Morphology of a new deep‐sea acorn worm (class Enteropneusta,phylum Hemichordata): A part‐time demersal drifter with externalized ovaries

Ten individuals of an enteropneust in the family Torquaratoridae were videotaped between 2,900 and 3,500 m in the Eastern Pacific—one drifting a few centimeters above the bottom, two exposed on the substrate, and seven partly burrowed, reflecting a bentho‐pelagic life style. Here, we describe a captured specimen (26 cm living length) as the holotype of Allapasus aurantiacus n. gen., n. sp. The small proboscis is dome‐shaped, and the collar is only slightly wider than deep; both of these body regions are more muscular than in other torquaratorids, which presumably facilitates burrowing. The proboscis complex, in contrast to that of shallow‐living enteropneusts, lacks a pericardial sac and is located relatively posteriorly in the proboscis stalk. The stomochord is separated from the main course of the gut by the intervention of a small, plate‐like proboscis skeleton lacking posterior horns. The most anterior region of the trunk houses the pharynx, in which the pharyngeal skeletal bars are not connected by synapticles. The postpharyngeal trunk comprises three intestinal regions: prehepatic, hepatic (with conspicuous sacculations), and posthepatic. On either side of the worm, a flap of body wall (lateral wing) runs the entire length of the trunk. The two lateral wings can wrap the body so their edges meet in the dorsal midline, although they often gape open along the pharyngeal region. The holotype is a female (presumably the species is gonochoric) with numerous ovaries located in the lateral wings along the pharyngeal region. Each larger ovary contains a single primary oocyte (up to 1,500 μm in diameter) and bulges outwards in an epidermal pouch attached to the rest of the body by a slender stalk. Such externalized ovaries are unprecedented in any animal, and nothing is yet known of their role in the reproductive biology of A. aurantiacus. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

Clypeo-cibarial apparatus, proboscis mobility, and cephalic foregut structure of Bombyliidae (Diptera)

The clypeus of Bombyliidae is subdivided into five morpho-functional parts: median centroclypeus, two lateral “membranes,” and two sclerotized zones. The centroclypeus and lateral membranes are formed by solid sclerotized cuticle and are apparently non-extensible. The dorsal hinge, around which the centroclypeus turns with respect to the head capsule, is an intraclypeal structure formed by rigid cuticle. The true membranes occur only in the distal portion of the clypeus and also at the bases of the labrum and labium. In the long-proboscid bombyliids, the labial base lies deep in the head cavity when the proboscis is in the resting position. As in the previously studied members of Stratiomyidae, the precerebral portion of bombyliid foregut shifts over a considerable distance during the movements of the proboscis. This makes the functioning of the radial pharyngeal dilators impossible, and these muscles disappear, while the typical sclerotized pharynx is modified into an oesophagus-like tube. The formation of the fulcrum is usually related to the development of the clypeus mobility. This process occurred repeatedly in several taxonomically distant dipteran groups.     

Organisation of the praesoma in Acanthocephalus anguillae (Acanthocephala, Palaeacanthocephala) with special reference to the muscular system

The organisation of the praesoma in the parasite Acanthocephalus anguillae was studied on the light and electron microscopic level, with emphasis on the morphology of the musculature. The study was compiled to add new data to the ground pattern of the Acanthocephala for analysis of the phylogenetic relationships within the Gnathifera. In A. anguillae the praesomal epidermis and lemnisci form a coherent syncytium, separated from the epidermis of the trunk. Hooks are seen to be derivatives of the subepidermal basal lamina and are covered by the praesomal epidermis. The praesomal circular body wall musculature forms a network of anastomosing muscle fibres that lines the proboscis; a praesomal longitudinal body wall musculature does not exist. The truncal circular and longitudinal body wall musculature rise up to the praesomal proboscis. The unpaired proboscis retractor, consisting of longitudinal circomyar fibres, forms an outer and an inner concentric tube; the latter extends through the entire praesoma and penetrates the receptacle wall. The sack-like receptacle is surrounded by a receptacle constrictor. The nervous system of the praesoma consists of a prominent cerebral ganglion, three nerves which extend anteriorly, ramify and end within the praesomal musculature, and two strong lateral posterior nerves. A. anguillae lacks an apical organ, lateral organs and a support cell. Many of the features present in the praesoma of A. anguillae can be assumed as ground-pattern characteristics of the Acanthocephala. Accepted: 22 January 2001     

