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1.
Exechia and Bibio have retained several plesiomorphic groundplan features of Diptera and Bibionomorpha, including a fully exposed and sclerotized head capsule, the transverse undivided labrum, the absence of movable premandibles, and undivided mandibles without combs. The fusion of the hypostomal bridge with the head capsule and largely reduced antennae are derived features shared by both taxa. The absence of teeth at the anterior hypostomal margin is a potential autapomorphy of Bibionomorpha. A basal position of Anisopodidae is suggested by a number of plesiomorphies retained in this family. Apomorphies of Bibionomorpha excluding Anisopodidae are the reduction of tentorial elements, the partial fusion of the labrum and clypeus, one-segmented antennae, the absence of a separate submental sclerite, the loss of the labial palpus, and the reduction of the pharyngeal filter apparatus. Head structures of Bibio are largely unmodified. The subprognathous orientation is one of few autapomorphic features. In contrast, the mouthparts of Exechia are highly modified in correlation with the specialized food uptake. The rasping counterrotating movements of maxillae and mandibles with teeth oriented in opposite directions are carried out by strongly developed extensors and flexors of the paired mouthparts. The modified labium mechanically supports the “drill head” formed by the mandibles und maxillae. The necessary stability of the head capsule is provided by the hypostomal bridge which also compensates the far-reaching reduction of the tentorium.  相似文献   

2.
The Acanthopteroctetidae are one of the first‐originated family‐group lineages within “tongue moths” (Lepidoptera‐Glossata). The purpose of this study is to provide a comprehensive account (based on whole mount preparations, serial sections, and Scanning electron microscopy) of the cephalic structure of an adult exemplar of the family, to supplement the sparse available information. Notable plesiomorphies include the retention of frontal retractors of the narrow labrum, a high supraocular index linked to strong development of cranio‐mandibular ad‐ and abductors, and perhaps the unusually short but still coilable (just ca. 1.5 turns) galeal “tongue.” Notable specializations (probably mostly family autapomorphies) include a complement of large sensilla placodea on the male antennae, an apical attachment of the long dorsal tentorial arm to the cranium, an extreme reduction of the single‐segmented labial palps, a particularly strong subgenal bridge and a surface structure of near‐parallel ridges on the ommatidial corneae. The presence of sizable saccular mandibular (type 1) glands opening into the adductor apodeme is unexpected, no counterparts being known from neighboring taxa. The same is true for ventral salivarium dilator muscles originating on the prelabium; and tentatively suggested to be homologues of the extrinsic palp flexors (the insertion shift being related to loss of original function due to palp reduction), rather than to the ventral salivarium muscles of more basal insects. A complete “deutocerebral loop”' may or may not be developed, as is true for a mutual appression of the optic lobe and circumoesophageal connective/suboesophageal ganglion, enclosing the anterior tentorial arm between them; a suboesophageal innervation of the retrocerebral complex was not observed. No characters bearing on the monophyly of the Coelolepida were identified. The scapo‐pedicellar articulation with a scapal process and a smooth intercalary sclerite is reminiscent of conditions in Neopseustidae, but remains debatable as a synapomorphy of the two families. J. Morphol. 275:153–172, 2014. © 2013 Wiley Periodicals, Inc.  相似文献   

3.
Despite several recent analyses on the phylogeny of Neuroptera some questions still remain to be answered. In the present analysis we address these questions by exploring a hitherto unexplored character complex: the tentorium, the internal cuticular support structure of the insect head. We described in detail the tentoria of representatives of all extant neuropteran families and the muscles originating on the tentorium using 3D microCT images and analyzed differences in combination with a large published matrix based on larval characters. We find that the tentorium and associated musculature are a source of phylogenetically informative characters. The addition of the tentorial characters to the larval matrix causes a basad shift of the Sisyridae and clearly supports a clade of all Neuroptera except Sisyridae and Nevrorthidae. A sister group relationship of Coniopterygidae and the dilarid clade is further corroborated. A general trend toward a reduction of the dorsal tentorial arms and the development of laminatentoria is observed. In addition to the phylogenetic analysis, a correlation among the feeding habits, the development of the maxillary muscles, and the laminatentoria is demonstrated.  相似文献   

