The one‐palped spider genera Tidarren and Echinotheridion in the Old World (Araneae, Theridiidae), with comparative remarks on Tidarren from America

The one‐palped spider genera Tidarren and Echinotheridion are more or less confined to the tropics and subtropics of the southern hemisphere, with occurrence in South America, continental Africa, Madagascar, Cape Verde Islands, Canary Islands, and Yemen. This comparative description of the taxonomy and biology of the one‐palped spiders in the Old World comprises altogether 22 species, of which 17 are new to science. Four new species of Tidarren are described from Yemen (T. dentigerum n. sp. ♂♀, T. gracile n. sp. ♂♀, T. konrad n. sp. ♀, and T. sheba n. sp. ♂♀), eight from mainland Africa (T. aethiops n. sp. ♀, T. afrum n. sp. ♀, T. circe n. sp. ♀, T. griswoldi n. sp. ♀, T. lanceolatum n. sp. ♀, T. perplexum n. sp. ♀, T. ubickorum n. sp. ♀, and T. usambara n. sp. ♀), and five from Madagascar (T. apartiolum n. sp. ♀, T. dasyglossa n. sp. ♀, T. ephemerum n. sp. ♂, T. horaki n. sp. ♀, and T. obtusum n. sp. ♂♀). Three new synonyms are established. Tidarren chevalieri nov. syn. is a junior synonym of T. cuneolatum. Theridion guineense Simon, 1907 nov. syn. and Theridion turrigerum Simon, 1899 nov. syn. are synonymized with Tidarren scenicum (Thorell, 1899) nov. comb. (transferred from Theridion). The following further species are reconsidered: Tidarren argo Knoflach and van Harten, 2001 from Yemen, with a first record also from Chad; T. cuneolatum (Tullgren, 1910) from mainland Africa, Yemen, and the Canary Islands; and T. levii Schmidt, 1957 and T. scenicum from mainland Africa. In addition, the three American species T. haemorrhoidale (Bertkau, 1880), T. mixtum (O. P.‐Cambridge, 1896), and T. sisyphoides (Walckenaer, 1842) are included for comparison. Thus all hitherto known Tidarren species of the world are covered. A key for the Yemeni species is provided (males and females) as well as a tentative key for the African species (females only; Madagascar included) and the American species. For T. dentigerum and T. gracile from Yemen we give a first description of the copulatory behaviour. The genus Echinotheridion, though represented by eight New World species, is known from only one species in the Old World, E. gibberosum (Kulczynski, 1899) from the Canary Islands and Madeira. Comparative notes on this endemic offshoot are given.


Introduction
The spider genera Tidarren Chamberlin andIvie, 1934 andEchinotheridion Levi, 1963 represent prominent taxa owing to numerous morphological and ethological characteristics Bonnet 1935;Branch 1942;van Harten 2000, 2001;Knoflach 2002aKnoflach , 2002bKnoflach , 2004Knoflach and Benjamin 2003;Levi , 1963Levi , 1980. They have a single male palp (Figures 4,6,8,10,(75)(76)(77)(78)(79)(80)118,132,190,293) and show striking sexual size dimorphism (Figures 1,(62)(63)(64)(65)(66)(67)(68)(69)103,104,(116)(117)(118)(119)(132)(133)(134)(135)(286)(287)(288)(289)(305)(306)(307). The minute males obligatorily self-amputate one of their palps after their penultimate moult (Figures 2-6). During copulation males die a sudden sexual death. Therefore, it seems unsurprising that sexual cannibalism is a regular phenomenon in these taxa (Knoflach and van Harten 2000). Moreover, in some species the males are emasculated by their mates during copulation (Figures 67-69, 307), the single male palp being amputated and acting independently of the male's body (Knoflach and van Harten 2001;Knoflach 2002aKnoflach , 2002b. These outstanding biological traits raise questions on their origin and evolution (e.g. Eberhard 1985;Ramos et al. 2004Ramos et al. , 2005. Nevertheless, the present taxonomic situation appears to conceal even more problems, which are fundamental for further research in evolutionary biology and phylogenetic analysis. This study provides a synoptic basis for the taxonomy, biology, and distribution of the onepalped spiders in the Old World, but probably still covers only a fraction of the species occurring in this region, especially in Africa. Currently 16 species of one-palped spiders are known worldwide from the tropics and subtropics of Africa and the Americas (seven species of Tidarren, nine species of Echinotheridion). Eleven of these occur in South and Central America (Platnick 2006). From Africa three representatives of Tidarren have so far been recorded: T. cuneolatum (Tullgren, 1910) from mainland Africa, the Canary Islands, and Yemen (Knoflach and van Harten 2000); T. levii Schmidt, 1957 from an import from Congo, and T. chevalieri from the Cape Verde Islands (Berland 1936;Schmidt and Krause 1994;, the last being considered here as a new synonym of T. cuneolatum. Another species has recently been described from Yemen, T. argo Knoflach and van Harten, 2001. In the course of our investigation of the theridiid fauna of Yemen, the genus Tidarren was found to be represented by a number of unknown species. Further private and museum collections from continental Africa and Madagascar revealed a surprising diversification of this genus. As a result, a total of 17 species of Tidarren can be described as new to science, four from Yemen and 13 from Africa. According to the regular occurrence of this genus in Africa it is assumed that older descriptions, especially in the large collective genus Theridion, could involve also Tidarren. This is shown for Tidarren scenicum (Thorell, 1899) nov. comb., being transferred from Theridion. Owing to their small size, males are under-represented in museum collections, hence many species comprise females only. Also in the closely related genus Echinotheridion seven out of nine species have been described from one sex only (Platnick 2006) and thus are still poorly known. Eight of these species occur in South America. The only Old World representative, E. gibberosum (Kulczynski, 1899), which is endemic to the Canary Islands and Madeira, appears to be best established. Descriptions and diagnoses of both genera were based upon material from the Americas Levi , 1963Levi and Levi 1962), whereas our knowledge of the fauna of the Old World has to be regarded as incomplete. It is the aim of this study to address this deficiency. in lateral (7,9) and frontal view (8, 10). (7, 8) Tidarren argo Knoflach and van Harten from Yemen. (9) T. cuneolatum (Tullgren) from Yemen. (10) Echinotheridion gibberosum (Kulczynski) from Tenerife. Note high and steep clypeus and site of palp amputation with trochanter still present. Scale bars: 0.2 mm.

Material and methods
Specimens were examined with a Wild M8 stereo microscope. For drawings of genitalia a Wild M20 compound microscope was used. Epigynes were immersed in Hoyers compound fluid. For measurements, palps and legs were separated from the body of the spider, prepared as permanent mounts and measured through a compound optical microscope. For species available only from one or a few museum specimens, leg measurements were taken from the whole animal through a stereo microscope. The latter method is indicated in the text to give an impression of accuracy of measurements. Number, position, and pattern of trichobothria have not been analysed in such cases.
SEM micrographs were made with a Zeiss DSM 950 by K. Pfaller (Institute of Anatomy, Histology and Embryology; Department of Histology and Molecular Cell Biology, University of Innsbruck). For [47][48][49][50][51]95, 96 the Auto-Montage system was used for assembling several images at different focal planes and producing one digital photo. Habitus photographs were done with an analogue Nikon camera. Photographs and figures are by B.K., unless indicated otherwise. Owing to their complexity, male and female genital organs are drawn in different perspectives. For space-saving reasons the orientation of the female epigynum in lateral view need not be in straight antero-aboral line, but may be rotated. Thus, direction of epigynal protuberance is discernible only on the habitus photos.
Spiders of all stages were kept alive separately in plastic boxes (76462.5 cm) at room temperature in Innsbruck and reared to adulthood. Copulations were observed with a stereo microscope with horizontal objective body (Nikon SMZ-2B), magnification up to 650.
The following abbreviations are used: male palp-bH, basal haematodocha; C, conductor; dH, distal haematodocha; E, embolus; f, fundus of sperm duct; S, subtegulum; T, tegulum; MA, median apophysis; mH, median haematodocha. Epigynum/vulva-CO, copulatory orifice; CD, copulatory duct; FD, fertilization duct; R, receptaculum seminis. Details of the generic diagnosis of Echinotheridion are based upon the only Old World representative E. gibberosum. The type species has not been examined. A key is provided for the Yemeni Tidarren species. The key for the African species has to remain provisional owing to the scarcity of material. Altogether, 25 species of Tidarren, grouped by geographic region, and one species of Echinotheridion are treated herein.

Etymology
Tidarren is a hybrid Gosiute and Greek name, meaning small male; from Gosiute Tidutsi5short, small and Greek arrhenos5male, gender neuter, see .
Shape of epigynal protuberance, cymbium, distal tegular rim, and conductor are diagnostic features. Unusually among Theridiidae, the cymbium shows marked speciesspecific features, such as shape, protuberances, ridges, and denticles. It provides the main functional contact, as it is placed over the epigynal protuberance during insertion (Knoflach and van Harten 2000).
Recently, the spermatozoa of T. argo have also been investigated (Michalik et al. 2005). They differ from those hitherto found in other spiders and appear to be highly specific, with a strongly elongated nucleus, a conspicuous implantation fossa, base of axoneme close to acrosomal complex, axoneme extending deeply into postcentriolar elongation, and proximal centriole unusually far away from the distal one. A thin secretion sheath encloses the encapsulated cleistospermium. Interestingly, mature spermatozoa are embedded in a copious secretion instead of being close-packed (Michalik et al. 2005).

