Diversity of aquatic and riparian Coleoptera of the Philippines: checklist,state of knowledge,priorities for future research and conservation

The diversity of the aquatic and strictly riparian beetles of the Philippines is reviewed. A checklist (with 317 species and subspecies) of the aquatic and riparian Coleoptera of the Philippines is provided for the first time herein. This checklist covers the following families (number of species and subspecies in parentheses): Hydroscaphidae (1), Gyrinidae (15), Dytiscidae (65), Noteridae (4), Haliplidae (2), Hydraenidae (15), Hydrochidae (2), Spercheidae (1), Hydrophilidae (72), Scirtidae (49), Elmidae (23), Dryopidae (2), Psephenidae (16), Eulichadidae (1), Limnichidae (20), Heteroceridae (5), Malachiidae (5), Lampyridae (15), Nitidulidae (4). A total of 201 species and subspecies (63%) are considered to be endemic to the Philippines. Aquatic/riparian beetle genera and families that are still unrecorded but likely to occur in this country are briefly discussed. Based on estimates at the genus level, it is concluded that about 850 aquatic and riparian species can be expected to occur in the Philippines. The implication of the Philippine biogeographic history for this diversity is discussed. The potential of aquatic/riparian beetles as bioindicators is briefly examined. The dilemma of aquatic invertebrate conservation is analysed and priorities for future research are outlined.     

The systematic position of Aspidytidae, the diversification of Dytiscoidea (Coleoptera, Adephaga) and the phylogenetic signal of third codon positions

Characters of the newly discovered larvae of the South African Cliff Water Beetle Aspidytes niobe were examined and integrated into a data matrix including all families of Dytiscoidea as well as Haliplidae. Fifty-three morphological characters of adults and larvae were analysed separately and combined with molecular data from six nuclear and mitochondrial genes. The phylogeny of the group is reconstructed for the study of the evolution of swimming behaviour and larval feeding habits, as well as the shift in diversification rates leading to the two most speciose lineages. The parsimony analysis of all equally weighted morphological and molecular characters combined resulted in a single well supported tree with the topology (Noteridae (Hygrobiidae ((Aspidytidae, Amphizoidae) Dytiscidae))), in agreement with the molecular data alone, but in contradiction to the morphological data, which favoured a topology in which Hygrobiidae is sister to Dytiscidae. The exclusion of third codon positions of the three protein coding genes resulted in a topology identical to that obtained with the morphological data alone, but the use of Bayesian probabilities or the amino acid sequence resulted in the same topology as that of the tree obtained with parsimony using all equally weighted characters. We concluded that interactions of third codon positions with the other data are complex, and their removal is not justified. There was a significant increase in the diversification rate at the base of the richest families (Noteridae and Dytiscidae), which could be associated with the development of simultaneous stroke and higher swimming performance, although data on the swimming behaviour of some basal groups of Noteridae are incomplete. The presence of larval mandibular sucking channels may have contributed to the diversification of Dytiscidae and the species-rich noterid genera Hydrocanthus and Canthydrus .     

Atlas of olfactory organs of Drosophila melanogaster: 1. Types,external organization,innervation and distribution of olfactory sensilla

