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Description of Sarax buxtoni (Gravely, 1915) (Arachnida: Amblypygi: Charinidae) and a new case of parthenogenesis in Amblypygi from Singapore

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Abstract

The type material of Sarax buxtoni (Gravely 1915) cannot be located and has to be considered as lost. Therefore, a description compiled from a population in Singapore is provided, including morphological and taxonomical details presented for the first time. Comparisons with closely related species are supplied. Furthermore, we describe the occurrence of parthenogenesis in a population of S. buxtoni, representing the first case of asexual reproduction in a member of the genus Sarax Simon 1892.

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... The largest amblypygid family, Charinidae, with approximately 95 species, is the only one to produce offspring from unfertilised eggs. So far, two species of Charinus Simon, 1892 (Armas 2005;Weygoldt 2005Weygoldt , 2007 and one of Sarax Simon, 1892 (Seiter and Wolff 2014) are known to reproduce parthenogenetically (which is 3% of the total number of species of the family). ...
... In Charinus ioanniticus Kristscher, 1959, from the Eastern Mediterranean, female-only populations are known from the islands of Rhodes and Kos (Greece) and it has been shown it can reproduce parthenogenetically in captivity (Weygoldt 2005(Weygoldt , 2007. Nonetheless, males of C. ioanniticus have been recorded in populations from Turkey (Kovařík and Vlasta 1996;Weygoldt 2005;Seyyar and Demir 2007), Israel (Rosin andShulov 1960), andEgypt (El-Hennawy 2002), showing that this species might be facultatively parthenogenetic (Weygoldt 2005;Seiter and Wolff 2014). Other populations of C. ioanniticus, such as those in continental Greece (Athens) (Agapakis and Miranda 2019), Italy (Colla et al. 2020), and Jordan (Shakhatreh et al. 2020), still require more studies to verify the exact type of reproduction, however, all known specimens are females. ...
... Other populations of C. ioanniticus, such as those in continental Greece (Athens) (Agapakis and Miranda 2019), Italy (Colla et al. 2020), and Jordan (Shakhatreh et al. 2020), still require more studies to verify the exact type of reproduction, however, all known specimens are females. The only Sarax species known to reproduce parthenogenetically is S. buxtoni Gravely 1915 from Singapore, whose evidence for parthenogenesis came from captivity experiments (Seiter and Wolff 2014). In addition to these species, many other charinids are known from females only (Giupponi and Kury 2002;Miranda and Giupponi 2011;Jocque and Giupponi 2012;Giupponi and Miranda, 2016;Ferreira 2016, 2017;Miranda et al. 2016aMiranda et al. , 2016b, which indicates that likely more species in the family are parthenogenetic. ...
Article
Parthenogenesis is documented in a few species of Amblypygi, but it is unknown how widespread in the order this reproductive behaviour is, and little has been researched regarding aspects of embryonic and post-embryonic development in the group. Here, we studied the parthenogenetic capacity of an Amazonian whip spider (Charinus guto) evaluating the time of egg and embryonic development and inter-moult period. We also provide a review on embryonic and post-embryonic development in Amblypygi, compiling and analysing data from 43 species in three families. Fifty-two females and 42 juveniles of C. guto were collected in fragments of a secondary forest in Belém (Brazil); specimens were kept in captivity and observed weekly from 2018 to 2020. Nineteen specimens were collected with and 32 without egg sacs. Fourteen of the nonovigerous females developed eggs in captivity, six of them moulted (i.e. lost stored sperm from previous contacts with males) before developing an egg sac, proving to be parthenogenetic. The mean time between the first day in captivity and moult was 96 days. In both adults and juveniles, a mean of 147 days passed between first day in captivity and first moult, and 125 days for a second moult. After moulting, a mean of 113 days passed for the females to develop an egg sac; the embryonic development took a mean of 59 days. Juveniles left the mother’s abdomen after 10 days of hatching from the egg and the mean number of live free-living protonymphs was five. Other amblypygids, especially charinids, have similar embryonic development and post-embryonic growth and a detailed discussion with all known information for whip spiders is presented. We also demonstrate a positive correlation between clutch size and female size across Amblypygi, in which larger females carry more eggs and have larger offspring regardless of climate and habitat.
... Whip spiders (Amblypygi) belong to the class Arachnida and are distributed globally. They are a very small group of arachnids with 190 currently known species (Harvey 2013;Armas et al. 2014;Seiter & Wolff 2014). They are nocturnal and cavernicolus predators, which are catching their prey with their raptorial pedipalps armed with numerous spines. ...
... A doubtful record from Greece is still listed in Harvey (2003Harvey ( , 2013: Sarax mediterraneus Delle Cave, 1986. However, this has been discussed as a case of confusion in the recent literature (Weygoldt 2005;Seiter & Wolff 2014). On the Philippines, only a few records of species of the genus Sarax are known. ...
... Parthenogenesis in Amblypygi is reported exclusively from one family, i.e. Charinidae: Charinus acosta (Quintero 1893), C. ioanniticus (Kritscher 1959) and S. buxtoni (Gravely 1915) (Armas 2000(Armas , 2005Weygoldt 2005Weygoldt , 2007Seiter & Wolff 2014). ...
Article
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A new species of the whip spider genus Sarax Simon, 1892 from Cebu Island in the Philippines is described: Sarax huberi sp. nov. With the description of this species, the diversity of the genus is increased to three species in the Philippines. Some additional data on their natural environment and their specific habitat are presented and compared with sibling species. The synonymization of Sarax mediterraneus Delle Cave, 1986 with Sarax buxtoni (Gravely, 1915) is carried out.
... Whip spiders belong to the order Amblypygi with its 220 extant species inhabiting mostly tropical and subtropical regions (Weygoldt 2000, Harvey 2013. Charinidae is the most diverse family and is the only one that has parthenogenetic species (e.g., Armas 2000, 2005, Weygoldt 2007, Seiter & Wolff 2014. ...
