Contribution of the maxillary muscles to proboscis movement in hawkmoths (Lepidoptera: Sphingidae)--an electrophysiological study

The role of the maxillary muscles in the uncoiling and coiling movements of hawkmoths (Sphingidae) has been examined by electromyogram recordings, combined with video analysis. The maxillary muscles of adult Lepidoptera can be divided into two groups, galeal and stipital muscles. The galea contains two basal muscles and two series of oblique longitudinal muscles, which run through the entire length of the galea. Three muscles insert on the stipes, taking their origin on the tentorium and on parts of the cranium and gena, respectively. Proboscis extension is initiated by an elevation of the galea base caused by the basal galeal muscles. The actual uncoiling of the proboscis spiral is accompanied by rapid compressions of the stipites which are caused by two of the stipital muscles. The study provides strong support for the hypothesis that uncoiling is brought about by an increase of hemolymph pressure by the stipites forcing hemolymph into the galeae. Recoiling is caused by the contraction of both sets of oblique longitudinal galeal muscles supported by elasticity of the galea cuticle. Finally, the remaining stipital muscle pulls down the galea base which brings the coiled proboscis back to its resting position where it is held in the U-shaped groove of the labium without further muscle activity.     

Proboscis musculature in the butterfly Vanessa cardui (Nymphalidae, Lepidoptera): settling the proboscis recoiling controversy

Krenn, H. W. 2000. Proboscis musculature in the butterfly Vanessa cardui (Nymphalidae, Lepidoptera): settling the proboscis recoiling controversy. —Acta Zoologica (Stockholm) 81 : 259–266 The proboscis of Vanessa cardui (Nymphalidae) contains two basal galeal muscles and two different series of numerous oblique muscles. Both muscle series extend from the proximal region up to the tip‐region; the individual muscles of each series run a constant course throughout the proboscis. In contrast to other butterflies, the knee bend region does not have additional types of muscles. The analysis of shock‐frozen proboscises reveals that the dorsal wall is arched outwardly in the uncoiled, feeding position whereas in the coiled, resting position the dorsal proboscis wall is flat or concave. This results in a significantly greater cross‐sectional area due to the significantly greater dorso‐ventral diameter in uncoiled proboscises. After freezing the proboscis in its distal region, it can still be uncoiled, however, it cannot be fully recoiled. These morphometric and experimental results indicate that the oblique proboscis muscles are responsible for recoiling the proboscis to the resting position.     

Vergleichende Morphologie des Schmetterlingsrüssels und seiner Sensillen — Ein Beitrag zur phylogenetischen Systematik der Papilionoidea (Insecta, Lepidoptera)

Comparative morphology of the butterfly proboscis and its sensilla — a contribution to the phylogenetic systematics of Papilionoidea (Insecta, Lepidoptera) The morphology of the proboscis was investigated in more than 70 European representatives of Papilionoidea using light microscopy and scanning electron microscopy. The composition of the proboscis wall, its surface structures, as well as the shape and distribution of the different types of sensilla are compared. Special attention is given to the tip region and the diversity of the sensilla styloconica. Plesiomorphic features of the proboscis of Papilionoidea were found to include vertically extended exocuticular ribs composing the galeal wall, cuticular spines restricted to the ventral side of the proximal galea, and two rows of fluted sensilla styloconica restricted to the tip region. Apomorphic features of the proboscis in Papilionidae are three rows of small sensilla styloconica. The presence of cuticular spines all over the galeae was identified as an autapomorphy of Pieridae. Possible apomorphies of Nymphalidae are oblique exocuticular ribs of the galeal wall and the great number and length of the sensilla styloconica (significant at p < 0.01, t-test). A possible synapomorphy of Lycaenidae and Riodinidae are cuticlar spines up to the distal galeae. Distinct transformation series of sensilla styloconica give evidence that divergent evolutionary trends led from fluted shafts to a multitude of other shapes in Papilionidae, Nymphalidae (sensu lato), and Lycaenidae. Long smooth-shafted, club-shaped sensilla styloconica, bearing apical spines, are found in Nymphalinae, Apaturinae and Limenitidinae. Highly derived sensilla styloconica evolved in Heliconiinae and Melitaeini, which are arranged in only one row in both taxa. Their shafts are smooth, flattened and bear an excentral sensory cone. Further apomorphic character states are dented flutes which evolved several times, independently from each other in Satyrinae, Lycaeninae and Riodinidae. The results are discussed in a systematical context and provide the basis for a better understanding of the function of different morphological structures of the proboscis in feeding.     