Proboscis sensilla in Vanessa cardui (Nymphalidae, Lepidoptera): functional morphology and significance in flower-probing

 Morphology and distribution of the proboscis sensilla in Vanessa cardui have been investigated in order to contribute to the understanding of flower-probing behaviour in butterflies. The proboscis has a bend region approximately one-third of the length from the base. A short tip region is characterized by rows of intake slits leading into the food canal. Along the dorsal, lateral and ventral sides of the proboscis, sensilla trichodea, sensilla basiconica and sensilla styloconica are distributed in varying patterns depending on their distance from the b ase. The medial food canal bears one longitudinal row of sensilla basiconica only. The bristle-shaped sensilla trichodea are longer in the proximal region of the proboscis and become gradually shorter towards the tip. They are most frequent in number near to the bend region and near the beginning of the tip region. Sensilla basiconica arranged in longitudinal rows increase in number the more distal they are on the proboscis. The tip region is characterized by rows of sensilla styloconica on the dorsal side whereas the sensilla trichodea are mostly restricted to the ventral side. The ultrastructure suggests that the aporous sensilla trichodea function as mechanosensilla while the uniporous sensilla basiconica act as contact chemosensilla. The sensilla styloconica are regarded as bimodal contact chemo/mechanosensilla since their sensory cones are equipped with a single terminal pore and a tubular body at the base. The mouthpart sensilla appear to provide tactile cues on the positioning of the proboscis and on the degree of its insertion into a floral tube. Furthermore, they receive chemical stimuli on the availability of nectar and on the immersion status of the food canal. Accepted: 12 September 1997     

Description and morphological variability of <Emphasis Type="Italic">Neoechinorhynchus beringianus</Emphasis> n. sp. (Acanthocephala: Neoechinorhynchidae) from north-eastern Asia

Neoechinorhynchus beringianus sp. n. is described from Pungitius pungitius L. in north-eastern Russia. Since 1986, when it was first found, it was reported as 'N. pungitius Dechtiar, 1971'. However, this new species differs from the latter in having an egg shell without a prolongation of the fertilisation membrane, a larger proboscis and proboscis hooks, a subterminal position of the female genital pore and a more slender trunk, and it occurs in a different site in the intestine. N. beringianus has a small, stout body with an asymmetrical position of the proboscis, which is located ventrally to and at an angle with the longitudinal axis of the body. The proboscis is wider than long, the hooks are of equal size in each circle but diminish in size posteriorly, whereas the lemnisci are subequal in length. It differs from those species of Neoechinorhynchus Stiles & Hassall, 1905 with somewhat similar characteristics in body length, proboscis size and proportions, proboscis hook lengths, egg size, size-ratio of the cement gland and testes, and the number of giant nuclei in the tegument and lemnisci. In different geographical populations of the new species, the sizes of both the proboscis and proboscis hooks exhibit some variation.     

Functional morphology of the feeding apparatus and evolution of proboscis length in metalmark butterflies (Lepidoptera: Riodinidae)

An assessment of the anatomical costs of extremely long proboscid mouthparts can contribute to the understanding of the evolution of form and function in the context of insect feeding behaviour. An integrative analysis of expenses relating to an exceptionally long proboscis in butterflies includes all organs involved in fluid feeding, such as the proboscis plus its musculature, sensilla, and food canal, as well as organs for proboscis movements and the suction pump for fluid uptake. In the present study, we report a morphometric comparison of derived long‐tongued (proboscis approximately twice as long as the body) and short‐tongued Riodinidae (proboscis half as long as the body), which reveals the non‐linear scaling relationships of an extremely long proboscis. We found no elongation of the tip region, low numbers of proboscis sensilla, short sensilla styloconica, and no increase of galeal musculature in relation to galeal volume, but a larger food canal, as well as larger head musculature in relation to the head capsule. The results indicate the relatively low extra expense on the proboscis musculature and sensilla equipment but significant anatomical costs, such as reinforced haemolymph and suction pump musculature, as well as thick cuticular proboscis walls, which are functionally related to feeding performance in species possessing an extremely long proboscis. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 291–304.     

