长蝎蛉幼期形态附生物学记述

【目的】蝎蛉科(Panorpidae)是长翅目(Mecoptera)最大的科,是重要的生态指示昆虫。然而,由于对环境条件要求苛刻,饲养困难,其幼期研究很不充分。【方法】本研究通过人工饲养成虫获得了长蝎蛉Panorpa macrostyla Hua的卵、幼虫和蛹等全部虫态,运用光学显微镜和扫描电子显微技术观察了其超微形态,并简要记载了其生物学特性。【结果】长蝎蛉每年发生1代,成虫发生于6月末至8月初。卵椭球形,卵壳表面覆盖一层隆起的网状结构。幼虫蠋型,具3对分4节的胸足和8对不分节的腹足;头壳高度骨化,具1对由26个小眼组成的复眼和1对3节的触角,口器咀嚼式;腹部第1-9节背面具有成对的背毛突,第10节仅有1根背毛突,腹部末端具有一个可伸缩的吸盘;呼吸系统为周气门式,具1对前胸气门和8对腹气门。幼虫共4个龄期,以预蛹期在土室内越冬。蛹为强颚离蛹,外形接近成虫,雄蛹腹部末端膨大。【结论】基于幼虫形态特征,长蝎蛉明显区别于新蝎蛉属Neopanorpa、华蝎蛉属Sinopanorpa、双角蝎蛉属Dicerapanorpa以及单角蝎蛉属Cerapanorpa幼虫。然而,长蝎蛉幼虫头部刚毛L2和SO2,腹部末节刚毛D2,SD1和SD2端部均膨大呈棒状,与蝎蛉属Panorpa其他种类区别明显,表明长蝎蛉的属级地位需要进一步研究。     

Leucokinin and callitachykinin immunoreactive neurons during postembryonic development of calliphora vomitoria (L.) (DIPTERA : CALLIPHORIDAE)

Neurons containing 2 types of myotropic neuropeptides were investigated by immunocytochemistry during postembryonic development of the brain and ventral nerve cord of the blowfly Calliphora vomitoria (Diptera : Calliphoridae). Antisera raised against the insect neuropeptides Callitachykinin II (CavTK II), Locustatachykinin I (LomTK I), and Leucokinin I (LK I) were used. Callitachykinin immunoreactive (CavTK–IR) neurons were detected from the 1st-instar larva throughout development to adult. The number of CavTK–IR cell bodies in the brain was 4–16 in larval stages, 10–84 in pupal stages, and over 140 neurons in the newly emerged fly. With the CavTK antiserum, the fibers of only 4 descending neurons were detected in thoracico–abdominal ganglia throughout development. The antiserum to LomTK displayed the same neurons as that to CavTK II as well as a small number of additional neurons. Notably, there were seen about 14–20 locustatachykinin-like immunoreactive (LomTK-LI) cell bodies in the thoracico–abdominal ganglia throughout development. Leucokinin-like immunoreactive (LK-LI) neurons were labeled throughout postembryonic development. In the brain, 2–4 LK-LI cell bodies were labeled from 1st-instar larva to 8-day-old pupa, and 6 LK-LI cell bodies were labeled in the adult brain. In the abdominal ganglia, 7 pairs of LK-LI cell bodies were labeled from 1st-instar larva to 96-h-old pupa, 8 pairs in 8-day-old pupa, and 9 pairs in newly emerged fly, respectively. The CavTK containing neurons in the brain displayed a drastic increase in numbers from larval stages to adult, which indicates an addition of functional roles for this type of peptide. During earlier pupal stages, the number of CavTK–IR neurons decreased. The LK-LI neurons, however, were strongly immunoreactive throughout postembryonic development. Only one additional pair of cells appeared in the brain and 2 additional pair of cells appeared in the abdominal ganglia of the adult as compared with larvae. The continuous high expression of LK-LI material may suggest a functional role for this type of peptide during development.     

