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Die Sekretdarbietung während des Paarungsverhaltens von Ischyropsalis C. L. Koch (Opiliones)
... As mentioned before, the copulatory behavior in P. thorellii is one of the longest found so far for the suborder (Matthiesen 1983;Gnaspini 1995;Machado & Oliveira 1998;Elpino-Campos et al. 2001;Machado & Macías-Ordóñez 2007;Nazareth & Machado 2009Buzatto et al. 2011) and within species of other suborders (Eupnoi: Macías-Ordóñez 1997, 2000Willemart et al. 2006;Dyspnoi: Pabst 1953;Martens 1969), and was characterized by tactile courtship (touches with legs I and II) like many other harvestmen (Machado & Macías-Ordóñez 2007). Copulatory position (face-to-face and forming a 908angle) is similar to what is observed in other harvestmen; the mutual chelicerae holding has not been reported for the suborder Laniatores. ...
... In females, cheliceral holding was observed in a few species of the genus Ischyropsalis C.L. Koch, 1839 (Dyspnoi) (see Table 12.1 in Machado & Macías-Ordóñez 2007). Females grab the base of male's chelicerae with her chelicerae, bringing them close to her mouth and maintaining that position until mating ends (Martens 1969). The fact that females actively participate in holding and maintaining mating position suggests they have a greater control of mating duration. ...
... The fact that females actively participate in holding and maintaining mating position suggests they have a greater control of mating duration. In fact, it was observed in these species and both P. thorellii and other gonyleptids that females are able to end mating (Pabst 1953;Martens 1969;Nazareth & Machado 2009). P. thorellii females lower their bodies, forcing males to withdraw the penis and release the chelicerae. ...
In order to study how sexual selection takes place during mating, it is necessary to have a clear knowledge of the interactions that occur throughout mating and which morphological and behavioral traits are involved. Available information about harvestman reproductive biology is mainly restricted to anecdotal field observations, most of them lacking a detailed description and quantification of mating behavior. In this paper, we study the reproductive behavior of the gonyleptid Pachyloides thorellii Holmberg, 1878 (Pachylinae) and provide quantitative and descriptive information about its sexual behavior. We observed 15 matings, measured females and males, and analysed our behavioral data in the context of individuals' sizes. We observed conspicuous pre-copulatory, copulatory and post-copulatory courtship. We also found that females have several strategies to reject males' mating attempts. Like most gonyleptids, males and females of P. thorellii mate in face-to-face position; however, we observed that both male and female clasp their chelicerae mutually. This behavior has not previously been reported for the suborder Laniatores. The information obtained through this study establishes the basis for further studies on this species' reproductive biology and supports the suitability of this species as a model to explore the importance of male copulatory courtship for female choice and sperm use.
... Fertilization is internal, and mature spermatozoa are immobile (Morrow 2004). Courtship before intromission is often quick and tactile, but in some cases males may offer a glandular nuptial gift before copulation (Martens 1969). Many studies also mention intense courtship during intromission and mate guarding after copulation (reviewed in Machado & Macías-Ordóñez 2007; see also Nazareth & Machado 2009). ...
... Only fi ve descriptions of copulation in the suborder Dyspnoi exist, but most of them are fairly detailed and show similarities and differences with Eupnoi. In at least one species of Paranemastoma (Nemastomatidae, Troguloidea) and two of Ischyropsalis (Ischyropsalididae, Ischyropsalidoidea) the bases of the male chelicerae are either offered or somewhat forced into the female's mouth, after which the female obtains a secretion from cheliceral glands (Martens 1969;Meijer 1972). Precopulatory interactions seem to be intense in some species, including male tapping on the female's back, and copulation occurs in a face-to-face position, much as in Eupnoi (Immel 1954;Martens 1969). ...
... In at least one species of Paranemastoma (Nemastomatidae, Troguloidea) and two of Ischyropsalis (Ischyropsalididae, Ischyropsalidoidea) the bases of the male chelicerae are either offered or somewhat forced into the female's mouth, after which the female obtains a secretion from cheliceral glands (Martens 1969;Meijer 1972). Precopulatory interactions seem to be intense in some species, including male tapping on the female's back, and copulation occurs in a face-to-face position, much as in Eupnoi (Immel 1954;Martens 1969). On the other hand, full intromission in Trogulidae occurs in a belly-to-belly position, and females may be able to reject the male by lowering the anterior end of the body (Pabst 1953). ...
... Finally, when males provide food resources to females in exchange for copulation or when males care for the offspring alone, females can choose males in terms of resource quality and/or direct or indirect benefits to the offspring (Borgia 1979;Hoelzer 1989). In harvestmen of the genera Ischyropsalis and Paranemastoma (Dyspnoi), for instance, males offer glandular secretions as a nuptial gift to females before intromission, but the role of this secretion for female mate choice is still unknown (Martens 1969;Meijer 1972). Harvestman species exhibiting exclusive paternal care, in turn, have received increasing attention in recent years, and some information on female mate choice is now available (Requena et al. 2013). ...