Behaviour
Palp amputation. In contrast to all other known spiders, adult males of Tidarren and Echinotheridion have only one palp, as a result of self-amputation (Branch 1942;van Harten 2000, 2001). Juveniles and newly moulted subadults still have both palps (Figures 2,3,5). The semi-emasculation is accomplished a few hours after the penultimate moult, when the male raises one palp and turns around in circles until the palp becomes caught in the threads at the moulting site. By further circling the palp becomes constricted and finally breaks off (Figure 4), regularly at the trochanter and femur joint (Figures 10,21,22). The amputation takes about 4-13 min in T. cuneolatum (see Knoflach and van Harten 2000), 2-6 min in T. argo (see Knoflach and van Harten 2001), 2-3 min in T. sisyphoides (see Branch 1942;Knoflach and Benjamin 2003), and involves 8-15 rotations of the male. Subsequently, the male feeds on its own detached palp (Knoflach and van Harten 2000; see also Figure 6, palp already sucked out). The last moult therefore involves only one-palped males. Among several hundreds of males there was no adult male with two palps.
Copulatory behaviour. Tidarren species show numerous outstanding behavioural traits during copulation van Harten 2000, 2001;Knoflach 2002aKnoflach , 2002bKnoflach , 2004Knoflach and Benjamin 2003). Males generally die from a sudden sexual death during insertion. They become exhausted with copulation, probably because a considerable part of the haemolymph is shifted from the male's body into the palpal organ (Figures 67,68,104,117,118,132,133,287,289). Moreover, the prosoma of the male becomes shrivelled (Figure 70), supposedly owing to irreversible contraction of the prosoma muscles. As a consequence, copulation comprises a single insertion only. In most species hitherto observed copulation is associated with sexual cannibalism of the male, either post-copulation or concurrent with sperm transfer. In T. cuneolatum the male is devoured after a few minutes of insertion (Knoflach and van Harten 2000), in T. dentigerum from Yemen likewise after some minutes ( Figure 119, see species description). Tidarren argo synchronizes sperm transfer and sexual cannibalism by emasculation, by tearing off the single male palp, which is accomplished within half a minute by the female immediately after successful genital contact (Figures 67-69;Knoflach and van Harten 2001;Knoflach 2002aKnoflach , 2002b. The separated gonopod remains fastened to the epigynum for several hours and presumably continues with sperm transfer independently of the male's body, while the female feeds on the palpless, emasculated male (Figures 68, 69). By this means genital contact is prolonged considerably. Furthermore, this procedure provides a temporary mating plug and thus may lengthen the interval between copulations of a female. Another new species from Yemen described herein shows an intermediate behaviour. In T. gracile n. sp. females start to consume their mates during copulation, without amputating the male gonopod (Figures 133,134). After a period of such consummatory sexual act the male is removed from the epigynum and finally completely sucked out (Figures 123,135). The American T. sisyphoides is an exception as regards mate consumption. Although copulation is also accompanied by the male's death, the female removes the dead male from the epigynum and casts him away without consuming him (Knoflach and Benjamin 2003;see also Figures 1b, 288).
Sex roles are evidently reversed in Tidarren. Males are monogynous, while females practise polyandry, which implicates higher potential reproductive success in females. This reversal is also indicated by the unusually active role of females during courtship. In many species, females actively court their mates throughout courtship.
In species with considerable size dimorphism, problems may arise concerning orientation of the partners in order to assume the appropriate copulatory posture. In Tidarren, this is solved by construction of a mating thread, where the male plucks rhythmically until the female approaches. Copulation then takes place on the mating thread. Obviously, it helps the male to locate the female genital organ. The female even cooperates by assuming a special posture, her legs III flexed and forming a circle, the tarsi close together on the mating thread (Figures 63,65,103). When the male approaches along the mating thread he is guided by her legs to the epigynum (Knoflach and van Harten 2000;Knoflach 2004). Only in T. sisyphoides does no web-spinning take place. Here the male climbs directly on to the female venter (Knoflach and Benjamin 2003;see also Figures 286, 287).

Inbreeding tolerance
In Tidarren argo females were observed to copulate with their offspring, apparently tolerating a certain amount of inbreeding in captivity.

Natural history
Egg-sac. Usually grey to brownish, with parchment-like envelope. Species produce different numbers of eggs according to differences in body size. The greatest fecundity is given in the large Tidarren sisyphoides, with 350-477 eggs per cocoon , far away from T. cuneolatum with about 70 on average (Knoflach and van Harten 2000).
Smaller species such as T. argo produce ca 40 eggs per sac (Knoflach and van Harten 2001) and T. gracile ca 20 (see below).
Web. The web consists of a retreat and a scaffold of threads extending to the side of a wall, not or rarely reaching the ground. Web dimensions depend on the space available, in the field large female webs may measure up to half a metre in height. The retreat is a densely spun hub into which mineral particles, exuviae, plant and prey remnants are incorporated. The webs of Tidarren haemorrhoidale and T. sisyphoides do not contain gum-footed lines or other viscid elements (Benjamin and Zschokke 2003).
Enemies. In Kansas Tidarren sisyphoides was severely preyed upon by the pirate spider Mimetus puritanus Chamberlin, 1923 (Mimetidae;. Other predators were the araneophagic theridiids Argyrodes trigonum (Hentz, 1850) and A. cancellatus (Hentz, 1850), which also invaded the webs of T. sisyphoides (see . The mantispid Mantispa viridis Walker, 1853 was encountered from egg-sacs of T. sisyphoides in Kansas (see . In Florida the eulophid hymenopteran Comastichus zopheros LaSalle, 1994 parasitized an egg-sac of T. sisyphoides (see . One juvenile female of the Yemeni T. argo was infested by a Mermithidae (Nematoda). One female of T. cuneolatum from South Africa had been parasitized by an acrocerid (Diptera), the primary larva being visible through the book lung cover. Another female from Tanzania had an ectoparasitic hymenopteran larva on the abdomen.

Distribution (Figure 51)
South and Central America, Africa, Yemen, Madagascar, Cape Verde, and Canary Islands. There are no records from northern Africa. Obviously, the genus is not present in SE Asia (Song et al. 1999). Most species are known from a few localities only. Two species are Figure 51. World distribution of the one-palped spider genera Echinotheridion (open circles) and Tidarren (closed circles and stripes). Occurrence in America based mainly upon Levi ( , 1963Levi ( , 1967. expansive and behave hemisynanthropically (T. cuneolatum in Africa and T. sisyphoides from southern USA to South America). For further details see discussion and species description.
Echinotheridion Levi, 1963 Type species: E. cartum Levi, 1963. In the Old World, the genus Echinotheridion is represented by only one species, E. gibberosum. As the type species from South America is not taken into consideration herein, a comparative analysis is not justified. For details of generic diagnosis see Levi (1963). Echinotheridion shares numerous features with Tidarren, sexual size dimorphism, general appearance and habits, web structure, palp amputation, copulatory behaviour, see also species description. In the following only differentiating characters are given.
The most striking and eponymous (Echinotheridion5''spiny Theridion'') character of this genus is shown by the females, which have coxal spurs. Posteriorly on the base of each coxa IV a conspicuous spur faces towards the other and towards the epigynal protuberance (Figures 23,24). It still is uncertain whether these spurs serve as additional holdfast structures or rather facilitate the female to terminate genital contact more easily (Figure 310). At least they are not essential for genitalia coupling. During insertion, contact between male palp and coxal spurs sometimes occurs and sometimes not.
Males have no coxal spurs. The male palp of Echinotheridion, especially of the three South American species known from the male, is apparently even more elaborate than that of Tidarren (figures given by Levi 1980;Buckup and Marques 1989;Ramirez and Gonzalez 1999). Cymbium and palpal organ exhibit numerous huge projections. These are comparatively less pronounced in E. gibberosum (Figures 295-298). Unlike in Tidarren, the embolus lacks distinct lobes at its base. The distal and presumably also the median haematodochae are vestigial, which is assumed to have similar effects on palpal application as in Tidarren. Basal haematodocha not modified (Figure 307-309). It appears that the epigynal protuberance is less prominent and less pointed (Figures 299,303; see also figures given by Levi 1963;Marques and Buckup 1989;Ramirez and Gonzalez 1999). Unlike Tidarren species, in E. gibberosum the copulatory ducts are long, coiled, and heavily sclerotized.

Behaviour
Palp amputation. Loss of one palp is achieved by self-amputation also in E. gibberosum males. The process closely resembles that in Tidarren. Two to three hours after the penultimate moult two males circled around their own appendage 16-17 times, thereby twisting off the palp within 6-11 min (Knoflach 2002a(Knoflach , 2002bKnoflach and Pfaller 2004).
Copulatory behaviour. Echinotheridion gibberosum follows the overall copulatory pattern of Tidarren (Knoflach 2002a(Knoflach , 2002b(Knoflach , 2004: copulation is induced via a mating thread, accompanied by the male's exhaustion and concludes with sexual cannibalism (Figures 305-307). As in T. argo the male is emasculated before being consumed. The female detaches the palp by continuous circling, but later than in T. argo, at least after some minutes of cataleptic insertion, during which the male appears to die of fatigue. Also here the palp remains on the female for many hours. However, the main holdfast structures for genital coupling are sclerites and not palpal membranes : the cymbium locks behind the epigynal protuberance in a fold of the integument, whereas the conductor adjoins the anterior side of the epigynum. Emasculation obviously also occurs in the South American E. cartum, as a female was collected with attached male palp (Ramirez and Gonzalez 1999). The protruding cymbium was found to even pierce the female's cuticle, thereby leaving copulatory scars.

Natural history
Web. Similar to that of Tidarren.
Enemies. In some retreats of Echinotheridion gibberosum from Tenerife the characteristic cocoons of the genus Ero (Mimetidae) were present, which may indicate possible predation by this araneophagic spider.