The third antennal segment (funiculus) and the maxillary palp are the main and accessory olfactory sense organs of Drosophila melanogaster. Cryofixed antennae and palps revealed a superior preservation of the sensory dendrites and other cellular details as compared to conventional chemical fixation. Extensive cross-section series through funiculus and palp were studied in order to obtain as complete an evaluation as possible of the sensillar complement on these appendages. About 75% of all sensilla on the male and female funiculus were individually studied and their position on the antennal surface mapped. Dimensions of the cuticular apparatus of the various types of sensilla are provided as well as the number of innervating receptor neurons. Particular attention has been paid to the cuticular pores important for stimulus transport and to the sensory dendrites. On the funiculus surface, all sensilla have multiple wall pores: sensilla (s.) trichodea and s. basiconica are single-walled, s. coeloconica are double-walled. The distribution of s. trichodea and s. basiconica follows opposing gradients along a diagonal axis parallel to the axis of the arista from proximo-medial to disto-lateral. In this disto-lateral direction the density of s. trichodea increases while that of the s. basiconica decreases. S. trichodea occur in three subtypes with one, two or three receptor neurons. Basiconic sensilla can be subdivided into three subtypes of large s. basiconica (with two or four receptor neurons), three subtypes of thin s. basiconica (with mostly two, rarely four neurons), and one subtype of small s. basiconica with two receptor neurons. Large s. basiconica occur only in the most proximal region (the ‘LB-zone’); thin s. basiconica are most common in a belt that borders the LB-zone distally, while small s. basiconica are most numerous even further distally along the mentioned diagonal axis in between the s. trichodea. S. intermedia are single-walled, multiporous sensilla which combine features of s. trichodea and s. basiconica; they are found in two subtypes with two or three receptor neurons, in the same region where s. trichodea abound. The s. coeloconica are irregularly distributed over the funicular surface, and occur in two subtypes with two or three receptor neurons. Sexual dimorphism on the antenna is moderate, the female funiculus is a bit longer and carries a larger number of small s. basiconica and large s. basiconica of the LB-I subtype; the male funiculus, however, has more s. trichodea than the female. On the maxillary palp, besides mechanoreceptive s. chaetica, there are only s. basiconica with two receptor neurons. According to the fine structure of their sensory dendrites, three subtypes can be discriminated which are distributed in a random pattern. The functional significance of the described structures and distribution awaits future immunocytochemical and electrophysiological experiments.     

松褐天牛触角感器电镜扫描和触角电位反应

应用扫描电镜对松褐天牛的触角感器进行扫描观察,并测定了触角对松树主要挥发物的电生理反应.松褐天牛触角上共有7种感器,即毛形感器、锥形感器、指形感器、棒形感器、瓶形感器、芽形感器以及刺形感器,以毛形和锥形感器的数量最丰富.其中毛形感器和锥形感器各有3个型,指形感器有2个型.毛Ⅱ、毛Ⅲ型和锥Ⅰ、锥Ⅱ型感器表皮结构具有深的纵刻纹,具有嗅觉感器的典型特征.同时对各种感器的数量、长度及分布特点等作了描述.电生理研究结果表明,几种供试化合物及其混合物均能引起松褐天牛雌、雄成虫较强的电生理反应;松褐天牛触角不同部位的EAG反应存在显著差异;剂量试验表明,松褐天牛成虫触角对不同浓度的α-Pinene的EAG反应存在一定的规律性.     

Comparative study of sensilla and other tegumentary structures of Myrmeleontidae larvae (Insecta,Neuroptera)

Antlion larvae have a complex tegumentary sensorial equipment. The sensilla and other kinds of larval tegumentary structures have been studied in 29 species of 18 genera within family Myrmeleontidae, all of them with certain degree of psammophilous lifestyle. The adaptations for such lifestyle are probably related to the evolutionary success of this lineage within Neuroptera. We identified eight types of sensory structures, six types of sensilla (excluding typical long bristles) and two other specialized tegumentary structures. Both sensilla and other types of structures that have been observed using scanning electron microscopy show similar patterns in terms of occurrence and density in all the studied species (with few exceptions). The sensilla identified are: coeloconica, placoidea, basiconica, trichodea type I, trichodea type II, and campaniformia. All these sensilla have mechano- or chemosensorial functions. Some regions of the larval body have been studied using SEM for the first time, such as the surface of the food canal, which bears sensilla coeloconica, and the abdominal segment X, that bears three types of sensilla: coeloconica, basiconica, and campaniformia. Sensilla placodea are newly reported on antlion larvae, being present on the mandibular base, pronotum, mentum, and cardum. Also, new locations of sensilla coeloconica (e.g., on rastra) and sensilla campaniformia (e.g., on odontoid processes) are noted. A novel porous texture with chemoreceptor function has been identified in the base of mandibles. A mechanism of dentate-notched surfaces that anchor maxillae and mandible, reinforcing the food canal, is detailed. All these sensorial structures, in addition to ocular tubercles for light caption and their great muscular system, confer to these larvae an extraordinary predation capacity to success hunting and living in such harsh environments.     