... As we stated before, pre-copulatory interactions in some species of Dyspnoi also include the transference of secretions, but in this case, they are produced in a pair of glands located dorsally on the first segment of the male chelicerae, which are either offered or somewhat forced into the female's mouth before intromission (Martens 1969; see also Fig. 12.3a in Machado and Macías-Ordóñez 2007). Although the composition, amount, and quality of the secretion offered by males in the suborders Eupnoi and Dyspnoi may influence the donors mating success and paternity, no study has investigated these questions in nuptial gift-giving harvestmen so far. ...
... The Dyspnoi ovipositor is shorter, and the seminal receptacles are smaller and highly variable in number. Detailed records of copulation for two species of Ischyropsalis (Dyspnoi) show a few short intromissions, while the female grasps and apparently feeds from the male's cheliceral glands (Martens 1969). The genitalic interaction during copulation, however, has never been described. ...
Harvestmen
belong to the order Opiliones
, and, unlike other arachnids, they are highly polygynandrous
, with both males and females mating multiply throughout the breeding season. In this chapter, we review the current information on sexual selection in the group, focusing mostly on intersexual interactions. Particularly, we provide an overview of harvestman mating systems, examine different temporal phases of male–female sexual interactions, and explore cases of sex role reversal
. Several traits in harvestmen make them unique in the context of most previous studies of sexual selection. First, they have evolved an intromittent organ
independently of other well-studied taxa, such as insects, spiders, and mammals. Second, the lack of long-range perception mechanisms reduces the window of opportunity for males and females to exchange information during the very short period between the first contact and intromission
. In some cases, however, acceptance or rejection of a mate may be based on information gathered before contact, such as the quality of the male territory or the presence of eggs in his nest. Regardless of the role of pre-copulatory interactions, actual fertilization success
is likely to be strongly dependent on the outcome of copulatory and post-copulatory processes. In this sense, the fact that most species are highly promiscuous and have sperm cells that lack flagella, which are stored near to the tip of the ovipositor and used to fertilize eggs immediately before oviposition, renders Opiliones a fertile ground to study the role of cryptic female choice, sperm competition, and sexual conflict.
... El género Ischyropsalis C.L.Koch, 1836, único representante europeo de la familia Ischyropsalididae, alberga dos docenas de especies, la mayoría siendo endemismos de muy escasa extensión geográfica (Martens 1969(Martens , 1978Schönhofer 2013). Se distribuyen por las regiones montañosas del sur de Europa, desde los Cárpatos hasta el noroeste de la península Ibérica (Schönhofer et al. 2015); casi todas las especies parecen capaces de mantener poblaciones en cavidades cársticas, y muchas especies son estrictamente troglobias. ...
... Son especies de gran tamaño e inmediatamente reconocibles a nivel genérico por el enorme tamaño y robustez de los quelíceros, más largos que el cuerpo y de color negro brillante (Martens 1969(Martens , 1978. Su hábitat es originalmente muscícola y únicamente se encuentran en ambientes con humedad muy elevada. ...
First observations of predation by troglobitic Ischyropsalis (Opiliones: Ischyropsalididae) on oligochaetes.
The first observations of predation on lumbricids by two troglobitic species of the genus Ischyropsalis C.L.Koch, 1839, I. magdalenae Simon, 1881 and I. sp. aff. dispar Simon, 1872, respectively from two caves in Bizkaia (Cueva de Pedro González in Galdames and Cueva de Armiñeta in Zeberio). And from Cueva del Sumidero (Puente Viesgo, Cantabria) another observation of the troglophilic species Ischyropsalis nodifera Simon, 1879 feeding on a lumbricid is reported.
... Paranemastoma (Nemastomatidae) release secretions in a pair of glands located dorsally on the first segment of their chelicerae, which are offered to the female before intromission (Martens 1969;Meijer 1972; Figure 5B). This is an interesting example of convergent evolution of nuptial gifts in harvestmen. ...
... Note that the female"s mouthparts are in contact with the base of the penis, probably feeding on glandular secretions (vertical arrow). (B) Mating pair of Ischyropsalis hellwigi (Dyspnoi: Ischyropsalididae) during pre-copulatory interactions: female on the right and male on the left (photo by Jochen Martens; original source: Martens, 1969). Note that the female"s chelicerae are in contact with the base of the male"s chelicerae (arrow), where a glandular secretion is released. ...
Harvestmen are a major arachnid order that has experienced a dramatic increase in biological knowledge in the 21 st century. The publication of the book Harvestmen: The Biology of Opiliones in 2007 stimulated development of many behavioral studies. Although the book is relatively recent, our understanding of the reproductive biology of harvestmen is already outdated due to the fast accumulation of new data. Our goal is to provide an updated review of the subject to serve as benchmark for the following years. In the pre-copulatory phase, we explore the evolution of facultative parthenogenesis, the factors that may affect the types of mating system, and the role of nuptial gifts in courtship. Regarding the copulatory phase, harvestmen are unique arachnids because they have aflagellate spermatozoa and a penis with complex morphology. We discuss the implications of these two features for sperm competition and cryptic female choice. In the post-copulatory phase, we connect oviposition site selection and climate conditions to the widespread occurrence of resource defense polygyny, alternative reproductive tactics, and sexual dimorphism in several clades of tropical harvestmen. Finally, we present the different forms of parental care in the order, and discuss the benefits and costs of this behavior, which can be performed either by females or males. Throughout the review, we indicate gaps in our knowledge and subjects that deserve further studies. Hopefully, the information synthesized here will stimulate researchers worldwide to embrace harvestmen as a study system and to improve our effort to unravel the mysteries of their reproductive biology.