Discussion
The one-palped spiders constitute a well-defined and conspicuous taxonomic group within Theridiidae, although discovered rather recently by Chamberlin and Ivie (1934) and Levi (1963). However, the generic separation of Tidarren and Echinotheridion has been discussed controversially (Wunderlich 1987(Wunderlich , 1992Schmidt 1990), since the proposed synonymy with Tidarren by Wunderlich (1987) has been withdrawn again (Schmidt 1990;Wunderlich 1992). Both genera appear to share a number of synapomorphic characters, such as palp amputation, overall copulatory behaviour accompanied by sexual cannibalism, sexual size dimorphism, and morphology of the male palp, e.g. the modified cymbium, the large development of the tegulum, and the enormous fundus of the sperm duct. Generic differentiation is based mainly on the female coxal spurs present in Echinotheridion (Figures 23,24) and on details of the male palp and epigynum. For a better understanding of these relationships a more profound knowledge of the species of Echinotheridion is required. Seven out of the nine species are known from one sex only. In recent taxonomic works the generic rank of Echinotheridion has not been doubted any more Ramirez and Gonzalez 1999).
In the comprehensive phylogenetic analysis of Theridiidae by Agnarsson (2004) Tidarren is placed in the derived subfamily Theridiinae owing to reduction of trichobothria on the male palpal tibia and absence of a colulus. For Echinotheridion a similar assignment was suggested. Interestingly, some of the copulatory features of the one-palped spiders are widespread in basal theridiids and apparently have been retained here, as copulation takes place via a mating thread and sperm induction is independent of copulation (Knoflach 2004). Close affinities evidently exist with Achaearanea in general morphology and habits, for its morphological and genetic conformity and close phylogenetic relationship see Agnarsson (2004) and Arnedo et al. (2004). Surprisingly, sister relationship of Echinotheridion and Tidarren is rejected in a recent phylogenetic study based on molecular and morphological data (Agnarsson forthcoming).
According to their world distribution the subtropical to tropical spider genera Tidarren and Echinotheridion pose a classical biogeographical problem owing to their disjunct distribution in South America and Africa. Their occurrence in the Old World includes continental Africa, Madagascar, Cape Verde Islands, Canary Islands, and Yemen ( Figure 51). There are no records from Asia and Australia. No fossil records exist for one-palped spiders (J. Wunderlich, personal communication). Anyway, their phylogenetic placement among Theridiinae (Agnarsson 2004) suggests that this lineage is too recent for a Gondwanan fragmentation. Oldest fossil Theridiinae trace back to the Dominican amber, being only 10-30 million years old (Wunderlich 1988(Wunderlich , 2004. Thus, one-palped spiders must have spread rather late, probably by ''oceanic dispersal' ' (de Queiroz 2005), resulting in the current cross-continental ''amphi-Atlantic'' distribution.
As a result of the present study, the genus Tidarren is known from 21 species in the Old World. Hot spots of species diversity are in particular in Africa and Yemen, whereas Echinotheridion seems to be largely vicariant in South America. Another remarkable gross vicariance pattern is evident when comparing the world distribution of the speciose genus Achaearanea. This genus shows its highest diversity in the tropics of America (about 40% of species) and SE Asia (about 30% of species; see Platnick 2006), whereas only two species (out of more than 140) are known from mainland Africa. There, it appears to be replaced by Tidarren, at least at large scale. This may be supported also by the personal collections of A.v.H. in mainland Yemen, where six Tidarren species were found, but no Achaearanea, while in the Socotra archipelago only Achaearanea turned up.
At present, lack of knowledge and sampling does not allow assessment of the distribution pattern at species level. Almost 40% of the Old World species are recorded from single localities, with allopatric occurrence being predominant. In only a few localities, e.g. in Yemen, two or three species occurred sympatrically. Tidarren cuneolatum shows the most widespread distribution area, as this species behaves hemisynanthropically. According to preliminary evidence the Malagasy representatives appear to be distinct from the continental African species. These regions have no species in common. The American species of Tidarren differ from the Old World representatives in genital morphology, the copulatory orifices being widely separated in the T. sisyphoides group and the conductor of the male palp forming a small, hyaline appendage. Moreover, sexual size dimorphism is much more pronounced in the American species, presumably owing to female giantism coupled with a decrease of male size (Hormiga et al. 2000). Hence these species separate unequivocally in a distinct group, which evidently mirrors biogeographical splitting.
The spider genera with a single male palp exhibit interesting morphological and behavioural traits, which still deserve further research in evolutionary biology, but they also hide numerous problems concerning biogeography, taxonomy, and phylogenetics. The present study should draw attention to these outstanding, though long-neglected spiders.  Provisional key to the Tidarren species of Africa mainland and Madagascar (females only)

Etymology
The specific name refers to the copulatory behaviour of this species, sharing some analogies with the cephalopod Argonauta argo L., 1758 (Knoflach and van Harten 2001).

Diagnosis
Tidarren argo is distinguished from most of its Yemeni congeners by having the abdomen rounded 73,75,76). Epigynal protuberance broad and strongly protruding 73), copulatory orifices fused to tiny atrium and copulatory ducts long. The male palp differs by the broad cymbium (Figures 50,(52)(53)(54)(55), with distal teeth, by the shape of conductor and distal rim of tegulum and in particular by the system of haematodochae when expanded (Figures 41,(67)(68)(69)72). Ethological traits distinct from other species: copulation with palp amputation performed by the female (emasculation) when contact of genitalia is achieved (Figures 66-69).  Tables I, II. Number of dorsal setae on tibiae I-IV 2/2/1/2. Trichobothria in retrodorsal/prodorsal row on tibia of female palp 2/1, of legs I-IV 2/2, 2/1, 2/2, 2/3 in female and IV 2/1, 2/1, 2/2, 2/2 in male (2", 2R examined).  Sternum without posterior knob. Abdomen rounded, without tubercle, higher than long. Carapace uniformly dark brown in female. In male carapace brown to dark brown with darkened margins and darkened median band from eye region to centre. Chelicerae light brown, slightly suffused with grey. Sternum, gnathocoxae, and labium dark brown. Legs and female palp pale yellow to yellow brown with dark patches and annnulations. Basal segments of male palp pale yellow, cymbium yellow brown. Abdomen yellow to light brown, speckled, dorsally and laterally with numerous dark spots of various extent and white guanin patches. Anterior half dark with some white patches, posterior half light with a few dark markings. A small dark patch in apical region. Aboral region with indistinct white median stripe from apex to spinnerets and dark transverse arches. Epigaster dark brown in both sexes, book lung covers pale yellow. Venter with two white paramedian patches between epigastric furrow and spinnerets. Further white patches encircle spinnerets. Spinnerets brown to dark brown.

Taxonomic remark
The specimen from Tibesti differs slightly by having its epigynal protuberance a little more slender ( Denis (1955) identified the female from Tibesti as T. cuneolatum with some doubt.

Copulatory behaviour (Figures 62-72)
Courtship. During courtship females continuously vibrate their bodies and move their legs II. The male carefully walks around the female and from time to time palpates the female's legs or body ( Figure 62). After ca 10 min of such mutual courtship, he installs a mating thread from the female's vicinity to a distance of 3-4 cm away. At the opposite end, the male then plucks intensively. This vibratory signal induces the female to approach him along the thread and to adopt the copulatory posture (Figures 63-65).
Copulation. The ensuing copulation proceeds in a fairly unusual manner (Knoflach and van Harten 2001). At the immediate beginning of insertion the male palp becomes torn off by the female. She turns quickly around the only contact zone between the pair, the palpepigynum connection, until the single palp of the male breaks off ( Obviously, there is no loss of haemolymph, and the volume of the haematodochal horns does not decrease noticeably in the detached palp. The female removes the gonopod usually after consuming the male, by either pushing it away with her hind legs or, indirectly, by circling again, so that the projecting palp becomes fixed to the threads of the web and finally is torn off. Thereafter it is sucked out at the haematodochal horns.

Natural history
From the fifth day after copulation onwards the female proceeds with the production of egg-sacs, altogether five to six at intervals of 1-2 weeks (Knoflach and van Harten 2001). Three females were observed to build their first cocoon even on the fourth day after copulation. The brownish, round egg-sacs are attached together and guarded by the female (Figures 73, 74). Each egg-sac contains about 40 hatchlings. Hatchlings appear about 2 weeks after egg-laying. Observations on development indicate rapid post-embryonic development. Males become adult after three moults within about 18 days after hatching, females after 36 days. The subadult males matured after 7-9 days. Adult females live for 3-5 months (Knoflach and van Harten 2001). The species apparently tolerates a certain amount of inbreeding. On a few occasions females were observed to copulate with their offspring.
Parasites. One juvenile female collected on 27 July 1999 in Khamis Bani Sa'd was infested by a mermithid (Nematoda), which emerged from the spider on 6 August.

Distribution, habitat (Figure 259c)
Tidarren argo is known from several localities in Yemen and from a single record from Tibesti. In the type locality it was collected from old stumps of banana up to 60 cm, in banana plantations in a valley bed with a permanent river at 500 m a.s.l. surrounded by rocky mountains 1000 m high. In Sana'a T. argo occurred in a garden, where T. cuneolatum also lives in large numbers. At Hammam 'Ali it was found under an old tree trunk, at ar-Rujum under a rock, in both cases in agricultural areas. The specimens from Tibesti came from a small grotto at 1000 m altitude, ''petite grotte sous la grande cascade de la source de Yerra'' (Denis 1955).

Etymology
The species name apparently reflects the epigynal shape; lat. cuneolatus is a diminutive of cuneatus, meaning attenuate.

Copulatory behaviour
Courtship and copulation proceed via a mating thread ( Figure 103). Courtship lasts 3.5 min on average and involves active female courting (Knoflach and van Harten 2000). Her twanging with legs II and body oscillations induce the male to spin the mating thread. At the starting point of this thread he performs intensive plucking sequences, which stimulate the female to approach him along the mating thread and to assume the copulatory posture ( Figure 103). Insertion lasts 4 min on average and is terminated by the female pulling off her mate. Males die during insertion. Sexual cannibalism takes place after copulation (Knoflach and van Harten 2000). This overall copulatory procedure has been observed many times in pairs from Yemen, but was the same in the few pairs from the Canary Islands (Figures 103, 104) and Cape Verde Islands. In one pair from the Cape Verde Islands courtship lasted 14 min, insertion 4.6 min, until the male was removed from the epigynum and sucked out by the female. Four females from the Cape Verde Islands were allowed to copulate with males from Yemen, as follows. Courtship lasted 0.5-13.0 min (mean 7.4, n54), insertions lasted 2.8, 4.8, and 5.9 min (n53). Three copulations proceeded typically. However, one female did not remove the dead male from the epigynum for 3.5 h and then she removed him also from the web, without eating him. This exceptional female behaviour possibly may be explained by her well-nourished state.

