On the distinction between Modisimus and Hedypsilus (Araneae, Pholcidae), with notes on behavior and natural history

Evidence is presented that the genera Hedypsilus Simon, 1893 and Modisimus Simon, 1893 are subjective synonyms. Precedence is given to Modisimus over Hedypsilus. The type species of both genera (M. glaucus Simon, 1893 and H. culicinus Simon, 1893) are redescribed. Hedypsilus lawrencei Lessert, 1938 is synonymized with Hedypsilus culicinus Simon, 1893. It appears to be a pantropical, synanthropic species. Preliminary notes on its natural history are presented. Two closely related new species are described, Modisimus simoni sp. n. (Venezuela) and Modisimus david sp. n. (Central America). Modisimus globosus Schmidt, 1956 (Colombia?) is redescribed and illustrated for the first time. Notes on the usefulness of the curvature of eye rows and of leg formulae for the systematics of pholcids are given.     

Two new species of litter-dwelling Metagonia spiders (Araneae, Pholcidae) document both rapid and slow genital evolution

The observation that genitalia, rather than other characters, are useful in distinguishing species has resulted in the statement that genitalia evolve relatively rapidly. In this paper we claim that relatively rapid evolution of genitalia may often be restricted to the shapes, numbers, and sizes of individual structures. In contrast, the more basic pattern, including the presence or absence of structures and their interrelationships (the bauplan), does not seem to evolve more rapidly than other (nongenital) aspects of morphology. We document this idea by contrasting two litter‐dwelling pholcid species, Metagonia petropolis sp. n. and M. paranapiacaba sp. n., with their mostly leaf‐dwelling congeners. Significant differences occur with regard to nongenital aspects of morphology as well as shapes of individual genital structures, but the bauplan of their genitalia is basically the same.     

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.     

Anti-predator defences of Psilochorus sphaeroides and Smeringopus pallidus (Araneae, Pholcidae), tropical web-building spiders

Anti-predator defence behaviours of Psilochorus sphaeroides from Queensland and Smeringopus pallidus from Sri Lanka were studied in the laboratory. Whirling, in which these web-building pholcid spiders move their bodies rapidly around in circles, with legs remaining on the silk, is the most frequent response of adults to disturbance. Experiments were carried out to determine what types of stimuli elicited whirling. Forceful tactile and air movement stimuli elicited whirling, but there was no evidence that chemical stimuli from potential predators were important. Tactile stimulation was more effective than air movement. The behaviour of these pholcids was compared to that of a cosmopolitan pholcid, Pholcus phalangioides .     

Slow genital and genetic but rapid non-genital and ecological differentiation in a pair of spider species (Araneae,Pholcidae)

Like most animals with internal fertilization, spiders tend to have species-specific genitalia, allowing closely related species to be identified by their reproductive morphology more easily than by non-genital characters. This implies that genitalia evolve on average more rapidly than non-genital morphological traits. Here we describe two putative species of Pholcidae from Taiwan (Pholcus pingtung, n. sp.; Pholcus chengpoi, n. sp.) that differ conspicuously in their microhabitat (rocks vs. leaves), coloration, color pattern, and body proportions, but have almost indistinguishable genitalia and cytochrome oxidase I (co1) sequences. The two species have identical yet highly unusual male cheliceral modifications, strongly arguing for sister species. Despite the almost identical genitalia and co1 sequences, we treat the two ‘morphs’ as species for three reasons: (1) they are easily distinguished by several characters; no intermediate specimens were found; (2) subtle yet consistent differences in genital (uncus) shape support the idea of reproductively isolated entities beyond the more conspicuous non-genital differences; (3) each locality provided both types of microhabitat but only one of the two species, arguing against environmental plasticity or polymorphism. We conclude that probably a very recent expansion into a novel microhabitat has led to speciation and rapid ecological and non-genital differentiation, with insufficient time to accumulate significant genital and genetic differences.     

