Head Capsule Research Articles

Factors responsible for estivation in spotted stem borer, Chilo partellus (Swinhoe)

Diapause is an important adaptive trait in certain groups of insects at a particular stage, which contribute in controlling insect ecology, phenology, and physiology. Considering the importance of various factors in appropriate combinations, we studied effects of different temperature and photoperiod conditions on larval morphometrics, development, induction, and termination of estivation, and overall mortality due to estivation in Chilo partellus (Swinhoe). Weight, length, and head capsule width of C. partellus larvae significantly increased with increase in temperature and photoperiod treatments upto certain age and duration of exposure. According to Dyar's ratio, overall head capsule width progressed geometrically with minimum constant ratio of 1.20, and second-degree polynomial regression equations were best fitted to all the test treatment conditions. The mortality of diapausing C. partellus individuals varied between 7.0% and 37.3% under different constant and varying temperature and photoperiod regimes, which at 38°C + 14L:10D reached upto 100.0%. However, the exposure to 30°C + 12.5L:11.5D and 32°C + 13L:11D along with dry food accelerated the process of estivation, wherein the diapause larval survival was significantly higher and overall mortality of individuals due to diapause was lower suggesting these conditions appropriate for induction of estivation in C. partellus. These studies have implications for better understanding the bioecology and population regulation system under varying climatic conditions and devising novel strategies for management of C. partellus.

Morphology of the tentorium in the ant genus Lasius Fabricius (Hymenoptera: Formicidae)

The tentorium is the internal skeleton of the head capsule of insects. Several studies have shown that the structure of the tentorium is an important factor not only for the morphology and systematics but also for the phylogeny and evolution. In ants, however, only three studies have reported tentorial morphology so far. We reveal the fundamental structure of the tentorium of the genus Lasius (Hymenoptera, Formicidae) and its minor variation among six species of the genus. Based on the results, we give new terminologies of the organ, presenting a schematic diagram of the tentorium. We clarify muscle attachment to the tentorium by constructing a three-dimension image of the tentorium and muscles. We then verify the attachment areas of the antennal muscles and maxillary adductor muscles on the tentorium. The results show that the muscular attachment areas are broader than previously thought. Our study indicates that the key to understanding the evolution of the tentorium is its functional morphology, in relation to the attachments of the muscles originating from the tentorium within the head capsule. This is the first report of the three-dimensional images of the ant tentorium and its attached muscles. The intra- and inter-specific variations of the tentorium is also reported for the first time.

Recognition of Five Larval Instars in Conotrachelus perseae (Coleoptera: Curculionidae)

Head capsule widths of Conotrachelus perseae Barber (Coleoptera: Curculionidae) larvae extracted from avocado, Persea americana Mill. cv. Fuerte, were measured by computerized imagery. Head capsule widths ranged from 212 to 1,562 µm and, according to an analysis of frequency distribution using the HCAP program and Dyar's rule, formed five well-defined modal groups which suggests five larval instars. The mean head capsule widths were 264, 428, 642, 1,009, and 1,493 µm for the first through the fifth instars, respectively, with a probability of misclassification of instar below 3%.

Aparatermes thornatus (Isoptera: Termitidae: Apicotermitinae), a New Species of Soldierless Termite from Northern Amazonia

The worker of Aparatermes thornatus is described as the fourth species of a widespread Neotropical termite genus, Aparatermes, from the worker caste. This species occurs in northern Amazonia. The enteric valve armature of A. thornatus and the unique coloration of its worker's head capsule distinguish it from its 3 congeners. Molecular data recovered the proposed new species as a sister to Aparatermes silvestrii. Type material is deposited at Coleccion Entomologica Forestal, Universidad Distrital “Francisco Jose de Caldas” (CEFUDFJC - 45 National Registry of Collections) in Bogota, Colombia, and the University of Florida, Termite Collection (UFTC) at the Fort Lauderdale Research and Education Center, Fort Lauderdale, Florida, USA.