Larval morphology of Panorpodes kuandianensis (Insecta,Mecoptera, Panorpodidae) and its evolutionary implications

Larval characters play a significant role in evolutionary and systematic studies of holometabolous insects. However, Panorpodidae, a derived family of Mecoptera, are largely unknown in their immature stages to date. Here, the first instar larva of the short-faced scorpionfly Panorpodes kuandianensis Zhong, Zhang & Hua, 2011 is described and illustrated using light and scanning electron microscopy. The larva of Panorpodes is remarkable for the absence of compound eyes on the head and the presence of seven small unpaired proleg-like processes along the midventral line on abdominal segments II–VIII. The homology of these unpaired appendage-like processes, their ecological adaptation, and the evolutionary implications of some larval characters of Panorpodidae are discussed.     

A new type of loriciferan larva (Shira larva) from the deep sea of Shatsky Rise,Pacific Ocean

Loricifera is a phylum of minute animals that live exclusively in marine sediments. A total of 33 species have been described so far in this phylum; however, several more are already known from preliminary observations. Loriciferans are characterised by a complex life cycle, which involves a succession of several adult and larval stages. Here, we describe a new type of loriciferan larval stage: the Shira larva. The gross morphology of this larva is generally similar to that of the most prominent larval type of Loricifera, the so-called Higgins larva. However, the Shira larva possesses a number of unique features, namely (1) a single pair of anteroventral setae is present in the most anterior region of the abdomen, (2) the bases of the anteroventral setae are very large and swollen, (3) the thorax and abdomen are thinner than the introvert and (4) the abdominal region is divided into five sub-regions. Accordingly, we described the new species, Tenuiloricus shirayamai gen. nov. et sp. nov. (incertae sedis). The new findings are discussed from a comparative perspective with the Higgins larva as well as with the fossil of a putative loriciferan larval stage from the Middle Cambrian.     

Morphology of the immature stages of Adoretus tenuimaculatus Waterhouse, 1875 (Coleoptera: Scarabaeidae: Rutelinae: Adoretini)

The immature stages of the brown chafers Adoretus tenuimaculatus Waterhouse, 1875 were investigated morphologically using scanning electron microscopy. The scarabaeiform larvae of A. tenuimaculatus are peculiar for bearing a pair of eye spots on the head capsule, devoid of palidium but numerous hamate setae on the raster. The larvae possess a crown-like apical sensory area on each antenna, maxillary and labial palp. The pupa possesses dorsally six pairs of dioneiform organs on the intersegment regions between the first seven abdominal segments. This study provides the first ultra morphological study of the larvae in Adoretini. Both advantages and disadvantages of the scanning electron microscopy in the larval taxonomy of Scarabaeidae were briefly discussed.     

Larval development of nemerteans of the genus Quasitetrastemma (Nemertea: Monostilifera)

Data on the larval development of Quasitetrastemma stimpsoni and Q. nigrifrons are presented. In both species, fertilization is external; the development passes through a free-swimming larval stage, the “hidden larva.” The larva has three pairs of eyes. After settling, the eyes of the second pair fuse with eyes of the first pair or are completely reduced. The basis and stylets are formed in 7–8 days after fertilization. Larvae of Q. stimpsoni settle on day 9–10 after fertilization; and Q. nigrifrons, on day 7–8.     

Morphogenesis of Pseudopallene sp. (Pycnogonida, Callipallenidae) II: postembryonic development

Pycnogonida (sea spiders) are bizarre marine arthropods that are nowadays most frequently considered as being the sister group to all other chelicerates. The majority of pycnogonid species develops via a protonymphon larva with only three pairs of limbs affiliated with the future head region. Deviating from this, the hatching stage of some representatives shows already an advanced degree of trunk differentiation. Using scanning electron microscopy, fluorescent nucleic staining, and bright-field stereomicroscopy, postembryonic development of Pseudopallene sp. (Callipallenidae), a pycnogonid with an advanced hatching stage, is described. Based on external morphology, six postembryonic stages plus a sub-adult stage are distinguished. The hatching larva is lecithotrophic and bears the chelifores as only functional appendage pair and unarticulated limb buds of walking leg pairs 1 and 2. Palpal and ovigeral larval limbs are absent. Differentiation of walking leg pairs 3 and 4 is sequential. Apart from the first pair, each walking leg goes through a characteristic sequence of three externally distinct stages with two intermittent molts (limb bud-seven podomeres-nine podomeres). First external signs of oviger development are detectable in postembryonic stage 3 bearing three articulated walking leg pairs. Following three more molts, the oviger has attained adult podomere composition. The advanced hatching stages of different callipallenids are compared and the inclusive term "walking leg-bearing larva" is suggested, as opposed to the behavior-based name "attaching larva". Data on temporal and structural patterns of walking leg differentiation in other pycnogonids are reviewed and discussed. To facilitate comparisons of walking leg differentiation patterns across many species, we propose a concise notation in matrix fashion. Due to deviating structural patterns of oviger differentiation in another callipallenid species as well as within other pycnogonid taxa, evolutionary conservation of characteristic stages of oviger development is not apparent even in closely related species.     