Article
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Whip spiders belong to the order Amblypygi with its 220 extant species inhabiting mostly tropical and subtropical regions (Weygoldt 2000, Harvey 2013, Miranda et al. 2018). Charinidae is the most diverse family and is the only one that has parthenogenetic species (e.g., Armas 2000, 2005, Weygoldt 2007, Seiter & Wolff 2014). Charinus ioanniticus (Kritscher, 1959) is one of the amblypygid species with all-female clonal reproduction, although a few males are known from collections (Weygoldt 2005, 2007, Miranda et al. 2016). The species is known from the Eastern Mediterranean including Egypt (Rosin & Shulov 1960, Kovařík & Vlasta 1996, El-Hennawy 2002, 2019, Miranda et al. 2016) and recently was recorded from Athens, Greece (Agapakis & Miranda 2019). Charinus ioanniticus was originally described from the Greek islands of Rhodes and Kos (Kritscher 1959). Further findings in different countries such as Turkey, Israel and Egypt make C. ioanniticus one of the species with the widest distribution in the family. Blick & Seiter (2016) summarised the distribution of the species and speculated on its occurrence in Jordan as well as in Lebanon and Syria. In this study, the order Amblypygi and Charinus ioanniticus are recorded for the first time from Jordan.
... Charinidae went through various taxonomical changes in recent years (summarized by ) and currently includes only the three above cited genera (Catageus Charinus and Sarax). Several papers described and redescribed species within these genera (Armas, 2006;Delle Cave et al., 2009;Giupponi and Miranda, 2012;Miranda et al., 2016a;Miranda and Giupponi, 2011;Miranda et al., 2016b;Miranda et al., 2016c;Seiter and Wolff, 2014;Seiter et al., 2015;Torres-Contreras et al., 2015;Vasconcelos and Ferreira, 2017;Vasconcelos et al., 2013Vasconcelos et al., , 2014Weygoldt, 2006), but none presented a thorough revision of species and genera or phylogenetic evidence for its monophyly. ...
Article
Sexual dimorphism is a common feature of life. Researchers have noted that some species of Amblypygi (Class Arachnida) exhibit male-biased sexual dimorphism, but a broad overview of the prevalence of sexual dimorphism in the order does not exist. In order to provide such an overview, we imaged and measured morphological characteristics of 390 amblypygids from 36 species. Our aim was to gain insight into general patterns of sexual dimorphism across the order. We found that, generally, males have larger pedipalps, longer antenniform legs, but smaller carapace widths relative to conspecific females for nearly all species studied. Species ranged from no sexual dimorphism (Phrynus exsul and Charinus insularis) to male pedipalps almost double the length of females in some species (Charon grayi, Euphrynichus amanica). Phylogenetic patterns remain unclear in Amblypygi and our data better match geographic patterns than evolutionary history. We thus suggest that sexual dimorphism is ancestral for the order, but has been reduced or lost in some species.
... Charinidae went through various taxonomical changes in recent years (summarized by ) and currently includes only the three above cited genera (Catageus Charinus and Sarax). Several papers described and redescribed species within these genera (Armas, 2006;Delle Cave et al., 2009;Giupponi and Miranda, 2012;Miranda et al., 2016a;Miranda and Giupponi, 2011;Miranda et al., 2016b;Miranda et al., 2016c;Seiter and Wolff, 2014;Seiter et al., 2015;Torres-Contreras et al., 2015;Vasconcelos and Ferreira, 2017;Vasconcelos et al., 2013Vasconcelos et al., , 2014Weygoldt, 2006), but none presented a thorough revision of species and genera or phylogenetic evidence for its monophyly. ...
Article
The modestly diverse order Amblypygi Thorell, 1883 includes five families, of which Charinidae Quintero, 1986 is the most diverse and with the widest geographical distribution. The family currently comprises three genera, Catageus Thorell, 1889, Charinus Simon, 1892 and Sarax Simon, 1892, the first known by one species from a single locality in Myanmar, the second with currently 74 species globally distributed, and the last with 17 species present in Southeast Asia and India. In this paper we describe and illustrate a new genus to accommodate the species Sarax davidovi Fage, 1946 based on unique characters. Weygoldtia gen. nov. (Laos, Vietnam and Cambodia) is supported by two synapomorphies: the presence of a straight crest anterior to the lateral eyes and the longitudinal orientation of the rod sensilla on tarsus of leg I. The new genus can be distinguished from Charinus and Sarax by the number of trichobothria on distitibia IV and the presence of one or two setae on the base of the cleaning organ on pedipalp tarsus. The enigmatic species Catageus pusillus Thorell, 1889 (the single species in the genus) is here synonymized with Stygophrynus cavernicola (Thorell, 1889) (family Charontidae Simon, 1892) and a neotype is designated. As Stygophrynus cavernicola is the type species of the genus, the synonymization of the two species results in the synonymy of the genera. Following the principle of priority, Catageus is maintained and all eight species of Stygophrynus Kraepelin, 1895 now have the following new combination: Catageus berkeleyi (Gravely, 1915), comb. nov., C. brevispina (Weygoldt, 2002), comb. nov., C. cavernicola, comb.nov., C. cerberus (Simon, 1901), comb. nov., C. dammermani (Roewer, 1928), comb. nov., C. longispina (Gravely, 1915), comb. nov., C. moultoni (Gravely, 1915), comb. nov., C. orientalis (Seiter and Wolf, 2017), comb. nov. and C. sunda (Rahmadi and Harvey, 2008), comb. nov.
... Parthenogenesis is well known among arachnids, including mites, harvestmen, true spiders, scorpions, whip spiders and possibly in some species of pseudoscorpions, schizomids and palpigrades (Christian and Christophoryová 2013;Dashdamirov and Golovatch 2005;Seiter and Wolff 2014;Tsurusaki 1986;Edwards et al., 2003;Zawierucha et al., 2013). In scorpions, parthenogenesis was previously recorded for seven buthid and one hormurid scorpions (review in Seiter et al., 2016). ...
... All other parthenogenetic species in Amblypygi belong to the family Charinidae, i.e. Charinus acosta (Quintero, 1983) from Cuba (Armas 2000), C. ioanniticus and Sarax buxtoni from Singapore (Seiter & Wolff 2014). This special reproduction positively results in establishing new populations and growth rates, but has negative effects, because all individuals are clones and genetically identical. ...
Article
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All records of the two amblypygid species occurring in the Western Palaearctic are mapped and both species (Charinus ioanniticus and Musicodamon atlanteus) are discussed. Charinus ioanniticus is known from the Eastern Mediterranean (Greece, Turkey, Israel and Egypt) from 10 localities and Musicodamon atlanteus is known from the Maghreb (Morocco, Algeria) from three localities. All records are mapped.
... to be the case with other arachnid groups (Eberhard 1985;Huber 2002 Weygoldt 2005Weygoldt , 2007Seiter & Wolff 2014). Given the relatively small number of observations on species-specific reproductive behavior, we lack a comprehensive overview of amblypygid mating system diversity and thus have an incomplete understanding of the selection pressures that might lead to variable mating systems. ...