Still greater disparity in basal planthopper lineage: A new planthopper family Yetkhatidae (Hemiptera, Fulgoromorpha, Fulgoroidea) from mid-Cretaceous Myanmar amber

A new family Yetkhatidae Song, Szwedo et Bourgoin fam. nov. from mid-Cretaceous amber of Myanmar is described. It is characterized by the tegmina with a triangular basal cell and in having the apex of the clavus closed, the female genitalia with robust endogonocoxal processes, the head capsule with trigones and foliated frontal margins, ridged lora (visible frontally) and by intermediate carinae on the mesonotum. Two new monotypic genera are described in this new family: Yetkhata Song, Szwedo & Bourgoin gen. nov., with Yetkhata jiangershii Song, Szwedo & Bourgoin sp. nov. and Parwaina Song, Szwedo & Bourgoin gen. nov. with Parwaina liuyei Song, Szwedo & Bourgoin sp. nov. A key to recognition of these genera is provided, and their morphological characters and disparity are briefly discussed.

The larva of Plectrocnemia renetta Malicky 1975 (Trichoptera, Polycentropodidae), including a discriminatory matrix to the larvae of Plectrocnemia Stephens 1836 species of Greece.

This paper describes the previously unknown larva of Plectrocnemia renetta Malicky 1975. Information on the morphology of the final instar larva is given and the most important diagnostic features are illustrated. A preliminary discriminatory matrix for the Greek larvae of Plectrocnemia Stephens 1836 is also provided. Plectrocnemia renetta and P. conspersa conspersa (Curtis 1834) belong to the group where the inner and outer dorsal secondary setae on abdominal segment IX are strongly different in length. These two species can be separated from each other by the arrangement of muscle attachment spots on the head capsule, number and length of setae on abdominal sternum IX, and by distribution patterns. With respect to zoogeography, Plectrocnemia renetta has been reported from Cyprus, Turkey, and from the Greek islands of Ikaria and Samos.

Notes on the first instar larvae of the genus Tipula (Diptera: Tipulidae).

The egg-larvae of 890 individuals in 8 species belonging to long palped crane flies of the genus Tipula (Diptera: Tipulidae) were obtained from 17 gravid females captured in Lithuania in 2011-2012. Depending on the species, it took from four days to more than eleven weeks for the eggs to hatch. Of the four instars of larvae for crane flies, the second, third and the last instar larvae are very similar in morphology, while the first instar or egg-larva differs radically. Descriptions and illustrations of the external morphology, chaetotaxy of abdominal segments, characters of head capsules and last abdominal segments are given for the previously unknown first instar larvae of T. (Lunatipula) fascipennis, T. (Pterelachisus) irrorata, T. (P.) pabulina, T. (P.) pseudovariipennis, T. (Yamatotipula) pruinosa and poorly known T. (Beringotipula) unca, T. (L.) vernalis T. (Tipula) paludosa. The differences of the head capsule and last abdominal segment among the first instar larvae of these species are more obvious than in the last instar. Characters of the head capsule such as the shape of the teeth of the mandible, shape of the basal segment of the antenna and number of sensillae on it, shape of hypostomium, arrangement of sensory structures on the labrum and frons, all differ among egg-larvae of Tipula. Sclerotisation and the arrangement of setae on the spiracular field are unique for each studied Tipula species. This is the first study to use head capsule characters to distinguish the larvae of Tipula.

Microplitis bicoloratus bracovirus modulates innate immune suppression through the eIF4E–eIF4A axis in the insect Spodoptera litura

Eukaryotic initiation factor 4E (eIF4E) is regulated during the innate immune response. However, its translational regulation under innate immune suppression remains largely unexplored. Microplitis bicoloratus bracovirus (MbBV), a symbiotic virus harbored by the parasitoid wasp, Microplitis bicoloratus, suppresses innate immunity in parasitized Spodoptera litura. Here, we generated eIF4E dsRNA and used it to silence the eIF4E gene of S. litura, resulting in a hallmark immunosuppressive phenotype characterized by increased apoptosis of hemocytes and retardation of head capsule width development. In response to natural parasitism, loss of eIF4E function was associated with similar immunosuppression, and we detected no significant differences between the response to parasitism and treatment with eIF4E RNAi. Under MbBV infection, eIF4E overexpression significantly suppressed MbBV-induced increase in apoptosis and suppressed apoptosis to the same extent as co-expression of both eIF4E and eIF4A. There were no significant differences between MbBV-infected and uninfected larvae in which eIF4E was overexpressed. More importantly, in the eIF4E RNAi strain, eIF4A RNAi did not increase apoptosis. Collectively, our results indicate that eIF4E plays a nodal role in the MbBV-suppressed innate immune response via the eIF4E–eIF4A axis.