4.
Female Heterobathmia have the segments behind VIII forming a compact ‘terminal unit’ with a large saddle-shaped dorsal plate and a membranous ventroposterior surface bearing the separate gonopore and anus. While females of most of the nine known species are overall similar, Heterobathmia valvifer is unique amongst lepidopterans in possessing paired ventral appendages (‘ovipositor valves’) arising from the intersegmental groove following segment VIII; evidence from musculature contradicts an interpretation of these appendages structures as ‘true’ ovipositor valves. The ventroposterior wall of the terminal unit in H. valvifer bears paired sclerites, possible homologues of the ‘ventral rods’ in basal Lepidoptera-Glossata. In Heterobathmia megadecella sclerites on paired longitudinal elevations in comparable positions probably are/include homologues of these sclerites. Their similarity with paired sclerotizations in the corresponding region of hydrobiosid caddisflies is noted. A prominent frame-like sclerotization in the genital chamber, located in front of the spermathecal duct origin, is present only in H. megadecella.Putative heterobathmiid autapomorphies include an enlarged ‘subgenital plate’ on venter VIII, absence of apophyses on segment VIII, shortened apophyses on the terminal unit, multilobed accessory glands (but their ‘type 1’ secretory epithelium is plesiomorphic at this level), a conspicuous papilla in the chamber cuticle bearing the opening of the ductus bursae on its apex, and inwards-pointing spines in the ductus bursae. A variably developed thickening of the anterior genital chamber intima is another putative family autapomorphy, while an extreme thickening of the posterior intima seen in Heterobathmia pseuderiocrania is not of general occurrence in heterobathmiids. A sistergroup relationship between Heterobathmiidae and Glossata is supported by their fully developed ‘2-compartment section’ of the spermathecal duct and losses of some likely lepidopteran groundplan muscles.  相似文献   

5.
6.
External and internal head structures of larval representatives of Raphidiidae are described. The obtained data were compared to characters of other neuropterid larvae and to larval characters of representatives of other endopterygote lineages. Characters potentially relevant for phylogenetic reconstruction are listed and discussed. The larvae of Raphidioptera differ distinctly from other neuropterid larvae in their morphology. They are mainly characterised by autapomorphic and plesiomorphic character states and few features indicate systematic affinities with other groups. Endopterygote groundplan features maintained in Raphidioptera are the complete tentorium, the free labrum, the full set of labral muscles, the presence of four extrinsic antennal muscles, the three-segmented labial palpi, the presence of a full set of extrinsic maxillary and labial muscles, the presence of a salivarium, and possibly the high number of stemmata. Apomorphies likely correlated with predaceous habits are the long gula, the protracted maxillae, the longitudinal arrangement of extrinsic maxillary muscles, and the elongated prepharyngeal tube. Highly unusual, potentially autapomorphic features are the presence of a dorsal ligament of the tentorium and paired gland-like structures below the pharynx. A prognathous or very slightly inclined head and slender mandibles without mola are features shared by larvae of all orders of Neuropterida. The parallel-sided head is a potential synapomorphy of Raphidioptera and Megaloptera. A fully prognathous head with anteriorly shifted posterior tentorial grooves and the presence of a parietal ridge and a distinct neck region are features shared with Corydalidae. Characters of the larval head are not sufficient for a reliable placement of Raphidioptera.  相似文献   

7.
Variation in structures of the posterior surface of the head in Hymenoptera is compared and interpreted according to theories of head capsule evolution, with focus on understanding previously baffling conditions in the superfamily Chalcidoidea. Features are investigated separately without first classifying subforaminal bridges into subcategories. In Proctotrupomorpha (including Chalcidoidea), Ceraphronoidea and some Ichneumonoidea, there are multiple posterior pits associated with the tentorium. In most examined Hymenoptera with a subforaminal bridge, there was a differentiated median area, typically with highly variable microtrichia. This area is elevated in Cephoidea and Pamphilioidea, but is not elevated in other Hymenoptera. Subforaminal bridges in Apocrita previously classified as hypostomal bridges are discussed in the context of A.P. Rasnitsyn's hypothesis that relative importance of adult feeding drives subforaminal bridge evolution.  相似文献   