小菜蛾成虫喙管感器的超微结构及其神经元中枢投射

【目的】明确小菜蛾Plutella xylostella成虫喙管感器的形态结构及感器神经元的投射。【方法】利用扫描电子显微镜观察小菜蛾成虫喙管结构和感器,利用神经回填技术和激光共聚焦显微镜观察喙管感器神经元在脑部的投射。【结果】小菜蛾成虫喙管上存在毛形感器(两种亚型)、腔锥形感器、锥形感器、刺形感器和栓锥形感器5种不同类型的感器。毛形感器表面光滑,分布于外颚叶外侧,可分为毛形感器Ⅰ型和Ⅱ型两种亚型,其中Ⅰ型比Ⅱ型长;锥形感器分布于喙管外表面,由一个感觉锥和一个短的圆形基座组成;腔锥形感器仅分布于食管内侧,只有一个粗短感觉锥而无基座;刺形感器由一个细长的感觉毛和一个圆形基座组成,表面无孔,分布于喙管的外表面;栓锥形感器是昆虫喙管上最典型的感受器,集中分布于喙管顶端区域,感器顶部凹腔伸出一个单感觉锥。此外,喙管上的感觉和运动神经元投射到初级味觉中枢咽下神经节。【结论】本研究阐明了小菜蛾成虫喙管感器的类型、分布和形态特征及其感器神经元在脑部的投射形态,为深入了解小菜蛾喙管感器的生理和功能奠定了基础。     

访花昆虫红腹毛蚊触角感器扫描电镜观察

【目的】本研究旨在明确访花昆虫红腹毛蚊Bibio rufiventris触角感器的类型和分布。【方法】通过扫描电镜(scanning electron microscope, SEM)观察红腹毛蚊雌、雄成虫触角感器的种类、数量和形态,比较雌雄个体间的差异。【结果】红腹毛蚊雌、雄成虫触角均包含3部分,分别为柄节、梗节和鞭节,其中鞭节由8个亚节组成。雌成虫触角平均总长度为862.556±78.662μm,雄成虫触角平均总长度为880.361±83.253μm,雌、雄成虫触角各亚节的长度几乎相似,只有鞭节第8亚节长度有显著差异。红腹毛蚊的触角感器共有4大类,即刺形感器、锥形感器、毛形感器和B?hm氏鬃毛。其中,雌性红腹毛蚊触角感器共有6种亚型,即刺形感器、毛形感器2型、锥形感器(1, 2和4型)和B?hm氏鬃毛;雄性红腹毛蚊触角感器共有5种亚型,即刺形感器、毛形感器1型、锥形感器(2和3型)和B?hm氏鬃毛。【结论】红腹毛蚊雌、雄成虫触角感器在种类、数量以及形态特征方面存在一定差异。本研究为进一步探究红腹毛蚊触角感器的生理功能及其行为活动的分子机制提供了形态学基础。     

疟疾媒介中华按蚊触角感器的扫描电镜观察

【目的】明确中华按蚊Anopheles sinensis雌成虫与幼虫触角感器的类型、形态和分布。【方法】利用光学显微镜观察中华按蚊成虫与幼虫触角的形态结构,利用扫描电镜观察触角上的感器类型、形态和分布。【结果】中华按蚊雌成虫触角由柄节、梗节和鞭节组成,鞭节有13个亚节。触角上共发现4种类型的感器,分别为毛形感器(锐型和钝型)、刺形感器(大型和小型)、锥形感器(Ⅰ型和Ⅱ型)和腔锥形感器(大型和小型)。雌成虫触角各类感器总计约1 135.67±86.75个,其中毛形感器数量最多(662.00±6.22个),随后是刺形感器(294.67±33.35个)和锥形感器(146.00±42.39个),腔锥形感器数量最少(36.50±5.90个)。毛形感器、刺形感器和锥形感器在鞭节的每个亚节均有分布,而大型腔锥形感器在第9-11亚节没有分布,小型腔锥形感器仅分布于第13节的顶端。幼虫触角的鞭节不分亚节,呈管状,触角末端有一个感觉锥,鞭节上分布有与成虫锥形感器相似的锥形凸起,初步定名为类锥形感器,其数量和大小随幼虫龄期的增长而显著增加,锥体表面的凹槽越来越明显,其功能还需要通过超微结构和电生理等研究进一步确定。【结论】本研究对中华按蚊幼虫和雌成虫触角感器的形态特征、类型、数量及其分布进行了观察和分析,结果为进一步研究中华按蚊感器的生理功能奠定了基础。     

