Evolution of arboreal tunnelling by larvae of Aenetus (Lepidoptera: Hepialidae)

Abstract

Aspects of larval feeding and tunnelling were described for seven species of the wood-boring genus Aenetus. The species were studied in Australia, New Caledonia, and New Zealand. A sequential pattern of larval feeding involving transfer from a fungal based diet to callus tissue in live trees, previously known for A. virescens, was confirmed for A. cohici and inferred for A. dulcis and A. paradiseus. A specialised “transfer” morph linking the two feeding stages in A. virescens was also confirmed for A. cohici and two unidentified species. The wood-boring habit involved entry into the host above ground level and the construction of a tunnel which extends into and then down the stem. Tunnels were often located on the lower surface of leaning branches or stems. Tunnel entrances were open to the host surface, but overlain by a silk- /frass web. Bark and underlying tissue were removed from around the tunnel entrance and a callus growth resulted, which the larva consumed. The evolutionary history of arboreal tunnelling in the Hepialidae was investigated using biogeographic analysis of generic distribution. It is argued that evolution and variation in arboreal tunnelling resulted from specialised recombinations of generalised ancestral characters. It is suggested that the study of arboreal tunnelling can provide a source of useful characters for systematic analysis of phylogenetic relationships within Aenetus. The potential systematic value of tunnel characters is illustrated for A. virescens.     

The suspensorium of Ctenopoma acutirostre: A comparative functional analysis with Anabas testudineus

The functional and structural aspects of the suspensorium of Ctenopoma acutirostre have been correlated with those of Anabas testudineus. The different parts of the suspensorium are described, as are the muscles that are functionally connected with the suspensorium. Functions were analyzed by observations on living specimens, and by measurements recorded from the movie films. The role played by various bones and muscles to carry out the functions (the respiration, the gulping, and the feeding) has been explained. The different bones and muscles have been considered as functional units which often are connected to form couplings. During the respiration in Ctenopoma the depression of the lower jaw is conducted by the levator operculiopercular apparatus-mandible coupling. The presence of this coupling is indicated by the presence of dorso-ventral movement of the operculum. A remarkable ventro-dorsal and antero-caudal movement in the urhyal during feeding shows in Ctenopoma the presence of the sternohyoideus-hyoid apparatus-interopercular-mandible coupling, which depresses the lower jaw. In Ctenopoma, the suspensorium takes part in respiration, gulping, and feeding, whereas in Anabas it is only involved in gulping and feeding. To carry out its functions, the suspensorium implies three articulations: palatocranial, craniohyomandibular, and quadratomandibular with the cranium and the lower jaw, respectively. Finally, the suspensorium has been analyzed as a part of the architectonic structure of the entire head by using a diagrammatic model (fig. 13) based on mutual influence, integration, and couplings.     

A reappraisal of the genus Megacaryon (Boraginaceae,Lithospermeae) based on molecular,morphological, and karyological evidence

The systematic position and phylogenetic relationships of Echium orientale, a rare endemic of the mountains of northern Turkey, were elucidated based on morphological, molecular, and karyological evidence.

Using nuclear and plastid DNA sequences, we found that this species is not included in Echium, but is rather at least as divergent from it as other related lineages, namely the South African Lobostemon and Echiostachys. Pollen characters revealed a strong affinity with Onosma, especially in the non-reticulate ornamentation of the tectum. Fruit morphology is unique in the genera of the Echium ‘alliance' (e.g. Echium, Pontechium, Lobostemon, and Echiostachys), because of the large size and the smooth, glossy nutlet surface as in most species of Onosma. Karyological observations show that this species is diploid with 2n = 12, and differing from both Lobostemon or Echium, where nearly all species have higher numbers (usually 2n = 14 and 2n = 16, respectively). Combined with a ‘megaherb' habit, the weak zygomorphism of the large flowers and a restricted range in the humid mountain forests of the Black Sea region, fruit, pollen and chromosome characters suggest an isolated position for E. orientale in the Echium alliance. We therefore propose its placement in the monotypic Megacaryon, a genus originally described by Boissier, but largely disregarded by later botanists. An original specimen is designated as generic lectotype.     

