Genital Morphology and Sperm Storage in Pholcus phalangioides(Fuesslin, 1775) (Pholcidae; Araneae)

The genital morphology of female Pholcus phalangioidesis examined to clarify the composition of the uterus externus and the place of sperm storage in this species. Two conspicuous pore plates serve as exits for glandular secretion that gets discharged into the uterus externus. The secretion accumulates close to the pore plates and to some extent in the region of the heavily sclerotized valve that separates the uterus externus from the uterus internus. During copulation, the male transfers spermatozoa and male secretions into the female genital tract where they are embedded and stored in the female secretion. As Ph. phalangioidesdoes not possess any separate sperm storage organs such as receptacula seminis, the glandular secretion serves to store and fix the sperm mass in a specific position within the uterus externus itself.     

Long-duration whirling of Pholcus phalangioides (Araneae, Pholcidae) is specifically elicited by Salticid spiders

Long-duration whirling (gyrating of the body during several hours a day) was shown by the pholcid spider Pholcus phalangioides to salticid spiders and hardly ever to predatory spiders from eight other families in laboratory arenas. Long-duration whirling has not been reported so far, in contrast to short-duration whirling lasting less than a few minutes. Long-duration whirling may have the anti-predatory function of disturbing continual visual fixation of prey in attacking salticids, in contrast to short-duration whirling that has been demonstrated to favour survival of pholcids in the presence of all sorts of predatory spiders.     

Anti-predator defences of Pholcus phalangioides (Araneae, Pholcidae), a web-building and web-invading spider

The behaviours used by Pholcus phalangioides (Fuesslin) (Araneae, Pholcidae) to evade its predators were studied with particular attention being given to a special defence behaviour, whirling. To whirl, this long-legged web-building spider swings its body around in a circle, with its legs remaining on the silk. Experiments were carried out to determine the types of stimuli that elicited whirling. Touching the spider or its web elicited whirling, as did air movement over the spider, but there was no evidence that chemical stimuli from potential predators were important. Small juveniles differed from adult females and larger juveniles by more often dropping from the web instead of whirling when confronted by a potential predator. Besides catching prey on its own web P. phalangioides invades other spiders' webs to catch the other spiders. By whirling in alien webs, P. phalangioides could deter attacks by the resident spider, but P. phalangioides was less inclined to whirl when in an alien than when in its own web.     

The biology of Pholcus phalangioides (Araneae, Pholcidae): predatory versatility, araneophagy and aggressive mimicry

Predatory versatility occurs in Pholcus phalangioides (Fuesslin). In addition to building prey-catching space webs, P. phalangioides invades webs of other spiders and feeds on the occupants. It acts as an aggressive mimic by performing specialized vibratory behaviours to which the prey-spider responds as it normally would to its own prey. Prey (spiders and insects) is attacked by wrapping. Prey that trips over lines at the edge of a web of P. phalangioides , but fails to enter the web, is successfully attacked: P. phalangioides leans out of its web to throw silk over the prey, keeping as few as two legs on the silk. However, P. phalangioides does not attack prey that is completely away from webs. Occasionally, P. phalangioides feeds on eggs of other spiders and on ensnared insects it encounters in alien webs. Experimental evidence indicates that vision is of little or no importance in the predatory behaviour of P. phalangioides . Although P. phalangioides invades diverse types of webs, in addition to using its own web, its efficiency as a predator varies with web-type. It is most efficient as a predator of spiders and, especially, insects on its own web, and least efficient as a predator of amaurobiids on their cribellate sheet webs. Sensory, locomotory and other factors which influence differential predatory efficiency are discussed. The behaviour of P. phalangioides is compared to that of Portia , an araneophagic web-invading salticid, and the results of this study are discussed in relation to hypotheses concerning salticid evolution.     