Thesis
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Behavioral researchers using Amblypygi have noted the regularity at which species engage in agonistic interactions. Despite this, why agonistic interactions occur and how they are resolved is unknown. I conducted paired interactions of the amblypygid Phrynus longipes in Puerto Rico to understand the dynamics of agonistic interactions. Through a series of analyses, I found that agonistic interactions are territory contests common across the demographic range of the species. Further, I decoded the strategy that opponents use to negotiate contests, and used resource contests to explain the peculiar pattern of cannibalism that this species exhibits. I used these results to identify the variation in contests and other behavioral phenotypes across cave and surface populations. This research broadens theory of resource contest evolution and behavioral variation by investigating phenomena in a non-model study system.
... 7. charinidae Quintero, 1986, type genus Charinus Simon, 1892, is the most diverse and largest family in the order Amblypygi (Arachnida), and this family-group name has been used in at least 25 works, published by at least 10 authors over a span of not less than 10 years (Article 3.9.1.2) (e.g. Kovarik, & Vlasta, 1996;Weygoldt, 2000Weygoldt, , 2002Weygoldt, , 2005Weygoldt, , 2006Weygoldt, , 2008Weygoldt, , 2009Weygoldt, , 2010Baptista & Giupponi, 2002;2003;Giupponi & Kury, 2002;El-Hennawy, 2002;Giupponi & Baptista, 2003;Harvey, 2003;Armas, 2005Armas, , 2006Armas, , 2010Armas et al., 2009Armas et al., , 2012Teruel, et al., 2009;Rahmadi, & Kojima, 2010;Rahmadi et al., 2010;Villareal, 2010;Miranda & Giupponi, 2011;Prendini, 2011;Giupponi & Miranda, 2012;Jocqué & Giupponi, 2012;Seiter, & Hörweg, 2013;Seiter & Wolff, 2014;Vasconcelos et al., 2014). ...
Article
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The purpose of this application, under Article 23.9.3 of the Code, is to suppress the little-used senior homonym charinidae Gray, 1849 in order to conserve the family-group name charinidae Quintero, 1986. charinidae Quintero, 1986 is a well-established name for the most speciose group of Amblypygi, while charinidae (originally charinini) Gray, 1849 has been little used since its original proposal; to resurrect this name would be disruptive to both arachnid and snake taxonomy. The suppression of charinidae Gray 1849 will help to conserve its widely used junior synonym ungaliopheinae McDowell, 1987. The name charinidae Cope, 1886 is declared nomen oblitum under Article 23.9.2 of the Code. The family-group name charinusidae Pyron, Reynolds & Burbrink, 2014 (Arachnida) proposed to replace the family-group name charinidae Quintero, 1986 is unavailable and unnecessary.
... longicornis; Weygoldt 1977a), however, mate guarding might be a better explanation for male presence. Lastly, parthenogenesis likely occurs in at least four species of Amblypygi, as evidenced by female-only populations or captive reproduction by virgin females (Armas 2000(Armas , 2005Weygoldt 2005Weygoldt , 2007Seiter & Wolff 2014). Given the relatively small number of observations on species-specific reproductive behavior, we lack a comprehensive overview of amblypygid mating system diversity and thus have an incomplete understanding of the selection pressures that might lead to variable mating systems. ...
Article
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Arachnologists have uncovered tantalizing details about amblypygid behavioral ecology—the study of the fitness consequences of their behavior. Thus, it is the aim of this review to position Amblypygi as a useful system in which to investigate the principles of animal behavioral ecology. We synthesize amblypygid habitat preference and navigation modalities; predator, prey, parasite, parasitoid, cannibal, and commensal interactions; resource contests and territoriality; mating systems and mate choice; parental investment and sociality; and genetics and genomics as they relate to behavioral ecology. We present ideas for future research in each of these areas and discuss future directions for Amblypygi behavioral ecology research as they relate to four areas of behavioral ecology: adaptation, evolutionary history, mechano-sensory control of behavior, and behavioral development. We conclude by identifying several avenues of Amblypygi behavioral ecology that we think have the highest potential for transformative discoveries.
... With the exception of mites in which various types of parthenogenetic reproduction are known to occur and parthenogenesis is prevalent in certain families (Oliver 1971;Sabelis 1985), asexual reproduction is quite rare in other arachnid orders (Bell 1982). Parthenogenesis is known to occur in several species of spiders (Edwards et al. 2003), harvestmen (Tsurusaki 1986), amblypygids (Armas 2000;Weygoldt 2005Weygoldt , 2007Seiter & Wolff 2014) and scorpions (review in Francke 2008;Lourenço 2008), and is strongly suspected to occur in some species of pseudoscorpions (Dashdamirov & Golovatch 2005), schizomids (Nedvěd et al. 2011;Zawierucha et al. 2013) and palpigrades (Christian & Christophoryová 2013) based on the observation of all-female populations or a pronounced scarcity of males. Although the majority of claims about parthenogenetic reproduction in scorpion species were proven by experimental evidence, some are based on field observations or uneven sex distributions in samples of collected species alone (Francke 2008;Lourenço 2008). ...
Article
Of all scorpion species described to date, only a small fraction are known to reproduce without fertilization by a male, instead producing offspring by parthenogenesis. Here we show that isolated females of the buthid Pseudolychas ochraceus (Hirst, 1911) are capable of parthenogenetic reproduction and we provide data on the postembryonic growth of this species.
... Its number of species exceeds only Opilioacarida, Holothyrida, Ricinulei, Palpigradi and Thelyphonida (Harvey 2003;Zhang 2013). Seiter et al. (2015) counted current 190 species of Amblypygi based on Harvey (2013), Armas et al. (2014) and Seiter & Wolff (2014). However, including two species of the genus Charinus from Brazil that were not added to the count (Vasconcelos et al. 2013(Vasconcelos et al. , 2014, plus the recent described species Charinus sillami Réveillion & Maquart 2015 from French Guiana (Réveillion & Maquart 2015), a total of 193 species have been recognized globally in the order. ...
Article
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The genus Charinus comprises eleven described species in Brazil. Herein we describe two new species, Charinus caatingae sp n. and Charinus iuiu sp n., from caves of the state of Bahia, Brazil. Charinus caatingae is threatened, requiring special attention to its conservation. Furthermore, we present an updated identification key and a table of characters for the genus in the country.