Speciation without changes in genital shape: A case study on Brazilian pholcid spiders (Araneae: Pholcidae)

Speciation in arthropods is usually coupled with marked changes in genital morphology, which explains the usefulness of genitalia in distinguishing closely related species. The present paper describes specimens that are assigned to separate species based on extreme size differences and colour pattern differences, but the shape of the genitalia is essentially identical. We argue that such cryptic species may be more common than currently assumed, but if marked morphological (non-genital) differences are missing, traditional taxonomic methodology is biased against discovering them. The two new species from the Brazilian Atlantic Forest are tentatively assigned to the genus Psilochorus: Psilochorus itaguyrussu n. sp. and Psilochorus ybytyriguara n. sp.     

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.     

Southern African pholcid spiders: revision and cladistic analysis of Quamtana gen. nov. and Spermophora Hentz (Araneae: Pholcidae), with notes on male−female covariation

The southern African pholcid spiders previously assigned to Pholcus and Spermophora are revised and their phylogenetic position within Pholcidae is analysed cladistically. Two distinct groups are recognized: the first is restricted to the Cape region and probably correctly placed in Spermophora ; it includes Spermophora peninsulae Lawrence and four new species. The second is more widely distributed in southern and eastern South Africa, and reaches as far north as Cameroon, Congo (Democratic Republic) and Uganda; it is assigned generic status ( Quamtana gen. nov .), and includes two species transferred from Pholcus [ Q. leptopholcica (Strand), Q. ciliata (Lawrence)] as well as 23 new species. A key to the pholcid spiders of South Africa is presented. A close correlation between a male character (distance between cheliceral apophyses) and a female character (distance between pockets on epigynum) is documented in Quamtana .  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 139 , 477−527.     

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.     

亚特巨幽蛛：1种泛热带物种在中国的新发现（蜘蛛目：幽灵蛛科）

本文重新描述了采自西沙群岛永兴岛的巨幽蛛属亚特巨幽蛛Artema atlanta Walckenaer,1837。描述标本保存在中国科学院动物研究所。亚特巨幽蛛Artema atlanta Walckenaer,1837（图1A-D,2A—D,3A-C）观察标本2♂,8♀,海南省西沙群岛永兴岛,2008-01—13—19,卜文俊 采集。     

Phylogeny of Habronattus jumping spiders (Araneae: Salticidae), with consideration of genital and courtship evolution

Abstract. DNA sequences from the mitochondrial (including ND1, 16S) and nuclear (EF‐1α) genomes of about ninety‐four species were obtained to reconstruct phylogenetic relationships of Habronattus jumping spiders. Maximum parsimony trees were sought with both separate (mitochondrial, nuclear) and combined analyses; maximum likelihood trees were sought with both separate (ND1, 16S, EF‐1α introns, EF‐1α exons) and combined (mitochondrial, nuclear) analyses. All analyses agreed on some fundamental aspects of the tree, including the monophyly of the previously recognized agilis, amicus, dorotheae and americanus species groups. The deep phylogenetic structure is well resolved, placing the agilis, amicus, tranquillus and dorotheae groups basally. Several other previously unrecognized clades were well supported, including a newly formulated decorus group. The large group of species with modified male first and third legs was supported as monophyletic except for the surprising placement elsewhere of three species of the group. The phenotypic similarities between these three and the others are so detailed and precise that convergence in ornamentation can probably be ruled out. There are hints of phylogenetically distant genetic introgression involving the coecatus group. The combination Habronattus paratus is restored based on the species falling within Habronattus. Regarding patterns of character evolution, there was consistent support for the basal placement of several species groups with a long embolus, suggesting that there were more evolutionary reductions in embolus length than postulated in a previous morphological phylogeny. This is in accord with the expectation that there is a bias to an overly conservative interpretation of a character's evolution if it is interpreted on a phylogeny based in part on that same character. In contrast, the molecular phylogeny did not suggest any instances of the evolutionary transformation of one complex style of courtship into another, a possibility that could have been difficult to detect using the morphological phylogeny because of the same bias to conservativism.     