... The sexual behaviour of Dicranolasma remains unknown; however Gruber (1996) reported a male and a female of D. scabrum (Herbst 1799) facing each other with the prosoma of one covering the other which corresponds to the typical frontal copulation position of many harvestmen including the closely related "dyspnoan" families Nemasomatidae (Immel 1954) and Ischyropsalididae (Martens 1969). Further circumstantial evidence for frontal copulation in Dicranolasma is provided by the presence, in all species of the genus except D. apuanum Marcellino 1970 (Gruber 1998), of "epigamic" or "pheromone" glands on chelicerae (Martens & Schawaller 1977) as well as glandular tissue in the thickened pedipalp patellae in the species of the D. scabrum group; Martens (1969) has shown that glands on male chelicerae of Ischyropsalis hellwigi (Panzer 1794) (Ischyropsalididae) play a role in "gustatory courtship" while partners are approaching one another front to front. ...
... The sexual behaviour of Dicranolasma remains unknown; however Gruber (1996) reported a male and a female of D. scabrum (Herbst 1799) facing each other with the prosoma of one covering the other which corresponds to the typical frontal copulation position of many harvestmen including the closely related "dyspnoan" families Nemasomatidae (Immel 1954) and Ischyropsalididae (Martens 1969). Further circumstantial evidence for frontal copulation in Dicranolasma is provided by the presence, in all species of the genus except D. apuanum Marcellino 1970 (Gruber 1998), of "epigamic" or "pheromone" glands on chelicerae (Martens & Schawaller 1977) as well as glandular tissue in the thickened pedipalp patellae in the species of the D. scabrum group; Martens (1969) has shown that glands on male chelicerae of Ischyropsalis hellwigi (Panzer 1794) (Ischyropsalididae) play a role in "gustatory courtship" while partners are approaching one another front to front. ...
Laboulbeniales are well known ectoparasites of insects. Among arachnids they were only known parasitizing mites. A new
genus of Laboulbeniales, with one species, Opilionomyces dicranolasmatis, is described for fungi parasitizing Dicranolasma harvestmen (Opiliones) collected in Turkey and Greece. The new genus is characterized by the uniseriate receptacle divided into two parts, below the perithecium as a pedicel and above as a row of cells adnate and following the dorsal side of perithecium. The three upper tiers of wall cells are equal in height but shorter than the lower tier. The new genus is accommodated in the subfamily Laboulbenioideae. Similarly to some other Laboulbeniales found on insects and millipedes, Opilionomyces shows site specificity, and we relate its restricted distribution on the pedipalps and chelicerae of the harvestmen to sexual transmission of the fungus. Although both Acari and Opiliones belong to the Arachnida subphylum within arthropods, the Laboulbeniales parasitizing the two orders show no morphological evidence of being closely related.
... The most likely mating system of species belonging to the suborders Cyphophthalmi and Dyspnoi, as well many species of the suborder Eupnoi, is scramble competition polygyny . Among these groups, which are mainly distributed in extratropical regions (Kury, 2013), females lay eggs on substrates that cannot be profitably monopolized by males, such as soil (Juberthie, 1972), the bark of trees, leaf litter (Edgar, 1971), cracks on rocky walls (Martens, 1969;Wijnhoven, 2011), and abundant snail shells (Pabst, 1953). Therefore, the spatial distribution of resources seems to play an important role in determining the type of mating system exhibited by different species . ...
... In several harvestman species with scramble competition polygyny, males offer nuptial gifts to their mates. These nuptial gifts are glandular secretions produced either on the base of male's chelicerae and delivered directly to the female's mouth before copulation (Martens, 1969;Meijer, 1972) or on the base of the penis, where females feed during penetration (Macías-Ordóñez et al., 2010;Burns et al., 2013). Although males of these species may fight for the access to receptive females, there is no evidence that males defend them. ...
... Bei allen Arten, die dieses Drtisenorgan besitzen, ist es auf das n~nnliche Geschlecht beschr/inkt; es fungiert als sekund~ires Geschlechtskennzeichen. Die Kombination dieser oft m/ichtig entwickelten Driise und ihre Zuordnung nur zu den d lielg auf eine Funktion im Sozialverhalten, vor allem bei der Fortpflanzung, schliegen. Diese Erwartung erftillte sich, denn ftir zwei Arten der Gattung Iscbyropsalis C.L. Koch konnte nachgewiesen werden, dag das Sekret im Paarungsverhalten -bei der ,,gustatorischen Balz" -eingesetzt wird (Martens, 1967(Martens, , 1969b(Martens, , 1975 Da schon die Austrittsstellen ffir das Sekret auf der Augenseite der Cheliceren (Form der Apophyse, Einzelporus, Porenplatte, Borstenfeld) auf eine bemerkenswerte Viel- gestaltigkeit des Drfisen-Komplexes schliegen lassen, erscheint es sinnvoll, vor weiteren feinstrukturellen Einzeluntersuchungen zuniichst lichtoptisch einen Oberblick fiber die wichtigsten Differenzierungen des Organs zu gewinnen. Zugleich soll der Frage nachgegangen werden, inwieweit evolutive Vorg~nge auf Funktion und Konstruktion des Organs eingewirkt haben. ...