Natural history
Egg-sac grey to light brown, with parchment-like envelope (Figure 121), containing 68 eggs on average (minimum-maximum 22-142, n512; population from Yemen). One female was observed to build three egg-sacs, the first 6 days after copulation. Development of T. cuneolatum from the Cape Verde Islands corresponds fully to the development of the Yemeni specimens, as observed by Knoflach and van Harten (2000). Juveniles hatch from the egg-sac in an early phase, and palp amputation and maturity moult take place outside the cocoon. Unlike in the rearings from Yemen, two virgin females from Cape Verde built two to three egg-sacs 2-3 months after they had reached maturity, which they devoured again after a few days of guarding.  assumed that males of ''T. chevalieri'' from Cape Verde would reach adulthood within the cocoon, which appears impossible according to our observations. We believe that the two one-palped males mentioned by these authors had stayed within a retreat and not hatched from an egg-sac, which is usually deposited in the retreat.
Parasites. One female from South Africa (Badplaas) had been infested by an acrocerid (Diptera), as the primary larva was visible through the book lung cover. One female from Tanzania had an ectoparasitic hymenopteran larva on the abdomen.

Habitat
In many localities T. cuneolatum behaves hemisynanthropically. Its preferred web sites are mainly stone walls and rock crevices, as well as house walls and window frames, and the stems of large, rough-barked trees. This may explain the synanthropic aspect of this species.

Distribution (Figure 259c)
Tidarren cuneolatum is widespread in continental Africa (South Africa to Kenya, Senegal, and Namibia) and mainland Yemen, and also occurs on the Cape Verde Islands. Berland (1920) reports this species also from Ethiopia. Apparently, it is not present in North Africa, the northernmost locality being Gran Canaria, where it obviously has been introduced.
There is no record up to now from Madagascar.

Etymology
Latin adjective derived from dens tooth and ger(um) to carry something, referring to the male cymbium, which bears several conspicuous teeth. The suffix agrees in gender of the genus name.

Measurements (mm)
["/R, n55/11, minimum-maximum ( examined) and 2/1, 2/1, 1/2, 2/2 in four males, 2/1, 2/1, 2/2, 2/2 in one male and 2/1, 2/0, 1/ 2, 2/2 in another male. Metatarsi I-III with one trichobothrium, position on I in female Somatic features, colouration (Figures 17,79,(116)(117)(118)(119)(120) In female, sternum shows a distinct posterior tubercle between coxae IV, bearing long hairs ( Figure 17); further tiny, marginal humps arise level with coxa. Male without sternal tubercle. Abdomen higher than long, with dorsal tubercle (Figures 79, 119, 120). Carapace uniformly dark brown in female. In male carapace yellow brown with dark longitudinal median band from eye region to centre, with dark to greyish clypeus and thin greyish margins. Chelicerae, labium, and gnathocoxae light brown to pale yellow. Chelicerae sometimes lightly suffused with grey. Sternum light brown with dark markings of various extent. Legs and female palp yellow brown to pale yellow with dark patches and annulations; in males dark markings less predominant. Basal segments of male palp pale yellow, cymbium yellow or reddish brown. Abdomen usually with large dark markings; abdominal pattern in male less distinct. Dorsum and sides dark brown from petiolus to tubercle; this dark anterior area may be interrupted by a longitudinal, lighter, whitish zone, ending subterminally. From subapical region a clear white stripe bends laterally and ventrally and forms borderline between anterior dark zone and an intermediate light zone. Apex dark anteriorly and light posteriorly. Aboral region with clear white median stripe from apex to spinnerets and dark longitudinal markings, sometimes rather large. Epigaster dark brown in both sexes, in females a small white membranous area behind epigynal protuberance; book lung covers pale yellow. Venter in female with dark and white marks, in male dark with two paramedian spots between epigastric furrow and spinnerets. Spinnerets brown. In two females the dark pattern was faded out, one female was very dark. Tibia not as wide as bulbus, tapering distinctly at base. Palpal organ 0.23-0.25 (0.24) mm wide (n55). Cymbium modified, protruding far beyond bulbus, ending as conspicuous, finger-like projection (Figures 79, 105-110), which bears several large sclerotized teeth. Number of teeth apparently variable (Figures 109, 110). This finger-like process was observed to be in contact with the epigastric furrow of the female during insertion. Distal  cymbial rim forms two small lobes facing palpal organ and shows ridges in between them. One of these lobes is close to conductor and bears strong hairs and tarsal organ. Tegulum very large. Subtegulum hidden between tegulum and cymbium. Distal rim of tegulum with short, rounded projection near embolar base. Retrolateral, membranous part of conductor 0.16 mm long, ending with pointed, sclerotized tip; prolateral, sclerotized part with a larger bilobed process and a small hump. Embolar base suboval, 0.10 long and 0.05 mm wide. Distal part of embolus 0.19 mm long. Haematodochae not modified (Figures 116-119).

Epigynum, vulva (Figures 17, 111-115)
Epigynal protuberance stout, short, about 0.1 mm long, strongly curved and therefore less protruding, heavily sclerotized, in side view hook-shaped ( Figure 111). Epigynal region around protuberance also sclerotized (0.3 mm diameter), posterior margin of this sclerotization behind epigynal protuberance excavated in midline, joining a small membranous area. Epigynal protuberance not clearly delimited from dark epigastric region, directed ventrad and therefore facing a similar direction as sternal knob (Figure 17). Both protuberances may serve as forceps for the male palp during genital coupling. Their distance apart, ca 0.2 mm, corresponds roughly with palpal width. In aboral view the protuberance is not clearly delimited from the surrounding integument ( Figure 115). Copulatory ducts short, 0.11-0.13 mm long, separate for most of their length, fused only at entrance. They bend laterally and enter receptacula at posterior inner side. Atrium with copulatory orifices situated in front of anterior border of epigynal protuberance (Figures 112-114), and in ventral view at posterior part of receptacula. Protuberance about half length of receptacula. Receptacula seminis 0.20-0.23 mm long and 0.13-0.14 mm wide.
Copulatory behaviour (Figures 116-119) Tidarren dentigerum follows the courtship and copulatory pattern of T. cuneolatum. Sexual cannibalism occurs post copulam. Only one out of two copulations was observed completely. Courtship lasted 6 and 7 min (n52). The female vibrated her body intensively and continuously and also plucked with her legs II. The male approached the female several times, palpated her, and installed a mating thread. At the starting point of this thread he performed a plucking sequence, which induced the female to assume the copulatory posture. Before successful insertion, the male made five attempts to insert his palp but did not achieve genital coupling. Insertion lasted 13 min (Figures 116, 117), then the female removed him from the epigynum ( Figure 118) and started to suck him out ( Figure 119). During copulation the male's legs became contracted, which obviously indicated his death ( Figure 117 versus Figure 116). After 4.5 h the female had completed her meal.

Distribution, habitat
Tidarren dentigerum has been collected from three localities in Yemen. At Ta'izz, T. dentigerum lived on small Lantana camara shrubs, up to 1 m high, at an agricultural research station. At Hammam 'Ali the species was found in mixed weeds at the margins of agricultural fields. Tidarren dentigerum seems to build its webs on the vegetation, whereas T. cuneolatum prefers brick walls, window frames, or large tree stems. At Ta'izz nearby buildings were closely inspected, but no webs were found.

Material examined
Only the types.

Etymology
Latin adjective gracile referring to the gracile appearance of this species. The suffix agrees in gender of the genus name.

Diagnosis
As in many Tidarren species from Yemen the abdomen ends in a dorsal tubercle (Figures 80, 122, 123, 132-135), and females lack a sternal knob. Tidarren gracile is the smallest representative in Yemen, female carapace width 0.6-0.8 mm. It differs by its pointed, hook-like and heavily sclerotized epigynal protuberance (Figures 128-131). Copulatory orifices separate, copulatory ducts short. The male is characterized by the  shape of the cymbium (Figures 124-127, 156), which is not projecting, and by the slender palpal tibia (Figure 127). Distal rim of tegulum with knob-like process (Figure 124), prolateral part of conductor with a few small humps and denticles ( Figure 125). Basal segments of palp and cymbium pale yellow.

Measurements (mm)
["/R, n55/7, minimum-maximum ( Sternum without posterior knob. Abdomen with dorsal tubercle (Figures 80, 122, 123), higher than long. Overall colouration in male and female similar; abdominal pattern in male less distinct. Carapace pale yellow brown with dark median band from eye region to centre; in females usually with broad dark margins, males not. Clypeus dark or greyish. Chelicerae light yellow brown, in females sometimes lightly suffused with grey. Gnathocoxae and labium pale yellow. Sternum usually uniformly pale yellow, sometimes with dark interrupted margins and some small dark spots. Legs pale yellow with dark patches and annulations; in females dark markings usually predominant. Female palp light yellow, with darkened distal part of tibia. Male palp pale yellow. Abdomen whitish to yellow brown with variable dark markings (Figures 122, 123, 132-135). Dorsum without longitudinal band, but showing two irregular paramedian dark bands at anterior sides, which meet at apex. Dark markings sometimes rather indistinct. Aboral area yellowish to whitish with white median stripe from apex to spinnerets and sometimes with two paramedian dark longitudinal markings. Epigaster yellow brown in female, greyish in male, book lung covers pale yellow. Venter in female with dark and white patches, in male dark with two paramedian spots between epigastric furrow and spinnerets. Spinnerets pale yellow to brown.