FMRfamide-like immunoreactivity in the ventral nerve cord of the larval eastern spruce budworm,Choristoneura fumiferana (clemens) (Lepidoptera : Tortricidae)

Distribution of FMRFamide-like immunoreactivity was examined in the larval ventral nerve cord of the eastern spruce budworm, Choristoneura fumiferana (Lepidoptera : Tortricidae). Indirect immunofluorescent methods revealed the existence of 3 groups of FLI neurons in each ganglion. The neurons are distributed in a bilaterally symmetrical fashion at the anterodorsal, midlateral and posteroventral regions of the ganglia. There are 4 FMRFamide-like immunoreactive fiber tracts on the dorsal surface of the ganglia to which the anterodorsal FLI neurons project ipsilaterally, while the midlateral pair projects both ipsi-, and contralaterally. The last abdominal ganglion (AG8) has 4 additional pairs of FLI neurons; and axons from some of these extend into the median abdominal nerve, which suggests some role for this neuropeptide in the control of posterior structures of the larva.     

Ultrastructure of the larval midgut of Bittacus planus (Mecoptera: Bittacidae) and Neopanorpa longiprocessa (Mecoptera: Panorpidae)

Bittacidae and Panorpidae are the two largest families in Mecoptera. The larvae of Bittacidae are different from those of Panorpidae in external morphology and habits, and have an interesting habit of spraying the body surface with soil through the anus. However, it remains unknown to date whether the larval midguts are different in structure between the two families. Here the ultrastructure of the larval midguts of the hangingfly Bittacus planus Cheng and the scorpionfly Neopanorpa longiprocessa Hua & Chou were compared using light, scanning, and transmission electron microscopy. The midguts of both species are simple tubes of single layered epithelia with digestive and regenerative cells but without diverticula. The basal plasma membrane of epithelial cells exhibits infolding in B. planus, but is closely apposed to its basal lamina in N. longiprocessa. Lymph spaces are present between adjacent epithelial cells in B. planus, but are absent in N. longiprocessa. The regenerative cells are scattered among the digestive cells in B. planus, but are aggregated in N. longiprocessa. The longitudinal muscle bands are compact in B. planus, but are sparse in N. longiprocessa. The compact longitudinal muscle bands are likely associated with their soil-spraying habit in Bittacidae.     

Ultrastructure of mediodorsal setae in biting midge larvae of the genus Atrichopogon Kieffer with notes on their biological significance (Diptera: Ceratopogonidae)

The ultrastructure of the strong mediodorsal setae in terrestrial stage IV larvae of Atrichopogon (Meloehelea) oedemerarum and A. (M.) meloesugans was examined using light, scanning and transmission electron microscopy. Serrated setae placed on prominent processes are distributed in pairs on all thoracic and abdominal segments. Setae are innervated by a single dendrite and their surface has no pores. The trichogen cell is not retracted from the setal lumen on completion of the hair-forming process but fills the mediodorsal seta also when the larval cuticle is fully sclerotised. Such a phenomenon was previously reported in terrestrial larvae of the genus Forcipomyia. We suggest that the mediodorsal setae described in Atrichopogon are plesiotypic mechanoreceptors for the subfamily Forcipomyiinae. They are preserved in the truly terrestrial larvae of Atrichopogon, but modified to secretory setae in the genus Forcipomyia. Both genera bearing distinct mediodorsal setae have developed functional tracheal gills, unknown in other biting midges.     