Fossil pollen grains of Asteraceae from the Miocene of Patagonia: Nassauviinae affinity

Fossil pollen grains with morphological features unique in the subtribe Nassauviinae (tribe Mutisieae, Asteraceae) occur in Miocene marine deposits of eastern Patagonia, southern South America. A new morphogenus and two morphospecies are proposed to assemble fossil pollen grains characterized by having a complex bilayered exine structure with delicate columellae, separated by an internal tectum. Subprolate specimens with Trixis exine type (ectosexine thinner than endosexine, straight internal tectum) are referred to Huanilipollis cabrerae. This species is similar to pollen of recent Holocheilus, Jungia, and Proustia. Suboblate specimens with Oxyphyllum exine type (ectosexine and endosexine equally thick, zigzag internal tectum) are referred to Huanilipollis criscii. This species is similar to pollen of recent Triptilion. The spore/pollen sequences in which Nassauviinae pollen types occur suggest a wide range of vegetation types varying from forest dominated during the Early Miocene (Chenque Formation) to virtually xerophytic ones during the Late Miocene (Puerto Madryn Formation). The subtribe Nassauviinae comprises 25 genera and ca. 320 species of vines, shrubs and low trees endemic to America with a wide range of ecological preferences; the nearest living relatives of the fossil types being mostly confined to humid landscapes. The unusual occurrence of these groups during the arid characterized Late Miocene time could be attributed to the complex interplay of the mountain uplift and global circulation patterns. These forcing factors would have created a mosaic of different habitats with both patches of forest and dry-adapted species developing in relatively small regions. This is the first fossil record of Nassauviinae and confirms that this subtribe of Asteraceae was already differentiated in the Miocene.     

Glass in the water: Molecular phylogenetics and evolution of Indian glassy perchlets (Teleostei: Ambassidae)

Glassfishes of the family Ambassidae, comprising around 50 species, are distributed in the Indo‐West Pacific where they inhabit marine, estuarine, and freshwater ecosystems. We investigated for the first time the molecular phylogenetic and evolutionary relationships of this group using a combined dataset of mitochondrial and nuclear genes, particularly focusing on the taxa occurring in the Indian subcontinent. Results revealed that marine and freshwater genera of Ambassidae diverged during the Paleocene (~62 mya). The enigmatic monotypic genus Chanda is nested within the larger clade currently recognized as Parambassis, indicating its paraphyly. Based on cleared and stained osteological preparations and phylogenetic placement of Chanda nama, we hypothesize that the elongated and protruding lower jaw is an autapomorphic character that might have evolved for the lepidophagous habit of the species. The southern Indian species of Parambassis, Parambassis dayi, and Parambassis thomassi, which formed a monophyletic group, probably diverged from other species of Parambassis and Chanda nama around the Eocene (~42 mya) and can potentially be recognized as a distinct genus in view of the apomorphic characters such as the presence of serration on the ventral fringe of interopercle, densely serrated palatine and ectopterygoid, and the presence of more than 30 serrations along the lower preopercle and the posterior edge. Our analysis provides new insights into the evolution and phylogenetic relationships of glassy perchlets, including detailed relationships among the Indian species within this family.     

Short communication. Food availability and diel feeding rhythms in the marine copepods Acartia grani and Centropages typicus

The effects of food availability on the daily feeding pattern of planktonic copepods was studied for two species: Acartia grani (cultured and wild specimens) and Centropages typicus (wild specimens). Both species showed clear diel feeding rhythms at high food concentrations, with significantly higher clearance rates during night hours. However, at limiting food conditions, while C.typicus maintained its daily feeding pattern, A.grani fed continuously, with similar day and night clearance rates. These results could be explained in relation to differences in predation risks and probability of experiencing starvation episodes in their respective habitats, as well as their capacity for dampening food fluctuations. On the other hand, the maintenance (although with less intensity) of the diel feeding rhythm in A.grani, when cultured in the laboratory for long periods of time (>10 generations) and in the absence of predators, suggests the importance of the endogenous control of copepod feeding patterns.      