... Die Funktion der Cheliceren-Drfisen, die allein den Schlfissel zum Verst/indnis der Konstruktion liefert, ist in grogem Rahmen bekannt: Fiir 2 Iscbyro- psalis-Arten ist nachgewiesen und fiir eine auch im Film belegt (Martens, 1975), dalg das Sekret im Paarungsverhalten eingesetzt wird. W~ihrend der Balz kommen die ~- Mundwerkzeuge mit der Bfirstenregion und mit dem austretenden Sekret der 5-Cheli- cere in Kontakt, und nur dann findet die Kopulation statt (Martens, 1969b(Martens, , 1975. Abb. ...
Most species of the families Dicranolasmatidae, Nemastomatidae, Ischyropsalididae, and Sabaconidae possess as a secondary male sex character an epidermal glandular organ which is situated in the proximal joint of the chelicerae, and the major parts of which are often shifted into the cephalothorax. The places of discharge of the secretion are always located on the proximal joint as a field of small pores or as one large pore. The field of small pores is usually exposed on a dorso-distal apophysis of the proximal joint.
The complex gland consists of the gland cells proper and of the enveloping cells. They connect the apices of the glandular cells with the secretion reservoir with at least two layers, and join it firmly with the cuticle.
Several types of glands occur. Cells are short and in this case restricted to the apophysis of chelicerae or if apophysis is absent they are concentrated in the proximal joint (type 1;Dicranolasma, Ischyropsalis, Sabacon with exceptions).
From narrow apophysis the bodies of the cells are discharged into the lumen of the proximal joint or even in the distal part of the cephalothorax. In such cases extremely long cell apices join the enveloping glands in the apophysis (type 2;Mitostoma, Nemastoma).
A short channel and a pointshaped discharging place for the secretion results from invagination of the pore field below the surface of the chelicerae; construction of cells as in type 2 (type 3;Carinostoma, Histricostoma).
Extreme prolongation of the channel into the cephalothorax and lampbrush like grouping of the shorter gland cells diminishes the problem of accomodating the gland in a different way (type 4;Paranemastoma).
The function of the gland in social behaviour (gustatory display) makes it possible to recognize enhanced efficiency in the different gland types from 1 to 4. This allows one to decide in which direction the evolution of cheliceral glands proceeded.
... Other sexually dimorphic structures are described for cyphophthalmid harvestmen but are generally known only for individual genera or species (see review in Willemart and Giribet, 2010). In several Dyspnoi taxa, males possess sexually dimorphic cheliceral glands from which females obtain a secretion before or during copulation (Martens, 1969Martens, , 1973 Martens and Schawaller, 1977). In the suborder Eupnoi, the cheliceral horns and longer pedipalps of males are thought to assist in both intrasexual (i.e., male–male contests) and intersexual (i.e., grasping the female) interactions (Willemart et al., 2006). ...
... Given that the basitarsal glands are sexually dimorphic and present only in males, it is possible that males play the role of attracting, signaling, or courting a mate through the use of pheromones rather than females, as it often would be in spiders. Secretions from sexually dimorphic glands used during courtship are known for several harvestmen species, for example, the nuptial gifts provided to some dyspnoid females by the cheliceral glands of males (Martens, 1969). Within the laniatorean lineage Grassatores, several different families are known to exhibit enlarged tarsomeres on leg I (Pinto-da-Rocha and Townsend et al., 2010). ...
Chemical communication is an important aspect of arthropod biology especially for those arthropods with limited abilities to detect visual and acoustic signals. Sexually dimorphic glands are often associated with the production of pheromones, which play a role in reproductive processes. In the family Manaosbiidae (Opiliones: Laniatores), males exhibit an enlarged, swollen, often fused, and spindle-like basitarsus on leg I. In this study, we provide a novel description of the morphology and ultrastructure of the glandular structures found in the proximal swollen tarsomeres of the male manaosbiid Rhopalocranaus albilineatus Roewer, 1932 and compare the external leg I morphology with that of two other manaosbiid harvestmen (Barrona williamsi Goodnight and Goodnight, 1942 and Cranellus montgomeryi Goodnight and Goodnight, 1947). The two proximal tarsomeres of the male R. albilineatus leg I contain two large, paired, acinar glands consisting of many glandular cells. Cells empty their secretory products into a large, branched epicuticular duct, which exits the leg via a pore on the ventral region. Thus, a total of four glandular structures are present within the two swollen tarsomeres and each possesses a conducting canal and pore. Finally, we discuss possible roles of these basitarsal glands in manaosbiid reproductive biology based on the present understanding of sexually dimorphic glands in other terrestrial arthropods (i.e., insects and spiders).
