MYSMENIDAE COMPARATIVE

Lopardo, Lara & Hormiga, Gustavo, 2015, Out of the twilight zone: phylogeny and evolutionary morphology of the orb-weaving spider family Mysmenidae, with a focus on spinneret spigot morphology in symphytognathoids (Araneae, Araneoidea), Zoological Journal of the Linnean Society 173 (3), pp. 527-786 : 658-713

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https://doi.org/ 10.1111/zoj.12199

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MYSMENIDAE COMPARATIVE
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MYSMENIDAE COMPARATIVE   MORPHOLOGY

Male palp (refer to characters 151–261)

The diversity and sclerite homologies of mysmenid male palps are only superficially understood. Mysmenidae   have been recognized and even diagnosed by the generalized shape of the cymbium, described as ‘apically twisted and with lobes’ ( Platnick & Shadab, 1978; Brignoli, 1980; Coddington, 1990; Wunderlich, 1995; Griswold et al., 1998; Schütt, 2003). Despite the fact that several relatively modern descriptions of mysmenids have included detailed illustrations of genitalic morphology (e.g. Kraus, 1967; Thaler, 1975, 1995; Saaristo, 1978; Baert & Maelfait, 1983; Baert, 1984a, 1990; Lin & Li, 2008; Miller et al., 2009), most mysmenid species remain poorly described. The details of the palp mor- phology are also insufficiently studied, especially in terms of explicit hypotheses of homology. For example, it has been suggested that a tegular conductor, median apophysis, and paracymbium are absent ( Coddington, 1990; Griswold et al., 1998; Miller et al., 2009); however, a paracymbium had previously been identified in Mysmenidae ( Kraus, 1967)   . The mysmenid male palp appears highly complex and morphologically variable, and in most species it is greatly translucent, so that the cymbium, conductor, and other sclerites are difficult to distinguish and to delineate precisely under light microscopy. In summary, mysmenid male palps have distinct cymbial structures, including a paracymbium, and can also have a tegular conductor. A median apophysis or any other tegular sclerites are lacking, however. As in the details of the differ- ent respiratory organs in Mysmenidae   , the diversity of palpal structures is great within the family, although each particular arrangement seems characteristic at the genus or sometimes subfamily level (this is simply a consequence of how taxonomists have circumscribed higher taxa in Mysmenidae   ). In the sections below, we address the large diversity of mysmenid palpal morphology.

Palpal femur, patella and tibia

Mysmenids lack any modifications on the palpal femur and patella. Conversely, and across symphytognathoids, varying shapes of the male palpal tibia can be found. A distally broad tibia (i.e. wider distally, usually more than two times its basal width) may be symplesiomorphic for symphytognathoids, as it also occurs in Theridiidae   . Distally broad tibiae occur in Theridiosomatidae   , Synaphridae   , and most Mysmenidae   ( Figs 4A View Figure 4 , 17D View Figure 17 , 28A View Figure 28 , 30E View Figure 30 , 32E View Figure 32 , 38A View Figure 38 , 42A View Figure 42 , 47B View Figure 47 , 63C View Figure 63 ). Within symphytognathoids, a flat tibia (i.e. flattened from the base, and usually with irregular, not circular, distal section outline) is synapomorphic for the clade comprising Symphytognathidae   plus Anapidae   ( Figs 91F View Figure 91 , 95C, D View Figure 95 , 102C View Figure 102 ). Within Mysmenidae   , broad male palpal tibiae are widespread, occurring in Trogloneta   , Isela   , and all Mysmeninae   . Moreover, a cylindrical tibia (distal width similar to or less than two times proximal width) is synapomorphic for Maymena   , and convergently present in Comaroma   and Tasmanapis   ( Figs 10A View Figure 10 , 81D View Figure 81 , 98B,C View Figure 98 ). A globose tibia is a synapomorphy of Mysmenopsis   ( Figs 55A, C View Figure 55 , 58A View Figure 58 , 60A View Figure 60 ).