The Balbiani body in the oocytes of a common cellar spider, Pholcus phalangioides (Araneae: Pholcidae)

Previtellogenic oocytes of a common cellar spider, Pholcus phalangioides, contain a single aggregation of organelles referred here to as the Balbiani body. It is a well defined ooplasmic structure predominantly composed of fine granular nuage, RNA rich material but comprising also mitochondria, vesicles of endoplasmic reticulum and stacks of Golgi cysternae. The Balbiani body originates early during previtellogenesis in the form of a cap-shaped mass in juxtaposition to one pole of the oocyte nucleus. During later stages of previtellogenic growth the Balbiani body translocates as a single body towards the ooplasm periphery. The results presented indicate that Balbiani body translocation is cytoskeleton independent. Balbiani body repositioning does not result in the localization of its components to any distinct, asymmetrically situated region of the ooplasm but, instead, ends up with their even dispersion in the oocyte cortex. The Balbiani body in Pholcus does not seem to be implicated either in germ cell determination or organelle inheritance. Its homology with similar organelle accumulations in the oocytes of other species is discussed.     

The genus Pholcus (Araneae, Pholcidae) in the Canary Islands

This paper offers a detailed taxonomic revision of all Canarian Pholcus species described before 2003, all of which are newly described and newly illustrated. Pholcus guadarfia sp. nov. is described, while a neotype for Pholcus malpaisensis Wunderlich, 1992 is also provided. In addition, we propose Pholcus gomerae Wunderlich, 1980 as a senior synonym for Pholcus gomeroides Wunderlich, 1987. More importantly, cladistic analysis based on the morphological characters of the Macaronesian Pholcus species was conducted for the first time. Parsimony analyses of 73 morphological characters revealed the close relationships between those species from the Canary Islands, Madeira and the Macaronesian enclave in Africa (between Agadir and Nouadhibou).  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 151 , 59–114.     

北京幽灵蛛的附肢自残

附肢自残是北京幽灵蛛（Pholcus beijingensis Zhu ＆ Song,1999）的习性之一。野外调查表明,约20％的个体至少失去1只步足;个体自残的发生率在2—7月间有显著的差异,同时第一对步足自残的比例最高（43．47％）;个体失去步足的比例随着年龄的增加而增加;左侧和右侧步足失去的比率接近（1．03：1）。野外调查发现,正在交配的个体与单独生活的个体相比,步足的完整性上并无显著差别。实验室研究表明,北京幽灵蛛附肢自残后没有再生现象。通过对自残幼体与非自残幼体的3龄与4龄龄期研究,发现各处理组间并无显著的差别。步足自残被认为是北京幽灵蛛在极端情况下的“避险策略”（Bet-hedging strategy）[动物学报54（6）：998-1004,2008]。     

Predator-prey interactions between jumping spiders (Araneae, Salticidae) and Phokus phalangioides (Araneae, Pholcidae)

Portia is a genus of specialized web-invading salticids that use aggressive mimicry. Some other salticids leap into webs to catch spiders but do not use aggressive mimicry. Pholcus phalangioides is a web-building spider with a special defensive behaviour—called whirling—in which it swings its body around in a circle while keeping its long legs on the silk. Pholcus phalangioides is preyed on by Portia and probably other salticid spiders in nature. Interactions between P. phalangioides and 13 species of salticids were studied in the laboratory to compare how effective salticids with different styles of predation were at catching the pholcids. Four species of Portia were studied and each was more efficient at catching P. phalangioides than were the other nine salticids tested. For one species—Portia fimbriata—individuals from three different populations were studied. The Queensland P. fimbriata used aggressive mimicry more consistently and were more efficient at catching P. phalangioides than were the other species of Portia and the other populations of P. fimbriata . The salticids that were the most efficient at catching pholcids were also better able to avoid setting off whirling by the pholcids. An experiment in which pholcids were artificially induced to whirl whenever the predator was near provided additional evidence that whirling is an effective defence of pholcids against predation by salticids.     

Sperm storage secretion of female cellar spiders (Pholcus phalangioides; Araneae): a gel-electrophoretic analysis

Glandular secretion in the genital cavity of female Pholcus phalangioides was removed and investigated gel-electrophoretically. Different staining techniques indicate that the secretion contains proteinaceous substances. Some protein fragments show a clear reaction to glyco- and lipoprotein staining procedures.
After copulation, female P. phalangioides keep the spermatozoa embedded in the secretion in the genital cavity. Since the spermatozoa are immobile, i.e. coiled and encapsulated, until shortly before oviposition, a nutritive function of the secretion during sperm storage is unlikely. The viscous quality of the material probably serves to retain the sperm mass in the sperm storage site. Lipoproteinaceous components are assumed to prevent desiccation of the sperm mass. Retaining the spermatozoa and maintaining them in a favourable environment are the most plausible functions of the secretion of female P. phalangioides. In spider species which possess receptacula seminis other functions can be attributed to sperm storage secretions.     