... The three phases of male-female sexual interactions can be easily observed and distinguished in species of the order Opiliones, commonly known as harvestmen or daddy long-legs (reviewed in Machado et al. 2015). Although the precopulatory phase is usually fast, involving mostly leg tapping and rubbing performed by the male on the female body (e.g., Nazareth & Machado 2009;Fowler-Finn et al. 2014), there are records of some species in which males can offer nuptial glandular secretions to females before intromission (Martens 1969). Copulatory interactions in harvestmen can also involve male leg tapping and rubbing on the female body, and complex genital interactions that may include tactile stimulation of the female genitalia and delivery of glandular secretions as nuptial gifts (e.g., Burns et al. 2013;Pérez-Gonza´lez & Werneck 2018). ...
Describing the signals involved in sexual interactions is crucial to understand how mating and fertilization success is achieved. We analyzed sexual interactions in the gonyleptid harvestman Pachyloides thorellii Holmberg, 1878 to test the possibility of associations between female and male behaviors. For that purpose, we recorded 21 sexual interactions of P. thorellii under laboratory conditions and performed fine scaled analyses of the videos. We found three female pre-copulatory behaviors (“Mouth parts protrusion,” “Genital operculum opening,” and “Ovipositor eversion”) that seem to be related to sexual receptivity and cooperation with mating occurrence, and four copulatory behaviors (“Bucking,” “Pulling,” “Body lowering,” and “Leg II movements”) that could be indicating to the male that further stimulation is required or that mating is about to end. We also found that males use multimodal courtship displays that include the exchange of tactile and possibly chemical signals between sexes. This study shows that courtship and copulation in P. thorellii include intense information flow between sexes and female evaluation from the beginning until the end of the sexual interaction.
... This male behavior could be interpreted as nuptial gift offering, but given the lack of further details, it is unclear. The only known cases of nuptial gifts in harvestmen are glandular secretions, which are produced by the males and transferred to the females via the chelicerae or male genitalia (Martens 1969;Wijnhoven 2011;Fowler-Finn et al. 2018). In the case of neopilionids, males have highly exaggerated chelicerae used for male-male competition (Painting et al. 2015;Powell et al. 2020). ...
... This gland plays an important role in courtship behavior. Its secretion is offered to the females as a nuptial gift (Martens 1969). In four Nemaspela species (borkoae, femorecurvata, ladae and prometheus) the apophysis and the gland are absent, no discharging pores are present nor is a tuft of fine setae surrounding the pores. ...
Two highly specialized endemic troglobiotic harvestman species of the genus Nemaspela Šilhavý, 1966 are described. N. melouri sp. nov. from Melouri Cave and N. prometheus sp. nov. from Prometheus Cave (Sataplia-Tskaltubo karst massif, Imereti region, western Georgia), respectively. Despite the fact that the entrances of the caves are positioned only 2.5 km apart, the new taxa differ from each other distinctly by presence vs. absence of male cheliceral apophysis, which is lacking in the second species. A key to the Caucasian species of the genus is provided. Relationships of Nemaspela species within the genus and with hypothetical epigean ancestors are discussed.
... Generally, the position of extrusion areas for the secretion on the male cheliceral apophysis differs among species and strongly varies among genera of nemastomatines. The secretion plays a role during courtship (Martens, 1969;Martens & Schawaller, 1977). Thus, alongside male genital morphology, the shape of this apophysis and the structure of the internal gland, which discharges its secretion via the cheliceral apophysis (Martens & Schawaller, 1977) plays a major role in systematics of Nemastomatinae. ...
Two species of the genus Starengovia Snegovaya, 2010 from Kyrgyzstan and Pakistan are characterized;
S. kirgizica Snegovaya, 2010 is redescribed and S. ivanloebli proposed as a new species. They belong to the easternmost representatives of the subfamily Nemastomatinae. Both species are separated by some 650 km in north to south direction; the Pakistan record extends nemastomatine distribution to the northwest of the Himalayas. Mediostoma pamiricum Staręga, 1986 probably belongs to Starengovia as well. Within Nemastomatinae Starengovia displays plesiomorphic characters (extremely short glans of penis, armed with inconspicuous robust spicules) and apomorphic ones (long, distinctly inflated base of penis; large lateral foliate wing-like structures on truncus penis). The foliate-wing character is unique among Nemastomatidae. Starengovia may represent a relict line in the early evolution of nemastomatine harvestmen.
... The placement of the secretion area on the male cheliceral apophysis varies among species and strongly varies among genera of nemastomatines. The secretion plays a role during courtship (Martens 1969, Martens & Schawaller 1977. ...
The easternmost Nemastomatinae species, Sinostoma yunnanicum n. gen., n. sp., from northern Yunnan, China is described. It extends the geographic distribution of Nemastomatinae by roughly 3000 km southeastwards. Within Nemastomatinae Sinostoma displays plesiomorphic characters, including the long, basic bulb of the truncus shaft and the extremely short glans of penis, armed with short robust spines. Sinostoma may represent a relict line in the early evolution of nemastomatine harvestmen.
... The placement of the secretion area on the male cheliceral apophysis varies among species and strongly varies among genera of nemastomatines. The secretion plays a role during courtship (Martens 1969, Martens & Schawaller 1977. ...