Copulatory behaviour (Figures 123, 132-135)
Tidarren gracile differs from the hitherto studied one-palped spiders in copulatory behaviour. Females start to consume their mates during copulation, without amputating the male gonopod (Figures 133, 134). After 10-20 min of such consummatory sexual act the male is removed from the epigynum and finally completely sucked out (Figures 123,  135). Males die during copulation. Three copulations were observed. During courtship the female shook and vibrated her body intensively, but did not pluck at all. Courtship lasted 2.0 min, 5.0 min and 7.8 min (n53). As typical of other Tidarren species, the male approached the female and installed the mating thread. At the starting point of this thread he performed a plucking sequence, which immediately induced the female to adopt the copulatory posture, perpendicular to the mating thread. Before insertion was successful, three, five and 15 insertion attempts were performed (n53). In one observation insertion lasted a very long time, 172.8 min. During the first 10 min of insertion the male pulsated his abdomen and along with this his prosoma became more and more shrivelled. Then he became motionless. After 10 min of insertion the female changed her inclined posture to a horizontal one so that the male's abdomen came to lie at her mouthparts (compare Figures 133 and 132). She then started to suck him out from the abdominal tubercle, while his palp remained inserted. In time with her sucking movements he was rhythmically see-sawed. In the 43rd minute of insertion the female tried to get rid of the male palp, but failed to loosen genital contact. She wrapped him with silk and continued consumption after changing her position. After 85 min of such synchronous sperm transfer and sexual cannibalism the male's abdomen was almost completely sucked out, its volume having become smaller than that of his prosoma (see Figure 134), whereas the volume of the palpal haematodochae did not decrease noticeably. Only after 150 min was haematodochal inflation observed to be smaller. During the last hour of insertion the female became more and more restless and sucked out the male from various parts of the body, changing from one leg to another and finally to the male's prosoma. She tried to pull off the male palp another two times, but did not succeed. Finally, after 172.5 min the remnants of the male were rapidly dislodged. Afterwards the female sucked out the male palp, which was still partially filled with haemolymph.
In the other two observations genital contact was much shorter, 12.5 and 21 min. The females started sexual cannibalism after 6.5 and 8.2 min, respectively, while palpal insertion was maintained. After removal the females continued to consume their mates for 4.5 and 5.5 h.

Natural history
Egg-sac white to grey, number of eggs low (20; n51). The sparse data on postembryonic development reveal fast growth. Males were found to be subadult 28 days and adult ca 35 days after egg-laying. Females show considerable longevity. One female was still alive 6 months after her maturation.

Distribution, habitat
Tidarren gracile is known only from the type locality in Yemen close to the Indian Ocean, from an agricultural area, where mainly fruit crops (mango, banana, guava), sorghum, cotton, sesame, and some vegetables are grown. Tidarren gracile was collected from a large shrub, which was densely covered with webs of a species of Pisauridae.

Material examined
Only the types.

Etymology
Proper noun in apposition and hence invariable. This species is named in honour of Prof. Dr Konrad Thaler (19 December 1940 to 11 July 2005) with whom B.K. had the good fortune to share a most pleasant, fascinating, and interesting period of life.

Somatic features, colouration (Figures 16, 141)
Sternum with posterior tubercle between coxae IV (Figure 16, arrow). Abdomen higher than long, with small dorsal tubercle ( Figure 141). Carapace uniformly brown to dark brown. Chelicerae, gnathocoxae, and labium light brown; sternum light brown with narrow dark margins or medium brown. Legs light yellow with extended dark patches and annulations. Pedipalps darker than legs. Posterior half of abdomen whitish to yellowish light brown, anterior half dark at sides. These different pigmentations join at tubercle. Dorsal band indistinct. In aboral whitish area a distinct white median stripe leads from tubercle straight to spinnerets (Figure 141b). Venter with dark and white patches. Spinnerets brown.

Epigynum, vulva (Figures 136-140)
Epigynal protuberance strongly protruding, ca 0.2 mm long, directed anteriorly towards posterior tubercle of sternum (Figures 16, 141a); more or less cone-shaped, in lateral view broad at base, but distinctly tapering at about half its length ( Figure 140). Posterior contour curved, anterior one only slightly arched. In aboral view protuberance clearly delimited from surrounding integument (Figure 139). Copulatory ducts fused at entrance for very short distance, altogether ca 0.13-0.14 mm long. They diverge posteriorly and enter receptacula posteriorly. Common copulatory orifice at anterior border of epigynal protuberance (Figure 137), in ventral view at posterior part of receptacula. Protuberance about as long as receptacula (Figure 137). Receptacula seminis 0.19 mm long and 0.12 wide.

Distribution, habitat
Tidarren konrad has been collected from two localities in Yemen in a habitat similar to that of T. dentigerum.

Material examined
Only the types.

Etymology
The species epithet is a noun in apposition, referring to its occurrence in Yemen, the ancient Sheba kingdom. The biblical Sheba (Arab: Saba') was the kingdom founded by the Sabaeans in the southwestern part of the Arabian Peninsula. Between the 10th and the 2nd century BC it was one of the most powerful and wealthy states of Arabia.

Diagnosis
Among the species from Yemen T. sheba shares with T. argo a rounded abdomen lacking a tubercle (Figures 77, 142). Female sternum also without knob. Tidarren sheba differs by the shape of the epigynal protuberance, which is inconspicuous, weakly sclerotized and scarcely projecting (Figures 142, 147-153). Copulatory ducts long, fused at entrance. Inner side of femur and tibia I of female speckled. The male is distinguished by the tapering cymbium (Figures 143-146, 155), which is not provided with teeth, but with ridges. Prolateral part of conductor with large spur-like process and several small denticles (Figures 144-146). Distal rim of tegulum forming two trianguloid processes (Figures 143, 145, 146). The embolus is rather long.

Somatic features, colouration (Figures 77, 142)
Sternum without tubercle. Abdomen rounded, without hump, about as high as long or only slightly higher. Colouration sexually dimorphic. Female carapace uniformly dark brown, in male pale yellow brown with dark margins and dark median area from eye region to centre. Chelicerae light brown, sometimes with dark longitudinal patches on inner margin. In female, sternum, gnathocoxae, and labium dark brown with a few small light spots. In male, sternum yellow brown, with dark margins and numerous dark spots; labium and gnathocoxae yellow brown. Legs light yellow brown with dark patches and annulations. In female, leg I with numerous small dark dots on inner side of femur and tibia. Female palp light yellow, with darkened tibia and tarsus. Male palp pale yellow. Abdomen whitish with dark markings of various extent. Dorsum lacks a longitudinal band, but shows two transverse, irregular dark bands, which may fuse at sides. Sides therefore dark in anterior half. Apical region with a small dark semilunar mark. White median stripe from apex to  spinnerets very inconspicuous or lacking. Aboral area yellowish to whitish with two to three irregular transverse bands which may fuse at sides in female. Abdominal pattern less distinct in male. Epigaster dark brown in both sexes, book lung covers pale yellow. Venter in female with dark and white patches, in male dark with two paramedian spots between epigastric furrow and spinnerets. Spinnerets brown to dark brown.
Male palp (Figures 143-146, 155) Tibia at distal rim almost as wide as bulbus (Figure 144), distinctly constricted at base. Palpal organ 0.24 mm wide. Cymbium rather inconspicuous and comparatively slender, about 1.2 times longer than wide (ca 0.3 mm long), distally tapering and projecting beyond bulbus, at end with some indistinct ridges and small scales. Distal rim of tegulum excavated and forming two trianguloid processes near embolar base; in the excavation between these processes lies the rudimentary distal haematodocha. Retrolateral, membranous part of conductor large, 0.2 mm long, and guides embolus. Prolateral, sclerotized part bears a large spur-like process and several small denticles. Embolar base circular, 0.06 mm in diameter. Distal part of embolus rather long, ca 0.3 mm.

Epigynum, vulva (Figures 147-153)
Epigynum embedded in dark epigastric region and therefore not clearly delimited. Epigynal protuberance inconspicuous, weakly sclerotized and scarcely projecting, less than 0.1 mm long; in ventral and lateral view it is a short, blunt, trianguloid process (Figures 147, 151, 153). In aboral view the protuberance gradually merges with the surrounding integument ( Figure 152). Tip of protuberance asymmetric in one female ( Figure 149). Copulatory ducts ca 0.19 mm long, fused for almost half their length. From this common part they diverge laterally and enter receptacula posteriorly. Copulatory orifice situated close to apex and below anterior border of epigynal protuberance (Figures 147, 150, 151, 153), in ventral view far behind receptacula. Protuberance about as long as receptacula or shorter (Figures 147-149). Receptacula seminis 0.14 mm long and 0.12 mm wide.

Distribution, habitat
Tidarren sheba is known from three localities in Yemen. It occupies a habitat similar to that of T. dentigerum.

Material examined
Only the types.

Etymology
The specific epithet is a noun in apposition and refers to the terra typica of the species (ancient Greek aethiops meaning burnt face).

Diagnosis
Tidarren aethiops is distinguished by its rounded and laterally incised epigynal protuberance (Figures 157-163). Position of copulatory orifices not at anterior border, but roughly at centre of epigynal protuberance. Male unknown.

Somatic features, colouration
Somatic features and colouration based on females from NMW (type specimens faded). Sternum without posterior tubercle. Abdomen higher than long, forming a dorsal hump ( Figure 172). Carapace with broad dark brown central area, broad dark margins, and interposed light paramedian bands. Chelicerae, labium, and gnathocoxae light brown. Legs and pedipalps light yellow with indistinct dark patches and annulations. Sternum either uniformly light yellow or with a few thin peripheral markings. Abdomen brownish, with dark pigmentation and white guanin markings; broad dorsal median area outlined by a pair of distinct white stripes, which branch laterally and end at tip of abdomen; lateral area behind stripes lighter than anterior area. From apex a distinct white median stripe leads straight to spinnerets, surrounded by a greyish zone and two dark longitudinal stripes. Epigynal protuberance flanked by dark epigastric region. Book lung covers light yellow. Venter dark, with a few white patches. Spinnerets light brown.

Distribution
Known from the Democratic Republic of Congo (Zaire).

Etymology
The specific epithet is derived from the Latin adjective afer, afra, afrum meaning African. The suffix agrees with the gender of the genus name.

Somatic features, colouration
Sternum without posterior small tubercle. Abdomen higher than long, ending in tubercle ( Figure 173). Carapace uniformly dark brown. Chelicerae brownish, gnathocoxae and labium light yellow or brown. Sternum light yellow, with distinct dark wavy margins, sometimes also with a few dark spots in centre. Pedipalps and legs light yellow with large dark annulations and patches. Abdomen brownish, with dark pigmentation and white guanin markings; whitish dorsal median area outlined by a pair of distinct white stripes, which diverge laterally; lateral area behind stripes lighter than anterior area. Aboral side with white median stripe from apex to spinnerets and transverse, upwardly curved dark pigmentation of various extent, sometimes forming two to three stripes. Epigastric region dark, book lung covers light yellow. Venter dark with white patches. Spinnerets light brown to brown.