Development of the sensory system in larvae and pupae of Chaoborus crystallinus (DeGeer, 1776; Diptera, Chaoboridae): sensory cells, nerves and ganglia of the tail region

Using various microscopical techniques, we have studied changes in the sensory equipment and architecture of the peripheral nervous system (PNS) around the first metamorphic molt from larva to pupa in the phantom midge Chaoborus. The transparent larvae and pupae of this dipteran with ancestral features allow us to investigate sensilla and their central projections from whole-mount preparations of complete groups of segments. Each sensillum on the posterior larval and pupal segments was identified using its external shape and position, and the morphology of the abdominal ganglia and segmental nerves was investigated. In addition, retrograde fills with the carbocyanine dye DiI were used to trace the axonal paths of most of the extero- and proprioreceptors. These findings were combined to produce maps of the sensory elements of larval and pupal abdomens that were analyzed at three levels: seriality (homonomy), ontogenetic changes of individual sensilla, and homology of the PNS between different species. Comparison of different segments shows for both stages that primarily there is a homonomous basic design of the PNS, but segment-specific modifications are evident in segments 8-10. Comparison of corresponding larval and pupal segments shows that many sensilla retain their internal structure and axonal projections. However, their external cuticular parts are changed in relation to the different life habits of larvae and pupae. Furthermore, some sensilla are completely reduced during the pupal molt, especially those of the tenth segment which appears as a distinct larval structure (caenogenesis). Comparison between species indicates that despite the varying types of sensilla their basic segmental arrangement and their axonal trajectories are conserved.     

Morphology of antennal,maxillary palp and labial palp sensilla in different larval instars of the Asian long‐horned beetle,Anoplophora glabripennis (Motschulsky) (Coleoptera: Cerambycidae)

The Asian long‐horned beetle (ALB) is one of the most important international trunk‐boring insect pests of forest trees, which has caused enormous losses in the forests of China, the USA and some countries in Europe. To investigate the morphology of the antennal, maxillary palp and labial palp sensilla in the five larval instars of the ALB, scanning electron microscopy was used to determine the typology, morphology, number and distribution of the sensilla. The antennae had two types of sensilla: the sensilla twig basiconica (s.t.b.) and sensilla basiconica. Three different types of sensilla were observed on the maxillary palps, which were the sensilla digitiformia, the sensilla chaetica and the s.t.b. The labial palps had two types of sensilla: the sensilla chaetica and the s.t.b. The s.t.b. had seven total subtypes, and the sensilla basiconica had two subtypes. The typology, quantity and distribution of sensilla on the antennae, maxillary palps and labial palps remained constant in the five larval instars, but the dimensions of all sensilla types increased throughout the development. We discussed the mechanoreception and the chemoreception of ALB larvae inside the host tree to provide a theoretical understanding and information for further research on the behaviour and the electrophysiology of this devastating forest pest.     

Comparative morphology of sensilla on antenna,maxillary palp and labial palp of larvae of Eucryptorrhynchus scrobiculatus (Olivier) and E. brandti (Harold) (Coleoptera: Curculionidae)