Comparative Ultrastructure and the Evolution of Nemertines

Nemertines are traditionally viewed as acoelomates with bloodvascular systems and a specialized cavity housing an eversibleproboscis, or, as coelomates with modified coelomic cavitiesforming the rhynchocoel, lateral vessels and gonadal sacs. Comparativeultrastructural data such as cell polarity, the occurrence ofpodocytes and mesodermal cells with rudimentary cilia for proboscis,rhynchocoel, lateral and dorsal vessels, and gonadal liningsare consistent with their interpretation as modified coelomiccavities as found, analogously, in the higher leeches. Theseand additional data regarding musculature, connective tissue,glia-like cells and gametes for 13 species affirm that nemertinesare intermediate between full-blown coelomate and acoelomategrades of organization. A comparison of ultrastructural details,however, indicates that their acoelomate attributes evolvedsecondarily from a more typical coelomate condition.     

The Allometry of Bee Proboscis Length and Its Uses in Ecology

Allometric relationships among morphological traits underlie important patterns in ecology. These relationships are often phylogenetically shared; thus quantifying allometric relationships may allow for estimating difficult-to-measure traits across species. One such trait, proboscis length in bees, is assumed to be important in structuring bee communities and plant-pollinator networks. However, it is difficult to measure and thus rarely included in ecological analyses. We measured intertegular distance (as a measure of body size) and proboscis length (glossa and prementum, both individually and combined) of 786 individual bees of 100 species across 5 of the 7 extant bee families (Hymenoptera: Apoidea: Anthophila). Using linear models and model selection, we determined which parameters provided the best estimate of proboscis length. We then used coefficients to estimate the relationship between intertegular distance and proboscis length, while also considering family. Using allometric equations with an estimation for a scaling coefficient between intertegular distance and proboscis length and coefficients for each family, we explain 91% of the variance in species-level means for bee proboscis length among bee species. However, within species, individual-level intertegular distance was a poor predictor of individual proboscis length. To make our findings easy to use, we created an R package that allows estimation of proboscis length for individual bee species by inputting only family and intertegular distance. The R package also calculates foraging distance and body mass based on previously published equations. Thus by considering both taxonomy and intertegular distance we enable accurate estimation of an ecologically and evolutionarily important trait.     

6.5-mm human embryo with a single nasal placode: cyclopia or hypotelorism?

A macroscopic study on the missing elements in cyclopia (a single eye or closely approximated eyes with all intergrades in a single orbit) with or without proboscis and hypotelorism was performed on 12 human fetuses and 2 human fetal skulls. In addition, microscopic investigations were carried out on the orbital contents of the cyclops with a single eye without proboscis, crown-heel length (C-HL) 37 cm, and on the 6.5-mm-crown-rump length (C-RL) human embryo with a single nasal placode localized in front of two eye cups. In the embryo and in all 14 fetal cases the midfacial region was more-or-less deficient. In the two cyclopia cases without proboscis the nasal placode(s) had not developed at all during the embryonic period. In cases with proboscis, consisting of a single tube localized above both eyes, and in the hypotelorismic specimens, there could only have been a single nasal placode during development: a situation evident in the 6.5-mm-C-RL human embryo. In this holoprosencephalic embryo the single nasal placode was undulated, as if formed from two fused nasal placodes, and flanked by the prospective areas for the lateral nasal processes. Caudally, it was bordered by the maxillary processes. In view of the position of the single placode in this embryonic face, as described above, it is most likely that this is a preliminary stage of hypotelorism. Moreover, both medial nasal processes with the internasal groove in between, i.e., the interplacodal area, were missing.(ABSTRACT TRUNCATED AT 250 WORDS)     

Flies and concealed nectar sources: morphological innovations in the proboscis of Bombyliidae (Diptera)

Bee-flies (Bombyliidae) have morphological adaptations of the mouthparts to particular floral traits. To investigate this the short, plesiomorphic proboscis of Hemipenthes morio was compared with the long, apomorphic proboscis of Bombylius major . A novel feeding position enables B. major to use flowers that open to the side as additional nectar sources. The new horizontal feeding position is enabled by the prolonged ventral base of the proboscis. Bombylius major exploits deep corolla tubes with an elongate proboscis, and an increased efficiency in both the suction pumps and the sealing mechanisms of the proboscis. The exploitation of narrow corolla tubes is made possible by the shift from a sponging feeding mode, exhibited by H. morio , to the exclusively sucking mode in B. major . Besides quantitative changes in the proportions of the different proboscis components, labellar movements as well as the structures of saliva distribution are changed along with this shift. The labial musculature of B. major does not significantly differ from the plesiomorphic state, since both examined species do not only feed on nectar, but also on pollen.     