Copulatory Mechanism in Holocnemus pluchei and Pholcus opilionoides,With Notes on Male Cheliceral Apophyses and Stridulatory Organs in Pholcidae (Araneae)

Huber, B. A. 1994. Copulatory mechanism in Holocnemus pluchei and Pholcus opilionoides, with notes on male cheliceral apophyses and stridulatory organs in Pholcidae (Araneae).—Acta Zoologica (Stockholm) 76: 291–300. The pholcid spiders Holocnemus pluchei (Scopoli, 1763) and Pholcus opilionoides (Schrank, 1781) are investigated with respect to functional morphology of their genital organs using freeze-fixation of spiders during copula in liquid nitrogen and subsequent preparation of histological serial sections of the copulatory organs in functional contact. Special attention is paid to the mode of male pedipalpal arrestation before copulation, which is achieved in two quite different ways: in Pholcus by contact of the lateral cheliceral apophysis with the pedipalpal trochanter-apophysis, in Holocnemus by locking the pedipalpal trochanter between chelicera and pedipalpal coxa. The condition in Pholcus is considered to be apomorphic and to present a synapomorphy of about a dozen genera for which the name “Pholcus-group” is proposed. The stridulatory apparatus of Holocnemus pluchei is described, its biological significance discussed and an overview of accounts on stridulation in Pholcidae given.     

Colonization and diversification of the spider genus Pholcus Walckenaer, 1805 (Araneae, Pholcidae) in the Macaronesian archipelagos: evidence for long-term occupancy yet rapid recent speciation

Macaronesian archipelagos stand apart from other oceanic islands reputed as laboratories for the study of evolution by their proximity to the mainland, lack of subsidence, and steep ecological gradients. The genus Pholcus Walckenaer, 1805, commonly known as daddy-long-leg spiders, is one of the most speciose arthropod groups in the region, with 25 endemic species. In the present study, we use information from four mitochondrial genes, along with morphological data, to examine the phylogenetic relationships and diversification patterns of the genus in the region. Phylogenetic analyses support monophyly of Macaronesian Pholcus including the Moroccan species Pholcus vachoni and hence a single colonization event in the archipelagos. Madeira colonizers most likely originated from the Canaries, and a back-colonization of the nearby mainland receives further support. Estimated lineage divergence times suggest a long-time presence of Pholcus in the region, but also reveal that most present-day species are the result of recent, and probably rapid, speciation events. Diagnostic characters among Macaronesian Pholcus are confined to structures involved in copulation. Coupled with the extremely high diversification rate, the highest recorded for spiders, these copulatory characters suggest that sexual selection has played a key role in the local diversification of Pholcus in Macaronesia.     

Ultrastructure of the Male Accessory Glands and Sperm Ducts of Cylindrotaenia hickmani(Cestoda,Cyclophyllidea)

Abstract The ultrastructure of unicellular accessory glands (= prostate glands) and external male ducts of the cestode Cylindrotaenia hickmaniare described. Accessory glands open into the lumen of the external common sperm duct (= external vas deferens). The gland cells contain abundant endoplasmic reticulum, Golgi bodies and secretory bodies, and have elongate necks that pierce the apical cytoplasm of the duct. Cell contact with the apical cytoplasm of the sperm duct is mediated by septate desmosomes. Accessory glands secrete spherical particles, with a diameter of approximately 70 nm, that adhere to spermatozoa. The roles of these accessory glands may relate to activity of the sperm or development of the female system after insemination. Paired sperm ducts arise from testes, and unite to form a common sperm duct. Each duct consists of a tubular anucleate cytoplasmic region which is supported by nucleated cytons that lie sunken in the parenchyma. The apical cytoplasm of the paired sperm ducts (= vasa efferentia) possesses apical microvilli and abundant mitochondria, but few other cytoplasmic features. The apical cytoplasm of the common sperm duct possesses sparse apical microvilli and numerous electronlucent vesicles. The male gonoducts form an elongate syncytium which is markedly polarized along the length of the ducts. The ducts also display apical–basal polarity in that sunken nucleated cytons support the apical cytoplasm which in turn has distinct basal and apical domains.     

