Visual stimuli that elicit appetitive behaviors in three morphologically distinct species of praying mantis

We assessed the differences in appetitive responses to visual stimuli by three species of praying mantis (Insecta: Mantodea), Tenodera aridifolia sinensis, Mantis religiosa, and Cilnia humeralis. Tethered, adult females watched computer generated stimuli (erratically moving disks or linearly moving rectangles) that varied along predetermined parameters. Three responses were scored: tracking, approaching, and striking. Threshold stimulus size (diameter) for tracking and striking at disks ranged from 3.5 deg (C. humeralis) to 7.8 deg (M. religiosa), and from 3.3 deg (C. humeralis) to 11.7 deg (M. religiosa), respectively. Unlike the other species which struck at disks as large as 44 deg, T. a. sinensis displayed a preference for 14 deg disks. Disks moving at 143 deg/s were preferred by all species. M. religiosa exhibited the most approaching behavior, and with T. a. sinensis distinguished between rectangular stimuli moving parallel versus perpendicular to their long axes. C. humeralis did not make this distinction. Stimulus sizes that elicited the target behaviors were not related to mantis size. However, differences in compound eye morphology may be related to species differences: C. humeralis’ eyes are farthest apart, and it has an apparently narrower binocular visual field which may affect retinal inputs to movement-sensitive visual interneurons.     

The functional morphology of the praying mantis forelimb (Dictyoptera: Mantodea)

The structure and function of the forelimbs of praying mantids are discussed. A large spine on the femur and the spine on the end of the tibia were shown experimentally to be important in the capture of unrestrained flies. These spines may function to narrow the gap through which a fly may escape while the tibia is closing on the femur. Some of the spines on the femur are hinged at their base; the structure, properties and possible functions of these spines are discussed.
Similar hinged spines are reported on the lorelimbs of Stomatopoda (CrustaceaI but were not found in the Mantispidac (Neuroptera:Insecta). The variation of forelimb structure within the Mantodea is briefly discussed. The theory that the optimum prev size can be predicted from the geometry of the mantid forelimb is critically discussed.     

Responses to computer-generated visual stimuli by the male praying mantis, Sphodromantis lineola (Burmeister)

We tested male mantis' responses to square, computer-generated visual stimuli of various sizes to determine the stimulus parameters that affect their striking behaviour. Males in a high-hunger group displayed a sharp preference for solid black 12×12-degree squares moving linearly against a white background, especially when the squares moved downward (versus horizontally). Males in a low-hunger group visually tracked but, with rare exception, did not strike at any linearly moving squares. In contrast, when solid black squares moved erratically around visual field centre, males struck regularly at them irrespective of hunger level, even when the squares were as large as 47×47 degrees. Males also recognized black-and-white Julesz-patterned square stimuli moving against a similarly patterned background, indicating that they recognized the synchronous movement of a group of stimulus elements as a single moving object. Finally, we compared allometric and life history data between male and females S. lineola to elucidate the possible reasons for the differences between male response patterns and previously published data on females. These comparisons suggest that males employ a different behavioural strategy than do females when faced with large, erratically moving visual stimuli.     

Revision of the Neotropical bark mantis genus Liturgusa Saussure, 1869 (Insecta,Mantodea, Liturgusini)