The easternmost Nemastomatinae species, Sinostoma yunnanicum n. gen., n. sp., from northern Yunnan, China is described. It extends the geographic distribution of Nemastomatinae by roughly 3000 km southeastwards. Within Nemastomatinae Sinostoma displays plesiomorphic characters, including the long, basic bulb of the truncus shaft and the extremely short glans of penis, armed with short robust spines. Sinostoma may represent a relict line in the early evolution of nemastomatine harvestmen.
... Interestingly, Zygopachylus albomarginis (Chamberlin, 1925) and Poassa limbata (Roewer, 1943) both exhibit a unique form of paternal care in which males build mud nests and provide care for eggs and nymphs (Mora, 1990; personal observations of P. limbata). The evolution of this unique form of paternal care may be related to the absence of sexually dimorphic tarsal glands, especially if the glands do in fact play some role in the reproductive biology of these harvestmen, for example, to produce secretions that are used as nuptial gifts, as was observed for some dyspnoid harvestmen with sexually dimorphic cheliceral glands (Martens, 1969). Cosmetidae, Gonyleptidae, Cranaidae, and Manaosbiidae are thought to be closely related, but the relationships among these families remain unclear (Sharma and Giribet, 2011). ...
In at least four closely related families of the diverse harvestmen lineage Gonyleptoidea, males may possess sexually dimorphic tarsal glands in the swollen tarsomeres of the basitarsus and/or metatarsus of leg I. The first histological and ultrastructural examination of the sexually dimorphic tarsal glands in leg I focused only on Manaosbiidae. In this study, we examine the morphology and ultrastructure of the sexually dimorphic glands, and their associated glandular openings, found in the basitarsus and/or metatarsus of leg I of males representing Cosmetidae, Gonyleptidae, and Cranaidae (glandular openings only). In cosmetids and gonyleptids, the tarsal glands are made up of 20–60 glandular units that form distinct groups within the prolateral and retrolateral half of the tarsomere. Each glandular unit consists of a pair of terminal secretory cells, an intercalary cell wrapped around the receiving canal, and a canal cell tightly wrapped around the length of the conducting canal. Cosmetidae, Gonyleptidae, and Cranaidae exhibit remarkably similar tarsal glands and gland openings although the location of the glands in the leg differs slightly among them. Males of these three families exhibit markedly different glands and glandular openings compared to males of the family Manaosbiidae. The sexually dimorphic tarsal glands may provide an important morphological character for determining phylogenetic relationships among gonyleptoid families. Finally, we provide morphological and ultrastructural data for the common tegumental glands. These data indicate that the sexually dimorphic tarsal glands are strikingly similar to, and may possibly be derived from, the tegumental glands.
... The most common reproductive mode in harvestmen involves copulation, although parthenogenesis may occur in some species (Phillipson, 1959; Tsurusaki, 1986). Mating in harvestmen is usually not preceded by courtship displays, but in some species the males may ®ght for access to females (Berland, 1949) while in others the courtship is very simple (see Martens, 1969). ...
Goniosoma longipes is a neotropical cavernicolous harvestman that exhibits parental care. Reproductive activity in G. longipes is more intense during the wet season. Mating lasts up to 3 min, and the whole oviposition process may take over 5 h. During oviposition the female may be reinseminated once or twice by the mating male. Females oviposit 60–210 eggs on the cave wall and guard egg batches for nearly 2 months, until the 1st-instar nymphs disperse. Four main factors can affect egg survival in G. longipes: dehydration, fungal attack, cannibalism, and interspecific predation. Frequency of fungal attack on egg batches was greater in the wet season and more intense near the river inside the study cave. Although egg-guarding by females failed to prevent fungal attack on eggs in G. longipes, the choice of a suitable oviposition site by the gravid female can reduce fungal attack within the cave habitat of this species. Guarding females successfully repel conspecific egg predators, but unguarded eggs are frequently consumed by adult and juvenile G. longipes, as well as cave crickets Strinatia sp. Field experiments in which females were removed from egg batches demonstrated that egg-guarding by the mother has an anti-predator role in G. longipes, with a significant positive effect on egg survival. Male G. longipes actively patrol their egg-guarding mates, and take over brood care for up to 2 weeks if the latter are experimentally removed. The degree to which male assistance can play a relevant role in parental care is still unclear for this species. Damage to the brood is regarded as a major force favouring the evolution of parental care in harvestman species. This field study provides the first experimental demonstration that egg-guarding by females affords protection against egg predation in a harvestman species.
... Especially the genus Ischyropsalis C.L. Koch, 1839 was thoroughly investigated by Martens (1969a). He defined biospecies status within the genus and identified reliable genital characters, as well as somatic ones based on a peculiar ethological trait in Ischyropsalis, a gustatory courtship during which, prior to copulation, a secretion is offered to the female (Martens 1969b). In most Ischyropsalis species this secretion is produced in a massive gland situated in the distal part of the proximal male cheliceral article (Martens & Schawaller 1977). ...