Variation
The single female from Uganda differs by having its epigynal protuberance more slender and pointed (Figures 168-171

Material examined
Only the type.

Somatic features, colouration
Sternum with posterior small tubercle. Abdomen higher than long, forming a large dorsal hump ( Figure 174). Carapace yellow brown with dark median area from centre to eye region and with faint greyish margins. Chelicerae, gnathocoxae, and pedipalps yellow brown. Sternum pale yellow with dark margins. Legs light yellow with slightly darkened patches and annulations. Abdomen whitish to yellowish brown, large white guanin areas predominate; on dorsum an irregular, white, broad band ends before tip of abdomen; on the sides a pair of distinct white stripes curve anteriorly; area behind these stripes whitish, lighter than anterior area. From apex a clear white median stripe leads straight to spinnerets. This stripe surrounded by large whitish aboral area. Venter whitish. Spinnerets light brown. Epigynum, Epigynal protuberance distinctly protruding (Figure 196), ca 0.1 mm long, directed anteriorly, almost touching posterior end of sternum ( Figure 174); its shape triangular, evenly narrowing to pointed tip. In aboral view protuberance clearly delimited from surrounding integument (Figure 198). Copulatory ducts fused at entrance (Figure 197), ca 0.15-0.18 mm long. Common copulatory orifice situated below anterior border of epigynal protuberance, in ventral view level with posterior end of receptacula. Copulatory ducts run anteriorly for a short distance, then abruptly diverge posteriorly and laterally , and enter receptacula posteriorly. Receptacula seminis 0.14 mm long and 0.10 mm wide. Epigynal protuberance about 0.7 length of receptacula ( Figure 197).

Distribution
Tidarren circe is known only from the type locality in Namibia. The single female was collected together with numerous females of T. cuneolatum.

Remarks
Since Tidarren fornicatum was described from a juvenile female, it is not possible to consider this species appropriately. Its general appearance, especially the large abdominal hump, agrees with the female of T. scenicum. However, T. fornicatum cannot be proposed as a senior synonym of T. scenicum without knowledge of the genitalia. Adult specimens from the type region, Khartoum, would probably help to clarify this problem.

Material examined
Only the types.

Etymology
The species is named in honour of Dr Charles Griswold (Californian Academy of Sciences, San Francisco), who collected numerous Tidarren specimens included in this study.

Somatic features, colouration
Sternum without posterior small tubercle. Abdomen higher than long, ending in tubercle ( Figure 175). Carapace uniformly dark brown. Chelicerae, gnathocoxae, and labium light brown or brown. Sternum dark, with light yellow central area surrounded by several light yellow spots. Pedipalps and legs light yellow with large dark annulations and patches. Abdomen brown to dark brown, dorsal and aboral white line pattern as in T. afrum. Sides with a few white guanin patches. Epigastric region dark, book lung covers light yellow. Venter dark with two small white spots near spinnerets. Spinnerets brown. Epigynum, Epigynal protuberance distinctly protruding (Figures 175, 200), ca 0.13-0.14 mm long, directed anteriorly; its shape triangular and pointed, in lateral view tip slightly bent ( Figure 200). In aboral view protuberance clearly delimited from surrounding integument ( Figure 202). Copulatory ducts fused at entrance (Figure 201), short, only 0.07 mm long. Copulatory orifices at anterior border of epigynal protuberance, in ventral and dorsal view close to posterior end of receptacula. Copulatory ducts run anteriorly for a short distance and abruptly diverge laterally. They enter receptacula posteriorly, at inner side. Receptacula seminis 0.15 mm long and 0.10 mm wide. Epigynal protuberance almost as long as receptacula (Figure 201).

Distribution
Known only from the type locality in Cameroon.

Material examined
Only the type.

Etymology
The species name refers to the lance-like shape of the epigynum, from Latin adjective lanceolatus. The suffix agrees with the gender of the genus name.

Diagnosis
In T. lanceolatum the epigynum is extremely protruding (Figure 176, 204-207), very long related to body size. This species has the longest epigynal protuberance among the representatives from mainland Africa. Male unknown.

Somatic features, colouration
Sternum without posterior tubercle. Abdomen higher than long, forming a large dorsal hump ( Figure 176). Carapace dark brown, central area slightly lighter with diffuse radial stripes. Chelicerae and gnathocoxae light brown. Sternum light brown with thin dark margins and small dark spots. Legs and pedipalps light yellow with dark patches and annulations. Abdomen whitish to greyish brown, with white guanin areas; on dorsum a pair of distinct white stripes outline an irregular, broad band, branching laterally and ending before tip of abdomen; area behind these stripes lighter than anterior area. From apex a clear white median stripe leads straight to spinnerets. Aboral area of abdomen whitish with two transverse, strongly upwardly bent dark stripes. Venter dark and white speckled. Spinnerets light brown.

Distribution
Tidarren lanceolatum is known only from the type region, Democratic Republic of Congo, without exact locality.

Other material examined
Democratic Republic of Congo (Zaire): R SMF 37459 and subadult " SMF 37592 (shrivelled and faded), imported with bananas, without exact locality.   Schmidt (1957). The following redescription is based mainly on non-type material, as this female is intact. The holotype female is shrivelled and faded; the epiygnum, which is preserved as a microscopic slide, is partially destroyed, epigynal protuberance and copulatory ducts are broken off and missing, only receptacles and fertilization ducts present. General appearance and dimensions are similar in the two females.

Diagnosis
Tidarren levii is a small-sized species with abdominal tubercle being very indistinct. Its epigynal protuberance is broad and pointed, clearly delimited from surrounding integument (Figures 208-211) and well sclerotized. Copulatory ducts diverge posteriorly. Male unknown.

Somatic features, colouration
Sternum without posterior tubercle. Abdomen higher than long, more or less rounded, tubercle very indistinct. Carapace uniformly dark brown. Chelicerae light brown, suffused with grey. Labium and gnathocoxae light brown. Sternum dark grey with several light spots. Legs and pedipalps pale yellow with dark patches and annulations. Coxae uniformly pale yellow. Abdomen dark, with interrupted white line pattern. White dots on dorsum are probably remnants of former outlines of a dorsal band. Sides with white arched stripe, aboral region with white stripe from apex to spinnerets. Venter dark with two pairs of paramedian white spots. Spinnerets brown.
Known only from an import from tropical Africa (Democratic Republic of Congo) without exact locality (Schmidt 1957).

Etymology
The specific epithet is derived from the Latin adjective perplexus meaning obscure. The name refers to the taxonomic situation among the inscrutable number of African Tidarren species. The suffix agrees with the gender of the genus name.

Diagnosis
According to genital characters T. perplexum closely resembles T. levii (Figures 212-215 versus Figures 208-211), but is differentiated by its ca 1.5 times larger size and by the presence of a distinct abdominal tubercle ( Figure 177). In contrast to T. levii the protuberance is not delimited from the surrounding integument in aboral view ( Figure 214) and is less sclerotized. Male unknown.

Epigynum, vulva (Figures 212-215)
Epigynal protuberance triangular, short, and pointed (Figures 177, 212), 0.08 mm long, 0.09 mm broad at widest part ( Figure 213). Anterior median part of protuberance less sclerotized. In side view, tip of protuberance slightly recurved, anterior contour straight, posterior one only slightly arched. In aboral view protuberance not delimited from surrounding integument (Figure 214), its sclerotization diffusing gradually. Copulatory orifices close to anterior border of epigynal protuberance, in ventral and dorsal view level with posterior half of receptacula; in aboral view close to end of protuberance. Copulatory ducts separate (Figure 213, 215), ca 0.07 mm long. They diverge laterally and enter receptacula at posterior inner side. Receptacula seminis 0.12 mm long and ca 0.1 mm wide. Epigynal protuberance about 0.7 length of receptacula ( Figure 213). The genital organ of the females from the Democratic Republic of Congo differs slightly in the shape of the epigynal protuberance. In side view, the protuberance is slightly more slender, the posterior contour less arched, and its tip is scarcely sclerotized. The copulatory orifices appear to be wider than in the types. Aboral view, position of copulatory orifices, course of the ducts, and overall dimensions are similar: epigynal protuberance 0.08 mm long and 0.07 mm broad at widest part. Copulatory ducts separate, 0.08 mm long. Receptacula seminis 0.12 mm long and 0.10 mm wide. Epigynal protuberance about 0.6 length of receptacula.

Geographic variation
The specimens from the Democratic Republic of Congo agree with the type females in general appearance, colour, pattern, and dimensions. Slight differences were found in the shape of the epigynal protuberance, see above. Without knowledge of males it is not yet possible to judge this minute variation.

Distribution
Known from the type locality in Cameroon and from the Democratic Republic of Congo without exact locality.
Somatic features, colouration (Figures 178-180) [Based on females from Cote d'Ivoire, Guinea-Bissau, and South Africa.] Sternum without posterior tubercle. Abdomen higher than long, forming a large dorsal hump (Figures 178,  180), so that the species name T. turrigerum chosen by Simon (1899) would be apposite. Carapace light yellow with broad dark margins and dark brown median band (Figure 179), which branches laterally at centre of carapace. Chelicerae light brown, labium and gnathocoxae light yellow. Sternum light yellow with dark patches at margins and some median spots. Legs and pedipalps light yellow with dark patches and annulations, especially femur, tibia, and metatarsus I covered with numerous fine dots. Abdomen brownish, with white guanin markings and some dark pigmentation; broad and light dorsal median area outlined by distinct white stripes, which branch laterally and end shortly before tip of abdomen; surrounding lateral area darker. Aboral field whitish, partially overlain by dark pigmentation, with distinct white median stripe from apex to spinnerets. Epigastric region light, book lung covers light yellow. Venter uniformly light brown or with a few white patches. Spinnerets light brown. The type of T. guineense is largely light yellow and perhaps partially faded (Figure 180), though Simon (1907) mentioned the light yellow colouration.

Distribution (Figure 259c)
Tidarren scenicum, although known from five females only, appears to be widespread in Africa.

