ventral lips are synapomorphies of the family. J. Morphol., 2011. © 2010 Wiley‐Liss, Inc.     

Morphology of the pupal heart,adult heart,and associated tissues in the fruit fly,Drosophila melanogaster

The early pupal heart of the fruit fly Drosophila melanogaster has recently been the subject of intense physiological and molecular work, yet it has not been well described, nor has it been compared with the heart of the adult fly. In the work reported here, the hearts of adults and early pupae of D. melanogaster were studied by scanning and transmission electron microscopy and by light microscopy. The hearts of adults and early pupae both consist of a tube of circular striated muscle one cell in thickness. The alary muscles, which suspend the heart, are more delicate in the adult compared to the early pupa. The pericardial cells in both early pupae and adults are connected to the heart by connective tissue radiating from the alary muscles or dorsal diaphragm. We confirm that four major changes occur in the heart during metamorphosis: 1) a conical chamber is formed de novo in the first and second abdominal segments; 2) the adult heart curves to conform to the contour of the abdomen; 3) a layer of longitudinal striated muscle appears on the ventral surface of the heart; 4) a fourth pair of ostia is added to the three already present in the early pupa; and note additionally that 5) the ostia appear as simple openings in the heart of the early pupa but are valve‐like in the adult. J. Morphol. 240:225–235, 1999. © 1999 Wiley‐Liss, Inc.     

Serial homology of the mandible and maxilla in the jumping bristletail Pedetontus unimaculatus Machida, based on external embryology (Hexapoda: Archaeognatha, Machilidae)

The development of the mandible and maxilla is examined with the scanning electron microscope in the Archaeognatha. Serial homology is discussed to elucidate the general construction of the hexapod mandible. The part comparable to the maxillary palp does not develop in the mandible. Thus, the mandible is coxopodal in origin, and not telognathic but coxognathic. The mandible proper is subdivided into two in late embryonic development, and the smaller proximal and larger distal parts are homologized with the maxillary cardo and stipes, respectively, being subcoxal and coxal in nature. The partition into the "mandibular cardo" in which the mandibular monocondyle is formed and the "mandibular stipes" is recognized as a cuticular ridge or the "mandibular basal ridge" in the postembryonic stages including the imaginal. The molar and incisor are comparable in position and homologized with the maxillary lacinia and galea, respectively. The lacinia and galea could be morphologically interpreted as being the endites of the maxillary coxae I and II, respectively, and the molar and incisor might represent the mandibular coxae I and II as their constituents or endites.     

Embryogenesis of the dipluran Lepidocampa weberi Oudemans (Hexapoda,Diplura, Campodeidae): External morphology

The external features of the embryo of the dipluran, Lepidocampa weberi Oudemans are described. The long germ band is formed, and blastokinesis is a simple flexion of the germ band. The primary dorsal organ is formed between the cephalic and abdominal ends by concentration of serosal cells. The mouth fold is formed by ventral extension of the intercalary, mandibular, and maxillary terga, through which entognathy is completed. The posteroventral region of the mouth fold develops into the admentum. Rotation of the labial anlagen is involved in labial formation, and the glossa, paraglossa, and labial palp acquire a tandem arrangement. The postmentum is formed by fusion of the labial subcoxae and is appendicular in origin. The styli and exertile vesicles are derived from the distal parts of bifurcated appendicular anlagen of the second to seventh abdominal segments. The columnar appendage of the first abdominal segment is serially homologous with the exertile vesicles of the following segments. The abdomen is composed of ten segments, and the cercus is the appendage of the tenth, last abdominal segment. Embryogenesis of Lepidocampa weberi resembles that of the rhabduran Campodea staphylinus (Uzel, 1898) as well as that of the dicelluratan Japyx major (Silvestri, '33). It may be emphasized that the rhabduran and dicelluratan diplurans share important features such as entognathy formation and abdominal organization, and the resemblance between them seems to be close enough to postulate their close affinity. Some embryogenetic features, which Diplura and Collembola share, are recognized as plesiomorphic and the manner of entognathy formation may significantly differ. J. Morphol. 237:101–115, 1998. © 1998 Wiley-Liss, Inc.     