Larvalentwicklung und Metamorphose vonPhoronis psammophila (Phoronida,Tentaculata)

The larval development ofPhoronis psammophila Cori is divided into 6 phases (on the basis of increasing pairs of larval tentacles); furthermore an initial and a ripe phase are distinguished. Specific aspects of the development are described: Formation and structure of larval tentacles; anlage of adult tentacles as a thickening in the larval tentacle base; late development of the metasome (larva with 4–6 tentacles); formation of the metasome pouch in the larva with 8 tentacles; enlargement of the apical plate; differentiation of the gut; differentiation of larval nephridia; formation of pigment particles in the larva with 6 tentacles (storage function of pigments and its significance for larval identification); different types of discoflagella in various regions of the body. The larval development shows the following tendencies: Improvement of locomotion; intensification of food filtration; anlage of adult organs in the larva leading to a shortening of metamorphosis duration. The larva ofP. psammophila is compared with those ofP. pallida, P. hippocrepia, andP. vancouverensis. Earlier larval determinations ofP. psammophila (e.g.Actinotrocha sabatieri, A. hatschekii) are shown to have been mistakes. Termination of the postembryonic phase (metamorphosis) can be induced experimentally by bacteria and also by cations. Pure or mixed bacteria cultures must be present at the beginning exponential growth phase. The bacteria density required is 20–94×10⁶ bact.ml^?1 for pure cultures and on the average 28×10⁶ bact. ml^?1 for mixed cultures. Metamorphosis initiation by cations can be induced with CsCl (0.06 M) and RbCl (0.035 M). Metamorphosis ofP. psammophila occurs in 6 phases: larva, ready for metamorphosis; larva, activated by bacteria or ions; evagination of the metasome diverticle, dislocation of gut; losing and swallowing of episphaere and larval tentacles; formation of the youngP. psammophila. All developmental phases are described and compared with those ofP. muelleri; imperfect metamorphosis is characterized and the youngP. psammophila compared with older stages and the adult Phoronis.     

The first instar larva of two species of Miltogramminae (Diptera: Sarcophagidae) from the Middle East – examples of peculiar morphology

The first instar larva is described for two species of Miltogramminae flesh flies representing the genera Miltogramma Meigen and Phrosinella Robineau‐Desvoidy. Using a combination of light microscopy and scanning electron microscopy, the habitus is documented along with all important exoskeletal and cephaloskeletal structures. The described first instar larvae show substantial modifications of their morphology in comparison with the saprophagous ground plan defined for calyptrate flies. Miltogramma (Miltogrammidium) efflatouni Rohdendorf possesses the general set of derived character states shared with other species of Miltogramma and shares larval synapomorphies with species representing the subgenera Anacanthothecum Rohdendorf, Cylindrothecum Rohdendorf and Miltogrammidium Rohdendorf. Unique character states of the first instar larva of this species are the spine‐like shape of antennal complex, laterally compressed apical part of mouthhooks and an extreme elongation of the sensilla of the maxillary palpus. Phrosinella (Asiometopa) ujgura (Rohdendorf) possesses a pair of retractable, robust processes on the ventral margin of the first thoracic segment and a very strong, elongated labrum. Both character states are present in other known first instar larvae of genus Phrosinella. A reduction of cuticular ridges on dorsal and lateral surfaces of abdominal segments supports a monophylum consisting of the subgenera Asiometopa Rohdendorf and Caspiomyia Rohdendorf within genus Phrosinella.     

条背萤的形态和生物学研究

首次在中国大陆发现水栖萤火虫条背萤Luciola substriata。形态观察发现,条背萤成虫橙黄色, 鞘翅末端灰黑色;发光器均为白色,雄虫发光器位于第5、6腹节,位于第5腹节的发光器呈带状,第6腹节的发光器呈“V”字形;雌虫的发光器呈带状,位于第5腹节;卵椭圆形,橙黄色。幼虫有两种形态,1～2龄具有7对呼吸鳃,3～6龄幼虫无呼吸鳃。幼虫具有一对发光器,位于第7腹节腹面;初蛹期仍保留幼虫形态的发光器,后呈现成虫的发光器,两种形态的发光器并存直至羽化。对条背萤生活史及习性调查发现,条背萤生活在水草较多的池塘、湖泊和流速缓慢的河流中。该虫1年发生1代,以幼虫在水中越冬,5月初老熟幼虫开始上岸化蛹。在25℃下,条背萤预蛹期平均为6.17天,蛹期平均为4.43天。成虫5月上旬至9月中旬发生。日落后的1 h内是条背萤成虫闪光求偶的最盛期。卵期平均12.5天。幼虫的猎物为静水椎实螺Lymnaea stagnalis,凸旋螺Gyraulus conwexiusculus等,天敌为克氏原螯虾Procambarus clarkii、中华绒毛蟹Eriocheir sinensis、草鱼Ctenopharyngodonidellus等。利用光谱仪对条背萤的发光光谱进行测定发现,条背萤的萤光光谱为425～603 nm,峰值为504 nm,颜色为黄绿混合色。     