Effects of flood disturbance on the cohort structure of the riverine mayfly Ephemera orientalis (Ephemeroptera: Ephemeridae) in the Han River in Seoul, Korea

AbstractThe aim of the present study was to investigate the adaptation of coexisting Ephemera orientalis (Ephemeroptera: Ephemeridae) cohorts to flood timing. Larvae were collected from the Han River (Seoul, Korea) from 2012 to 2014, and their life history was assessed by measuring head capsule width and by comparing the emergence period of the insects in relation with the flood season. Two slow‐growing cohorts (S1 and S2) and one fast‐growing cohort (F) were identified. The S1 cohort was found to hatch in June or July and to emerge the following summer. Whereas, the S2 cohort was found to hatch in August or September and emerge the following August. Early‐hatching larvae from the S1 cohort became the F cohort through the process of cohort splitting and subsequently merged with the S2 cohort. The F cohort, however, was not observed when huge floods occurred or under low temperature conditions. Furthermore, floods that resulted from >80 mm precipitation significantly reduced larval abundance; therefore, flooding affects cohorts both directly and indirectly. Most cohorts emerged before and after the flood season (July 10 to August 10). The results of the present study indicate that the emergence period of E. orientalis has evolved to avoid the flood season.

Evolutionary history of Polyneoptera and its implications for our understanding of early winged insects

Polyneoptera represents one of the major lineages of winged insects, comprising around 40,000 extant species in 10 traditional orders, including grasshoppers, roaches, and stoneflies. Many important aspects of polyneopteran evolution, such as their phylogenetic relationships, changes in their external appearance, their habitat preferences, and social behavior, are unresolved and are a major enigma in entomology. These ambiguities also have direct consequences for our understanding of the evolution of winged insects in general; for example, with respect to the ancestral habitats of adults and juveniles. We addressed these issues with a large-scale phylogenomic analysis and used the reconstructed phylogenetic relationships to trace the evolution of 112 characters associated with the external appearance and the lifestyle of winged insects. Our inferences suggest that the last common ancestors of Polyneoptera and of the winged insects were terrestrial throughout their lives, implying that wings did not evolve in an aquatic environment. The appearance of the first polyneopteran insect was mainly characterized by ancestral traits such as long segmented abdominal appendages and biting mouthparts held below the head capsule. This ancestor lived in association with the ground, which led to various specializations including hardened forewings and unique tarsal attachment structures. However, within Polyneoptera, several groups switched separately to a life on plants. In contrast to a previous hypothesis, we found that social behavior was not part of the polyneopteran ground plan. In other traits, such as the biting mouthparts, Polyneoptera shows a high degree of evolutionary conservatism unique among the major lineages of winged insects.

Life Cycle and Development of Coptoborus ochromactonus (Coleoptera: Curculionidae: Scolytinae), a pest of balsa.

Coptoborus ochromactonus Smith and Cognato is a recently described xyleborine ambrosia beetle pest associated with balsa, Ochroma pyramidale (Cav. Ex Lam.) Urb., in Ecuador. This pest has caused significant loss of cultivated balsa in Ecuador, but little is known of its biology and ecology. Based on examination of multiple gallery systems, this study describes the gallery pattern and life cycle of C. ochromactonus and confirms the generic identity of the symbiotic fungus. Females initiated attack, excavating a gallery perpendicular to the bole. The primary tunnel branched into a secondary tunnel at a mean 3.13 mm. This first secondary tunnel was excavated in a horizontal plane between the phloem and xylem for a mean 18.0 mm. Along its length, two tertiary tunnels were constructed on each side, measuring a mean 26.0 and 20.0 mm, respectively. Conidiophores and conidia of an unidentified Fusarium sp. grew on the tunnel walls and were fed upon by the beetle and her progeny. Coptoborus ochromactonus exhibits sexual dimorphism. The female is 2.6 mm long and possesses a round pronotum, whereas the male is 2.2 mm long and has a quadrate pronotum. The developmental stages last a mean 6, 14, 6, 4 and 30 d for egg, larva, pupa, teneral adult, and adult, respectively, when reared at 25°C. Three larval instars were present, with mean head capsule widths of 0.23, 0.31, and 0.42 mm, respectively. In general, the life cycle of C. ochromactonus is similar to those described for other xyleborine ambrosia beetle species.