Article
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The whip spider family Charinidae Quintero, 1986 is the most speciose and widely distributed in the arachnid order Amblypygi Thorell, 1883. It comprises three genera and 95 species distributed across all tropical continents and the eastern Mediterranean. Despite recent advances in the taxonomy of the family, a global revision of all its species, necessary to advance understanding of its systematics, biogeography and evolution, has never been conducted. In the present contribution, the family is revised in its entirety for the first time, including all previous names and 33 new species, 24 in the genus Charinus Simon, 1892: C. alagoanus sp. nov., C. apiaca sp. nov., C. carinae sp. nov., C. carioca sp. nov., C. carvalhoi sp. nov., C. cearensis sp. nov., C. diamantinus sp. nov., C. euclidesi sp. nov., C. goitaca sp. nov., C. guayaquil sp. nov., C. imperialis sp. nov., C. loko sp. nov., C. magalhaesi sp. nov., C. miskito sp. nov., C. mocoa sp. nov., C. monasticus sp. nov., C. palikur sp. nov., C. perquerens sp. nov., C. puri sp. nov., C. renneri sp. nov., C. sooretama sp. nov., C. souzai sp. nov., C. susuwa sp. nov., C. una sp. nov.; eight in the genus Sarax Simon, 1892: S. bilua sp. nov., S. dunni sp. nov., S. gravelyi sp. nov., S. indochinensis sp. nov., S. lembeh sp. nov., S. palau sp. nov., S. rahmadii sp. nov., S. tiomanensis sp. nov.; and one in the genus Weygoldtia Miranda et al., 2018: W. consonensis sp. nov. Taxonomic keys to the 132 species (excluding four nomina dubia) are presented and several taxonomic rearrangements implemented. Four subspecies are elevated to species level: Charinus cavernicolus Weygoldt, 2006, C. elegans Weygoldt, 2006, C. longipes Weygoldt, 2006, and Sarax bispinosus (Nair, 1934). Sarax batuensis Roewer, 1962 is removed from synonymy with Sarax buxtoni (Gravely, 1915) and S. buxtoni newly synonymized with Sarax rimosus (Simon, 1901). Stygophrynus moultoni Gravely, 1915 is transferred to Sarax, resulting in Sarax moultoni (Gravely, 1915) comb. nov. Ten species are transferred from Charinus to Sarax, resulting in new combinations: S. abbatei (Delle Cave, 1986) comb. nov., S. bengalensis (Gravely, 1911) comb. nov., S. dhofarensis (Weygoldt, Pohl & Polak, 2002) comb. nov., S. ioanniticus (Kritscher, 1959) comb. nov., S. israelensis (Miranda et al., 2016) comb. nov., S. omanensis (Delle Cave, Gardner & Weygoldt, 2009) comb. nov., S. pakistanus (Weygoldt, 2005) comb. nov., S. seychellarum (Kraepelin, 1898) comb. nov., S. socotranus (Weygoldt, Pohl & Polak, 2002) comb. nov. and S. stygochthobius (Weygoldt & Van Damme, 2004) comb. nov.
Article
Full-text available
The whip spider family Charinidae Quintero, 1986 is the most speciose and widely distributed in the arachnid order Amblypygi Thorell, 1883. It comprises three genera and 95 species distributed across all tropical continents and the eastern Mediterranean. Despite recent advances in the taxonomy of the family, a global revision of all its species, necessary to advance understanding of its systematics, biogeography and evolution, has never been conducted. In the present contribution, the family is revised in its entirety for the first time, including all previous names and 33 new species, 24 in the genus Charinus Simon, 1892: C. alagoanus sp. nov., C. apiaca sp. nov., C. carinae sp. nov., C. carioca sp. nov., C. carvalhoi sp. nov., C. cearensis sp. nov., C. diamantinus sp. nov., C. euclidesi sp. nov., C. goitaca sp. nov., C. guayaquil sp. nov., C. imperialis sp. nov., C. loko sp. nov., C. magalhaesi sp. nov., C. miskito sp. nov., C. mocoa sp. nov., C. monasticus sp. nov., C. palikur sp. nov., C. perquerens sp. nov., C. puri sp. nov., C. renneri sp. nov., C. sooretama sp. nov., C. souzai sp. nov., C. susuwa sp. nov., C. una sp. nov.; eight in the genus Sarax Simon, 1892: S. bilua sp. nov., S. dunni sp. nov., S. gravelyi sp. nov., S. indochinensis sp. nov., S. lembeh sp. nov., S. palau sp. nov., S. rahmadii sp. nov., S. tiomanensis sp. nov.; and one in the genus Weygoldtia Miranda et al., 2018: W. consonensis sp. nov. Taxonomic keys to the 132 species (excluding four nomina dubia) are presented and several taxonomic rearrangements implemented. Four subspecies are elevated to species level: Charinus cavernicolus Weygoldt, 2006, C. elegans Weygoldt, 2006, C. longipes Weygoldt, 2006, and Sarax bispinosus (Nair, 1934). Sarax batuensis Roewer, 1962 is removed from synonymy with Sarax buxtoni (Gravely, 1915) and S. buxtoni newly synonymized with Sarax rimosus (Simon, 1901). Stygophrynus moultoni Gravely, 1915 is transferred to Sarax, resulting in Sarax moultoni (Gravely, 1915) comb. nov. Ten species are transferred from Charinus to Sarax, resulting in new combinations: S. abbatei (Delle Cave, 1986) comb. nov., S. bengalensis (Gravely, 1911) comb. nov., S. dhofarensis (Weygoldt, Pohl & Polak, 2002) comb. nov., S. ioanniticus (Kritscher, 1959) comb. nov., S. israelensis (Miranda et al., 2016) comb. nov., S. omanensis (Delle Cave, Gardner & Weygoldt, 2009) comb. nov., S. pakistanus (Weygoldt, 2005) comb. nov., S. seychellarum (Kraepelin, 1898) comb. nov., S. socotranus (Weygoldt, Pohl & Polak, 2002) comb. nov. and S. stygochthobius (Weygoldt & Van Damme, 2004) comb. nov.