The praying mantis genus Liturgusa Saussure, 1869 occurs only in Central and South America and represents the most diverse genus of Neotropical Liturgusini (Ehrmann 2002). The genus includes bark dwelling species, which live entirely on the trunks and branches of trees and run extremely fast. All species included within the genus Liturgusa are comprehensively revised with a distribution stretching from central Mexico, the island of Dominica to the southeastern regions of Brazil and southern Bolivia. All known species are redescribed to meet the standards of the new treatment of the genus (11 species). Three new genera are described including Fuga gen. n., Velox gen. n., and Corticomantis gen. n. for species previously included in Liturgusa as well as Hagiomantis. Liturgusa mesopoda Westwood, 1889 is moved to within the previously described genus Hagiomantis Audinet Serville, 1838. A total of 19 species are newly described within Liturgusa, Fuga, and Velox including L. algorei sp. n., L. bororum sp. n., L. cameroni sp. n., L. cura sp. n., L. dominica sp. n., L. fossetti sp. n., L. kirtlandi sp. n., L. krattorum sp. n., L. manausensis sp. n., L. maroni sp. n., L. milleri sp. n., L. neblina sp. n., L. purus sp. n., L. stiewei sp. n., L. tessae sp. n., L. trinidadensis sp. n., L. zoae sp. n., F. grimaldii sp. n., and V. wielandi sp. n. Four species names are synonymized: Liturgusa peruviana Giglio-Tos, 1914, syn. n. = Liturgusa nubeculosa Gerstaecker, 1889 and Hagiomantis parva Piza, 1966, syn. n., Liturgusa sinvalnetoi Piza, 1982, syn. n., and Liturgusa parva Giglio-Tos, 1914, syn. n. = Mantis annulipes Audinet Serville, 1838. Lectotypes are designated for the following two species: Liturgusa maya Saussure & Zehntner, 1894 and Fuga annulipes (Audinet Serville, 1838). A male neotype is designated for Liturgusa guyanensis La Greca, 1939. Males for eight species are described for the first time including Liturgusa cayennensis Saussure, 1869, Liturgusa lichenalis Gerstaecker, 1889, Liturgusa guyanensis La Greca, 1939, Liturgusa maya Saussure & Zehntner, 1894, Liturgusa nubeculosa Gerstaecker, 1889, Fuga annulipes (Audinet Serville, 1838), Corticomantis atricoxata (Beier, 1931), and Hagiomantis mesopoda (Westwood, 1889). The female of Fuga fluminensis (Piza, 1965) is described for the first time. Complete bibliographic histories are provided for previously described species. The spelling confusion surrounding Liturgusa/Liturgousa is resolved. Full habitus images for males and females are provided for nearly all species. Habitus and label images of type specimens are provided when possible. Diagnostic illustrations of the head and pronotum for males and females are provided for all species when possible. Illustrations of male genital structures are provided for all species for which males are known. Measurement data, including ranges and averages, are provided for males and females of all species. Combined male and female genus and species level dichotomous keys are provided with a Spanish translation. A complete table of all examined specimens lists label data, museum codes, repositories, and other specimen specific information. A KML file with all georeferenced locality records is downloadable from mantodearesearch.com for viewing in Google Earth. Natural history information is provided for species observed by the author.     

A new genus and species (Cornucollis gen. n. masoalensis sp. n.) of praying mantis from northern Madagascar (Mantodea,Iridopterygidae, Tropidomantinae)

An examination of Malagasy specimens accessed within the Muséum national d’Histoire naturelle, Paris, France, produced a praying mantis (Insecta: Mantodea) of an undescribed genus and species. An investigation of the internal and external morphology, in addition to its collection locality, revealed that this specimen belongs to the Iridopterygidae subfamily Tropidomantinae. Furthermore, the specimen’s unique combination of characters justified the creation of a new genus. Geographic distributional records and external morphological character evidence are presented for Cornucollis gen. n. masoalensis sp. n. We provide a dichotomous key of the Tropidomantinae and Nilomantinae genera distributed within Madagascar. High-resolution images, illustrations of morphological characters, natural history information, and measurement data are presented.     

Spermatophore feeding and mating behaviour in praying mantids (Mantodea: Liturgusidae)

In a number of insects, males transfer sperm to females via an externally attached spermatophore, which females subsequently remove and consume. Here I report the first study of spermatophore feeding in a praying mantid. While studies of praying mantid mating behaviour have largely focused on sexual cannibalism, this behaviour is not known in the genus Ciulfina . The general patterns of mating behaviour and spermatophore feeding are described for four species: Ciulfina rentzi , Ciulfina klassi , Ciulfina biseriata and Ciulfina baldersoni . Copulation duration and postcopulatory spermatophore attachment duration were found to vary both inter- and intraspecifically. Ciulfina rentzi exhibited a considerably longer mean copulation duration and a considerably shorter mean postcopulatory spermatophore attachment duration than the other Ciulfina species. Smaller males copulated for longer durations in C. rentzi and C. klassi . For C. baldersoni , both copulation duration and postcopulatory spermatophore attachment duration increased with female mass. The genus Ciulfina is highlighted as an intriguing new system in which to test hypotheses surrounding the function of spermatophore feeding and the significance of copulation duration.     

Stenophylla lobivertex,a new species of Stenophyllinae from Amazonia (Insecta,Mantodea)

Stenophylla lobivertex, a new species of Stenophyllinae from Ecuador and Peru, where this genus has never been found before, is described. This new species differs from Stenophylla cornigera Westwood, the only species known for this genus, by the remarkable size of the median process of the vertex and by the shape of the male genitalia. The systematic position and some biogeographical remarks about the genus are made.     

Cephalic morphology of Hymenopus coronatus (Insecta: Mantodea) and its phylogenetic implications

External and internal head structures of the mantodean Hymenopus coronatus are examined and described in detail. The results are elaborately compared with the literature. Strong crests on the anterior tentorial arms that articulate with the subantennal suture, a parietal suture and glossae and paraglossae with anteriorly bent tips are proposed as new potential apomorphies for Mantodea while a head capsule being wider than long, enlarged compound eyes, the presence of a frontal shield or scutellum, lateral lobes in the anterior tentorial arms, the presence of a transverse and an interantennal suture and the reduction of the mentum are confirmed as apomorphies, As potential apomorphies for Dictyoptera the reduction of Musculus tentoriobuccalis lateralis (M. 49) is newly presented and a “perforate” tentorium, lacinial incisivi that are located in a galeal pouch and the presence of a postmola are confirmed. The present study shows the value of cephalic morphology for phylogenetic analysis but also points out that further studies including evolutionary key taxa are essential for resolving the evolutionary adaptations among dictyopterans.     