Most mysmenid palpal tibiae are small, shorter than cymbium, as in other symphytognathoids ( Figs 10A View Figure 10 , 27B View Figure 27 , 30D View Figure 30 , 47A, B View Figure 47 , 63B, C View Figure 63 , 71D View Figure 71 , 106C View Figure 106 , 110E View Figure 110 ). A large tibia is synapomorphic for Mysmenopsinae   ( Figs 1A View Figure 1 , 4A View Figure 4 , 55A, C View Figure 55 ). In addition, no distinct tibial processes occur in Mysmenidae   , except for a prolateral extension in Mysmeniola   ( Fig. 134D View Figure 134 ; see also Thaler, 1995: figs 5, 7) and an apical ventral (sometimes ventroretrolateral) excavation usually bearing spurs in Mysmenopsis   ( Figs 53A View Figure 53 , 58A View Figure 58 , 60A View Figure 60 ). Mysmenopsines have modified setae distally on the tibiae, such as spurs in Mysmenopsis   (as mentioned above, Figs 53E View Figure 53 , 55H View Figure 55 , 58E View Figure 58 , 60B View Figure 60 ), or spine-like, strong setae in Isela   ( Figs 1A, B View Figure 1 , 4A, E View Figure 4 ). Mysmenid tibial rim setae are longer than remaining tibial setae, and are arranged distally in one or two rows ( Figs 4E View Figure 4 , 10D View Figure 10 , 18A View Figure 18 , 32E View Figure 32 , 36D View Figure 36 , 42B View Figure 42 , 45C View Figure 45 , 53B View Figure 53 , 55C View Figure 55 ), except in Trogloneta   and a few other mysmenids where the setae are equally short and dispersed in an irregular conformation ( Figs 63C View Figure 63 , 66A View Figure 66 ). Cymbium: general morphology

(see Fig. 126 View Figure 126 for reference)

The size of the cymbium and bulb (relative to the size of the carapace, in lateral view) varies across mysmenid taxa. Although small male palps are common in adult spiders, medium-sized palps are widespread within symphytognathoids, including mysmenids (i.e. the cymbium-bulb complex is about half the size of the carapace; Figs 27F View Figure 27 , 66A View Figure 66 , 141K View Figure 141 ). The cymbium bulb becomes secondarily small (about one-fifth the size of the carapace) in Maymena mayana   and mysmenopsines, however ( Figs 2B View Figure 2 , 140E View Figure 140 , 141D View Figure 141 ). A very large cymbium- bulb complex (i.e. as large as the prosoma) has evolved independently in some distal clades within Microdipoena   and Mysmena   , including Mysmena leucoplagiata   ( Figs 19B View Figure 19 , 142A View Figure 142 ), in Theridiosomatidae   , and the synaphrid genus Cepheia   . Superficially, there appears to be a tendency towards an increase in palpal size, which suggests a correlation with the reduction of body size in symphytognathoids; however, most symphytognathoids are equally minute in body size, regardless of their familial placement or size of the palp. For example, most Microdipoena species   (and also theridiosomatids) with huge palps are as small as any other mysmenine or even larger than members of Symphytognathidae   , which possess medium-sized palps. Taphiassa   , a small micropholcommatine anapid, also has small palps.

The cymbium in most mysmenids is uniquely oriented ventrally or prolatero-ventrally in the palp ( Figs 38A,B View Figure 38 , 42B View Figure 42 , 66A View Figure 66 ). The cymbium evolved independently to a prolateral position in Mysmenopsis   , Mysmeniola   , and the mysmenine MYSM-023-MAD ( Fig. 50B View Figure 50 ), or to a retrolateral–dorsal position in some Maymena species   ( Fig. 15B View Figure 15 ).