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The present contribution addresses the phylogeny and biogeography of the pantropical whip spider family Charinidae Quintero, 1986, the most species-rich in the arachnid order Amblypygi Thorell, 1883, based on morphology and multilocus DNA sequences, analysed simultaneously using parsimony, maximum likelihood and Bayesian inference. The morphological matrix comprises 138 characters, scored for four outgroup taxa and 103 ingroup terminals representing all genera and 64% of the species of Charinidae. The multilocus dataset comprises sequences from two nuclear and three mitochondrial gene loci for four outgroup taxa and 48 ingroup representing 30 (23%) taxa of Charinidae. Charinidae are monophyletic, with Weygoldtia Miranda et al., 2018 sister to a monophyletic group comprising Charinus Simon, 1892 and Sarax Simon, 1892, neither of which are reciprocally monophyletic. Charinidae diverged from other amblypygid families in the Late Carboniferous, c. 318 Mya, on the supercontinent Pangaea. Weygoldtia diverged from the common ancestor of Charinus and Sarax during the Late Permian, c. 257 Mya, when changes in climate reduced tropical forests. The divergence of Charinus and Sarax coincides with the fragmentation of Pangaea, c. 216 Mya. Sarax colonized South-East Asia via Australia. The charinid fauna of New Caledonia originated before the Oligocene, when the island separated from Australia, c. 80 Mya.
Chapter
This chapter deals with the diversity of terrestrial cave invertebrates from the viewpoint of its contribution to global diversity and of the variety of morphological traits and life forms associated with cave habitats. We stress the low diversity of cave compared to non-cave fauna, its uneven taxonomic and geographic distribution, differences in its composition between temperate and tropical bioclimates, and its high originality, due to its richness in endemics, in large radiations and in relictual species. The main morphological traits linked to subterranean life as well as their relevance as markers of troglomorphy are discussed. Life forms classically used for describing morphological features of cave organisms are redefined in a larger context. The diversity patterns of the different subterranean groups are summarized in turn in the same perspective, with a focus on the magnitude of their taxonomic richness, of recent progress in their knowledge, and on life forms.
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Resumen: Se actualiza la distribución geográfica conocida del amblipigio Charinus acosta (Quintero, 1983) en el archipiélago cubano. Se registran siete nuevas localidades que incluyen sus primeras citas para las provincias de Pinar del Río, Matanzas y Granma, así como ampliaciones considerables para las de Artemisa y Guantánamo. Se presenta además un amplio complemento fotográfico que ilustra los individuos y las condiciones ecológicas en que éstos habitan. Abstract: The known geographical distribution of the whip spider Charinus acosta (Quintero, 1983) in the Cuban archipelago is updated. Seven new localities are recorded, which include its first findings in the provinces of Pinar del Río, Matanzas and Granma, as well as remarkable extensions for those of Artemisa and Guantánamo. A thorough photographic complement is presented, in order to show the individuals and ecological conditions where they live in those populations.
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Resumen Se revisan los amblipígidos de República Dominicana (La Española, Antillas Mayores). Se registra, por primera vez, la familia Charinidae y se describen tres especies nuevas del género Phrynus Lamarck, 1801 (Phrynidae) y una de Charinus Simon, 1891 (Charinidae). Otra especie de este último género, muy parecida a Charinus cubensis (Quintero, 1983), no pudo ser identificada debido a la falta de adultos. Sólo una de las especies nuevas de Phrynus es endémica de la Española, pues una de ellas también habita en el extremo oriental de Cuba y otra en la Isla de Mona, Puerto Rico. Se registran nuevas localidades para Phrynus longipes (Pocock, 1893) y P. marginemacula-tus C. L. Koch,1840, se aportan datos sobre la historia natural y la biogeografía de las especies involucradas y se ofrece una clave para separar los taxones dominicanos. También se describen algunas anomalías de los apéndices. Abstract The whip spiders of the Dominican Republic (Hispaniola, Greater Antilles) are revised. The family Charinidae is recorded for the first time. Three new species of the genus Phrynus Lamarck, 1801 (Phrynidae) and a new species of Charinus Simon, 1891 (Charinidae) are described. Another species of Charinus, very close to C. cubensis (Quintero, 1983), remains unidentified because only young specimens were available. Only one of the three new Phrynus species is a Hispaniolan endemism, because one of the others inhabits the easternmost part of Cuba, and another one has been found in Mona Island, Puerto Rico. New localities are recorded for Phrynus longipes (Pocock, 1893) and P. marginemaculatus C. L. Koch, 1840 and data on natural history and biogeography of all these species are given. A key for the Dominican amblypygids is included, too. Also, some appendicular anomalies are described. Taxonomy: Phrynus eucharis sp. nov. Phrynus hispaniolae sp. nov. Phrynus kennidae sp. nov. Charinus dominicanus sp. nov. Introducción Los amblipígidos constituyen un pequeño grupo de arácnidos de hábitos nocturnos y de amplia distribución en los países tropicales y subtropicales del mundo. Durante el día permanecen refugiados en sitios sombríos y húmedos, por lo general debajo de piedras, troncos caídos y objetos similares, aunque también se introducen en algunas plantas epífitas (Bromeliaceae) y debajo de la corteza semidesprendida de los árboles. Varias especies son habitantes facultativos (troglófilos) u obligatorios (troglobios) de las cuevas o grutas. Aunque el observador lego a veces las toma por arañas, sus diferencias con éstas son notables, pues entre otras características, no poseen órganos productores de seda ni de veneno. Su parecido, sin embargo, no es simple coincidencia, pues ambos órdenes se hallan evolutivamente muy emparentados (Weygoldt, 1980; Shultz, 1990; Wheeler & Hayashi, 1998). En la actualidad, la posición filogenética de los amblipígidos se debate entre dos hipótesis: (1) Amblypygi como grupo hermano de Araneae y (2) Amblypygi como grupo hermano de Uropygi (Uropygi = Thelyphonida + Schizomida). Ambas presentan evidencias que las soportan (Weygoldt, 2000), por lo que futuros estudios deberán aportar nuevas evidencias que permitan discriminar la hipótesis correcta.
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Resumen: Se determinó que Charinus acosta (Quintero, 1983) ovoposita entre marzo y agosto. En los meses más cálidos (junio-agosto), la formación del saco ovígero ocurrió 35-88 días con posterioridad a la ecdisis o a la ovoposición anterior. Pero si la ecdisis ocurría a finales del verano (septiembre u octubre), entonces el saco ovígero se formaba 178-187 días después. La cantidad de recién nacidos en el laboratorio, varió entre tres y nueve por camada (promedio = 4,2), en relación directa con el tamaño de la madre. La duración (en días) de los primeros estadios fue el siguiente: preninfa = 7; protoninfa = 60-131; deu-toninfa = 74-149; tritoninfa = 74. Abstract: It was found that Charinus acosta (Quintero, 1983) oviposits from March to August. In the warmest month (June-August) the oviposition occurred 35-88 days latter that either the ecdysis or the anterior oviposition. But if ecdysis occurred ending the summer (September or October), the formation of an ovigerous sac occurred after 178-187 days. In the laboratory, size of the brood varied between three and nine (average = 4.2), in a directly relationship with mother size. The first postembry-onic states lasted (in days) as follows: prenymph = 7; protonymph = 60-131; deutonymph = 74-149, tritonymph =74.