8.
Larval head structures of Xyela sp. are described in detail. The characters are compared to conditions found in larvae of other groups of Hymenoptera and Endopterygota. Like other symphytan larvae the immature stages of Xyelidae are mainly characterized by presumably plesiomorphic features of the head. The head sutures are well developed and all parts of the tentorium are present. The labrum is free and a complete set of labral muscles is present. The maxillae are in a retracted position. In contrast to other hymenopteran larvae Xyela possesses a clypeofrontal suture, a comparatively long antenna and three well‐developed antennal muscles. Apomorphic features of Xyela are the absence of muscles associated with the salivarium and the complete absence of Musculus craniocardinalis. A clade comprising Orussidae and Apocrita is supported by the unsegmented maxillary and labial palps and the absence of the lacinia. Six potential autapomorphies for the Hymenoptera were revealed: (1) the caudal tentorial apodeme, (2) the bifurcated tendon of Musculus craniomandibularus internus, (3) the lateral lobe of the cardo, (4) the origin of M. tentoriohypopharyngalis from the posterior head capsule, (5) the exceptionally strong prepharyngo‐pharyngeal longitudinal muscle and (6) the longitudinal muscle of the silk press. The maxillolabial complex, the vestigial M. craniocardinalis and a distinctly developed labio‐hypopharyngeal lobe bearing the opening of the salivary duct are potential synapomorphies of Hymenoptera and Mecopterida. The globular, orthognathous head capsule, the modified compound eyes, the occipital furrow and the X‐shaped tentorium are features with unclear polarity shared by Hymenoptera and Mecoptera.  相似文献   

9.
10.
The tentorium, the anterior sulci of the head capsule (epistomal, subgenal, subantennal, circumantennal, and circumocular sulci), and the extension of the anterior tentorial pit were studied in 26 species of Blattaria (representing most principal lineages), 4 species of Mantodea (including the basal Mantoida schraderi), and 1 species each of Isoptera (the basal Mastotermes darwiniensis) and Mantophasmatodea (Austrophasma caledonense). The morphology of these head structures is compared with literature data on other insect orders, mainly Phasmatodea, Orthoptera, Dermaptera, Embioptera, and Plecoptera, and partly Odonata and Zygentoma. Characters are defined, presented in a matrix, and evaluated with regard to phylogenetic implications and homoplastic evolution. The structural relationships of the subantennal sulcus to the subgenal, circumocular, and circumantennal sulci, which are highly variable and strongly homoplastic (depending much on the size of the compound eyes) are a focal issue; several types of subantennal sulci are defined. The presence of an anterior transverse bridge in the tentorium (“perforated tentorium”) of all Dictyoptera here studied confirms the monophyly of this group. Mantophasmatodea lacks this element.  相似文献   

11.
Platypsyllus castoris is closely associated with beavers and displays a unique set of structural specializations. We document the morphology of adults with modern techniques, and interpret evolutionary changes linked with the specific life style. The small subfamily Platypsyllinae has evolved an entire suite of features correlated with a more or less close association with mammals, for instance a flattened body, a dorsal cephalic shield, flightlessness, eye reduction, and depigmentation. Within this small group, Platypsyllus displays numerous autapomorphic features, correlated with a close association with the beaver. Essential is a combination of mechanical stabilization and firm anchorage on the host, and efficient forward movement in the fur. Exo- and endoskeletal structures of the head and thorax are reinforced by vertical cuticular columns and by an array of internal ridges. The antennae are shortened and strongly modified, the mandibles distinctly reduced and flattened, unsuitable for cutting, scraping or grinding. The musculature of the mouthparts is simplified, whereas an enhanced set of prepharyngeal and pharyngeal dilators forms an efficient sucking pump. The prothoracic musculature is strongly developed. In contrast, the pterothoracic muscle system is distinctly simplified, even though leg muscles are strongly developed. Using the legs, the flattened beetles move sideways through the dense fur of the beaver, using posteriorly directed groups of setae and ctenidia to prevent being pushed backwards by the densely arranged hairs. In contrast to the anterior body, the cuticle of the abdomen is thin, and the entire tagma flexible, with thin layers of segmental muscles. The hind gut is not connected with the mid gut. The beetles probably consume liquid, possibly with emulgated minute skin debris. As the morphology of the mouthparts excludes damage to the skin of the host, the association should not be addressed as ectoparasitic but as commensalism.  相似文献   