Further notes on the natural history of the South American pepper frog, Leptodactylus labyrinthicus (Spix, 1824) (Anura, Leptodactylidae)

Leptodactylus labyrinthicus tadpoles reach a large size in the nest through consumption of trophic eggs. We previously suggested that the trophic eggs are laid just after amplexus has finished, but our new data do not support this hypothesis. We also present further details on the natural history of the species with regard to breeding activity, spawning site, retreats and the ability of tadpoles in preying upon fully-growth heterospecific tadpoles. We also show that the tadpoles are mainly nocturnal and take diurnal refuges. We collected the data in Brazil in three localities within the Cerrado Biome. We examined burrows used by L. labyrinthicus males, verified if females still contained mature eggs just after released from amplexus, and tested the ability of tadpoles in preying fully-growth heterospecific tadpoles. Field observations and experiments were conducted on tadpole activity time, hiding behaviour and level of susceptibility to predation by the bird leaf-scrapers in four sheltering situations. Reproduction could start before the first rains; this may be advantageous by allowing the tadpoles to exploit eggs of other frogs. We found one floating nest built in a temporary pool. The nest of the species is normally circumscribed in an excavated basin beside the water body. Adult males were found during the day with their head-out of the entrance of underwater burrows, which were perforations through dense root mats beside calling/spawning sites. Probably, these burrows in permanently water-filled soil are actively excavated by males. Females released all their eggs during the amplexus, so trophic eggs are not produced by the currently-accepted mechanism. Fully-grown heterospecific tadpoles were not preyed upon by L. labyrinthicus tadpoles, which can prey only slow-moving newly hatched ones. Field tadpoles took shelter under mud/dead leaves during daylight and became exposed on the bottom at night. Free-ranging leaf-scrapers removed dead leaves from a pool with their beaks and preyed upon tadpoles. In the experiments, the tadpoles sheltered under gravel/leaves during daylight, but they were exposed at night. Leaf-scrapers ate all exposed tadpoles, but no tadpole of the gravel/leaves trays was consumed. Hence the nocturnal habits and use of diurnal refuges may protect the tadpoles from visual predators, such as the leaf-scrapers.     

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.     

Predator-prey interactions between web-invading jumping spiders and a web-building spider, Holocnemus pluchei (Araneae, Pholcidae)

Three species of web-invading salticid spiders, with different predatory strategies, were tested with Holocnemus pluchei in the laboratory: Portia fimbriata and Portia labiata , which practise aggressive mimicry, and Euyattus sp., which leaps from outside on to spiders in webs but does not practise aggressive mimicry. Portia was shown to be more efficient than Euryattus at catching H. pluchei. Portia fimbriata from Queensland was more consistent at using aggressive mimicry than was Portia labiata and was also more efficient at catching H. pluchei . The web-invaders that were more efficient at catching H. pluchei were also better able to avoid setting off bouncing, a special defence behaviour used by H. pluchei . An experiment, in which H. pluchei was artificially induced to bounce whenever the predator was near, provided additional evidence that bouncing is effective in defending H. pluchei against web-invaders.     

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.     

Cobweb management and control of the spider Holocnemus pluchei (Araneae: Pholcidae) on buildings

The spider Holocnemus pluchei (Scopoli) (Araneae: Pholcidae) is a nonnative species commonly found in the southwestern United States. In urban areas around homes and other structures, it is often the dominant spider species on outside surfaces; requests for control of spiders and their cobwebs are common for the commercial pest management industry. We tested two physical (brushing and vacuuming) and two low-impact chemical (permethrin and botanical insecticide) spray control methods for H. pluchei on outbuildings on the University of California-Riverside campus. Forty sites containing at least two H. pluchei spiders along a 5-m section of eave or overhang were chosen for treatment. Treatment was applied within 2 d of a pretreatment census. The number of spiders per section was counted at 2, 4, 8, 12, 16, and 52 wk posttreatment. Untreated sections showed an increase in spiders through summer and a decrease in autumn. Sites treated with permethrin significantly reduced spiders >95% and had strong residual effect throughout the remainder of the sampling period. Compared with untreated controls, the three other treatments did not significantly reduce spiders; these three were similar throughout the season and were intermediate between untreated and permethrin-treated sites. In comparison with the before and 2-wk posttreatments, spiders at the untreated sites showed no difference in body sizes, whereas those in the nonpermethrin treatments showed a decrease, indicating that larger spiders were eliminated. We also tested H. pluchei spiders in the laboratory against the two insecticides used in the field studies to determine the minimum lethal concentrations for controlling H. pluchei.