Phylogeny of Berosini (Coleoptera: Hydrophilidae,Hydrophilinae) based on larval and adult characters,and evolutionary scenarios related to habitat shift in larvae

Abstract A phylogenetic analysis of Berosini including 15 taxa, 11 of them belonging to Berosini (ingroup), was performed. Of the 58 characters used, 32 derive from the larval stages, and 26 from the adult stage. Two well‐supported clades are recognized, one comprising Hemiosus and Berosus, and the other comprising Derallus, Regimbartia and Allocotocerus. Several larval evolutionary trends are discussed: shifts to benthic and cryptic habits, morphological modifications and adaptations related to these habits, and morphological changes of the clypeolabrum and head appendages. A comparative table for the larval stages of the five genera is included.     

Megapanorpa,a new genus with a single anal horn in males from Oriental China (Mecoptera: Panorpidae)

A new genus of Panorpidae, Megapanorpa gen. nov. , is erected for five new species, Megapanorpa grandis sp. nov. (type species), M. absens sp. nov. , M. gaokaii sp. nov. , M. jiangorum sp. nov. and M. wanghongjiani sp. nov. This genus is similar to Cerapanorpa Gao et al., 2016 in a single anal horn on the sixth tergum of male, but can be differentiated from the latter by the following characters: the seventh abdominal segment of male is constricted and stalk‐like at base, the subgenital plate of female with a pair of lateral plates, and the medigynium of female with a concealed axis. Keys to genera of Panorpidae and species of Megapanorpa gen. nov. are presented. The morphology and function of anal horns in different genera and species have been discussed.     

New data concerning postembryonic development in Antarctic Ammothea species (Pycnogonida: Ammotheidae)

In this paper, new data on larval and postlarval stages after newly collected and museum-deposited material of six Ammothea species is provided and compared with previously known information. Different developmental stages attached to the ovigerous legs of adult males for each species were found: four stages [protonymphon (Ptn), postlarval instar 1 (PL-1), postlarval instar 2 (PL-2), and postlarval instar 3 (PL-3)] for A. carolinensis; just one (Ptn) for A. clausi and A. minor; three stages (Ptn, PL-1, PL-2) for A. bicorniculata and A. spinosa; and other three (Ptn, PL-2, PL-3) for A. longispina. In the present contribution, the external morphology of each larval and postlarval instar is described, illustrated, and discussed. The larval and postlarval development of Ammothea bicorniculata, A. carolinensis, A. longispina, and A. spinosa is characterized by (1) the eggs hatch as a protonymphon larva; (2) the larvae and subsequent postlarval stages have yolk reserves and a relatively large size (0.5–0.85 mm in length for the protonymphon); (3) the postlarvae remain on the ovigerous legs of males during several moults; (4) the spinning spine is absent; and (5) the development of walking legs is sequential. The protonymphon larva of A. clausi and A. minor is the only stage on the ovigerous legs of males, and this stage is characterized by: (1) there is no yolk reserve and it has a relatively small size (0.22–0.3 mm in length); (2) the spinning spine is present; and (3) all larval appendages have a relatively large size.     

Serotonin immunoreactivity in the nervous system of the Pandora larva, the Prometheus larva, and the dwarf male of Symbion americanus (Cycliophora)