In some symphytognathoids the cymbium is sometimes modified relative to the typical cymbium of most araneoids, which is scoop-shaped and round to oval in dorsal view (e.g. Griswold et al., 1998: figs 16A, 18F). This is especially so in mysmenids. Without taking into account any other cymbial structures (such as cymbial conductors, apophyses, paracymbia, expansions, etc.; see comments below), an oval cymbium is plesiomorphic in Mysmenidae   ( Figs 10A View Figure 10 , 14A View Figure 14 ), and it has independently evolved in MYSM-005-ARG ( Mysmena   ; Fig. 28B View Figure 28 ). A cymbium as long as wide occurs in mysmenopsines and independently in a clade comprising most mysmenines ( Figs 1A View Figure 1 , 4C View Figure 4 , 22F View Figure 22 , 30D View Figure 30 , 41B View Figure 41 , 55B View Figure 55 ), whereas a distinctly flat and tapering cymbium is unique to Trogloneta   ( Fig. 63C View Figure 63 ; convergent in some symphytognathids).

The cymbium of mysmenids is greatly modified when compared with other symphytognathoids. There seems to be a pattern of shared modifications, such as grooves, processes, and modified setae, which can be recognized in the cymbium ( Fig. 126 View Figure 126 ). Not all mysmenids have all the cymbial structures that we describe. Furthermore, different combinations of these features usually vary among (and are distinctive of) each genus. A summary of these cymbial features is presented below (refer to Fig. 126 View Figure 126 ).

Primary and secondary cymbial conductors (CyC1 and CyC2): Up to two apical grooves, which seemingly interact with the distal portion of the embolus, can occur in mysmenid cymbia. Both structures are here considered cymbial conductors.

The ‘primary cymbial conductor’ (CyC1) is located internally (i.e. closer to the bulb; Figs 4G View Figure 4 , 10C, G View Figure 10 , 14A, B, D View Figure 14 , 22C View Figure 22 , 27A View Figure 27 , 30F View Figure 30 , 43C View Figure 43 , 47C View Figure 47 , 63B, C View Figure 63 ), and it can bear the cymbial fold (CyF, see below). This internal conductor is a synapomorphy of Mysmenidae   , although it is secondarily absent in Mysmenopsis   ( Figs 53D View Figure 53 , 60D View Figure 60 ). Usually, the CyC1 is pointed apically, which is the plesiomorphic condition in Mysmenidae   ( Figs 4G View Figure 4 , 30F View Figure 30 , 40B View Figure 40 , 43C View Figure 43 , 47C View Figure 47 ); however, the CyC1 evolved independently into different shapes. A ‘half-circle’ shaped conductor is characteristic of Trogloneta   ( Fig. 63B, C, F View Figure 63 ). A CyC1 consisting of prolateral, retrolateral, and apical projections occurs in the clade comprising Microdipoena   and Mysmeniola   , independently occurring in Mysmena   (= Calomyspoena   ) santacruzi   ( Fig. 27A View Figure 27 ). A spiral cymbial conductor occurs in Microdipoena   s.s. ( Fig. 22C View Figure 22 ). In Maymena   , the characteristic CyC1 comprises a particular apical cymbium that is bent over the ventral side ( Fig. 10D, G View Figure 10 ).

The ‘secondary cymbial conductor’ (CyC2) is external, located on the edge or dorsally on the cymbium (e.g. Figs 30F View Figure 30 , 31C View Figure 31 , 43C View Figure 43 ). This external conductor is plesiomorphically absent in Trogloneta   , Maymena mayana   , and in most Mysmenopsis species   ( Fig. 63C View Figure 63 ). Within Mysmenidae   , the CyC2 has evolved convergently in Maymena   (excluding M. mayana   ), Isela   , Mysmenopsis penai   , and Mysmeninae   ( Figs 4G View Figure 4 , 10G View Figure 10 , 31C View Figure 31 , 40A View Figure 40 , 41D View Figure 41 , 43C View Figure 43 , 60D View Figure 60 ). The CyC2 has been secondarily lost in the clade comprising Microdipoena   and MYSM-019-MAD ( Fig. 22F View Figure 22 ).