Spermatozoa and spermiogenesis of Holocnemus pluchei (Scopoli, 1763) (Pholcidae, Araneae)

Until now, the knowledge on pholcid spermatozoa is based on two species, Pholcus phalangioides and, incompletely, Holocnemus pluchei. To complete this knowledge and to reveal more potential phylogenetic characters, we have investigated sperm ultrastructure and spermiogenesis of H. pluchei. We found that the sperm cells of this species are clearly different from those of P. phalangioides with respect to: (1) the lack of specialization in the cylindrical acrosomal vacuole; (2) a nuclear canal which is located in the periphery and not in the center of the nucleus; (3) a more prominent postcentriolar elongation of the nucleus; (4) the presence of "inner microtubules" in the implantation fossa in early and mid-spermatids; (5) the absence of a helical band of nuclear material; (6) the proximal centriole which is not prolonged; (7) the types of secretion in the seminal fluid (only two types in H. pluchei). Similarities in the spermatozoa of both species concern: (1) a large implantation fossa which contains large amounts of glycogen in mature spermatozoa; (2) absence of a centriolar adjunct; (3) an axonemal basis located in the posterior part of the implantation fossa; (4) the formation of the so-called cleistospermia in the vas deferens. Our results strongly support systematic relationships within Pholcidae placing these two species in different subgroups.     

Sexual Selection on Accessory Glands, Genitalia and Protarsal Pads in the Whirligig Beetle Dineutus nigrior Roberts (Coleoptera: Gyrinidae)

Sexual selection is a potent force in the evolution of morphology in sexually reproducing species. When large size in a trait is favored by sexual selection the trait often exhibits positive allometry. Mating behavior in whirligig beetles consists of males attempting to grasp reluctant females using enlarged protarsi (protarsal pads). Here we use allometry and a mating experiment to investigate sexual selection pressures on accessory glands, intromittant genitalia (aedeagus), and protarsal pads in males of the whirligig beetle Dineutus nigrior Roberts. Accessory gland size exhibited positive allometry and males with larger accessory glands were more likely to copulate suggesting that larger size in this trait is favored by sexual selection. Males with larger accessory glands attempted to copulate more often but did not exhibit fewer failed mating attempts before copulating. This suggests that the increased probability of mating in males with large accessory glands is due to higher mating attempt frequency and not to increased ability to overcome female resistance. The length of the aedeagus exhibited negative allometry and males with a longer aedeagus did not have increased mating success. This is consistent with stabilizing selection favoring an intermediate size in this trait. The allometric slope of the protarsal pad did not differ from isometry and males with larger protarsal pads did not have increased mating success. This suggests that larger protarsal pads are not favored by sexual selection.     

Commensal Nematodes in the Glands,Genitalia, and Brood Cells of Bees (Apoidea)

Seven species of bees from the eastern United States, representing four families in the Apoidea, were dissected and examined for nematode associates. Dufour''s glands in females of Halictus ligatus, Augochlora pura mosieri, and Augochlorella gratiosa (Halictidae) from Florida were infested with dauer juveniles of Aduncospiculum halicti (Diplogasteridae). The Dufour''s glands of Colletes thoracicus (Colletidae) females from Maryland were infested with dauer juveniles of a new species of Koerneria sp. (Diplogasteridae), and abdominal glands of females of Andrena alleghaniensis (Andrenidae) from New York were infested with dauer juveniles of another new species of Koerneria. The lateral and median oviducts, Dufour''s glands, and poison sacs in females of Anthophora abrupta (Anthophoridae) from Maryland and Alabama were infested with dauer juveniles of a new species of Bursaphelenchus sp. (Aphelenchoididae). Cross sections of the nematode-infested poison sacs of A. abrupta revealed two types of humoral host defense reactions.