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.     

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.     

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.     

Importance of Queen Bumble Bees as Pollinators Facilitating Inter-morph Crossing in Primula sieboldii

Abstract Flower-visiting insects, the distribution of pollen on an insect's body, and fruit and seed sets of Primula sieboldiiE. Morren were investigated in a maritime deciduous forest habitat of the Hidaka region in Hokkaido. Queens of Bombus diversus tersatus Smith (Hymenoptera: Apidae), Bombus schrencki albidopleuralis Skorikov, and Bombus deuteronymus deuteronymus Schulz were observed to visit the flowers of the species. Bumblebee claw marks were found on the flower petals of 68% of the flowering ramets of the species. Scanning-electronmicroscopy revealed that pin and thrum pollens of P. sieboldii were deposited in different positions along the proboscis of a B. diversus tersatus queen (n=38400 grains). On the glossa, all the observed pollen was of the pin type, while more than half of the pollen grains observed on the upper part of the proboscis, galea, stipes, and maxillary palpus were of the thrum type. In the field, high fruit and seed sets were recorded for both pin and thrum morphs, indicating the occurrence of sufficient legitimate pollination during the flowering season. In addition to the effectiveness of queen bumblebees as pollinators for P. sieboldii suggested in this study, phenological and morphological matches, i.e., early spring flowering during the queen bee emergence season and the similar lengths of the corolla tube and the bee proboscis, also support the inference that queen bumble bees play an important role in legitimate pollination of P. sieboldii.     

Paralineopsis taki gen. et sp. nov., a littoral heteronemertean from Japan,provided with special proboscideal,circulatory and sensory organs of significance to nemertean systematics

Some morphological features with major systematic significance are recorded in the heteronemertean Paralineopsis taki gen. et sp. nov. from Onomichi, Japan as follows: horizontal band of specialized epithelium extends from near apex to the opening of the cerebral organ canal on either side of the head; precerebral region filled with gelatinous (hyaline) connective tissue in which longitudinal muscles are absent; body wall muscles do not accompany rhynchodaeal invagination; rhynchodaeum initially only epidermal; inner longitudinal muscles of ventral wall of cephalic blood lacuna become intimately associated with rhynchodaeum forming a dorsal saddle over it; cerebral organs do not penetrate inner longitudinal muscles, and do not contact blood vascular system; proboscideal diaphragm post-cerebral; outer longitudinal muscles absent throughout body; longitudinal muscles of proboscis derived from inner longitudinal musculature. The systematic relationship of P. taki and Paralineus elisabethae (Schütz, 1911) from Villefranche, France is discussed.     

Food and feeding behavior of the hoplonemertean Oerstedia dorsalis

The monostiliferous nemertean Oerstedia dorsalis was collected from eelgrass (Zostera marina) beds located along the coast of New Jersey, and feeding responses to amphipods and isopods were observed in the laboratory. Tests with 46 worms showed that they fed suctorially on Ampelisca vadorum, Ampithoe longimana, Corophium acherusicum and C. tuberculata. Corophiids were preferred. Upon contact with an amphipod, the proboscis is everted and strikes the prey on the ventral side, immobilizing it in a few minutes. The worm probes the sternal region with its head and everts its proboscis one or more times during the process. The exoskeleton is eventually penetrated by the head, and the stomach is everted into the hemocoel as a flattened funnel-like structure. Peristaltic undulations of the body signify the suctorial action that removes the living contents from the exoskeleton. The actual feeding process (from head penetration to removal of the head) takes about 7 min. O. dorsalis is only the third species within the Prosorhochmidae for which the feeding behavior has been documented. The other two are terrestrial species, and are also suctorial.     

Locomotory and other movements of the proboscis of Bonellia viridis (Echiura, Bonelliidae)

An account is given of the various movements of the proboscis of Bonellia viridis. When at rest, the proboscis is coiled up in front of the trunk. The proboscis lobes progress with a velocity of the order of 1 mm/sec by means of powerful cilia situated on the dorsal surface of the leading edge. The lobes passively drag and uncoil the stem of the proboscis which is further uncoiled by muscular contractions taking place along the uncoiled part of the stem. Proboscis retraction takes place by means of contraction of the numerous longitudinal muscle strands of the stem. The terminal lobes can attach to the substrate and pull the trunk by contraction of the proboscis stem. The proboscis often ties itself into a knot which it can untie.     