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We present data and remarks on the history and contents of the whip spider collection housed in the Natural History Museum of Vienna, Austria. The collection comprises a total of 167 specimens from 4 families, 10 genera and 27 species. It includes types of four species: Charinus ioanniticus (Kritscher, 1959), Damon brachialis Weygoldt, 1999, Phrynus parvulus (Pocock, 1902) and Paraphrynus mexicanus (Bilimek, 1867). Short notes on interesting objects and former curators are provided as well as an appendix with a list of species kept alive by Michael Seiter.
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Se describe una especie nueva del género Phrynus Lamarck, 1801, sobre la base de numerosas hembras y machos procedentes del occidente de Guatemala (departamentos de Escuintla y Sololá) y del occidente de Honduras (departamento Fran-cisco Morazán). Se distingue de todos sus congéneres por presentar una espina supernumeraria entre Pd-5 y Pd-6 (patela del pe-dipalpo, superficie dorsal). Además, se aportan nuevas localidades y datos de historia natural para otros amblipigios de Guatemala y Honduras, incluido otro caso de depredación de amblipigios por un escorpión (previamente registrado, pero pasado por alto). Abstract: A new species of the genus Phrynus Lamarck, 1801, based upon several females and males, is described from western Guatemala (Escuintla and Sololá departments), and western Honduras (Francisco Morazán department). It differs from its conge-ners by having a supernumerary spine between Pd-5 and Pd-6 (dorsal surface of the pedipalp patella). Also, new localities and data on natural history are given for other Guatemalan and Honduran whip spiders, including another case of whip spider predation by scorpions (previously recorded, but overlooked).
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Five species of the whip spider genus Sarax are recognized from Borneo, with the following four species newly described: Sarax yayukae sp. nov. from Sabah (Malaysia), West and Central Kalimantan (Indonesia), and three species from East Kalimantan, S. cavernicola sp. nov., S. sangkulirangensis sp. nov., and S. mardua sp. nov.. Sarax mardua and S. cavernicola have pale coloration, reduced eyes and elongate legs suggesting troglomorphic adaptations to cave environments. The characters diagnosing the family Charinidae and the genus Sarax are discussed and revised. The distribution patterns of Sarax species in Southeast Asia, especially in Borneo Island, are discussed in relation to their habitat preferences. The generic status of Stygophrynus moultoni Gravely 1915 (Charontidae) is briefly discussed.
Book
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This authoritative catalogue will greatly assist readers in finding the correct taxonomic name for any given family, genus or species within each of the six arachnid orders treated. It contains a valuable summary of bibliographic information, enabling readers to access the worldwide literature for these smaller orders. The catalogue presents full bibliographic data on each of the taxa named thus far, treating over 1600 species. It contains the most current classification system for each group, some of which have not been catalogued on a world scale for over 70 years. A summary of taxonomic changes is included. This quality reference will be of immense value to arachnologists, systematists, taxonomists, ecologists and biodiversity professionals, especially those interested in tropical rainforest communities.
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The genus Heterophrynus is for the first time recorded from Transandean areas. Heterophrynus boterorum sp. nov. and Heterophrynus silviae sp. nov. are described respectively from Tolima and Valle del Cauca departments, Colombia, based on material from the 2006 Arachnological Expedition of Museu Nacional to Colombia. Heterophrynus nicefori Amado & Morales, 1986, from Meta department is newly considered a junior subjective synonym of Phrynus batesii Butler, 1873 (currently in Heterophrynus). Heterophrynus is currently known from Amazon forest, Brazilian Cerrado, Littoral Ridge of Venezuela and Andean forests. A revised terminology is proposed for the constituent parts of male and female gonopods of Heterophrynus.
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Charinus ioanniticus (Kritscher 1959) (Charinidae) is recorded for the first time from Egypt. Two specimens were collected from Burg El-Arab near the Mediterranean coast, north of Egypt.
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A new species of Charinus is described from 2 caves in Serra do Ramalho, Carinhanha, Bahia, Brazil. Hitherto six other species were known from this country. The new species is the first troglobiont Charinus from Brazil and the largest of all known species. It has no median eyes, a small whitish spot at each side of the carapace in place of the lateral eyes, a reduced tritosternum, not projecting in a long apophysis, the ventral corner of the prolateral face of femora II-IV prolonged in a spiniform apophysis, elongated spines and legs, thin cuticle and pale coloration.
Article
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A new species of Charinus is described from the Olhos d'Água cave, in Itacarambi, Minas Gerais, Brazil. Hitherto seven other species were known from this country. The new species is the second troglobiont Charinus from Brazil. It has only tiny median eye spots (sometimes absent), reduced lateral eyes, the ventral corner of the prolateral face of femora II-IV prolonged in a spiniform apophysis, elongated spines and legs, thin cuticle and pale coloration. Resumen Se describe una nueva especie de Charinus que habita en la cueva Olhos d'Água, Itacarambi, Minas Gerais, Brasil. Hasta ahora se conocían otras siete especies de este país. La nueva especie constituye el segundo Charinus troglomórfico de Brasil. Entre sus características morfológicas sobresalen: Ojos medianos ausentes (en su lugar existen diminutas manchas oculares, ausentes en algunos ejemplares), ojos laterales reducidos, esquina ventral de la cara prolateral de los fémures II-IV prolongada formando una apófisis espiniforme, espinas y patas alargadas, cutícula delgada y despigmentada.
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A new species of the genus Sarax Simon, 1892 is described from Panay Island, Philippines. Sarax curioi sp. n. is the second species of the genus from the country and can be distinguished from the other Philippine species (Sarax brachydactylus Simon, 1892) by the sclerotized granules of the pedipalp surface, the spines of the pedipalp distitibia, the number of denticles of the chelicerae claw and the shape of the denticles of the chelicerae basal segment. Sarax newbritainensis Rahmadi and Kojima, 2010 is newly recorded from New Ireland Island, Papua New Guinea.