Groundplan Anatomy of the Proboscis of Butterflies (Papilionoidea, Lepidoptera)

The anatomy of the proboscis was studied in representatives of all major subfamilies of Papilionoidea and several outgroup taxa which included Hesperiidae, Hedylidae and Geometroidea. In all species the cross-sectional outline of the tapering proboscis continuously changes from proximal to the tip while the central food canal, formed by the concave medial galeal walls, retains its oval shape. Each galea contains three types of muscles, a branching trachea, nerves, sensilla, and at least one longitudinal septum. We focused on the varying arrangement and distribution of the intrinsic galeal muscles from the basal galeal joint to the tip region. The plesiomorphic condition of the galeal composition of Papilionoidea is regarded to include one basal intrinsic muscle in the basal joint region and two series of intrinsic muscles, i.e. the lateral intrinsic galeal muscles and the median intrinsic galeal muscles, both series extending from the proximal region to the tip region. The plesiomorphic arrangements of the intrinsic muscle series are found in all representatives of Papilionidae, in one species of Lycaenidae (sensu lato), in many Nymphalidae (sensu lato), and in all outgroup species. Three apomorphic character states are distinguished regarding the presence and extension of the median intrinsic galeal muscles. (1) Present up to 35% of the proboscis length and absent distally in Pieridae, Lycaeninae (Lycaenidae), Satyrinae (Nymphalidae), and Danainae (Nymphalidae). (2) Present in the proximal third of the proboscis and again near the tip between 80 and 90% of the proboscis length in the examined Heliconiinae (Nymphalidae). (3) Completely absent, as in one lycaenid species from the subfamily Riodininae.     

The Proboscis Armature of Turbellaria-Kalyptorhynchia,a Derivative of the Basement Lamina?

The fine structure analysis of the proboscis teeth in the genus Carcharodorhynchus Meixner and of the proboscis hooks in the genus Diascorhynchus Meixner revealed that these structures are not derived from local cuticularizations of the epidermis, but rather are closely associated with the basement lamina of the proboscis sheath epithelium. In Diascorhynchus the hooks are specializations within the basement lamina enclosing the proboscis. The proboscis teeth in Carcharodorhynchus seem to be either an intracellular differentiation within the modified proboscis sheath epithelium or, as in Diascorhynchus , a derivative of the basement lamina. The latter is supported by the finding of similar, dermal teeth clearly derived from the basement lamina in the body wall of several species of the genus Carcharodorhynchus. It appears from this study that most "cutic-ular" differentiations of the body wall in Turbellaria are either derivatives of the basement lamina or of the cytoplasm of the epidermis cells. These considerations are discussed in the light of possible evolutionary pathways leading from a ciliated epidermis (e.g. free-living acoelomates) to a cuticularized hypodermis (e.g. Aschelmimhes and Articulata).     

Social organization and ecology of proboscis monkeys (Nasalis larvatus) in mixed coastal forest in Sarawak

Data are presented from a 16-month study of proboscis monkeys in an area of mixed coastal forest in Sarawak. The population density, social organization, and feeding and ranging behavior are described in detail. Results are compared with those from other primates in an attempt to understand why females of certain species (including proboscis monkeys) transfer between social groups. The scarcity of available food and reasons for the limited habitat preferences of proboscis monkeys are also discussed.     

Morphological fine tuning of the feeding apparatus to proboscis length in Hesperiidae (Lepidoptera)

The form and function of the hesperiid feeding apparatus was studied in detail. The butterflies in the family Hesperiidae are of particular interest because the longest proboscis ever recorded in Papilionoidea was found in the Neotropical genus Damas. We focused on the functional morphology by comparing proboscis morphology as well as size and composition of both the stipes pump and the cibarial suction pump in skippers with short and extremely long proboscis. Results revealed that all studied Hesperiidae have the same proboscis micromorphology and sensilla endowment regardless of the proboscis length. However, the numbers of internal muscles of the proboscis, the morphology of the stipes pump as well as the pumping organs for nectar uptake are related to the proboscis length. We conclude that the low number of tip sensilla compared to proboscis length is responsible for remarkably longer manipulation times of long‐proboscid species during flower visits. The organs for proboscis movements and nectar uptake organs are well tuned to the respective proboscis length and are accordingly bigger in species with a proboscis that measures twice the body length.     