Cystacanths of Acanthocephala in notothenioid fish from the Beagle Channel (sub-Antarctica)

The morphology of relaxed cystacanths of polymorphid acanthocephalans collected from notothenioid fishes in the Beagle Channel (Magellanic subregion of sub-Antarctica) is described. A parasite of birds, Andracantha baylisi (Zdzitowiecki, 1986), was found in Patagonotothen longipes and Champsocephalus esox. It has: a proboscis 0.82–0.89 mm long; a proboscis hook formula of 16 rows of 9/10–10/11, including 4–5 basal hooks; distal hooks with the longest blades; a fore-trunk not separated from the hind-trunk by a constriction; large somatic spines arranged in two zones separated by a zone of small, loosely dispersed spines; and only the anterior 36–40% of ventral side of the trunk is covered with spines. One male specimen of Corynosoma sp. was found in Patagonotothen tessellata. It differs from A. baylisi in that the distal proboscis hooks are similar in length to the prebasal hooks, it has a smaller proboscis (0.77 mm) and in the distribution of the somatic spines, which are contiguous with the genital spines on the ventral side of the trunk and lack a zone of small spines between zones of larger spines. A parasite of seals and fur seals, Corynosoma evae Zdzitowiecki, 1984, was found in P. longipes and Champsocephalus esox. It has: a proboscis 0.61–0.78 mm long; a proboscis hook formula of 20–22 rows of 12–13, including 3/4–4 basal hooks; prebasal hooks with the longest blades; a trunk divided into fore-trunk and hind-trunk; somatic spines covering the anterior 64–74% of the ventral side of the trunk; genital spines present only in males; and a terminal genital opening in both sexes. Corynosoma beaglense n. sp. was found in Champsocephalus esox. It has: an almost cylindrical proboscis (length 0.52–0.56 mm); a proboscis hook formula of 16 rows of 9/10–10/11, including 4–4/5 basal hooks; distal hooks shorter than the prebasal hooks; a fore-trunk not separated from the hind-trunk by a constriction; somatic spines contiguous with the genital spines on the ventral side of the trunk of the male and covering the entire length of the ventral side of the female trunk, and the presence of genital spines surrounding the terminal genital pore of the male. The definitive host of this species is unknown.     

Presomal morphology and development of Prosthorhynchus formosus,Prosthenorchis elegans,and Moniliformis dubius (Acanthocephala)

Larvae removed at one-day intervals from laboratory infected intermediate hosts provided material for a comparative study of presomal development in Prosthorhynchus formosus (Van Cleave, '18) Travassos, '26, Prosthenorchis elegans (Diesing, 1851) Travassos, '15, and Moniliformis dubius Meyer, '33. Acanthellae begin development soon after entering intermediate hosts' hemocoels, and by the 18th day all three species possess three nuclear masses representing primordia of the proboscis, proboscis receptacle and ganglion, and trunk musculature and genitalia. Presomal development of P. formosus and P. elegans results in structures concurring with morphology of other adult palaeacanthocephalans and archiacanthocephalans. Development of M. dubius, however, differs from that of other archiacanthocephalans in that the muscular receptacle wall lines the entire surface of the nonmuscular sheath, failing to form a ventral cleft characteristic of other archiacanthocephalans. Unlike receptacle protrusor muscles of other archiacanthocephalan species, those of M. dubius spiral around the receptacle as they extend posteriad to attach individually to a pouchlike, muscular thickening at the receptacle's base. These protrusor muscles are distinct from the receptacle wall, as attested by their development alongside neck retractor muscles, not from the receptacle primordium, and the manner in which they are left to trail behind the receptacle when it is drawn anteriorly into the proboscis during larval development. The proboscis receptacle of M. dubius should not be thought of as being double-walled, as envisioned by previous workers.     

Restricted diet breadth of the larvae of Antheraea assamensis and the role of the labrum‐epipharynx and galeal sensilla

The silkworm, Antheraea assamensis Helfer (Lepidoptera: Saturniidae), grows primarily on Persea bombycina and Litsea polyantha. To understand if the restricted diet breadth is due to the specific role of gustatory sensilla of the larvae of A. assamensis, the same fifth instar larvae retaining only labrum‐epipharynx or galeal sensilla were subjected to food choice tests. The foods used were leaves of two host‐plant and two non‐host‐plant species. Mean per cent consumption and per cent of choosing larvae were used as parameters for drawing conclusions. The finding indicated involvement of the labrum‐epipharynx for acceptance and galeal sensilla for rejection of a non‐host‐plant species. Scanning electron microscope studies revealed the presence of two sensilla on the galea, one lateral and one medial sensilla styloconicum and two gustatory sensilla in the epipharynx of A. assamensis. The study revealed the key role of galeal sensilla in the restrictive diet‐breadth of A. assamensis.     