12.
External and internal head structures of Nannochorista species were examined and described in detail. The characters are discussed with regard to their functional and phylogenetic implications. The structure of the mouthparts indicates that adults of Nannochorista feed on fluids. The loss of the mandibular muscles and the precerebral pharyngeal dilators are presumptive autapomorphies of the genus. A possible clade comprising Nannomecoptera, Siphonaptera and Diptera is supported by the presence of a labral food channel, the absence of the galea, a sheath for the paired mouthparts formed by the labium, very strongly developed labial palp muscles and cibarial dilators, and the presence of a well‐defined postcerebral pharyngeal pumping chamber. Closer affinities of Nannomecoptera with Diptera are suggested by the presence of a unique sensorial groove on the third maxillary palpomere. Further potential synapomorphies are the presence of a frontal apodeme and a primarily lamelliform mandible without teeth. The presence of a salivary channel on the laciniae and a subdivided labrum are shared derived features of Nannochorista and Siphonaptera. A derived condition present in Mecoptera including Boreidae but excluding Nannochoristidae is the secretion with a strongly developed intrinsic muscle of the salivary duct. The loss of the lateral labral retractor, the cranial muscle of the cardo, and of two of the three premental retractors, and the absence of transverse epipharyngeal muscles are potential autapomorphies of Antliophora. The formation of a maxillolabial complex is a possible synapomorphy of Hymenoptera and Mecopterida.  相似文献   

13.
Mouthparts of unfed larvae Piona carnea (Koch, 1836) (Acariformes: Pionidae) were studied on whole-mount preparations, semi-thin sections and with TEM and SEM methods. The mouth apparatus is incorporated within the pseudotagma, gnathosoma, composed of the infracapitulum and of the chelicerae resting on the roof of the latter. The gnathosoma inclines to the long axis of the body and is inserted at its base into the idiosoma by the circumcapitular fold. The basal cheliceral segments are long and fused. An anterior projection, the proposed fused fixed digits, protrudes from the distal end of the basal cheliceral segment forward between the movable digits. The movable digits are always found in protruded position, strongly curved upward and show a groove on their inner sides. The ventral wall of the infracapitulum is made of the mentum posterior and the malapophyses anterior to the palp articulation. The malapophyses are squeezed between the large palps and envelope the distal portion of the chelicerae from the sides. The ventral portion of the fused malapophyses are provided with a characteristic ventral cuticular fork of unknown function. Each malapophysis terminates by a flexible lateral lip provided by several rigid jags looking posterad. The palps face downward and backward, and bear on the tibia the large curved palpal claws turned laterad. The palp femur bears on the ventral aspect a characteristic wide spade-like projection provided with its own muscles originating on the dorsal wall of the femur. The labrum is a thick cuticular arrow-like structure protruding forward into the preoral cavity, whereas the cervix is a thin weakly sclerotized plate. The particular labral valve projects forward from the dorsal basis of the labrum into the preoral cavity. The labrum and the cervix are provided by their own small labral and cervical muscles originating on the cervical apodemes. The pharynx is totally separated from the ventral wall of the infracapitulum and is devoid of ventral dilators. The dorsal pharyngeal dilators originate on the thick and sclerotized capitular apodeme and, posteriorly, on the paired cuticular branches, capitular apodemes, which end freely in the body cavity and are combined with the common salivary duct. The short sigmoid pieces serve for origin of the levator muscles of the chelicerae. Retraction of the gnathosoma and the chelicerae is mediated by several sets of muscles originating on the dorsal plate.  相似文献   