Appetitive Responses to Computer-Generated Visual Stimuli by Female Rhombodera basalis, Deroplatys lobata, Hierodula membranacea, and Miomantis sp. (Insecta: Mantodea)

We used computer-generated stimuli to assess the stimulus parameters eliciting prey-directed behaviors in four mantis species: Rhombodera basalis (60 mm body length), Deroplatys lobata (70 mm), Hierodula membranacea (80 mm), Miomantis sp (38 mm). All responded appetitively to erratically moving disks subtending 10° or more. R. basalis responded more consistently to checkered (versus solid black) disks; only D. lobata displayed a size preference (14–23 deg disks); Miomantis sp. struck at disks as large as did the largest species; only Miomantis sp. displayed approaching behavior. H. membranacea and R. basalis preferred black disks on a white background (versus the reverse), and red disks (versus blue or green) on a brightness-matched grey background. All preferred elongated stimuli moving parallel (versus perpendicular) to their long axes.     

Purification and characterization of a serine protease with fibrinolytic activity from Tenodera sinensis (praying mantis)

Mantis egg fibrolase (MEF) was purified from the egg cases of Tenodera sinensis using ammonium sulfate fractionation, gel filtration on Bio-Gel P-60 and affinity chromatography on DEAE Affi-Gel blue gel. The protease was assessed homogeneous by SDS-polyacrylamide gel electrophoresis and has a molecular mass of 31500 Da. An isoelectric point of 6.1 was determined by isoelectric focusing. Amino acid sequencing of the N-terminal region established a primary structure composed of Ala-Asp-Val-Val-Gln-Gly-Asp-Ala-Pro-Ser. MEF readily digested the Aalpha- and Bbeta-chains of fibrinogen and more slowly the gamma-chain. The nonspecific action of the enzyme results in extensive hydrolysis of fibrinogen and fibrin releasing a variety of fibrinopeptide. The enzyme is inactivated by Cu2+ and Zn2+ and inhibited by PMSF and chymostatin, yet elastinal, aprotinin, TLCK, TPCK, EDTA, EGTA, cysteine, beta-mercaptoethanol, iodoacetate, E64, benzamidine and soybean trypsin inhibitor do not affect activity. Antiplasmin was not sensitive to MEF but antithrombin III inhibited the enzymatic activity of MEF. Among chromogenic protease substrates, the most sensitive to MEF hydrolysis was benzoyl-Phe-Val-Arg-p-nitroanilide with maximal activity at pH 7.0 and 30 degrees C. MEF preferentially cleaved the oxidized B-chain of insulin between Leu15 and Tyr16. D-Dimer concentrations increased on incubation of cross-linked fibrin with MEF, indicating the enzyme has a strong fibrinolytic activity.     

Telmapsylla gen.n., an unusual psyllid from black mangrove in Florida and Costa Rica (Insecta, Homoptera, Psylloidea)

Telmapsylla minuta . gen. et sp.n., a diminutive species of psyllid collected from black mangrove, Avicennia species (Verbenaceae), is described from Florida and Costa Rica. Telmapsylla is particularly significant in that it possesses characters currently used to define both the families Psyllidae and Calophyidae. Its taxonomic position and host-plant relationships are discussed. Telmapsylla is most probably a primitive calophyid.     

Identification of acoustic stimuli that mediate sexual behavior inDrosophila busckii (Diptera: Drosophilidae)

We have shown that D. busckiimales and females, unlike other drosophilids that have been analyzed in this regard, court and copulate as well in relatively dim red light as they do in bright white light. We have also shown that males and females of this species flutter their wings during courtship and that wing fluttering in both sexes is associated with acoustic stimuli. Wingless males perform vigorous courtship but are incapable of mating, suggesting that females must perceive male song to be receptive to copulation. When they are tested with normal males, wingless females stimulate vigorous courtship, but their copulation frequencies are significantly lower than winged females. This observation suggests that perception of the female's song by either or both sexes facilitates mating.     

Morphological adaptations to an unusual defensive strategy in the mantid Orthoderella ornata (Insecta: Mantidae)

The behaviour of Orthoderella ornata Giglio Tos, 1897, a cryptically coloured mantid, is described with reference to some morphological adaptations that conceal the typical insect outline.
The modification of the typical morphology of the head in mantids is analysed with special mention of the internal structure, the tentorium, and by comparison with the disposition of two other species with a different defensive strategy.
The hypothesis that O. ornata has short legs in correlation with its resting position is supported by a morphometric comparison with five other species of mantids which show different behaviour and positions in their cryptic strategy.     