The larval head of Agathiphaga (Lepidoptera, Agathiphagidae) and the lepidopteran ground plan

Abstract. The larval head of Agathiphaga vitiensis is described. There is a complete hypostomal bridge but no hypostomal ridges. Adfrontal ridges and distinct ecdysial lines are absent. There are two vestigial stemmata (without lenses) on each side. The antenna is one-segmented. All ‘typical lepidopteran’ head setae have been identified. The corporotentorium is very slender; dorsal tentorial arms are present. Intrinsic labral muscles are lacking. The mandible has retained a tentorial muscle. The maxilla is without a discrete cardo and has but a single endite lobe; ‘intrinsic maxillary muscles’ and the ‘cranial flexor of the dististipes’ are lacking. The postlabium is undivided and without setae, the labial palp is one-segmented and the lateral prelabio-hypopharyngeal sclerotization is continued into an oral arm. Some of the ventral pharyngeal dilators arise on the tentorium; mouth-angle retractors and dorsal post-cerebral pharynx dilators are absent. The two brain lobes have almost parallel long axes and are united by a narrow (almost pure neuropile) bridge. The corpora cardiaca and callata are contiguous. The aorta is an open gutter in front of the retrocerebral complex. Available evidence on the ground plan structure of the lepidopteran larval head is reviewed. The ancestral head supposedly was prognathous and was autapomorphic in having the cranio-cardinal articulation far behind the mandible; it had a complete hypostomal bridge but neither hypostomal nor adfrontal ridges, its tentorium was probably stout and with dorsal arms. Paulus & Schmid (1978, Z. zool. Syst. EvolForsch. 16) described a lepidopteran/trichopteran synapomorphy in stemma structure. A tentative table of homologies between cranial setae in Lepidoptera and Trichoptera is presented; it differs considerably from the scheme of Williams & Wiggins (1981, Proc. 3rd Symp. Trichopt.). The mouth parts and their musculature must have been overall very primitive for a panorpid larva, but the number of maxillary palp segments was reduced (three). The ‘dististipes’sensu Hinton is considered to consist of complexly fused parts of the stipes and basal palp segments. The cephalic stomodaeum must have possessed all primitive groups of extrinsic muscles. The incomplete available information on Micropterigidae impedes reconstruction of some details of the lepidopteran ground plan. Larval head structures support the monophyly of an entity comprising the Agathiphagidae + Heterobathmiidae + Glossata. There is one suite of derived characters shared by Heterobathmiidae and Agathiphagidae only and another shared by Heterobathmiidae and the Glossata only; one of these must represent parallelisms.     

Role of galeal sensilla in host recognition and feeding behaviour of the Colorado potato beetle

Abstract. Close-range interactions with plants and the early stages of feeding behaviour of adult Colorado potato beetles, Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae), were studied using beetles with and without various mouthpart sensilla. Representative host, potato (Solarium tuberosum) , and non-host, tomato (Lycopersicon esculentum) , leaves were treated very differently by beetles with and without their galeal or palpal sensilla. Galeal sensilla were particularly important for beetles interacting with these two plants. Compared with normal beetles, fewer beetles without galeae fed on potato leaves, and those that did feed spent more time assessing the plant and took more bites before feeding. On tomtato, more beetles without galeae fed, spent less time assessing the plant and took fewer bites before feeding. Beetles without galeae also had lower consumption rates on potato and higher consumption rates on tomato. Beetles were very reluctant to feed on tomato if the galeae were present so removal of maxillary sensilla was studied only on potato. Without the sensilla on the maxillary palpi only the time between first touch and first bite was affected (lengthened). The importance of galeal sensilla in host recognition by this species and the implications for chemosensory studies are discussed here and in a companion paper.     