The rowing-to-flapping transition: ontogenetic changes in gill-plate kinematics in the nymphal mayfly Centroptilum triangulifer (Ephemeroptera, Baetidae)

Comparative studies encompassing a wide range of aquatic animals have shown that rowing is exclusively used at low Reynolds numbers ( Re  < 1), whereas flapping is predominantly used at Re  > 100, although few studies have been undertaken to document the transition in individual species that traverse the intermediate Re regime using a single set of appendages. Thus, it is not generally known whether a gradual increase in Re within a system results in a gradual or sudden shift between rowing and flapping. In the present study, we document ventilatory kinematics of a nymphal mayfly Centroptilum triangulifer that develops using a serial array of seven pairs of abdominal gill plates and operates at Reynolds numbers in the range 2–22 during ontogeny. We found that some kinematic variables (stroke frequency and metachronal phase lag) did not change during ontogeny but that others changed substantially. Specifically, gill kinematics in small instars used strokes with large pitch and stroke-plane deviations, whereas larger instars used strokes with minimal pitch and minimal stroke-plane deviation. Gills in larger instars also acquired an intrinsic hinge that allowed passive asymmetric movement between half strokes. Net flow in small animals was directed ventrally and essentially parallel to the stroke plane (i.e. rowing), whereas net flow in large animals was directed dorsally and essentially transverse to the stroke plane (i.e. flapping). The change in whole-gill kinematics from rowing to flapping occurred across a narrow Re range (3–8), which suggests a possible hydrodynamic demarcation between rowing and flapping.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 540–555.     

Morphology of locomotor appendages in Arachnida: evolutionary trends and phylogenetic implications

The morphological diversity of locomotor appendages in Arachnida is surveyed lo reconstruct phylogenetic relationships and discover evolutionary trends in form and function. The appendicular skeleton and musculature of representatives from the ten living arachnid orders ate described, and a system of homology is proposed. Character polarities are established through comparison with an outgroup. Limulus polyphemus Xiphosura). Cladistic analysis suggests that Arachnida is monophyletic and that absence of extensor muscles is a primitive condition. Extensors are primitively absent in Araneae. Amblypygi, Uropygi, Palpigradi, Ricinulei and Acari. Most similarities in the appendages of these orders are symplesiomorphic so that phylogenetic relationships among the 'extensorless' groups cannot be resolved solely on the basis of appendicular characters. Extensor muscles appear to have evolved once, and their presence is considered a synapomorphic feature of Opiliones, Scorpiones, Pseudoscorpiones and Solifugae. Solifugae lack extensors, but a parsimonious interpretation of other characters indicates that this is a secondary, derived condition. The phylogenetic relationships among these four orders are clarified by modifications of the patellotibial joint. Cladistic analysis indicates that Opiliones may be the sister group of the other three orders and that Scorpiones is the sister group of Pseudoscorpiones and Solifugae. Conclusions concerning phylogenetic relationships and evolutionary morphology presented here differ substantially from those of earlier studies on the locomotor appendages of Arachnida.     

Morphology of locomotor appendages in Arachnida: evolutionary trends and phylogenetic implications

The morphological diversity of locomotor appendages in Arachnida is surveyed lo reconstruct phylogenetic relationships and discover evolutionary trends in form and function. The appendicular skeleton and musculature of representatives from the ten living arachnid orders ate described, and a system of homology is proposed. Character polarities are established through comparison with an outgroup. Limulus polyphemus Xiphosura). Cladistic analysis suggests that Arachnida is monophyletic and that absence of extensor muscles is a primitive condition. Extensors are primitively absent in Araneae. Amblypygi, Uropygi, Palpigradi, Ricinulei and Acari. Most similarities in the appendages of these orders are symplesiomorphic so that phylogenetic relationships among the ‘extensorless’ groups cannot be resolved solely on the basis of appendicular characters. Extensor muscles appear to have evolved once, and their presence is considered a synapomorphic feature of Opiliones, Scorpiones, Pseudoscorpiones and Solifugae. Solifugae lack extensors, but a parsimonious interpretation of other characters indicates that this is a secondary, derived condition. The phylogenetic relationships among these four orders are clarified by modifications of the patellotibial joint. Cladistic analysis indicates that Opiliones may be the sister group of the other three orders and that Scorpiones is the sister group of Pseudoscorpiones and Solifugae. Conclusions concerning phylogenetic relationships and evolutionary morphology presented here differ substantially from those of earlier studies on the locomotor appendages of Arachnida.     