Cymbial fold (CyF): The external cuticle of the cymbium is usually hirsute, the internal one is glabrous. Although the external cuticle can also cover the internal side of the cymbium, both cuticles are delimited by a well-defined border (e.g. Figs 86G View Figure 86 , 102D View Figure 102 ). In some mysmenids the delimitation or border between external and internal cuticles (or cymbial areas) is rather clear, often corresponding to the outline of the cymbium; however, the internal cuticle on the tip of the cymbium bears setae (and can also bear the tarsal organ), and it frequently appears flattened against the outer cuticle. The internal cuticle can also be modified into a primary cymbial conductor. This suggests that the inner cymbium, at least apically, might be composed of part of the external cuticle. This condition is here referred to as a ‘fold’, and it is different from a twisted tip of the cymbium, where the same external cuticle is bent inwards, i.e. ventrally (compare with Figs 10G View Figure 10 , 14B View Figure 14 ). The ‘cymbial fold’ is a synapomorphy of Mysmenidae   ( Figs 4G View Figure 4 , 18E View Figure 18 ), secondarily and independently lost in Maymena mayana   , Mysmenopsis   , and Microdipoena   (= Mysmenella   ) illectrix   ( Fig. 60D View Figure 60 ).

On the cymbial fold cuticle, a distinct row of setae can be present, usually associated with the primary cymbial conductor (CyFs). Fold setae arise independently in Isela   , the clade comprising Brasilionata   and Microdipoena   , and in a large clade within Mysmena   ( Figs 4G View Figure 4 , 43E View Figure 43 ). The row setae can be similar to the surrounding setae at the tip of the cymbium ( Figs 40D View Figure 40 , 43E View Figure 43 ), or can be distinctly minute ( Figs 4G View Figure 4 , 17A View Figure 17 , 18E View Figure 18 , 22C View Figure 22 , 132A, D View Figure 132 , 134A View Figure 134 ).

Cymbial tarsal organ (to): In most spiders, and basally in Mysmenidae   , the tarsal organ is located externally on the cymbium (e.g. Figs 10I, J View Figure 10 , 44D View Figure 44 , 58C View Figure 58 ). An internal tarsal organ, located within the cymbial fold, optimizes as independently synapomorphic in Trogloneta   , Isela   , Mysmena- MYSM-015-MAD ( Mysmena   ), and the mysmenine MYSM-020-MAD ( Figs 4I View Figure 4 , 40B, C View Figure 40 , 63B, F View Figure 63 ).

Cymbial groove (CyG): A diagonal groove of varying depth can occur dorsally on the cymbium of some mysmenids ( Fig. 126 View Figure 126 ). The cymbial groove can be either a shallow and wide irregular depression ( Figs 36E View Figure 36 , 51A, B View Figure 51 ) or a narrow and deep furrow ( Figs 18E View Figure 18 , 22F View Figure 22 , 28B View Figure 28 , 30B, C View Figure 30 , 45A View Figure 45 , 134D View Figure 134 ). Besides the depth and width of this groove, its position and length appear to be correlated: apical grooves are always shorter than medial or basal grooves ( Figs 36E View Figure 36 , 45A View Figure 45 ). The latter are longer, extending sometimes into the prolateral basal expansion of the cymbium ( Figs 18E View Figure 18 , 22F View Figure 22 , 28B View Figure 28 , 30A, C View Figure 30 ; see below).

Cymbial process (CyP) ( Kraus, 1967: ‘Kegeldorn des paracymbium’; Baert & Maelfait, 1983: ‘cymbial thorn’): In most mysmenids there is a process, often pointed, on the dorsal surface of the cymbium. The process is located often apically, retrolateral to the cymbial tip ( Figs 1A, E View Figure 1 , 4C View Figure 4 , 40F View Figure 40 , 51A View Figure 51 , 63C View Figure 63 , 133G View Figure 133 , 134G, H View Figure 134 ), or basally and prolaterally, at the end of the cymbial groove ( Figs 45A View Figure 45 , 132D, E View Figure 132 ), or it can be located apically, but prolateral to the cymbial tip ( Fig. 43C, D View Figure 43 ).