Article
Charimus ioanniticus (KRITSCHER 1959) has been reported from the countries around the eastern border of the Mediterranean, Greece, Turkey, Israel, and Egypt. The Greek population, from the islands Rhodes and Kos, is an all-female population that most likely reproduces parthenogenetically. Redescriptions of C. ioanniticus, C. bengalensis GRAVELY 1911 from India and of C. abbatei DELLE CAVE 1986 from Somalia are presented, and Charinus pakistanus n. sp. from Pakistan (holotype male in MHNG, Geneva) is described. Morphologically, C. ioanniticus is most closely related to C. socotranus WEYGOLDT, POHL & POLAK 2002 and C. bengalensis. These species are members of the natural C. bengalensis species group or Charinides group, which ranges from India over Pakistan and Turkey to Greece and Socotra. C. dhofarensis WEYGOLDT, POHL & POLAK 2002 from the Arabian Peninsula and C. abbatei from Somalia are not related to this group. The question of the enigmatic Sarax mediterraneits DELLE CAVE 1986 is also addressed: it may finally turn out to be based on mislabelled specimens and be a junior synonym of one of the earlier described species of Sarax.
Article
Whip spiders use stalked spermatophores for sperm transfer. These are complex structures, and their morphology varies among genera and families. Usually, the paired sperm masses hidden within the spermatophores are small, and there has been a co-evolution of spermatophores and those parts of the female genitalia which are used to pick up the spermatozoa and to store spermatozoa. These are structures like specialized sclerotizations, glands or, in a few species, seminal receptacles which are hidden inside the genital atrium (or uterus externus). In most species there are paired erectile bodies, homologous to genital appendages, which are attached to the dorsal side of the genital operculum which also is part of an appendage homologon. All these structures vary among genera and families. The comparison of spermatophores and genitalia of different species belonging to most genera and families suggest that the female gonopods consist primarily of paired cushion-like structures, each equipped with a small finger-like appendage vestige. These appendage vestiges are retained in many species, particular in the Charinidae and Charontidae. They are erectile by increase in blood pressure, and they are thereby probably bent in characteristic ways and thus can pull off the sperm masses from the spermatophore. In some Charinidae, and in some species of Damon and Phrynichus (Phrynida, Phrynichidae) these appendage vestiges are totally lost. In the Phrynidae, on the other hand, they have become sclerotized and hard. They form the well-known claw-like sclerites, and an invagination at the base of each sclerite has been shaped to form a true seminal receptacle. Similar genitalia have evolved convergently in the genus Trichodamon (Phrynida, Phrynichidae). Spermatophores and the corresponding female genitalia and their mechanisms of a number of genera from most families are described and illustrated.
Article
The kingdom Animalia is here estimated to have a total of 1,659,420 described species (including 133,692 fossil species) in 40 phyla. Among these, the most successful phylum Arthropoda alone represents 1,302,809 species, or about 78.5% of the total. The second largest phylum, Mollusca (118,061 species), is <10% of Arthropoda in diversity, but it is still much more diverse than other successful invertebrate phyla Platyhelminthes (29,488 species), Nematoda (25,043 species), Echinodermata (20,550 species), Annelida (17,426 species), Cnidaria (16,363 species), Bryozoa (11,474 species) and Porifera (10,876 species). The phylum Craniata, including the vertebrates, represents 85,432 species (including 19,974 fossil species): among these, 35,644 species of "fishes", 7,171 species of amphibians, 15,507 species of reptiles, 11,087 species of birds, and 16,014 species of mammals.
Article
Reproductive biology and spermatophore morphology of four species of Amblypygi, belonging to two different families, are described. Basically, courtship and mating in Sarax buxtoni, S. brachydactylus, Stygophrynus brevispina nov. spec. and Charon cf.grayi are similar to those of many other whip spiders. However, each species has its own peculiarity. Like in other species, the female is not passive during courtship; she plays an active role with her own courtship signals and may even initiate courtship behaviour. The comparison of the spermatophores of S. buxtoni and S. sarawakensis with S. brachydactylus show two different types of spermatophores. At present, all three species should be included in the same taxon and Phrynichosarax should be considered a junior synonym of Sarax as long as there are no synapomorphies known. Mating behaviour and spermatophore morphology of C. cf.grayi from two different localities suggest both represent different species. Their behaviour and spermatophores and that of Stygophrynus brevispina supports the placement of the genera Stygophrynus and Charon in the taxon Charontidae. The presence of lateral horns on the spermatophore head of Charon raises the question of whether these structures are synapomorphic for the Neoamblypygi or even Euamblypygi. This question cannot be answered conclusively at present.
Article
Abstract— This paper reports results from a cladistic analysis of the 11 Recent arachnid orders. The polarities of 64 newly discovered and traditional characters were determined through outgroup comparisons that included Eurypterida, Xiphosura, Trilobita and Crustacea. A branch-and-bound algorithm was used to discover a single tree (consistency index 0–59). The relationships suggested by this analysis differ substantially from previous interpretations of arachnid phylogeny, and a new taxonomic system is introduced to accommodate these results. This analysis suggests that Arachnida is monophyletic and composed of two principal lineages, Micrura and Dromopoda. Possible synapomorphies of Micrura include a pygidmm, tntosternum, six principal lateral eyes, poorly sclerotized postgenital appendages, coxal gland orifices near leg 1, an array of micxotubules associated with the spermatozoan nucleus, and absence of coxal endites on the walking legs. The micruran orders appear to have the following relationships: (Palpigradi (Araneae (Amblypygi (I helyphonida, Schizomida)))) (Ricinulei, Acari). Possible synapomorphies of Dromopoda include transverse carapaeal furrows, greatly reduced prosomal sternum, prosomal endosternite with two segmental components, stomotheca, bicondylar femoropatellar and patellotibial joints and extensor muscles. The dromopodan orders appear to have the following relationships: Opiliones (Scorpioncs (Pscudo-scorpiones, Solifugae)).
Article
The available evidence on the ecological factors favoring the existence and origin of natural parthenogenesis is evaluated. Analysis of the geographical distributions of the well-known cases of animal parthenogenesis in nature reveals (i) that most of these species exist in natural disclimax communities and (ii) that within these communities they exist in isolation from closely related congeneric species. Parthenogenesis can only evolve in areas devoid of the generating bisexual species, because such species would prevent newly formed unisexuals from establishing clones due either to hybridization or competition. Furthermore, the two unique features allowing parthenogenetic species to invade and occupy open habitats faster than bisexuals are (i) a double intrinsic rate of increase and (ii) the ability of one individual to establish a new colony.