A multivariate analysis of morphological variation in the Hemizygia bracteosa complex (Lamiaceae, Ocimeae)

The variation and recognition of taxa within the Hemizygia bracteosa complex was examined using multivariate techniques. Morphological characters were sampled on 197 herbarium specimens. Phenetically H. bracteosa and H. welwitschii overlap in many floral characters. However differences in leaf characters and habit maintain their morphological distinctness from each other and they are therefore upheld as species. Hemizygia ornata, synonymized under H. welwitschii in recent treatments of the genus Hemizygia, differentiates from H. welwitschii on the basis of differences in leaf and floral characters and its reinstatement at specific level is here proposed. The concept of H. linearis is revised on account of the type specimen separating from all other elements of the taxon used in the analyses. The latter form a coherent group with H. petrensis and H. canescens thus negating any attempt to give any of them specific recognition. It is proposed that they should be synonymized under the earliest name H. canescens. The specific boundary of H. petiolata is revised but its specific status maintained.     

A STUDY OF THE PROPOSED GENUS OBREGONIA (CACTACEAE)

The monotypic genus Obregonia Frič was compared with Ariocarpus, Lophophora, Strombocactus, and certain other cactus genera. Obregonia and Ariocarpus are similar in characters of seeds, seedlings, habitat, fruits, and general habit. They differ in time of flowering, point of flower origin, areole structure, presence or absence of druses and a mucilage system, and presence or absence of spines. Obregonia and Lophophora are similar in characters of seeds, habitat, basic areole structure, and fruits. They differ in seedling form, habit of adult plants, presence or absence of spines, and chemical analysis. Strombocactus resembles Obregonia in few ways except in basic areole structure and some aspects of anatomy. The author concludes that Obregonia is an intermediate form between Ariocarpus and Lophophora and deserves generic rank. A formal taxonomic treatment of the genus follows the conclusions.     

Micro-computed tomography of spicule networks in three genera of dorid sea-slugs (Gastropoda: Nudipleura: Doridina) shows patterns of phylogenetic significance

Dorid nudibranchs (Gastropoda: Nudipleura) are a key taxon for studying the evolution and interaction of chemical defence, colour patterns and feeding specialization, but we lack a robust phylogeny for hypothesis testing. To provide new morphological characters, we investigated the extensive interior spicule networks of several dorid taxa. We compared traditional staining methods versus micro-computed tomography (μCT) of 31 specimens representing 10 species of Aldisa, Cadlina and Onchidoris. We found that μCT offered a nondestructive view of sufficient resolution to study the gross morphology of networks, although images of fine structures were too grainy to allow robust comparisons. Network form did not vary within species and was consistent within genera. The three genera varied in several obvious characters, such as the presence of a pleural sinus channel, large dorsal spicules and multispicular tracts, as well as in relative size, shape and orientation of spicules. These characters, combined with those from the literature, supported recent molecular phylogenies that group Cadlina with Aldisa, and that question the monophyly of the Cryptobranchia. This suggests that these network forms will prove a fruitful source of phylogenetic characters with at least genus-level resolution.     

Occlusal pattern of cheek teeth in extant Spermophilus: A new approach to the identification of species

Discrete characters of the occlusal surface (additional cusps) have been studied to elaborate a new approach to the identification of the Ground Squirrel species Spermophilus odessanus, S. suslicus, S. pygmaeus, S. citellus, and S. xanthoprymnus. Data on the presence/absence of the additional cusps have been represented as star plots and, in addition, have been studied using discriminant function analysis. The species‐specific sets of the characters (patterns of bunodonty) have been revealed and are of high diagnostic value. The Citellus‐set is defined by the presence of mesostyles and the rareness of the metastylids, paraconules and metaconules, hypostyles and protostyles. The Pygmaeus‐set is characterized by the presence of additional cusps in the lower cheek teeth. The Odessanus‐oriented set is found in the Spermophilus pygmaeus, S. odessanus, and S. suslicus. The relatively high frequency of additional cusps of the metaloph and the paraloph is characteristic for this set. The Plesiomorphic‐set (characters shared by all the studied species and for this reason regarded herein as ancestral) is found in S. xanthoprymnus. The patterns of bunodonty serve as diagnostic criteria only as a whole: the shape of a star plot (relations among the character frequencies), rather than certain character values, is indicative. An optimal level of identification of species is possible based on the combination of the discrete characters mentioned and on the size parameters of the third upper molar. The occlusal sets are intended to remain stable during the time of species existence and seem to correspond to trends in specialization. The functional meaning of the sets can be explained by the dependence between the presence/absence of the discrete characters and the shape of the crown and its main lophs. Each pattern is likely to correspond to a trophic niche, and this niche corresponds to the species. J. Morphol. 277:814–825, 2016. © 2016 Wiley Periodicals, Inc.     