Rhinoglena frontalis (Rotifera,Monogononta): a scanning electron microscopic study

Females and males of Rhinoglena frontalis (Monogononta, Epiphanidae) are observed by SEM and their external morphologies are compared. The two sexes differ in size and shape of the body. The female body is fusiform with a short, conical foot, while the male body is more slender and has a rather long foot. The rotatory apparatus (or corona) of both sexes is similar with only minor differences and consists of rows and tufts of cilia arranged around the mouth opening. The corona is made of two paired lobes lateral to the mouth and of a third prominent dorsal lobe, usually called proboscis. The three lobes are lined externally by dense rows of cilia, which constitute the cingulum, used for swimming. The central surface of the proboscis is covered with numerous longitudinal rows of cilia bent towards the mouth. The lateral lobes show, on their central surfaces, two concentric arcs of cirri (made of tightly packed cilia) bent towards the mouth. The similar organization of the rotatory apparatus of both sexes is related to the fact that the male, in this species, is able to feed and has a developed mastax and digestive system. The trophi of both sexes are illustrated and compared.     

The effect of proboscis and corolla tube lengths on patterns and rates of flower visitation by bumblebees

Summary The rates at which bumblebees of different proboscis lengths forage on flowers of a series of corolla tube lengths were determined. The results indicate significant correlations between proboscis length and time spent by bees on flowers. Bumblebees of long proboscis length can forage significantly faster than bees of shorter proboscis length on flowers with long corolla tubes. There is also evidence which suggests that bumblebees of short proboscis length prefer and are more efficient on short corolla tubes. These results support the use of proboscis length as a morphological indicator of resource utilization in bumblebees.     

Mouthpart separation does not impede butterfly feeding

The functionality of butterfly mouthparts (proboscis) plays an important role in pollination systems, which is driven by the reward of nectar. Proboscis functionality has been assumed to require action of the sucking pump in the butterfly's head coupled with the straw-like structure. Proper proboscis functionality, however, also is dependent on capillarity and wettability dynamics that facilitate acquisition of liquid films from porous substrates. Due to the importance of wettability dynamics in proboscis functionality, we hypothesized that proboscides of eastern black swallowtail (Papilio polyxenes asterius Stoll) (Papilionidae) and cabbage butterflies (Pieris rapae Linnaeus) (Pieridae) that were experimentally split (i.e., proboscides no longer resembling a sealed straw-like tube) would retain the ability to feed. Proboscides were split either in the drinking region (distal 6–10% of proboscis length) or approximately 50% of the proboscis length 24 h before feeding trials when butterflies were fed a red food-coloring solution. Approximately 67% of the butterflies with proboscides split reassembled prior to the feeding trials and all of these butterflies displayed evidence of proboscis functionality. Butterflies with proboscides that did not reassemble also demonstrated fluid uptake capabilities, thus suggesting that wild butterflies might retain fluid uptake capabilities, even when the proboscis is partially injured.     

Proboscis length and resource utilization in two Uruguayan bumblebees: Bombus atratus Franklin and Bombus bellicosus Smith (Hymenoptera: Apidae)

Bumblebees (Bombus sp.) are eusocial insects with an annual life cycle whose use as pollinator of crops has gained great importance in modern agriculture. Several authors have found that resource use in Bombus species is usually based on the correlation between the proboscis length of the bumblebees and the corolla depth of the flowers. The aim of this study was to determine proboscis length of Bombus atratus and B. bellicosus, two Uruguayan bumblebees, and verify the resource exploitation testing two cultivated species, the red clover and the bird's foot trefoil. Bumblebee foraging activity was recorded in two culture conditions: in a red clover and bird's foot trefoil mixed meadow, and in contiguous plots of these legumes, and the proboscis length of collected foragers was determined. Both species of bumblebees exploited red clover and bird's foot trefoil although they did it in different proportions in all instances tested. The results indicated that the choice of the resources in B. atratus and B. bellicosus was influenced by their proboscis length. Bombus atratus has a longer proboscis and preferably visited red clover, possibly obtaining nectar easier and faster than B. bellicosus, which has a shorter proboscis. Bombus bellicosus used both resources without any clear preference.