Ischyropsalis dentipalpis Canestrini, 1872 is re-defined, and a neotype from the type locality in the Aosta Valley, Italy, is assigned. I. helvetica Roewer, 1916 (sensu Martens 1978) is shown to be conspecific and therefore is synonymised with I. dentipalpis. A population from Bergamo Province, Italy, formerly assigned to I. entipalpis (sensu Martens 1978), is here regarded as a different species and described under the name Ischyropsalis lithoclasica sp. n. Discrimination of males is possible by genital characters and by the shape of the cheliceral and pedipalpal apophyses. Females are distinguished by peculiarities of cheliceral spination. The distribution of I. lithoclasica n. sp. is allopatric to that of I. dentipalpis.
... Das $, das allein diese Drüse besitzt, wirbt um das 0, indem es ihm das Distalende der Cheliceren-Grundglieder entgegenstreckt, auf dem das Sekret austritt. Erst dann, wenn das 9 das Sekret mit den Mundwerkzeugen berührt, kann die Kopulation stattfinden (Martens, 1969). Die Drüse produziert somit eine Substanz, die als Pheromon wirkt. ...
1. Nemastoma dentigerum Canestrini possesses as a secondary male sex character an apophysis on the first member of chelicerae. The apophysis contains a gland. 2. The single gland cells always join as groups of three cells whose apices are embraced by three enveloping cells. These six cells operate as a unit within a complex gland organ. Every gland cell is divided into three parts: 1. a basal section containing the nucleus; 2. a central very long section of the gland cell tube, and 3. an apex of the gland cell with a cavity containing microvilli. 3. Two types of cells of a gland unit secrete: the gland cells proper and the inner enveloping cell (H1). Secretion of both types of cells flows in a common reservoir surrounded by H1 and further through the bent channel to the surface of the apophysis by pressure of hemolymph. 4. The enveloping cells are compared with those of hair sensilla of insects and arachnids. A common function of H1 in both organs is pointed out. Central (H2) and outer enveloping cell (H3) work in a different way: secretion of hair and follicle (hair sensilla) and probably only supporting function (cheliceral gland).
The scientific life of Prof. Dr Jochen Martens (Germany: Mainz) is illuminated on occasion of his 80th birthday. Facts and impressions are given as well as lists of his publications (329), taxa he has described (2 families, 29 genera, 296 species) and that have been dedicated to him (11 genera, 219 species, 1 subspecies). Jochen Martens is a renowned specialist for birds (Aves) and for harvestmen (Opiliones). So far, he travelled to 27 countries in 80 journeys with Nepal and the Himalayas as one geographic focus.
The catalogue of Harvestmen from France including mainland and overseas territories (Delfosse, 2017) is completed and corrected.
A new harvestman (Arachnida: Opiliones) from Baltic amber (Palaeogene: Eocene; ca. 44–49 Ma) is described as Piankhi steineri n. gen., n. sp. This enigmatic fossil expresses long, slender pedipalps without a tarsal claw, which is characteristic for the suborder Dyspnoi. The chelicerae are notably enlarged and the dorsal body surface is formed from a carapace with a separate prosomatic tergite (metapeltidium), plus a large opisthosomal scute (or scutum parvum). However these characters, combined with the distinctly spiny limbs and further rows of spines across the fossil's opisthosoma, have no parallel among the modern dyspnoid harvestmen that we are aware of. The fossil resolves features reminiscent of modern members of the dyspnoid families Ceratolasmatidae, Nipponopsalididae, Ischyropsalididae and Sabaconidae, but does not show unequivocal apomorphies of any one particular family. We must entertain the possibility that this is an extinct body plan from the Eocene of north-central Europe, and we tentatively refer the fossil to a new genus in an unresolved position among the Ischyropsalidoidea (Dyspnoi). An amorphous triangular structure behind the anal region is assumed to be faecal matter, rather than part of the original anatomy.
doi:10.1002/mmng.201200007
There are records of glands that produce sexual pheromones that are released into the environment or applied directly on sexual partners. Within Opiliones (Arachnida), several harvestmen in the suborder Laniatores have sexually dimorphic glands on legs I and IV, the mode of use of which is recorded only in two species but their function is unknown: While walking, males rub the glands against the substrate or against their body. Here we test an alternative and non-exclusive hypothesis that the glands present on the legs of male Gryne perlata (Cosmetidae) produce contact pheromones used in mating. We predicted that males would touch the females with the gland openings or with other male body parts previously rubbed by these glands. We also predicted that there are chemoreceptors on those parts of the females where males touch them. We analyzed 13 videos of G. perlata mating, a species in which the males have glands on legs I and IV of unknown function. We also analyzed 14 videos of Discocyrtus pectinifemur (Gonyleptidae) mating as a control, a species that lacks these glands. Finally, we l ooked for chemoreceptors on the legs of female G. perlata using a scanning electron microscope. During copulation, males of both species rubbed the legs of females with their first pair of legs, but not with the regions of these legs where the openings of the glands are. The fourth pair of legs were only used to support the body. Rubbing other body parts of the female by males with their glands was not observed during mating. Setae on the legs of the female did not have tip pores and therefore do not seem to be chemoreceptors. We therefore did not find any evidence that these sexually dimorphic glands in G. perlata release contact pheromones during mating. © Institute of Entomology, Biology Centre, Czech Academy of Sciences, Ceské Budejovice.