Feeding plasticity of Trematomus newnesi (Pisces, Nototheniidae) in Terra Nova Bay, Ross Sea, in relation to environmental conditions

Forty-three and 49 specimens of the Antarctic fish Trematomus newnesi were collected in the coastal waters of Terra Nova Bay, Ross Sea, during December 1994 and February 1998, respectively. The dietary composition of the two fish samples was determined by means of stomach content analysis and then compared. In general, as reported in most of the previous studies, planktivory appeared to be the main feeding habit of T. newnesi. However, the different environmental conditions occurring in the study area in the two periods investigated, such as the degree of the sea-ice coverage and the related amount of light available below the ice, influenced the food composition of T. newnesi. In December 1994, the whole area investigated was covered by a thick layer of sea ice and the diet of T. newnesi consisted of few prey taxa. Some species that characterize the so-called “cryopelagic habitat”, such as the euphausiid Euphausia crystallorophias and the copepod Metridia gerlachei, were by far the most important prey, followed by amphipods and the pteropod Limacina helicina. Conversely, the ostracod Alacia belgicae and larval stages of fish (Pleuragramma antarcticum) were consumed occasionally. In late summer (February 1998), the lack of sea-ice coverage and changes in the associated fauna influenced the feeding habit of T. newnesi. In this period the food spectrum appeared to be more diverse, revealing an active feeding search in the water column. The bulk of food was composed of several prey groups, such as E. crystallorophias, hyperiid amphipods (Hyperiella dilatata), copepods, L. helicina and several species of fish larvae. Present data provide evidence of a marked feeding plasticity of T. newnesi, in response to diverse environmental conditions that characterize the High-Antarctic Zone. Accepted: 17 July 1999     

New Middle Cambrian bivalved arthropods from the Burgess Shale (British Columbia,Canada)

The morphology of two new bivalved arthropods, Loricicaris spinocaudatus gen. et sp. nov. and Nereocaris briggsi sp. nov. from the middle Cambrian (Series 3, Stage 5) Burgess Shale Formation (Collins Quarry locality on Mount Stephen, Yoho National Park, British Columbia, Canada), is described. The material was originally assigned to the genus Branchiocaris, but exhibits distinctive character combinations meriting its assignment to other taxa. Loricicaris spinocaudatus possesses an elongate and spinose abdomen comparable to the contemporaneous Perspicaris and Canadaspis, as well as chelate second head appendages and subtriangular exopods, comparable to Branchiocaris. Nereocaris briggsi possesses a laterally compressed carapace, elongate and delicate appendages and a medial eye located between a pair of lateral eyes on a rhomboidal eye stalk. Although undoubtedly congeneric with Nereocaris exilis from a slightly younger horizon of the Burgess Shale Formation, N. briggsi differs in overall proportions and segment number, warranting assignment to a new species. The newly described taxa were coded into an extensive cladistic analysis of 755 characters, and 312 extinct and extant panarthropods, including a variety of Cambrian bivalved arthropods from both the Burgess Shale and the Chengjiang Lagerstätten. Cambrian bivalved arthropods consistently resolved as a paraphyletic assemblage at the base of Arthropoda. Important innovations in arthropod history such as the specialization of the deutocerebral head appendages and a shift from a nekton‐benthic deposit feeding habit to a benthic scavenging/predatory habit, the symplesiomorphic feeding condition of Euarthropoda (crown‐group arthropods), were found to have occurred among basal bivalved arthropods.     