Cycliophora is a recently described phylum of enigmatic metazoans with a very complex life cycle that includes several sexual and asexual stages. Symbion pandora and Symbion americanus are the only two cycliophoran species hitherto described, of which morphological and genetic knowledge is still deficient to clarify the phylogenetic position of the phylum. Aiming to increase the database on the cycliophoran neural architecture, we investigated serotonin immunoreactivity in the free swimming Pandora larva, the Prometheus larva, and the adult dwarf male of S. americanus. In the larval forms, serotonin is mainly expressed in a ring-shaped pattern at the periphery of the antero-dorsal cerebral ganglion. Additionally, several serotonergic perikarya emerge from both sides of the cerebral ganglion. Thin neurites project anteriorly from the cerebral ganglion, while a pair of ventral longitudinal neurites emerges laterally and runs along the anterior-posterior body axis. Posteriorly, the ventral neurites fuse and extend as a posterior projection. In the dwarf male, serotonin is found mainly in the commissural neuropil of the large anterior cerebral ganglion. In addition, serotonin immunoreactivity is present in the most anterior region of the ventral neurites. Comparative analysis of spiralian nervous systems demonstrates that the neuroanatomy of the cycliophoran larval stages resembles much more the situation of adult rather than larval spiralians, which may be explained by secondary loss of larval structures and heterochronic shift of adult components into the nervous system of the Pandora and the Prometheus larva, respectively.     

The Chemoreceptive Lattice Organs in Cypris Larvae Develop from Naupliar Setae (Thecostraca: Cirripedia, Ascothoracida and Facetotecta)

Lattice organs are peculiar chemoreceptors found only in the Crustacea Thecostraca (Facetotecta, Ascothoracida, Cirripedia). In these taxa, five pairs occur in the head shield (carapace) of the terminal larval instar (y-cyprid, ascothoracid larva, cyprid), which is the settlement stage. Lattice organs represent an autapomorphy for the Thecostraca but their evolutionary origin and possible homologues in other Crustacea remain obscure. We have used scanning electron microscopy to describe the setation pattern of the head shield in late nauplii of one species of Ascothoracida, one species of Facetotecta and several species of the Cirripedia Thoracica, Acrothoracica, and Rhizocephala. The naupliar head shield always carries two pairs setae situated anteriorly near the midline. Each of these setae carry a single pore, and positional, structural and ontogenetic evidence show that these setae are homologous in all the examined species and that they represent precursors of the two anterior pairs of lattice organs of the succeeding larval stage, viz., the ascothoracid larva (Ascothoracida), y-cyprid (Facetotecta), and cyprid (Cirripedia). This leads us to infer that lattice organs are among the most highly modified sensilla in all Crustacea and they have in most cases lost all external resemblance to a seta. The nauplii of the Rhizocephala carry an additional three pairs of setae situated more posteriorly on the head shield and they could be precursors of the three posterior pairs of lattice organs. All other species examined lack these posterior setae, except the Facetotecta which have one posteriorly situated pair.     

Expression of synapsin and co-localization with serotonin and RFamide-like immunoreactivity in the nervous system of the chordoid larva of Symbion pandora (Cycliophora)

Abstract. Cycliophora is one of the most recently described metazoan phyla and hitherto includes only two species: Symbion pandora and Symbion americanus . With a very complex life cycle, cycliophorans are regarded as an enigmatic group with an uncertain phylogenetic position, although they are commonly considered lophotrochozoan protostomes. In order to extend the database concerning the distribution of immunoreactive substances in the free-swimming chordoid larva of S. pandora , we investigated synapsin immunoreactivity using fluorescence-coupled antibodies in combination with confocal laserscanning microscopy. Moreover, we analyzed the co-localization patterns of synapsin, serotonin, and RFamide-like immunoreactivity in the chordoid larva by 3D imaging technology based on the confocal microscopy image stacks. Synapsin is expressed in large parts of the bilobed anterior cerebral ganglion including anterior and dorsal projections. Two pairs of ventral neurites run longitudinally into the larval body of which the inner pair shows only weak, scattered synapsin immunoreactivity. In addition, a lateral synapsin immunoreactive projection emerges posteriorly from each ventral longitudinal axon. Double immunostaining shows co-localization of synapsin and serotonin in the cerebral ganglion, the outer and the inner ventral neurites, and the anterior projections. Synapsin and RFamide-like immunoreactivity co-occur in the cerebral ganglion, the outer ventral neurites, and the dorsal projections. Accordingly, the cerebral ganglion and the outer ventral neurites are the only neural structures that co-express the two neurotransmitters and synapsin. The overall neuroanatomical condition of the cycliophoran chordoid larva resembles much more the situation of adult rather than larval life cycle stages of a number of spiralian taxa.