Paracymbium (PC): As in most araneoids (except Theridiidae   ), a retrolateral paracymbium is present in all symphytognathoid families, except for Anapidae   ( Figs 71F View Figure 71 , 92A View Figure 92 ). Although the loss of the paracymbium is synapomorphic for Anapidae   , it has secondarily evolved in Comaroma   ( Fig. 81B View Figure 81 ). Almost all mysmenids have a paracymbium; within this data set, it is secondarily absent in Maymena rica   and Isela   ( Fig. 4D, E View Figure 4 ). Although in recent studies the paracymbium was considered to be absent in mysmenids ( Coddington, 1990; Griswold et al., 1998; Miller et al., 2009), it had been previously reported as present by some authors (e.g. Kraus, 1967).

Our phylogenetic hypothesis suggests that the mysmenid paracymbium evolved from a basal hookshaped paracymbium into a characteristic shape and position (e.g. as in Mysmena tasmaniae   , Fig. 51B View Figure 51 ). In mysmenids the paracymbium is flat and rounded, with a uniform transition to the cymbium, and it is located medially, not basally (i.e. as a medial flat extension of cymbial edge; Figs 18B View Figure 18 , 22G View Figure 22 , 27B View Figure 27 , 30E View Figure 30 , 32A View Figure 32 , 36A View Figure 36 , 45B View Figure 45 , 63A View Figure 63 ). A flat, rounded paracymbium evolved independently as synapomorphic for Mysmenidae   , but also in Iardinis mussardi   and in Synaphris   . The paracymbium becomes secondarily basal in Trogloneta   ( Fig. 63A View Figure 63 ), and secondarily hook-shaped in Mysmenopsis   , where it is a thick (i.e. not flat), short distinct process, usually as long as wide ( Figs 53D View Figure 53 , 60D View Figure 60 ). Furthermore, in Mysmenopsis   , the paracymbium is bent inwards and is seemingly interacting with a tegular groove located dorsally on the bulb ( Figs 53F View Figure 53 , 55F View Figure 55 , 58D View Figure 58 , 60D View Figure 60 ). The interaction is here considered tentative. The dorsal tegular groove does not appear to have a ‘conductor’ function related to the embolus, and the paracymbium– bulb interaction as a locking mechanism is not evident, as in the case of theridiids ( Levi, 1961; Saaristo, 1978; Agnarsson, 2004; and references therein).

Prolateral basal expansion: This prolonged cuticle on the prolateral basal edge of the cymbium was originally observed in Theridiosomatidae   , and has been termed the ‘prolateral basal paracymbium’ by Schütt (2003). The term ‘paracymbium’, however, has been proposed and long used for the classical araneoid retrolateral process on the cymbium (e.g. Comstock, 1910; Coddington, 1986b, 1990; Griswold et al., 1998; and references therein). Here, this prolateral structure is simply referred to as ‘prolateral basal expansion’. Furthermore, both the paracymbium and the prolateral basal expansion can co-occur in the palp of some species. Such conjunction refutes the homology statement among the two structures. The prolonged cuticle of the basal expansion surrounds the bulb ventrally in varying degrees and occurs in theridiosomatids and most mysmenids ( Figs 4B View Figure 4 , 27A View Figure 27 , 30B, C View Figure 30 , 36C View Figure 36 , 47B View Figure 47 , 66D View Figure 66 ), but is absent in Maymena   and most mysmenopsines ( Figs 10B, C View Figure 10 , 55A View Figure 55 ).

A

B

Mysmenid bulb: general morphology

The median apophysis is absent in all mysmenids. In Trogloneta   and independently in the clade comprising Brasilionata   , Mysmeniola   , and Microdipoena   , a tegular groove housing the embolus can occur ( Figs 27A View Figure 27 , 63E View Figure 63 , 66C View Figure 66 ; see below).