On Intraspecies Forms of the Mongolian grayling (<Emphasis Type="Italic">Thymallus brevirostris</Emphasis>Kessl.) from Hoton Nur Lake (Western Mongolia)

Morphological and ecological traits of the Mongolian grayling Thymallus brevirostrisfrom Hoton Nur Lake are investigated with consideration of the assumption for a possible presence in this species of two morpho-ecological forms. The data on growth, feeding, sex, and age composition are presented. Plastic and meristic characters, distribution of fish in the water body, and feeding are analyzed. Within the age range of 3+ to 6+, in the population there are two groups of specimens differing in growth rate, diet, and some plastic characters of the head. These two groups may be designated as “predatory” and “benthos-feeding” forms. The predatory form lacks specimens of junior age groups; the benthos_feeding form lacks specimens older than seven years. Validity of these morpho-ecological groups is discussed.     

Early Developmental Stages of the Green Tiger Prawn, Penaeus semisulcatus de Haan (Crustacea, Decapoda, Penaeidae)

Gravid females of Penaeus semisulcatus were spawned in the laboratory by natural means. The embryos were documented and the larvae were reared from hatching to postlarval stage at 28.2–30.0 °C and 33.5–34.5 g kg⁻¹ salinity for about 10 days (223 h 55 min). Six naupliar stages, three protozoea stages, three mysis stages and the first postlarval stage were described and illustrated. The larvae were fed only with microalgae Tetraselmis tetrathele and Chaetoceros gracilis from first protozoea until the second mysis, with about 90% survival rate; from the third mysis until the first postlarva they were fed with similar microalgae coupled with rotifer Brachionus plicatilis and Artemia nauplii. The embryonic and larval stages of P. semisulcatus are generally similar to those of other closely related species in the family Penaeidae, such as Melicertus canaliculatus, Fenneropenaeus merguiensis, and Marsupenaeus japonicus, except for the size and structure of diagnostic characters, setation of appendages and duration of metamorphoses. The change in the feeding habit during ontogeny was related to morphological transformation of the feeding apparatus of larvae and postlarvae. This paper is the first comprehensive and complete account of the early developmental stages of P. semisulcatus.     

Defensive strategies of soft corals (Coelenterata: Octocorallia) of the Great Barrier Reef

Summary The relationship between ichthyotoxicity and predation-related defensive functional morphology was examined in alcyonacean soft corals of the central and northern regions of the Great Barrier Reef (GBR), Australia. Approximately 170 specimens were assessed encompassing a number of genera within three families: 1) the Alcyoniidae (Lobophytum, Sarcophytum, Sinularia, Cladiella, Parerythropodium, and Alcyonium); 2) Neptheidae (Lemnalia, Paralemnalia, Capnella, Lithophyton, Nephthea, Dendronephthya, Scleronephthya, and Stereonephthya), and 3) Xeniidae (Anthelia, Efflatounaria, Cespitularia, Heteroxenia, and Xenia). Ichthyotoxicity data were derived from earlier studies which used Gambusia affinis Baird and Girard (Vertebrata, Pisces) as a test organism. These data were compared to morphological data collected from specimens in the field and laboratory. Three sets of statistical analyses were performed, each considering a progressively narrower group of taxa. The first included 68 specimens and considered 16 morphological characters in each, falling into the general categories of gross colony form, colony texture, presence of mucus, colony color, polyp retractility, and sclerite morphology and distribution. These were tested for independence against ichthyotoxicity data. The second set of analyses involved a more restricted morphological data set derived from 28 species of Sinularia in combination with 28 species within the Nephtheidae, comparing them to their respective toxicity ranks. The third analysis considered the previous two taxonomic groups separately in relation to their toxicity levels.The attempt to consider many morphological characters in a taxonomically diverse collection did not reveal any general association in the Alcyonacea between defensive morphology and toxicity, and those associations which did emerge were clearly erroneous. The second analysis, considering only Sinularia spp. and nephtheids, demonstrated a negative association between ichthyotoxicity and the morphological characters of a) polypary armament, b) microarmament of the individual polyp, and c) strong mineralization of the coenenchyme. The third analysis revealed that the negative association found between toxicity and the first two characters was derived entirely from the nephtheids while the association detected between toxicity and the third character was restricted to Sinularia. It is concluded that a relationship between toxicity and morphology can be demonstrated, but it is heavily dependent upon which specific morphological characters are being considered and at what level of taxonomic resolution the analysis is being performed. An approach utilizing many characters over many taxa is unlikely to yield significant, reliable, or meaningful results.Australian Institute of Marine Science Contribution Number 383     