14.
The larval trunk morphology including chaetotaxy, locomotory structures, and trunk musculature of Heterobathmia pseuderiocrania, Eriocrania cicatricella, and Acanthopteroctetes unifascia is described using conventional light, polarization, and scanning electron microscopy. The ground plan morphology of the lepidopteran larva and neolepidopteran caterpillar is discussed in light of the life history succession from free soil dwelling organism to endophagous and finally to a primarily free living, angiosperm associated organism. I suggest that the larval morphology is argued to be strongly influenced by the shift in number of surfaces present in the larval environment. Especially the environment of the endophagous species, where the upper surface of the leaf mine is linked to the presence of dorsal locomotory structures such as the retractable calli and dorsal friction patches is proposed to have had a significant impact on the morphology and locomotory mechnism of the lepidopteran caterpillar. The chaetotaxy of the lepidopteran ground plan is found to be simple, consisting only of primary and secondary tactile setae and segmental proprioceptors. The presumption of Gerasimov ([1935] Zool Anz 112:177–194) that MXD1 of the prothorax is a shifted mesothoracic MD setae is supported. I suggest that the serial arrangement of the proprioceptors MD1, present on all trunk segments except the prothorax, and a trisetous MV group on all the thoracic segments is part of the lepidopteran larval ground plan. The absence of apodeme structures associated with trunk musculature in the nonglossatans suggests that this is an autapomorphic character of the Lepidoptera and it is further found to have been influential in the evolution of the typical caterpillar trunk. The attachments of the thoracic muscles directly to the trunk integument, suggest that the apodemal structures ancestral to the Amphiesmenoptera have been reduced in the Lepidoptera. Within the non‐Neolepidoptera, the lifehistory shift may have resulted in reduction of the dorsal locomotory structures, such as calli. The abdominal musculature and structural similarities further suggest that the ventral calli are structural predecessors to the crotchet bearing proleg of the “typical caterpillar.” J. Morphol. 274:1239–1262, 2013. © 2013 Wiley Periodicals, Inc.  相似文献   

15.
We elucidate the configuration of the tentorium and the sclerites of the hypopharynx–prementum complex in selected spore‐ (pollen‐) and non‐spore‐feeding Aleocharinae (Staphylinidae) by presenting the first comparative 3D reconstructions of these structures for 19 staphylinoid beetle species (six outgroups, 13 Aleocharinae). General organization of the tentorium follows the groundplan previously proposed for adult Staphylinidae, although some taxa have reduced or lost the dorsal (all Aleocharinae studied, Agathidium mandibulare [Leiodidae]) or anterior (Omalium rivulare [Omaliinae], Anotylus sculpturatus [Oxytelinae]) tentorial arms. All species investigated have premental and hypopharyngeal sclerites that are partly homologizable across taxa. We clarified that Musculus praementopalpalis externus originates from the premental sclerite, resolving its unclear origin reported in our previous publications. Eight of 13 investigated Aleocharinae species are spore/pollen feeders, six obligatorily. Three of these six (Eumicrota, Gyrophaena fasciata, G. gentilis) have grinding pseudomolae and a fully developed hypopharyngeal suspensorium with posterior bridge and anterior elongations; the remaining three (Oxypoda, Pagla, Polylobus) lack pseudomolae and suspensorial bridge, but have the suspensorium elongated anteriorly. The dorsolateral side of the hypopharyngeal sclerite interacts with the pseudomola. Obligate sporophagy/pollinivory apparently arose at least three times in Aleocharinae, not always involving the pseudomola–hypopharynx grinding mechanism.  相似文献   

16.
Musculature of the head and cervical region in eight lepidopteran species from the families Cossidae, Sesiidae, Limacodidae, and Zygaenidae (Cossiformes) was studied. Besides, musculature of the anterior cervical region in Brachodes appendiculata Esper (Brachodidae) is discussed. The data on musculature are analyzed and compared with the authors’ earlier results on a relatively primitive ditrysian moth from the family Tineidae (Korzeev, 2004). The absence of the labro-epipharyngeal musculature in Cossiformes is confirmed and the presence of a new, indirectly acting secondary labral extensors CMx.5 in Zygaenidae is revealed. These secondary extensors are strongly developed in Papilionomorpha. Cossiformes and Tineidae have been shown to possess several common plesiomorphic traits in the structure of labrum, while the dilatators of salivarium within Cossiformes are reduced in number from two pairs to a single pair. The present study shows that the musculature of the cervical region has remained relatively stable in the evolution of Lepidoptera despite the loss of some muscles in different taxa. The groundplan of the head musculature is reconstructed for Glossata, Papilionomorpha, Cossiformes, and the “higher” Lepidoptera from the Geometriformes-Noctuiformes complex of superfamilies. In addition, the groundplan of the musculature of the cervical region is depicted for Myoglossata, Papilionomorpha, Cossiformes, and the Geometriformes-Noctuiformes complex of superfamilies.  相似文献   