Eucryptorrhynchus scrobiculatus (Olivier) and E. brandti (Harold) are two wood boring pests of Ailanthus altissima (Mill.) Swingle (tree of heaven) and the variety Ailanthus altissima var. Qiantouchun. These beetles attack healthy trees and bore into roots and trunks during the larval stage. We studied the typology, distribution and morphostructure of the sensilla on the antennae, maxillary palps and labial palps of E. scrobiculatus and E. brandti larvae using scanning and transmission electron microscopy. The results showed the following: (i) the antennae of the two weevil larvae had two types of sensilla, sensilla basiconica (S.b.1 and S.b.2) and sensilla twig basiconica (S.tb.1‐S.tb.3), with S.tb.4 observed only on the antennae of E. brandti larvae; (ii) the maxillary palps had three types of sensilla, S.b.2, S.tb. (S.tb.2, S.tb.3 and S.tb.5) and digitiform sensilla; (iii) the labial palps had two types of sensilla, S.b.2 and S.tb. (S.tb.2, S.tb.3 and S.tb.5); (iv) the quantity and distribution of sensilla on the antennae, maxillary palps and labial palps remained constant between E. scrobiculatus and E. brandti larvae; and (v) sensilla basiconica had distinct sidewall pores, an apical pore was observed on sensilla twig basiconica, and digitiform sensilla were oval in shape, with a distinct apical pore. Based on the microstructure of the cuticle wall and dendrite, we hypothesized that these sensilla functioned as olfactory, gustatory and hygro‐/thermo‐receptors, respectively. We discuss the relationships among types of sensilla and the types of damage caused by the larvae inside the host tree to understand olfactory and gustatory receptor mechanisms. The results of this study will provide a firm basis for future electrophysiological studies.     

Fine structure of the antennal sensilla of the millipede Orthomorphella pekuensis (Polydesmida: Paradoxosomatidae)

The fine structural characteristics of various sensory receptors on the antenna of a millipede, Orthomorphella pekuensis, were observed with field emission scanning electron microscopy. The antenna of this millipede has eight segments, called articles. On the surface of the antenna, there are a variety of sensory receptors, including olfactory and mechanical receptors. According to their morphological and fine structural characteristics, we could identify four basic types of antennal sensillum: chaetiform sensilla (CS), trichoid sensilla (TS), basiconic sensilla (BS) and apical cone sensilla (AS). The BS are divided further into three subtypes: large basiconic sensilla (BS₁) on the 5th and 6th articles; small basiconic sensilla (BS₂) on the 5th article; and a distinct type of basiconic spiniform sensilla (BS₃) on the 7th article. The most prominent sensilla are four large AS on the distal tip of the 8th segment. Based on our results, we conclude that the main function of the CS and TS are related to mechanical reception, and that the BS and AS are likely to function in olfactory reception of volatile odors of plants, as these sensilla have base and apex pores, respectively.     

Morphology of the second- and third-instar larvae of Dermatobia hominis by scanning electron microscopy

Larvae of Dermatobia hominis 10–27 days old were collected from experimentally infected rats and their morphology was studied by scanning electron microscopy. The moult from the second to third instar occurs at 18 days, with emergence from the host at 30 days post-infection. The second-instar larvae bear on the pseudocephalon, antennae (coeloconic sensilla), and coeloconic and basicoconic sensilla on the maxillary sensory complex. The thoracic segments bear small backwardly-directed spines anteriorly and ventral trichoid and campaniform sensilla. The first four abdominal segments have small and large backwardly-directed spines that are absent on segments five and six. The seventh and eighth abdominal segments have medium-sized forwardly-directed spines. Abdominal segments are encircled by campaniform sensilla. The terminal end of the eighth abdominal segment bears the anus, prominent anal lobes and two spiracular openings on each spiracular plate. Spiracular plates show a radial sun ray pattern. The rear abdomen also bears an ecdysal aperture, several pores and eight coeloconic sensilla. Although there are slight morphological differences, the spines (predominantly flat and thorn-like) and sensilla (campaniform and coeloconic) of the third-instar larvae show a similar arrangement to that of second-instar larvae. Thoracic trichoid sensilla are not seen in third-instar larvae. A perispiracular gland aperture is situated above each posterior spiracular opening. These morphological features are compared with those of other cuterebrid larvae.     

Cuticular sensilla on newly hatched larvae of Gasterophilus intestinalis and Oestrus ovis

Abstract. Cuticular sensilla on newly hatched larvae of Gasterophilus intestinalis De Geer (Diptera: Gasterophilidae) and Oestrus ovis (L.) were studied by scanning electron microscopy. Two types of trichoid sensilla, two types of coeloconic sensilla and a pit sensillum were present on the thoracic and abdominal segments of G. intestinalis larvae. Sensilla on larvae of O. ovis were similar although only one type of trichoid sensillum was present. Total number of sensilla were higher for O. ovis than for G. intestinalis (248 v . 214). Variation in numbers of sensilla is consistent with the concept that increasing numbers of sensilla are associated with increasingly complex searching behaviour required to locate suitable habitats for development.     