Embolus: The general shape of the embolus varies greatly within Mysmenidae   , and no general pattern can be proposed. The embolus of mysmenids can be thin (or filiform) and coiled ( Figs 47B View Figure 47 , 134G View Figure 134 ), or thick (and flattened) and either coiled ( Figs 4H View Figure 4 , 27A View Figure 27 , 36B View Figure 36 , 131H View Figure 131 , 132B, D, E View Figure 132 ), or straight ( Figs 10E View Figure 10 , 28D View Figure 28 , 60F View Figure 60 , 63B, E View Figure 63 , 131A View Figure 131 ). Thick and straight emboli occur in Trogloneta   , Maymena   , and Mysmenopsis   , whereas thick and coiled emboli occur in Microdipoena   , Isela   , and some Mysmena species.   An apical switch in the coiling direction of the embolus is characteristic of the clade comprising Brasilionata   and Microdipoena   [secondarily absent in Microdipoena   (= Anjouanella   ) comorensis   ] ( Figs 18C, F View Figure 18 , 27C View Figure 27 , 132A–F View Figure 132 ). In addition, the embolus surface can be smooth ( Figs 10H View Figure 10 , 18F View Figure 18 , 27C View Figure 27 , 60F View Figure 60 ) or ridged ( Figs 28C View Figure 28 , 32G View Figure 32 ).

As in most symphytognathoids, the embolus of mysmenids is often long (i.e. much longer than the bulb, Figs 4A View Figure 4 , 10E View Figure 10 , 27A View Figure 27 , 63E View Figure 63 , 132E View Figure 132 ), usually tapering apically without further modifications ( Figs 1D View Figure 1 , 36B View Figure 36 , 60F View Figure 60 , 63B View Figure 63 ). Short emboli occur in Mysmenopsis   ( Figs 60F View Figure 60 , 131A–C View Figure 131 ) and in Trogloneta granulum   . In some Maymena   , Trogloneta   , and a few other mysmenids, distal modifications of the embolus can occur, such as a distal apophysis ( Figs 10H View Figure 10 , 18F View Figure 18 ) or a distal irregular membrane ( Fig. 27A, C View Figure 27 ).

The embolic base can be simple, or it can be lobed and bearing an apophysis, as in Mysmenopsis   and Trogloneta   ( Figs 55G View Figure 55 , 60F View Figure 60 , 63E View Figure 63 , 66E View Figure 66 , 131A View Figure 131 ). In Mysmeninae   , the basal or medial embolic trajectory has a pars pendula ( Comstock, 1910), a membrane that houses the spermatic duct before entering to the embolus ( Figs 32H View Figure 32 , 36B View Figure 36 , 132C, E, F View Figure 132 , 133C View Figure 133 ). Therefore, the spermatic duct enters the embolus not at its origin but further distally, meaning also that the embolus is actually longer than the embolic portion of the ejaculatory duct. Although found in other spider families (e.g. in some theridiids, linyphiids, cyatholipids, and agelenids), the pars pendula is an ambiguously optimized synapomorphy for Mysmeninae   (ambiguously optimizes at its node because of missing information on the basal clade of this group), and it is secondarily absent in MYSM-005-ARG ( Mysmena   ).

Tegular conductor: As in most symphytognathoids, most mysmenids have a conductor. Mysmenid conductor is distinctly voluminous and membranous, and originates subterminally from the tegulum, close to the embolic base ( Figs 17A View Figure 17 , 18D View Figure 18 , 27A, B View Figure 27 , 30E View Figure 30 , 36C View Figure 36 , 41D View Figure 41 , 63B, D View Figure 63 ). This structure has been named ‘bulbal shield’ (e.g. Baert, 1984a; Schütt, 2003), and other than the embolus, it appears to be the only tegular sclerite. In mysmenids, the conductor often embraces and even covers the embolic base ( Figs 41C View Figure 41 , 43A, B View Figure 43 ). In some species there is a groove on the conductor surface housing the basal portion of the exposed embolus ( Figs 36B, C View Figure 36 , 47E View Figure 47 ); however, a groove housing the distal portion of embolus is absent in the tegular conductor of mysmenids, and the tip of the embolus is instead housed by one of the two conductors on the cymbium (see above). Within symphytognathoids, the occurrence of a tegular conductor is symplesiomorphic and widespread, with few independent losses of this sclerite occurring in anapids, symphytognathids, and mysmenids. Within Mysmenidae   , the conductor is lost two times: in the clade that includes Maymena   and Mysmenopsinae   , and in Mysmena   MYSM-005-ARG ( Figs 4A View Figure 4 , 10E View Figure 10 , 28D View Figure 28 ).