Archips machlopis (Meyrick) (Tortricidae: Lepidoptera)-Occurrence on garlic

The present observations on the occurrence of Archips machlopis (Tortricidae: Lepidoptera) on garlic (Allium sativum) is a new host record and it was observed during rabi 2018 at Pune. Larvae were observed tying the adjacent leaves of plants and feed from within, and hence referred to as leaf-tiers. Larva has a brown prothoracic shield, and a narrow white line that demarcates the head capsule. Adult moths are cryptically coloured, with forewings grey, brown, rust, or tan coloured, have a prominent round-pointed apex with wings held in a characteristic a flattened roof fashion at repose. Pupa dark brown to black with cremaster elongate and tapered, enclosed within a dense web of silk. Mild to moderate level of infestation (5–8% damage) with typical leaf-tying and scraping symptom was observed.

The Developmental Characteristics for the Head Capsule Width ofMonochamus alternatus(Coleoptera: Cerambycidae) Larvae and Determination of the Number of Instars

The objective of this study was to determine the number of instars of Monochamus alternatus Hope (Coleoptera: Cerambycidae) larvae by comparing their head capsule widths (HCW) published in previous studies, as well as additional laboratory experiments. Larvae of M. alternatus showed repeated molting in the laboratory. Most larvae ceased their development at the 10th instar stage. Frequency distributions of HCW for the first, second, and third instar larvae were clearly separated while those of the fourth through 11th instar larvae largely overlapped between successive instars in our results. The HCW values for the first, second, and third instar larvae directly measured for each instar in our study indicated that they were more precise than those of previous reports based on field-collected HCW which might have missed HCW of the first instar larvae or wrongly determined HCW for some instars. Unlike the reports of four instars of previous studies, M. alternatus larvae passed five instars in the field, which was confirmed by the discovery of five pairs of mandibles in the feeding gallery and pupal chamber. Also, the comparative study for the frequency distributions of HCW revealed that most M. alternatus larvae passed five instars. Consequently, the average sizes of HCW for their first, second, and third instar larvae are newly suggested to be 0.896 ± 0.069, 1.291 ± 0.131, and 1.707 ± 0.165 mm (mean ± SD) .

Larval Morphology of Armostus ohyamatensis Hoshina and Satô (Coleoptera: Hydrophilidae: Megasternini)

The larval morphology, including chaetotaxy of the head capsule and head appendages, of Armostus ohyamatensisHoshina and Sato, 2006 is described in detail. This is the first description of larval morphology for the genus Armostus Sharp, 1890. Adults and larvae were associated using DNA barcoding of a molecular marker, 658 bp of mitochondrial cytochrome c oxidase I. Larvae of Armostus Sharp resemble other megasternine larvae, but they have a unique feature within the known hydrophilid larvae, i.e., incompletely cylindrically sclerotized maxillary palpomere 2.

Effects of microsporidiosis and food availability on the two-spotted lady beetle, Adalia bipunctata L., and convergent lady beetle, Hippodamia convergens Guérin-Méneville.

Two species of microsporidia have been described from lady beetles that are commercially available for biological control: Nosema adaliae from the two-spotted lady beetle, Adalia bipunctata L., and Tubulinosema hippodamiae from the convergent lady beetle, Hippodamia convergens Guérin-Méneville. These pathogens delay larval development under controlled conditions, but little is known regarding the effects that microsporidia cause when their hosts are subjected to stressful conditions that are often experienced in nature. In this study, the combined effects of microsporidiosis (N. adaliae on A. bipunctata and T. hippodamiae on H. convergens) and irregular food availability were observed on host fitness (larval development and mortality, sex ratios, and adult morphometrics). For each beetle species, 24 h-old larvae were provided either an uninfected or microsporidia-infected conspecific egg. After the egg was eaten, some larvae were provided an abundance of aphids daily, whereas others were provided aphids on an irregular basis. Development was delayed significantly for larvae that consumed a microsporidia-infected egg, and for those fed irregularly. For A. bipunctata, a significant interaction was observed between infection status and food availability. This suggests that N. adaliae-infected A. bipunctata larvae that have an irregular supply of aphids undergo further developmental delays than those with a generous food supply. This interaction was not observed for T. hippodamiae-infected H. convergens. For both species, larval mortality and sex ratios did not differ significantly, regardless of infection status or food availability. Adults that were fed daily as larvae were significantly larger than those fed irregularly. However, the elytra of N. adaliae-infected A. bipunctata were significantly larger than the elytra of their uninfected cohorts, and T. hippodamiae-infected H. convergens had wider pronota and head capsules than uninfected H. convergens. Because N. adaliae and T. hippodamiae prolong larval development of their respective hosts under controlled conditions, one would expect these pathogens to cause more profound effects when their hosts experience stressful conditions. The results from this study indicate that this was the case for A. bipunctata, but not for H. convergens.