Moth pollination of Metaplexis japonica (Apocynaceae): pollinaria transfer on the tip of the proboscis

Asclepiad pollinaria (including pollen masses) attach to diverse body parts of flower visitors in many ways. In this paper, we observed nocturnal moths (Lepidoptera: Pyralidae and Noctuidae) transporting the pollinaria of the Japanese species Metaplexis japonica (Thunb.) Makino (Apocynaceae: Asclepiadoideae) on the tip of the proboscis. Flowers of this species may induce nectar-feeding moths to pull out the proboscis along a guide rail (anther slit), thus clipping the pollinaria onto the tip of the proboscis and transferring the pollinaria to the next flower. The transfer of pollinaria on the unique vector of a moth proboscis tip is an interesting pollination mechanism among previously reported entomophiles.     

The proboscis of eye-frequenting and piercing Lepidoptera (Insecta)

The external structures of the proboscis are investigated in eye-frequenting species of Noctuidae, Geometridae and Pyralidae by means of scanning electron microscopy. They are compared with non-eye-frequenting representatives of these families. In Noctuidae, highly specialized fruit-piercing, skin-piercing blood-sucking, and sweat-feeding representatives have been included. All hemi- and eulachryphagous species have a soft proboscis tip which is characterized by few sensilla and strongly elongated, dentate plates of the dorsal galeal linkage. The latter structures leave broad gaps between them that lead into the food canal at the tip. This arrangement permits the uptake of fluid suspensions such as lachrymal fluid, wound exudates and pus. The modified dorsal galeal linkage is regarded as an adaptation for this highly derived feeding habit. The rough surface of the proboscis is likely to cause irritation and possible mechanical damage to the conjunctiva and cornea which results in an increased lachrymal flow and production of pus. In contrast to fruit-piercing and skin-piercing Noctuidae, there are no erectile structures on the proboscis of eye-frequenting species.—The comparison with related non-eye-frequenting species demonstrates that the particular morphology of the proboscis tip in lachryphagous moths evolved convergently in different families of Leipdoptera.     

The larval head anatomy of Rhyacophila (Rhyacophilidae) with discussion on mouthpart homology and the groundplan of Trichoptera

The external and internal features of the larval head of Rhyacophila fasciata (Trichoptera: Rhyacophilidae) were described in detail. Anatomical examinations were carried out using a multimethod approach including histology, scanning electron microscopy, confocal laser‐scanning microscopy, microcomputed tomography, and computer‐based three‐dimensional reconstructions. Additionally, the information on the larval head of Limnephilus flavicornis (Limnephilidae) and Hydropsyche angustipennis (Hydropsychidae) available in the literature were reinvestigated. These anatomical data were used to address major questions of homology and terminology, that is, the ventral closure of the head capsule, the sclerites, and appendages of labium and maxilla and their muscles. These topics were discussed by summarizing the main hypotheses present in the literature and a critical inclusion of new findings. Consequently, the inner lobe of the maxilla very likely represents the galea. The distal maxillary sclerite (palpifer) is an anatomical composite formation at least including dististipes and lacinia. Based on these homology hypotheses several potential groundplan features of the larval head of Trichoptera were reconstructed. The head of Rhyacophila shows several presumably plesiomorphic features as for instance the prognath orientation of the mouthparts, the well‐developed hypocranial bridge, the triangular submentum and eyes composed of seven stemmata. Derived features of Rhyacophila are the reduced antennae, the anterior directing of three stemmata and the shift of the tentorio‐stipital muscle to the mentum. J. Morphol. 276:1505–1524, 2015. © 2015 Wiley Periodicals, Inc.     

Ultrastructure of Male Sexual Apparatus of Scutellonema brachyurum

Electron micrographs of serial sections show that the male sexual apparatus of Scutellonema brachyurum includes two morphologically identical spicules. Each is composed of a swollen tubular head, crescentic shaft, and leaf-like blade with membranous velum expanded from the central trunk. The spicules are concave and grooved on the ventral side and convex on the dorsal side near the trunk. The trunk is continuous with the shaft and head. Nerve tissue occupies the core of the spicule and includes a dendritic process which gains access to the exterior via a small pore on the lateral side of the spicule tip. Three protractor and two retractor muscles are associated with each spicule. A sensory accessory piece connects with the tip of the gubernaculum and protrudes from the lower side of the opening of the spicular pouch; it protracts and retracts with the muscularized gubernaculum. The gubernaculum varies from bow-shaped in the distal part to boat-shaped in the mid region. A sac exits beneath the accessory piece as a buffer for its movement. A cuticular guiding bar originating from the dorsal wall of the spicular pouch has a tongue. The ventral surface of the tongue is sclerotized to separate the two spicules. It is mobile by muscles of the protractor gubernaculi, retractor gubernaculi, and seductor gubernaculi.