Spermatic duct trajectory (SDT; refer to Fig. 127 View Figure 127 and characters 220–228): As with most of the aforementioned palpal features, there does not seem to be a consistent pattern in the trajectory of the spermatic duct across the family, although there is some regularity within clades. The spermatic duct usually travels clockwise from the fundus (in left palp). If a switchback (SB) occurs, it alters this direction to travel counterclockwise. Usually, a counter-switch also occurs to return the duct trajectory to its original clockwise direction. Therefore, when SBs are present, they usually occur in pairs of switchbacks. The trajectory of the spermatic duct in Trogloneta   differs from all other mysmenids in that the pair of switchbacks that occur before the spermatic duct completes one loop from the fundus (i.e. SB I and II, see Fig. 127 View Figure 127 ) are absent, therefore the trajectory is completely spiralling or has one complete loop before the ‘second’ pair of switchbacks (SB III and IV) occurs ( Fig. 131E, F View Figure 131 ). Although the general arrangement of the spermatic duct in all other mysmenid species examined in this study (except Trogloneta   ) is not necessarily similar among clades, all have the first pair of switchbacks on their trajectories (SB I and II; Figs 131I View Figure 131 , 132B, D View Figure 132 ).

The position of switchback I (SB I) varies within Mysmenidae   also. A distal SB I (apart from basal fundus, on the opposite area of the bulb) occurs in most mysmenines, most Maymena   , and in Isela okuncana   ( Figs 131H, I View Figure 131 , 132B, E View Figure 132 , 133A, B View Figure 133 , 134B, C View Figure 134 ). A basal SB I that does not reach the distal part of the bulb can occur in other mysmenines, however ( Figs 133H, I View Figure 133 , 134F– H View Figure 134 ); whereas in most mysmenopsines and Maymena mayana   the SB I occurs close to the fundus, after the spermatic duct has reached the distal wall of the bulb (i.e. ‘beyond distal’; Fig. 131C, G, J View Figure 131 ).

In most mysmenids, the portions of spermatic duct forming the SB I are divergent, and SB II occurs relatively close to SB I ( Figs 131B, H, J View Figure 131 , 133A, B View Figure 133 , 134D–G View Figure 134 ). In Microdipoena   and most Mysmena species   , the portions of spermatic duct comprising SB I run close to each other, and the SB II occurs close to the midpoint between SB I and the fundus, or even closer to the fundus ( Figs 131I View Figure 131 , 132B–D View Figure 132 , 133D, E View Figure 133 , 134A–C View Figure 134 ).

The plesiomorphic condition in Mysmenidae   is to have two or more ascending loops in the last portion of the coiling reservoir before entering the embolus. This occurs basally within Mysmeninae   ( Figs 133D, E View Figure 133 , 134E, H View Figure 134 ). Within the family, the number of loops decreases independently in distal clades (no loops or less than one entire loop, Figs 131B, E, H, I View Figure 131 , 132F View Figure 132 , 133A, B View Figure 133 , 138C, G View Figure 138 , 139E View Figure 139 ; or about one and one and a half loops, Figs 131D View Figure 131 , 139C View Figure 139 ).