Egg structure and outline of embryonic development of the basal mantodean, Metallyticus splendidus Westwood, 1835 (Insecta,Mantodea, Metallyticidae)

The egg structure and outline of the embryonic development of Metallyticus splendidus of one of the basal Mantodea representatives, Metallyticidae, were described in the present study. The results obtained were compared with those from the previous studies, to reconstruct and discuss the groundplan of Mantodea and Dictyoptera. In M. splendidus, the egg is spheroidal, it has a convex ventral side at the center in which numerous micropyles are grouped, and it possesses a conspicuous hatching line in its anterior half. These are the groundplan features of mantodean eggs and the “grouped micropyles in the ventral side of the egg” are regarded as an apomorphic groundplan feature of Dictyoptera. A small circular embryo is formed by a simple concentration of blastoderm cells, which then undergoes embryogenesis of the typical short germ band type. Blastokinesis is of the “non-reversion type” and the embryo keeps its original superficial position and original orientation throughout embryonic development. During the middle stages of development, the embryo undergoes rotation around the egg's anteroposterior axis. These features are a part of the groundplan of Mantodea. It is uncertain whether sharing of the “non-reversion type” of blastokinesis by Mantodea and blaberoidean Blattodea can be regarded as homology or homoplasy.     

Epaphroditidae <Emphasis Type="Italic">sensu novo</Emphasis>, an Endemic Caribbean Family of Morphologically Divergent Praying Mantises (Insecta,Mantodea)

Three endemic Caribbean praying mantis genera with a complex taxonomic history were recently discovered to be part of a lineage that colonized the Caribbean region during the Cretaceous period (Svenson & Rodrigues, Proc R Soc B Biol Sci 284, 2017). In all classification systems proposed up to now, the three genera, Callimantis, Epaphrodita, and Gonatista, were never considered as close relatives, a reflection of their divergent morphology. More recently, the genus Brancsikia was placed with Epaphrodita in a family based on the similarity of camouflage-related morphology. To address recent phylogenetic results that do not track current classification, we compared the morphology of the three Caribbean genera with each other and representative members of traditional or current family groups. Our morphological analysis of external and male genital characters provides strong support for the Caribbean lineage despite the divergent morphological evolution present in the three genera. We raise this Caribbean lineage to family status by employing a precedent family-group name, Epaphroditidae Brunner de Wattenwyl, 1893 sensu novo. We remove Brancsikia from our new concept of Epaphroditidae, rendering the genus incertae sedis.     

Visual system of the stalk-eyed fly,Cyrtodiopsis quinqueguttata (Diopsidae,Diptera): an anatomical investigation of unusual eyes

Diopsid flies have eye stalks up to a centimeter in length, displacing the retina laterally from the rest of the head. This bizarre condition, called hypercephaly, is rare, but has evolved independently among several insect orders and is most common in flies (Diptera). Earlier studies of geometrical optics and behavior have led to various hypotheses about possible adaptive advantages of eye stalks, such as enhanced stereoscopic vision while other hypothesis suggest that eye stalks are an outcome of sexual selection. Here, we focus on how these curious distortions of head/eye morphology are accompanied by changes in the neural organization of the visual system of Cyrtodiopsis quinqueguttata. Histological examinations reveal that the optic lobes, lamina (La), medulla (Me), lobula (Lo), and lobula plate (LP) are contained entirely within the fly's eye bulbs, which are located at the distal ends of the eye stalks. We report that the organization of the peripheral visual system (La and Me) is similar to that of other Diptera (e.g., Musca and Drosophila), but deeper visual areas (Lo and LP) have been more strongly modified. For example, in both the lobula and lobula plate, fewer but larger giant collector neurons are found. The most pronounced difference is the reduction in the number of wide-field vertical cells of the lobula plate, where there are only four relatively large fibers, as opposed to 11 in Musca. The “fewer but larger” neural organization may enhance the conduction velocities of these cells, but may result in a loss of spatial resolution. At the base of the eye bulb, axon bundles collect and form a long optic nerve that extends the length of the eye stalk. We suggest that this organization of the diopsid visual system provides evidence for the costs of possessing long eye stalks. © 1998 John Wiley & Sons, Inc. J Neurobiol 37: 449–468, 1998     

New mayfly nymphs (Insecta: Ephemeroptera) from Yixian Formation,China

Three new species of mayfly nymphs, Clavineta excavata sp. nov., Clavineta brevinodia sp. nov., and Siberiogenites branchicillus sp. nov., from the Yixian Formation (Late Jurassic-Early Cretaceous) in western Liaoning Province, China, are described. These mature nymphs are small-sized, with the body less than 16.0 mm long. The systematic position of the genera Clavineta and Siberiogenites, which are recorded for the first time in the Yixian Formation, is briefly discussed.