Pollen morphology of JapaneseSymplocos (Symplocaceae)

The pollen morphology of all 23 Japanese species ofSymplocos (1 of subgenusEusymplocos, and 22 of subgenusHopea) was comprehensively studied using LM, SEM and TEM, and found to be classifiable into two types (Type I and II) primarily on the basis of wall structure. Type I, characterized by a thick tectum lacking a supratectal structure and reduced columellae, occurs in subgenusEusymplocos, while Type II, characterized by a thin tectum with a supratectal structure and generally distinct columellae, occurs in subgenusHopea. Resemblances in wall structure suggest a closer relationship of subgenusEusymplocos to subgenusMicrosymplocos as well as of subgenusHopea to subgenusEpigenia. This fact contradicts the earlier proposed infrageneric classification, and may require its revision. Within Type II, five subtypes are recognized primarily on the basis of sculpture combined with other pollen characters. Systematic value of sculpture (i.e., subtype), aperture morphology and “globules”, as well as certain species relationships based on those characters, are also discussed.     

Hidden taxonomic characters in otoliths of blenniid fishes (Teleostei: Blenniidae) from the Iranian coasts of the Persian Gulf and Oman Sea ecoregions: A scanning electron microscopy approach

In the present work, the morphological characteristics of sagitta were described and analysed for 10 species of combtooth blennies (Blenniidae) inhabiting the intertidal region of the Persian Gulf and Makran zone of Oman Sea to see the taxonomic usefulness of otoliths, especially their hidden characters for species identification. Based on the obtained results, sagitta morphology could be considered as diagnostic characters distinguishing the genera Alticus, Antennablennius, Istiblennius, Omobranchus and Parablennius showing differences related to general shape, that is, elliptic/oval to triangular; shape of antirostrum and its degree of development; the degrees of convexity of ventral and dorsal margins, and shape of ostium. The common/general otolith characters shared in the all studied combtooth blennies, is the heterosulcoid condition, which is a shared character with Perciformes and Gobiiformes. The type of sulcus is ostial, and it can be considered as a plesiomorph character shared with a basal group of actinopterygian fishes. The obtained results could be useful in taxonomical, paleontological, ecological and food and feeding habits studies.     

Feeding mechanics and dietary implications in the fossil sloth Neocnus (Mammalia: Xenarthra: Megalonychidae) from haiti

Haitian species of the extinct ground sloth genus Neocnus (Mammalia: Pilosa: Megalonychidae) have previously been hypothesized to have a much reduced jugal bone and a correspondingly reduced masseter musculature but a paucity of specimens has prevented further investigation of this hypothesis. Recent discovery of jugal bones belonging to Haitian specimens of Neocnus within the University of Florida Museum collections enables the element to be more accurately described. The discovery also makes it possible to explore mastication in these sloths. Osteological characters related to feeding were examined, along with comparative estimations of bite force with the extant tree sloths, Bradypus and Choloepus, and their known dietary habits as a means to infer aspects of the paleodiet of Neocnus. There is a significant difference in moment arm calculations for m. masseter between predicted and actual jugals, but the overall significance for bite force is lost and hampered by small sample size. Neocnus demonstrates a variety of characters that are similar to those of Bradypus and not to Choloepus, which is a close phylogenetic relative. The masticatory musculature of Neocnus enabled a chewing cycle emphasizing a grinding combination of mesiodistal and linguobuccal movements of the molariform dentition. The orientations of m. masseter and m. temporalis are estimated to produce relatively high bite force ratios that imply a masticatory system with stronger versus faster components. Because of the similarity of bite forces and jaw mechanics to those of Bradypus, in addition to a number of osteological adaptations indicative of herbivorous grazers (elevated mandibular condyle, large and complex masseter, and robust angular process), the Haitian forms of Neocnus are considered to have been selective feeders with a folivorous diet. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.