Republic of Singapore: 1 L adult (wild caught), Singapore, South West District, outskirts of Singa-pore, near Jurong Bird Park
  • Sarax
Sarax singaporae: Republic of Singapore: 1 L adult (wild caught), Singapore, South West District, outskirts of Singa-pore, near Jurong Bird Park, 1u 199 7.340N, 103u 429 23.190E, 14 September 2010, leg. and det. M. Seiter (NHMW 21893);
52 (as Phrynicosarax [sic] buxtoni); Speijer 1937: 173; Weygoldt 1994: 244. Sarax batuensis Roewer 1962: 519–520, Figs
  • Mello
  • Leitã
Mello-Leitã o 1931: 52 (as Phrynicosarax [sic] buxtoni); Speijer 1937: 173; Weygoldt 1994: 244. Sarax batuensis Roewer 1962: 519–520, Figs. 3a–b (syn. by Kraus 1970: 178).
Amblypygi: Phrynidae) de Guatemala y Honduras
  • L F Armas
  • C De
  • R E Viquez
  • Trujillo
Armas, L.F. de., C. Viquez & R.E. Trujillo. 2013. Nueva especie de Phrynus Lamarck, 1801 (Amblypygi: Phrynidae) de Guatemala y Honduras. Revista Ibé de Aracnologí 23:25–31.
Catalogue of the smaller arachnid orders of the world: Amblypygi
  • M S Harvey
Harvey, M.S. 2003. Catalogue of the smaller arachnid orders of the world: Amblypygi, Uropygi, Schizomida, Palpigradi, Ricinulei and Solifugae. CSIRO Publishing Huntingdon, Collingwood, Victoria, Australia.
Whip spiders of the World, version 1.0. Western Australian Museum
  • M S Harvey
Harvey, M.S. 2013. Whip spiders of the World, version 1.0. Western Australian Museum, Perth. Online at http://museum.wa.gov.au/ catalogues-beta/whip-spiders [accessed 06 March 2014].
Genitalmorphologie und Systematik der Amblypygi (Arachnida)
  • O Kraus
Kraus, O. 1970. Genitalmorphologie und Systematik der Amblypygi (Arachnida). Bulletin du Muséum National d'Histoire Naturelle, Paris 41:176-180.
Pedipalpos do Brasil e algumas notas sobre a ordem
  • C Mello-Leito
Mello-Leitã o, C. 1931. Pedipalpos do Brasil e algumas notas sobre a ordem. Archivos do Museu Nacional 33:7-72.
Revision of the amblypygid spiders of Cuba and their relationships with the Caribbean and continental American amblypygid fauna. Studies on the Fauna of Curaç and other Caribbean Islands
  • D Quintero
  • Jr
Quintero, D. Jr. 1983a. Revision of the amblypygid spiders of Cuba and their relationships with the Caribbean and continental American amblypygid fauna. Studies on the Fauna of Curaç and other Caribbean Islands 65:1–54.
Einige Arachniden aus den Batu Caves in Malaya
  • C.F Roewer
Roewer, C.F. 1962. Einige Arachniden aus den Batu Caves in Malaya. Pacific Insects 4:517-520.
Representatives of the order Amblypygi (Arachnida) found in Israel
  • R Rosin
  • A Shulov
Rosin, R. & A. Shulov. 1960. Representatives of the order Amblypygi (Arachnida) found in Israel. Bulletin of the Research Council of Israel 9B:167–168.
Two new troglobitic Amblypygid of the genus Tarantula from Mexican Caves (Arachnida)
  • J.M Rowland
Rowland, J.M. 1973. Two new troglobitic Amblypygid of the genus Tarantula from Mexican Caves (Arachnida). Bulletin of the Association for Mexican Cave Studies 5:123-128.
A new locality for Charinus ioanniticus (Kritscher, 1959) (Amblypygi: Charinidae) in Turkey
  • O Seyyar
  • H Demir
Seyyar, O. & H. Demir. 2007. A new locality for Charinus ioanniticus (Kritscher, 1959) (Amblypygi: Charinidae) in Turkey. Serket 10: 109–111.
A collection of pedipalps from the Raffles Museum
  • E.A.M Speijer
Speijer, E.A.M. 1937. A collection of pedipalps from the Raffles Museum. Bulletin of the Raffles Museum 13:171-175.
  • L F Armas
  • De
Armas, L.F. de. 2000. Parthenogenesis in Amblypygi (Arachnida). Avicennia 12/13:133-134.
  • P Weygoldt
Weygoldt, P. 1994. Amblypygi. Pp. 241–247. In Encyclopaedia Biospeologica. (C. Juberthie & V. Decu, eds.). Société de Biospéologie: Moulis and Bucarest.
Notes on Pedipalpi in the collection of the Indian Museum. I. New Pedipalpi from Ceylon
  • F.H Gravely
Gravely, F.H. 1911. Notes on Pedipalpi in the collection of the Indian Museum. I. New Pedipalpi from Ceylon. Records of the Indian Museum 6:33-36.
First record of Amblypygi (Charinidae: Charinus ioanniticus) from Turkey
  • F Kovarik
  • D Vlasta
Kovarik, F. & D. Vlasta. 1996. First record of Amblypygi (Charinidae: Charinus ioanniticus) from Turkey. Klapalekiana 32:57–58.
Ergebnisse der von Dr. O. Paget und Dr. E. Kritscher auf Rhodos durchgefuhrten zoologischen Exkursionen, II Pedipalpi (Amblypygi)
  • E Kritscher
Kritscher, E. 1959. Ergebnisse der von Dr. O. Paget und Dr. E. Kritscher auf Rhodos durchgefuhrten zoologischen Exkursionen, II Pedipalpi (Amblypygi). Annalen des Naturhistorischen Museums Wien 63:453-457.
Revision of the amblypygid spiders of Cuba and their relationships with the Caribbean and continental American amblypygid fauna
  • D Quintero
Quintero, D. Jr. 1983a. Revision of the amblypygid spiders of Cuba and their relationships with the Caribbean and continental American amblypygid fauna. Studies on the Fauna of Curaçao and other Caribbean Islands 65:1-54.
Bifid spines in Paraphrynus azteca (Pocock) (Amblypygi: Phrynidae)
  • D Quintero
Quintero, D. Jr. 1983b. Bifid spines in Paraphrynus azteca (Pocock) (Amblypygi: Phrynidae). Journal of Arachnology 11:99-100.
Parthenogenesis in Amblypygi (Arachnida)
  • L.F de Armas
Online at <http://museum.wa.gov.au/catalogues-beta/whip-spiders> accessed 06 March 2014.
  • M.S Harvey
Los amblipgos de Repblica Dominicana (Arachnida: Amblypygi)
  • L.F de Armas
  • A.P Gonzlez
  • Gravely F.H