Egg envelopes of Baetis rhodani and Cloeon dipterum (Ephemeroptera, Baetidae): a comparative analysis between an oviparous and an ovoviviparous species

A comparative ultrastructural investigation on the eggshell (vitelline and chorionic envelopes) has been carried out in the nymphs of two mayfly species encompassed into the Baetidae, namely Baetis rhodani and Cloeon dipterum. During oocyte differentiation in the meroistic telotrophic ovarioles, gametes are connected to discrete nurse cells by trophic cords. In B. rhodani, each ovariole contains several eggs arranged in sequence, whereas in C. dipterum each contains a single egg. Follicle cells are competent for vitelline and chorionic envelope synthesis. Baetis rhodani is an oviparous species and the chorion is fairly thick, formed by an alveolate endochorion and a fibrillar exochorion delimited by a honey‐comb roof. Cloeon dipterum stands out among Ephemeroptera for being ovoviviparous. In B. rhodani, ovulation starts in the older nymphs with dark wing‐pads, whereas in C. dipterum choriogenesis ends in the imaginal stage. Here the chorion is very thin and laid eggs hatch almost immediately, allowing the larvulae to move out. The maturation of a single egg per ovariole is synchronized with the achievement of the adult stage. The acquisition of ovoviviparity has led to remarkable changes in the ovariole organization along with a simplification of the eggshell structure.     

The male postabdomen and reproductive system of Bibio marci Linnaeus, 1758 (Hexapoda: Diptera: Bibionidae)

The male postabdomen and the internal parts of the male genital system of Bibio marci (Bibionomorpha) were examined and reconstructed 3‐dimensionally. Several features differ from the presumptive dipteran groundplan. The bases of the gonopods are fused with each other and with tergite IX. The penis is not tube‐shaped and only sclerotized on the ventral side. The vasa deferentia are S‐shaped, and two pairs of accessory glands are present. In contrast to these characteristics, the arrangement of the internal parts is probably close to the ancestral condition. With its specific shape, the penis is well suited for the transfer of a spermatophore. The dorsal sclerite of the copulatory organ probably represents the medially fused parameres. A cladistic analysis of 27 characters of the postabdomen yielded two most parsimonious trees, with the strict consensus as follows: Nannochoristidae (outgroup) + (Culicidae [Culicomorpha] + ((Nymphomyiidae + (Tipulidae + Trichoceridae)) + (Tabanidae [Brachycera] + (Bibionidae, Anisopodidae, Axymyiidae [Bibionomorpha])))). Potential synapomorphies of Bibionomorpha (including Axymyiidae) and Brachycera are the fusion of sternum IX with the gonocoxites, the fusion of the parameres forming the dorsal sclerite and the presence of an entire series of postabdominal muscles (M4, M20, M23, M26, M27, M31, M35 and M37). The results of the analysis are preliminary as it is based on a single‐character system with a limited taxon sampling. However, the main result – a clade Bibionomorpha + Brachycera – is fully compatible with current hypotheses on dipteran phylogeny.     

New and earlier unnoted mealybugs and felt scale (Homoptera: Coccinea: Pseudococcidae,Eriococcidae) from Morocco

Two new species of mealybugs (Pseudococcidae), Puto (Ceroputo) chetverikovi n. sp. and Fonscolombia toubkalensis n. sp., are described and illustrated. Thirteen other species are noted for the first time to fauna of Morocco or to Africa as a whole. Additionally, one species of closely related family Eriococcidae, Acanthococcus ericae (Signoret, 1875), is also reported for the first time to Morocco.     

A survey of scale insects in soil samples from Europe (Hemiptera,Coccomorpha)

In the last decades, several expeditions were organized in Europe by the researchers of the Hungarian Natural History Museum to collect snails, aquatic insects and soil animals (mites, springtails, nematodes, and earthworms). In this study, scale insect (Hemiptera: Coccomorpha) specimens extracted from Hungarian Natural History Museum soil samples (2970 samples in total), all of which were collected using soil and litter sampling devices, and extracted by Berlese funnel, were examined. From these samples, 43 scale insect species (Acanthococcidae 4, Coccidae 2, Micrococcidae 1, Ortheziidae 7, Pseudococcidae 21, Putoidae 1 and Rhizoecidae 7) were found in 16 European countries. In addition, a new species belonging to the family Pseudococcidae, Brevennia larvalis Kaydan, sp. n. and a new species of Ortheziidae, Ortheziola editae Szita & Konczné Benedicty, sp. n. are described and illustrated based on the adult female stage. Revised keys to the adult females of Brevennia and Ortheziola are presented.