Synonymy
Both Tidarren turrigerum Simon, 1899 andT. guineense Simon, 1907 correspond well with the type of T. scenicum Thorell, 1899 in body dimensions, shape of abdomen and of epigynum, and are therefore considered as synonyms. Tidarren fornicatum was described from a juvenile female from Sudan (sub Theridion fornicatum; Simon 1884). General appearance, especially the large abdominal hump, agrees with the female of T. scenicum. Nevertheless, this species should not be proposed as a senior synonym without knowledge of the genitalia. Otherwise, T. fornicatum would have priority. Further material from the type region would probably clarify this problem. Theridion turrigerum Simon and T. scenicum Thorell are clearly synonyms, but were both described in 1899. The exact date of publication should decide on the priority. Thorell's paper was almost certainly printed in 1899. It is the first part of Bihang till Kongliga Svenska Vetenskaps-Akademiens Handlingar, Volume 25, Section IV, Zoology. This volume contains six independently paginated papers, each with a separate title page, papers 1-3 having the printing year 1899 and papers 4-6 1900 (T. Kronestedt, in litt.). The ''communicatum'' date of the reprint, 8 February 1899, has nothing to do with the printing date but indicates that the paper was presented to the academy early in 1899 and thus was almost certainly printed in the same year (T. Kronestedt, in litt.). The footnote on the reprint of Simon's work, ''Imprimerie nationale-Janvier 1900'' indicates that the actual date of publication was later. For this reason, T. scenicum has to be considered the valid name. Localization of the type locality of T. turrigerum appears to be arguable. According to Simon (1899) it was collected from ''Soudan français'', which would be partially consistent with Mali (Platnick 2006). Yet, when trying to locate the ''frontier post'' Zo at the river Cavally, it appears more likely to match Cote d'Ivoire. Simon (1907) indicated a close relationship between T. turrigerum and T. scenicum, but his T. scenicum now turns out to be a female of T. cuneolatum (from Guinea-Bissau, MNHN AR 2370), see above. Therefore, it is not surprising that he distinguished T. turrigerum by its larger size, although being as large as the proper T. scenicum, and by the obtuse epigynal shape.

Etymology
The species epithet is dedicated to the collectors Drs Darell and Suzanne Ubick (Californian Academy of Sciences, San Francisco).

Diagnosis
Tidarren ubickorum is the only species from mainland Africa hitherto known, which has a rounded abdomen (Figures 181,182,193). Its epigynal protuberance is slender in side view and rather small (Figures 222-225   wide. Sternum about as long as wide, 0.45-0.58. Leg formula 1423, see Table XXII. Somatic features, colouration (Figures 181,182,193) Abdomen without tubercle. Sternum without posterior tubercle, projecting between coxae IV and separating coxae IV by one diameter. Carapace uniformly dark brown, in one female with yellow posterior and lateral parts. Sternum dark brown with irregular light median area or stripe. Chelicerae light brown with dark longitudinal patches at margins. Legs and palps light yellow with dark patches and annulations. Abdomen dorsally with irregular, dark median band, which ends at the highest point ( Figure 193). This band is surrounded by a zone of white guanin inclusions, in which dark spots are also embedded. Each side is flanked by a large dark patch within a light intermediate area. Aboral part of abdomen with clear white stripe from highest point to spinnerets, on a background of irregular dark pigmentation. Spinnerets brown, surrounded by a dark ring. Book lung covers yellow, epigaster dark brown, venter with white mark between epigastric furrow and spinnerets. Abdominal markings in one female faint. Epigynum , Epigynal protuberance in side view slender and finger-like (Figures 181, 222), 0.09 mm long, rather small and tapered. In aboral view protuberance not clearly delimited, gradually merging with surrounding integument (Figure 224). Copulatory orifices close together, apparently separate ( Figure 223, 225), situated anterior to epigynal protuberance, in ventral view close to posterior end of receptacula. Copulatory ducts separate, short, 0.07 mm, diverging laterally and entering receptacula posteriorly. Receptacula seminis 0.13 mm long and 0.10 mm wide. Epigynal protuberance about half as long as receptacula (in ventral view, Figure 223).

Geographic variation
Body dimensions of the female from Zimbabwe are slightly larger than those from South Africa (in mm: total length 3.21, carapace length 1.09, width 0.98), and legs are considerably longer (in mm: length femur I 2.01, tibia I 1.19).

Distribution
Known from the type locality Mpumalanga in South Africa and from a suburban garden in Zimbabwe.

Material examined
Only the types.

Etymology
Noun in apposition referring to the type locality.

Distribution
Known only from the type locality Usambara Mtns at Mazumbai. The species was collected together with Tidarren cuneolatum.

Material examined
Only the type.

Etymology
The specific epithet is a diminutive and indicates that the species is apart from the others, and also refers to the APART-grant from the Austrian Academy of Sciences (Austrian Programme for Advanced Research and Technology), by which B.K. was supported.
Somatic features, colouration (Figures 187,194) Sternum without posterior tubercle. Abdomen higher than long, ending in tubercle ( Figure 187). Carapace dark brown. Chelicerae brown, with dark longitudinal stripes at  margins. Sternum, gnathocoxae, and labium dark brown. Legs and palps light yellow with dark patches and annulations. Abdomen dark, except for some white stripes and light brown patches; without dorsal longitudinal median band or longitudinal white outlines ( Figure 194); instead, two short white transverse bars. At sides a distinct white stripe arches downwards; anterior lateral area darker than posterior one. Aboral area of abdomen with white median stripe from apex to spinnerets, surrounded and partially overlain by dark and reddish brown pigmentation. Venter and epigaster dark, book lung covers light. Two lateral white spots next to anterior spinnerets. Spinnerets brown.

Distribution
Known only from the type locality in Madagascar.

Etymology
The species name is derived from ancient Greek dasys, meaning hairy or villous, and from glossa, meaning tongue; compound adjective and noun in apposition, hence invariable. It refers to the tongue-shaped epigynum, which bears numerous hairs on its posterior side.

Diagnosis
Tidarren dasyglossa is the only species hitherto known from Madagascar with a small tubercle on posterior end of sternum. As in T. horaki the abdomen is rounded. The species has a characteristic tongue-shaped, hairy epigynal protuberance, which occupies most of the epigastric region (Figures 188,(234)(235)(236)(237). Copulatory ducts are fused at entrance and run posteriorly in parallel. Adult male unknown. It is expected to have a large palp.   Epigynal protuberance more than twice as long as receptacula ( Figure 235).

Distribution
Tidarren dasyglossa is known from three localities in Madagascar.

Matching of sexes
One subadult male is assigned to T. dasyglossa owing to its well-developed palp (ca 0.6 mm long, see Figure 189, 238), which corresponds to the large female genital organ. The male has an indistinct posterior tubercle on the sternum and on the abdomen (Figure 189).

Material examined
Only the type.

Etymology
The species name refers to the ephemeral presence of males in all Tidarren species, since they die with copulation, and also to the possible ephemeral state of this species, as the male may turn out to belong to either T. apartiolum n. sp. or T. horaki n. sp.

Diagnosis
The male is distinguished by the slender, tapering cymbium, which lacks distal lobes ( Figure 239-242), but bears apical teeth. Prolateral part of conductor with large sickleshaped process and several conspicuous teeth arranged like a crown ( Figure 240). Distal rim of tegulum without process. Female unknown. It cannot be excluded that T. ephemerum is the male of either T. apartiolum or T. horaki.

Somatic features, colouration
Sternum without tubercle. Abdomen higher than long, ending in tubercle. Carapace pale yellow with dark median band from clypeus to eye region and to centre. Chelicerae light yellow with greyish inner margin. Gnathocoxae and labium pale yellow. Sternum pale yellow, with broad dark wavy margins and some small dark spots. Legs pale yellow with dark patches and annulations, coxae pale yellow. Palp pale yellow. Abdomen whitish to yellow brown with dark markings on anterior half. Aboral area whitish with indistinct white median stripe. Epigaster greyish, book lung covers pale yellow. Venter dark with two paramedian spots between epigastric furrow and spinnerets. Spinnerets pale yellow at base, distally brown.

Male palp (Figures 239-242)
Tibia at widest part ca 0.6 width of palpal organ. Distal projection of tibia lobe-like and slender, ca 0.3 width of palpal organ ( Figure 240). Palpal organ 0.20 mm wide. Cymbium inconspicuous and less modified than in the other species, apparently without distal lobes, but tapering ( Figure 241), with several apical teeth; about 1.3 times longer than wide, projecting slightly beyond bulbus. Distal rim of tegulum without process. Retrolateral, membranous part of conductor 0.08 mm long, its tip pointed and hyaline, with furrow guiding embolus; prolateral, sclerotized part with large sickle-shaped process and several conspicuous teeth arranged like a crown (dorsal view, Figure 240) or like a claw (side view, Figure 239). Embolar base suboval, 0.06 long and 0.04 mm wide, with two small basal lobes. Distal part of embolus 0.11 mm long.

Distribution
Known only from the type locality in Madagascar.

Material examined
Only the types.

Etymology
This species is dedicated to our enthusiastic fellow arachnologist Dr Peter Horak (Graz).

Measurements (mm)
[R, n55, minimum-maximum (mean Sternum without posterior tubercle. Abdomen higher than long, more or less rounded, tubercle inconspicuous (Figures 184, 185). Carapace uniformly dark brown. Chelicerae light brown, suffused with greyish longitudinal stripes at margins. Gnathocoxae and labium dark brown, lighter at margins. Sternum uniformly dark. Legs and palps light yellow with large dark patches and annulations. Coxae light yellow, distally with large dark patches or at least darkened. Abdomen dark in most specimens, with several white spots ( Figure 195). In one well-nourished female with numerous white and dark spots on light brown background. Dorsum without longitudinal band, but with four white paramedian patches; from posterior one an indistinct white line turns laterally. Aboral area dark (in one female dark and white speckled), with interrupted white median stripe from apex of tubercle to spinnerets. Epigastric region strongly protruding. Epigaster dark to greyish, book lung covers light. Venter dark, with two white paramedian spots. Spinnerets dark brown.