17.
18.
The internal head anatomy (and the peculiar integumental structure of the epicranial notch region) of Heterogynis penella larvae are described; special attention is paid to the skeleto‐muscular and nervous systems and to the cephalic glands. Transverse ligaments connect the apodemes of the mandibular adductor muscles of both sides and the anterior maxillo‐labial articulations of both sides. The two ligaments are linked to each other by a thin, apparently acellular membrane. An accessory, trilobed mandibular gland is present. A putative stretch receptor, connecting the oblique dorsal cibarial dilators of both sides, is described for the first time in a lepidopterous larva and its importance in assessing the homology of these muscles is discussed. The presence of cibarial sensilla, previously predicted in other caterpillars on the basis of behavioural experiments and observations of the nerve pattern, is confirmed. The structural diversity of larval head anatomy in ditrysian Lepidoptera is discussed, with particular emphasis on the innervation of the corpora cardiaca and corpora allata and of the sensilla of the head capsule.  相似文献   

19.
Leiodidae are the second largest subterranean radiation of beetles at family rank. To explore morphological trends linked with troglobiontic habits and characters with potential phylogenetic significance, the head of the cave-dwelling species Troglocharinus ferreri (Cholevinae, Leptodirini) was examined in detail. Overall, the general pattern is similar to what is found in Catops ventricosus (Cholevini). Shared apomorphic features include a fully exposed anterolateral concavity containing the antennal socket, a distinct bead above this depression, a bilobed lip-like structure anterad the labrum, a flat elevated portion of the ventral mandibular surface, and a ventral process at the proximomesal edge of this mandibular area. The tentorial structures are well-developed as in C. ventricosus, with a large laminatentorium and somewhat shortened dorsal arms. The mouthparts are largely unmodified, with the exception of unusually well-developed extrinsic maxillary muscles. Features of T. ferreri obviously linked with subterranean habits are the complete lack of compound eyes, circumocular ridges, and optic lobes. A series of characters is similar to conditions found in other genera of Leptodirini: the head capsule completely lacks a protruding ocular region, a distinct neck is missing, the transverse occipital crest is indistinct, and the antennae are elongate and lack a distinct club. Two different trends of cephalic transformations occur in troglobiontic Leptodirini, with some genera like Troglocharinus and Leptodirus having elongated head capsules and antennae, and others having broadened, more transverse heads. In contrast, the modifications are more uniform in the closely related Ptomaphagini, with a pattern distinctly differing from Leptodirini: the head is transverse, with a distinctly protruding ocular region, a distinct transverse occipital crest, and a very narrow neck region.  相似文献   

20.
The invertebrate collection of the Australian Museum revealed the existence of three new species of the genus Uromunna (family Munnidae) from New South Wales, Australia. Uromunna tenagoika sp. nov. from Batemans Bay has the head anterior margin concave, without simple setae; eyes with few ommatidia, eye lobes with both margins parallel, in male posterior margin directed forward; pereonite 7 as wide as pereonite 6, subequal to pleotelson width; pleotelson distal margin pointed, suburopodal shelf present; mandible palp absent; pereopod I propodus robust setae absent; pereopods relatively short; pleopod IV exopod distal tip with one pappose seta. Uromunna rhamnda sp. nov. from Batemans Bay has a head anterior margin concave, without simple setae; eye lobes with both margins parallel, in male posterior margin parallel to frontal margin; pleotelson distal margin pointed, suburopodal shelf absent; mandible palp present; pereopod I propodus robust setae absent; pleopod IV exopod distal tip with one pappose seta. Uromunna eora sp. nov. was found at Southern Creek, located north of Sydney and is the first species of the genus described from a fully freshwater environment with no marine water input. It has a relatively elongate body (length 3.4× width), anterior margin concave, eye lobes with both margins converging distally, in male posterior margin directed forward; pereonite 7 as wide as pereonite 6, wider than pleotelson width; pleotelson distal margin truncate, suburopodal shelf present; mandible palp present; pereopod I propodus robust setae present; pleopod IV exopod distal tip with two pappose setae. This work raises the number of Uromunna species found in Australia to a total of six.

http://zoobank.org/urn:lsid:zoobank.org:pub:D877FC47-BC6E-4827-BA89-3A75AFDE1124  相似文献   


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