Predicting Larval Stage Distribution of Lobesia botrana (Lepidoptera: Tortricidae) at Three Constant Temperatures.

European grapevine moth, Lobesia botrana (Denis and Schiffermüller), is an invasive pest species subject to regulatory control under the International Plant Protection Convention. An extensive infestation of this insect detected in California vineyards in October 2009 resulted in the initiation of postharvest research intended to ensure continuing trade of California table grapes by preventing the unintentional introduction of L. botrana to uninfested export destinations. Having the ability to accurately predict and confirm the particular stadia of larval development of insects obscured from direct observations, such as those being mass-reared in diet for use in phytosanitary control experiments, is necessary to be able to ensure that the most tolerant life stage is accurately targeted. Lobesia botrana were reared from egg to eclosion at three different temperatures: 16, 20, and 24°C (60% RH; 16:8 [L:D] h). Daily samples (≥15) were taken of larvae and pupae to determine the chronological distribution of life stages until daily samples comprised 100% pupae, after which only adults were counted. Head capsule measurements of all sampled larvae were taken (n = 2,820) and analyzed using nonlinear least squares (NLLS) fitting to Gaussian curves to describe stadia-specific head capsule demarcations and the probability of misclassifying instars due to overlapping life stage distributions.

Two unusual new genera of cavernicolous Hydrometridae (Insecta: Heteroptera) from eastern Brazil

Two new genera of cavernicolous Hydrometridae are described from limestone caves in northeastern Brazil: Spelaeometra n. gen., with type species S. gruta n. sp., from the Lapa D’Água do Zezé, Lapa do Nestor, Lapa do Saco Velho, and Lapa do Caboclo caves in Minas Gerais state; and Cephalometra n. gen., with type species C. pallida n. sp., from the Natividade cave in Tocantins state. Both new taxa are flightless, possess reduced eyes, and exhibit other reductional character states linked to life in underground ecosystems. The new taxa are interpreted to be highly aberrant members of the family Hydrometridae, based on the possession of eyes set well forward on the head capsule, the posterior pair of cephalic trichobothria arising from distinct tubercles, the presence of an apical invagination on the fourth antennal segment, and the structure of the male parameres. They are provisionally placed in the subfamily Hydrometrinae based on the apical articulation between the first and second antennal segments. On the basis of a preponderance of shared apomorphic character states these new genera appear most closely allied to Veliometra schuhi Andersen, which is also endemic to Brazil, although currently placed in a different subfamily. Color habitus photographs and line drawings of key characters are provided for both new taxa, accompanied by maps and photographs of the caves in which the species occur.

The head morphology of Micromalthus debilis (Coleoptera: Micromalthidae) - an archostematan beetle with an unusual morphology and a unique life cycle

Cephalic features of Micromalthus debilis were examined and described in detail for the first time. The head displays several seemingly plesiomorphic features compared to other extant species of Archostemata, especially representatives of Cupedidae and Ommatidae. Cephalic protuberances characteristic for species of these two families are missing and antennal grooves are also absent. The surface of the head capsule is largely smooth, without the characteristic tubercles found in stemgroup beetles and ommatid and cupedid species. Cuticular scales, probably ancestral for Archostemata and possibly for Coleoptera, are also completely absent. The arrangement of three mandibular teeth in a vertical row and an immobilized labrum are derived features shared with Ommatidae. The maxillary endite lobes are absent, as in the very small Crowsoniella relicta (Crowsoniellidae). Like in all other examined archostematan species, mandibular molae and prosthecae are missing. The simplified maxillae apparently play no role in the food uptake but rather function as accessory “ventral antennae”. Derived features include the partly reduced maxillary musculature and lack of extrinsic labial muscles. Apomorphies of the digestive tract include the sclerotized median protuberances of the anterior epipharynx and hypopharynx, and the presence of a vertical anterior pharyngeal loop and a subcerebral postpharyngeal pouch. The tentorium is strongly reduced. Consequently, all antennal muscles originate from the head capsule. A very unusual and possibly plesiomorphic feature is the presence of a short salivary tube and two associated muscles. This is a unique condition in Coleoptera as far as known at present. Structural features suggest that Micromalthus probably feeds on wood infested with fungi. A robust phylogenetic evaluation of anatomical features is presently not possible due to the lack of data for Crowsoniella (Crowsoniellidae) and Sikhotealinia zhiltzovae (Jurodidae). Moreover, phylogenetic and evolutionary interpretations are impeded by possible effects of vestigialization of adults possibly resulting from endosymbionts (e.g. Wolbachia, Rickettsiales).