苹果蠹蛾头部感器的电镜扫描结构

本文使用扫描电镜系统观察并描述了苹果蠹蛾Cydia pomonella(L.)成虫及幼虫触角上及口器上的感器。主要研究结果如下:1)苹果蠹蛾成虫触角背面密布鳞片,感器很少,腹面和侧面鳞片稀疏,具大量感器;2)触角上的感器大部分分布于鞭节各节,少部分分布于柄节和梗节;3)雄虫触角上着生10种感器,雌虫触角上着生9种感器,与雄虫相比,雌虫缺少鳞形感器;4)雄虫口器具鳞形感器和刺形感器,雌虫口器仅具刺形感器;5)幼虫触角3节,基节无感器,第2节具2刺形感器且其分布位置存在个体差异,端节端部具呈三角状排列的3个栓锥感器;6)幼虫口器具一定数量和不同形态的感器。     

Structure and sensilla of the antennae and mouthparts of Loxocephala perpunctata Jacobi (Hemiptera: Fulgomorpha: Eurybrachidae)

The ultramorphology of the antennae and mouthparts of the adult Loxocephala perpunctata Jacobi was studied through a scanning electron microscope. Seven types of sensilla were found on antennomeres, including a Böhm bristle on the scape, sensillum trichoideum and plaque organ on the pedicel, two subtypes of sensilla chaetica and two subtypes of sensilla campaniformia on these two antennomeres; and Bourgoin's organ with sensory pegs and sensilla basiconicum on the basal bulb of the flagellum. The mouthparts of L. perpunctata are of the typical piercing-sucking type, similar to mouthparts found in other hemipteran insects. In general, six types of sensilla (i.e., four subtypes of sensilla chaetica, sensillum basiconicum, subapical labial sensillum, uniporous peg-like sensillum, multiporous peg-like sensillum and two subtypes of bristle-like sensilla) were detected on different locations of the labium, with the last three, and numerous cuticular processes, present on the labial tip. The potential functions of these sensilla are discussed.     

Morphology and ultrastructure of the chemosensory sensilla of larval antennae in the Colorado potato beetle, Leptinotarsa decemlineata Say (Coleoptera: Chrysomelidae)

The allocation, external morphology, and ultrastructure of various types of sensilla on the antennae have been studied in the larvae of the Colorado potato beetle (Leptinotarsa decemlineata Say) using scanning and transmission electron microscopy. It has been found that the larval antenna bears 11 sensilla of four morphological types: trichoid (five), basiconic (four), styloconic (one), and antennal cone (one). It has been shown that the sensilla are innervated by 2?C7 receptor cells: trichoid, by 2?C4 receptor cells; basiconic, by 4?C7 receptor cells; and styloconic, by 6 receptor cells. Judging by the specifics of their ultrastructural organization, the function of these sensilla is gustatory. The antennal cone is innervated by two receptor cells, and its function is olfactory.     

Hygroreceptors in the larva of Libellula depressa (Odonata: Libellulidae)

Ultrastructural and electrophysiological (single-cell recordings) investigations were carried out on the coeloconic sensilla borne by the apical antenna of the larvae of Libellula depressa (Odonata: Libellulidae). These sensilla appear as pegs located in pits. One of them is a compound sensillum constituted of two fused pegs in a common pit and the other two are single pegs located in separated pits close to each other. Coeloconic sensilla show position and ultrastructural details very similar to those described in insect hygroreceptors. The electrophysiological recordings on the apical antennae of the last larval instar of L. depressa clearly show the presence of moist and dry cells responding antagonistically to humidity changes. This study gives the first evidence of hygroreceptors in dragonfly larvae and represents the first electrophysiological approach to larval sensilla of aquatic insects. The presence of hygroreceptors in L. depressa larvae is in agreement with the hygropositive response shown by these insects in laboratory and field behavioural experiments.