Harvestmen (Arachnida: Opiliones) are a highly diverse group found in extremely different environmental conditions. For this and other reasons described in this chapter, they offer a novel and unique opportunity to explore hypotheses regarding the effect of abiotic environmental conditions on several life-history traits, and thus on their mating systems. Here we review the current knowledge about harvestman mating systems, and then elaborate on how environmental factors may influence their life cycles and reproductive biology. Next, we use comparative methods to show that the occurrence of maternal care in the order, as well as the length of mating seasons, may be influenced by an interaction between environmental temperature and precipitation. We discuss how these influences might extend to the types of mating systems found in the species of the order, and then suggest potential avenues to obtain and analyze the data required to improve our understanding of the macroecology of harvestman mating systems.
We estimated a multigenic molecular phylogeny and reconstructed biogeographic history for the European harvestman genus Ischyropsalis C.L. Koch 1839 (Dyspnoi). To reconstruct historical biogeographic patterns we conducted an algorithmic VIP analysis which revealed patterns consistent with a vicariance-dominated history. The existing morphology-based systematic framework for Ischyropsalis is mostly inconsistent with molecular phylogenetic results, and a new informal system is established that recognizes three main clades and several sub-clades. Species-level analyses revealed two non-monophyletic species (I. pyrenaea Simon 1872 and I. luteipes Simon 1872); subspecies of I. pyrenaea are distant relatives, and are formally elevated to species (I. pyrenaea pyrenaea to I. pyrenaea and I. pyrenaea alpinula to I. alpinula). A preference for cryophilic microhabitats has favored the diversification of high-altitude and cave-dwelling Ischyropsalis species; molecular phylogenetic data suggest that cave-dwelling species have evolved multiple times independently.
A new harvestman (Arachnida: Opiliones) from Baltic amber (Palaeogene: Eocene; ca. 44–49 Ma) is described as Piankhi steineri n. gen., n. sp. This enigmatic fossil expresses long, slender pedipalps without a tarsal claw, which is characteristic for the suborder Dyspnoi. The chelicerae are notably enlarged and the dorsal body surface is formed from a carapace with a separate prosomatic tergite (metapeltidium), plus a large opisthosomal scute (or scutum parvum). However these characters, combined with the distinctly spiny limbs and further rows of spines across the fossil's opisthosoma, have no parallel among the modern dyspnoid harvestmen that we are aware of. The fossil resolves features reminiscent of modern members of the dyspnoid families Ceratolasmatidae, Nipponopsalididae, Ischyropsalididae and Sabaconidae, but does not show unequivocal apomorphies of any one particular family. We must entertain the possibility that this is an extinct body plan from the Eocene of north-central Europe, and we tentatively refer the fossil to a new genus in an unresolved position among the Ischyropsalidoidea (Dyspnoi). An amorphous triangular structure behind the anal region is assumed to be faecal matter, rather than part of the original anatomy.
doi:10.1002/mmng.201200007
All species of Sironidae possess as a secondary male sex character an epidermal gland organ in the last tarsal joint of the fourth leg. The glandular secretion emanates from a conical apophysis, the adenostyl, which is located dorsally on the joint.
InSiro duricorius (Joseph, 1868) the complex organ consists of numerous functional units, the secretion of which flows into the common channel opening on the adenostyl.
Every gland unit contains 5 cells: 3 gland cells (DZ), 1 enveloping cell (HZ), 1 duct cell (KZ). The enveloping cell connects the gland cells with the duct cell like a sleeve. A marginal fold of the gland cell apex provides tight connection of gland cells and enveloping cell.
Gland cells and enveloping cell are equipped with microvilli, and both cell types secrete in a common reservoir.
The duct of the duct cell is divided into two parts: 1) a distal fibrillaceous-spongy one and 2) a proximal massive one. Both are accompanied only by a single duct cell.
Besides this complex gland organ a similar one is located in the ventral part of the tarsus. Its collecting channel opens into the channel of the main gland.
Isolated functional gland units are scattered over the epidermis of the tarsus; their secretion flows immediately on the tarsal surface.
The gland cells proper produce a protein, the fact is indicated by the electronmicrographs. The secretion of the enveloping cell is a different one, but at this time cannot be attached to a chemical group.
Sexually dimorphic glands often release sexual pheromones both in vertebrates and invertebrates. Species of Laniatores (Arachnida, Opiliones) seem to depend on chemical communication but few studies have addressed this topic. In this study, we review the literature for the Phalangida and present new data for 23 species of Laniatores. In 16 taxa, we found previously undescribed sexually dimorphic glandular openings on the femur, patella, metatarsus, and tarsus of legs I and metatarsus of legs III and IV. For the other species, we provide scanning electron micrographs of previously undescribed sexually dimorphic setae and pegs located on swollen regions of the legs. We also list additional species in which males have swollen regions on the legs, including the tibia, metatarsus, and tarsus of legs I, trochanter and tibia of legs II, femur, metatarsus, and tarsus of legs III, and metatarsus and tarsus of legs IV. The function and biological role of the secretions released by these glands are discussed.
Aus: Zoologische Jahrbücher. Abt. f. Systematik. Bd 82. 1953 Mainz, Naturwiss. F., Diss. v. 20. Febr. 1953 (Nicht f. d. Aust.).
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