Epigynum, vulva (Figures 243-246)
Entire epigastric region swollen, rounded and strongly projecting together with epigynal protuberance, ca 0.3 mm beyond venter (Figures 184, 185, 243-246), epigynum itself 0.12-0.14 mm long in side view. Protuberance triangular, directed anteriorly, in ventral view ca 0.2 mm long, with gradually tapering outlines; posterior, aboral side clearly delimited from surrounding integument, but surrounding aboral region also sclerotized ( Figure 245). Copulatory orifice at anterior border of epigynal protuberance, in ventral view closer to posterior end of receptacula. Copulatory ducts separate, though with common orifice (Figures 244, 245), ca 0.1 mm long; they immediately diverge posteriorly and laterally and enter receptacula at posterior inner side. Receptacula seminis 0.19 mm long and 0.12 mm wide. Epigynal protuberance about as long as receptacula (in ventral view, Figure 244). Female (Figures 186, 192). Carapace yellow with dark median area from eye region to centre, sometimes also with dark margins. Chelicerae yellow, rarely suffused with greyish longitudinal stripes at margins. Gnathocoxae and labium pale yellow. Sternum with variable pattern, from pale yellow with indistinct greyish margins and undulations to distinct pattern, covering posterior half of sternum, or sternum even mainly dark with several light spots. Legs and palps light yellow with dark patches and annulations of various extent. Coxae light yellow. Abdomen whitish to yellowish, with white line pattern and some dark pigmentation. Dorsum with broad whitish longitudinal band, ending on tubercle and surrounded by dark pigmentation. One short anterior and one longer posterior white line curve downwards laterally. Aboral area yellowish, with clear white median stripe from apex to spinnerets and one or two interrupted dark transverse bands. Epigastric region yellowish, epigynal area sometimes darkened, book lung covers pale yellow. Venter yellowish, sometimes with a few dark and white spots. Spinnerets light brown.
Male (Figures 190,191 Figures 190, 191) and a smaller inner one close to tip of membranous part; their distal surfaces bear some warts. Embolar base circular, 0.05 mm in diameter, with two small basal lobes. Distal part of embolus 0.2 mm long.

Matching of sexes
It cannot be definitely assured that the male is conspecific with the females, as different species may occur together in the same region. However, the single male was collected from exactly the same locality as two females. Also the yellowish colouration of the carapace agrees with that of the females.

Distribution (Figure 259c)
Tidarren obtusum occurs in several places on Madagascar.
Tidarren sp. (Figure 259a,  This male specimen eluded a clear assignment to other Madagasy species, as its male palp is expanded and its details are hard to compare (Figure 259a, b). However, it does not agree with T. ephemerum. Shape of conductor also differs from T. obtusum. Probably, it represents the male of another species.

Material examined
Costa Rica: 2R, 1juv, NMW Inv. No. 720, San José, La Caja, 9u589N, 84u079W, leg. E. Schmidt, 1930, det. Reimoser (sub Theridion fordulum). Venezuela: 2R, MNHN 11236 AR 10708, Carabobo, La Cumbre, leg. Simon, 1888, det. Levi, 1960. 6R 2sadR, MNHN 11664, without AR number, La Guaira, leg. E. Simon, det. Levi, 1960 (sub T. fordum). 1R, MNHN 11299 AR 10774, Aragua, Tovar, leg. Simon, 1888, det. Levi, 1960. 8R 5juv, MNHN AR 2411, Caracas, in vial of ''Theridium taeniatum'' together with several Achaearanea females, which at least partially appear to represent Achaearanea taeniata    Female (Figure 260a). Carapace yellow with broad dark margins and with dark median band from eye region to centre, tapering posteriorly and sometimes extending laterally. Clypeus greyish. Chelicerae, gnathocoxae, and labium light brown or yellow. Sternum light brown to yellow, with thin dark margin and frequently with distinct dark stripes opposite each coxa; sometimes with numerous fine dots instead. Legs and palps light brown to yellow with dark patches and annulations of various extent. Numerous fine dots present especially on femora I-II and tibia I-II. Coxae light yellow, only in few specimens coxae I and IV with a greyish spot. Abdomen whitish to light brown, with white line pattern and variable dark pigmentation. On dorsum two dark, rounded median patches just before apex, outlined by clear white lines; from these two white stripes curve to each side. Further white and dark dots on anterior part of dorsum. Sides sometimes with large areas of dark pigmentation. In some specimens this abdominal pattern is very indistinct or even almost absent. Aboral area whitish to brownish with clear white median stripe from apex to spinnerets and with two or three dark transverse, upwardly curved bands. Epigastric region greyish, region in front of epigynal protruberance yellowish, so that receptacula are visible through integument. Book lung covers yellow. Venter with dark and white patches. Spinnerets light brown. Male (Figure 260b). Carapace pale yellow with dark median area from eye region to centre, gradually tapering, and sometimes also with dark margins. Clypeus greyish. Chelicerae yellow, often suffused with grey. Gnathocoxae and labium yellow. Sternum yellow with dark undulated margins as in female. Legs pale yellow with dark patches and annulations of various extent. Coxae uniformly pale yellow. Male palp uniformly pale yellow. Abdomen yellowish to whitish. Dark pigmentation, if present at all, on anterior half of dorsum and sides. Posterior half light. Epigaster greyish or dark, book lung covers yellow. Venter dark, with two small white paramedian spots. Further white spots surround spinnerets. Spinnerets light brown suffused with grey.
Male (Figures 1b, 286-289). Carapace light yellow to reddish brown with dark margins and dark median area from eye region to centre, tapering posteriorly. Clypeus light greyish. Chelicerae, gnathocoxae, and labium uniformly light brown. Sternum light brown, with fine dark margins. Legs uniformly yellow to reddish brown, sometimes suffused with grey. Male palp yellow brown from coxa to tibia. Distal parts of cymbium reddish brown. Abdomen uniformly reddish brown, with indistinct greyish longitudinal band, ending in apical region. Spinnerets brown.
Haematodochae not modified (Figures 1b, 287). Palps and legs yellowish (sometimes reddish) with dark patches and annulations of various extent. Coxae yellow. Abdomen with whitish dorsal median band and posterior white lines, which branch laterally. Aboral area with distinct white stripe from apex to spinnerets. Epigastric region and venter of variable colouration according to degree of overall pigmentation, book lung covers light brown. Epigynal protuberance heavily sclerotized in contrast to surrounding integument.
Male ( Figure 293, 305). Carapace, chelicerae, gnathocoxae, labium, and sternum uniformly dark or reddish brown. Legs light to reddish brown, with some dark patches and annulations, sometimes largely suffused with grey. Distal parts of male palp dark to reddish brown. Abdomen usually dark or reddish brown, with a few white patches on dorsum, aboral area whitish with white stripe from apex to spinnerets. Epigaster dark to reddish brown, book lung covers yellow. Venter dark to reddish brown with two small white paramedian patches. Spinnerets dark to reddish brown. Figure 310. Echinotheridion gibberosum (Kulczynski) from Tenerife. Functional contact of male palp after emasculation. Palp comes to rest close to female coxal spurs, but without direct contact. Male palp (Figures 10,(295)(296)(297)(298)304,308,309) Tibia ca 0.5 width of palpal organ at widest part (dorsal view); its base ca 0.3 width of distal rim ( Figure 296). Male palp pincer-shaped, owing to strongly protruding cymbium and conductor (Figures 10, 295, 308, 309). Cymbium ends in a hook-like, apical process and a hairy, narrow lobe, bearing tarsal organ and adjoining broad furrow of conductor ( Figure 297, 309 arrow). Its apical margin is strongly incised (Figure 297). On retrolateral side another, less distinct, hairy projection (Figures 297, 298). Bulbus ca 0.4 mm long (including conductor) and 0.33 mm wide. Subtegulum and tegulum appear to be partially fused, though hard to recognize, connecting median haematodocha presumably reduced or at least inconspicuous. Distal rim of tegulum sclerotized and incised, with two small rounded, but indistinct projections (Figures 295, 296). Conductor strongly developed, curved, forming a lamella with broad, shallow furrow, its inside curved surface covered with tiny scales (Figures 295-298). Base of embolus slender and apparently lacking the two basal lobes present in all Tidarren species (Figure 296), gradually passing into distal part.
Copulation involves single usage of the male palp and regularly ends in exhaustion of the male, in emasculation and subsequent sexual cannibalism (Knoflach 2002a). Copulation proceeds via a mating thread. Unlike T. argo, emasculation does not take place immediately after application of the male palp. At first, there is a normal insertion for about 4 min on average, during which both partners are completely motionless (Figures 305, 306). The male seems to be dead, the female is cataleptic. When she awakes from her cataleptic state, she entangles the male and starts to turn around in circles. Emasculation takes more than 3 min, and the female has to try several times until she succeeds in breaking off the palp. The breaking point of the palp is usually between the tibia and tarsus, as in T. argo. Holdfast structures of the palp are sclerites, there is no haematodochal modification. These palpal sclerites clasp the epigynum like forceps, the hook-like apex of the cymbium locks on the membranous folds of the posterior declivity of the epigynum, and the conductor locks in front of the epigynal protruberance (Figures 307-309). The male palp remains fastened to the epigynum for 5 h on average (up to 11 h) while the female is occupied with mate consumption (Figure 307). The function of the coxal spurs on legs IV (Figures 23, 24) of the female remains uncertain, as they seem to not be necessary for genital coupling ( Figure 310). Remarkably, the first observation on the copulatory behaviour of E. gibberosum was mistaken as a heterospecific copulation between two genera (Schmidt 1980).
Natural history (Figures 3, 4, 290, 291) In Tenerife, the webs of Echinotheridion gibberosum were found at a height of about 1-2 m on tree trunks and between large branches. The spiders are concealed in their large retreats, which consist mainly of debris. The egg-sac has a parchment-like light brown envelope (Figures 290,291). In some of the retreats cocoons of the pirate spider Ero (Mimetidae) were also found, which may indicate possible predation by this araneophagic spider. For palp amputation (Figures 3, 4) see generic diagnosis and Knoflach (2002a).

Distribution
Echinotheridion gibberosum is known only from laurel forests or mixed pine and laurel forests in Madeira and Canary Islands and appears to be the only representative of the genus in the Old World.