Chlorantraniliprole impact on survival and progeny quality of the pupa of the parasitoid Palmistichus elaeisis (Hymenoptera: Eulophidae)

AbstractThe insecticide chlorantraniliprole is recommended for integrated pest management programmes of caterpillar (Lepidoptera) pest species. This insecticide is considered harmless to parasitoids, but its selectivity may vary with the species of natural enemy, few of which were subjected to testing. Furthermore, in addition to the active ingredient, formulation adjuvants may improve insecticide efficiency, but also its impact on nontargeted organisms. Here the chlorantraniliprole effect and its interaction with adjuvant on the survival and qualitative parameters of progeny of the nontarget parasitoid Palmistichus elaeisis (Delvare and LaSalle) (Hymenoptera: Eulophidae) were studied. The treatments used were: chrolantraniliprole (T1), chlorantraniliprole+surfactant (T2), and deionised water (T3, control). Pupae of Diaphania hyalinata Linnaeus (Lepidoptera: Crambidae) were subjected to these treatments for 24 hours and offered to the adult parasitoids. Chlorantraniliprole and chlorantraniliprole+surfactant did not elicit direct toxicity to female parasitoids. However, chlorantraniliprole and its combination with surfactant reduced the head capsule width of the female progeny. This insecticide alone also reduced emergence and the progeny metatibia length. Chlorantraniliprole did not induce mortality of P. elaeisis adults, but its negative impact on the emergence and progeny size of this parasitoid suggests an important sublethal effect to be considered.

A new Vietnamese species of Euconnus (Napochus) with modified antennal club (Coleoptera: Staphylinidae: Scydmaeninae).

Napochus Thomson, 1859 is the largest subgenus of Euconnus Thomson, 1859, currently comprising over 300 nominal species (Newton Franz 1998; Jałoszyński 2015, 2016, 2017a, b). The subgenus was recently redefined and its adult morphological structures were illustrated and described in detail (Jałoszyński 2015). Moreover, the subgenera Neonapochus Machulka, 1929, Paranapochus Franz, 1986, Smicrophus Casey, 1897, Cephaloconnus Franz, 1998 and Cerviconnus Franz, 1962, and a genus Protoscydmus Franz, 1992 were placed as junior synonyms of Napochus (Jałoszyński 2015, 2016, 2017a, b). Currently Napochus is defined mainly by the subtrapezoidal pronotum, which is always broadest behind middle (typically at base or shortly in front of base) and strongly narrowed anterad, so that its anterior margin is much shorter than the posterior margin; and by the tetramerous antennal club. The latter, however, may be indistinctly tetramerous in some species, with the antennomere VIII only slightly larger than VII; antennae in such beetles may appear gradually broadened or even trimerous. The problem of antennal variability within Napochus requires further studies, and it is possible that the number of antennomeres forming the club should be excluded from the diagnosis, as too variable. Until recently, species of Euconnus similar to the type species of Napochus and with unmodified antennomeres were placed in Napochus, and one species, in which males have the antennal club modified, in a subgenus Neonapochus. Also beetles resembling Napochus but showing a sexual dimorphism in the head capsule (with cavities, impressions or unusual concavities in males) were placed not in Napochus, but in Cephaloconnus. Jałoszyński (2016, 2017b) demonstrated that beetles showing such modifications do not differ from Napochus in any important characters, and synonymized Neonapochus and Cephaloconnus with Napochus.