A pair of switchbacks (SB III and IV) can occur either after SB II, or if SB I and II are absent, after a complete loop of the spermatic duct (e.g. as in Symphytognatha picta   ; Fig. 139C View Figure 139 ). In this data set, an absence of pairs of extra switches (i.e. SB III and IV) is plesiomorphic for both symphytognathoids and Mysmenidae   ( Figs 131D, E, J View Figure 131 , 133A, B, D, E, H, I View Figure 133 , 134E View Figure 134 ). Switchbacks III and IV evolve independently in Microdipoena   , Mysmena   MYSM-007- MEX, Isela okuncana   , and Trogloneta cantareira   ( Figs 131F, H View Figure 131 , 132B–E View Figure 132 ). Furthermore, a distinct trajectory of the spermatic duct where several pairs of switchbacks occur evolved convergently in the mysmenines MYSM-020- MAD and MYSM-023-MAD ( Figs 134F–H View Figure 134 ).

Epigynum (refer to characters 59–87)

Mysmenids are entelegyne spiders. That is, they have fertilization ducts leading from the spermathecae to the uterus externus and copulatory ducts connecting external openings to the spermathecae (e.g. Wiehle, 1967; Uhl, 2002). No comparative study of mysmenid female genitalia has ever been performed. In most female mysmenids, particularly mysmenines, the ducts and sometimes even the spermathecae are extremely membranous, almost invisible under light microscopy. A few authors describing mysmenid species have published detailed illustrations and have attempted to identify the different components of female genitalia in as much detail as possible (e.g. Kraus, 1967; Loksa, 1973; Thaler, 1975; Baert, 1984a, b, 1986, 1988; Snazell, 1986; Baert & Murphy, 1987). Given the membranous and almost undetectable nature of mysmenid female genitalia, the interpretation of these structures is often difficult. Furthermore, the great diversity of mysmenid female genitalic morphology (see below) makes diagnosis of the family a challenging task if only based on this system of characters. Our interpretations and homology statements of female genitalic structures are based on light and SEM microscopy data.

External genitalia

The epigynum, as a sclerotized modification of the cuticle, is absent in most of the members of the subfamily Mysmeninae   . In addition, the copulatory openings are located within the epigastric furrow (i.e. the epigynal area containing the copulatory openings is hidden within it; Fig. 24A View Figure 24 ). This latter trait has convergently evolved in most anapids. All representatives of Mysmeninae   here examined have a membranous atrium ( Figs 59H View Figure 59 , 129A, E, G View Figure 129 , 130B View Figure 130 ). The atrium has been defined as a ‘widened cavity into the copulatory ducts’ ( Sierwald, 1989: 2), and can occur independently of the location (external or internal) of the copulatory openings. In most mysmenids, however, the seemingly epigynal area located centrally between the copulatory openings (here regarded as the ‘dorsal plate’ sensu Millidge, 1984; ‘middle field’ on Sierwald, 1989) is projecting, i.e. it is exposed or protruding from the epigastric furrow ( Figs 11B, C View Figure 11 , 12D, E View Figure 12 , 37C View Figure 37 , 42C View Figure 42 , 44A View Figure 44 , 49D View Figure 49 , 67A, B View Figure 67 ). The dorsal plate is secondarily internal (i.e. neither exposed nor projecting) in Mysmenopsinae   , Maymena rica   , Microdipoena   , and few other mysmenines (e.g. Fig. 24B View Figure 24 ).

Trogloneta   , Maymena, Mysmenopsinae   , and the mysmenine MYSM-023-MAD have a modified copulatory area or epigynum in the form of a sclerotized plate or a protruding modification of the cuticle, usually bearing setae and the copulatory openings ( Figs 5A, B View Figure 5 , 11B, C View Figure 11 , 12D, E View Figure 12 , 49D, E View Figure 49 , 59H View Figure 59 , 61A View Figure 61 , 67A, B View Figure 67 , 140D, I View Figure 140 , 141G, H View Figure 141 , 142C View Figure 142 ). The copulatory openings are exposed caudally (i.e. posteriorly) in the epigynal area ( Figs 49D View Figure 49 , 59H View Figure 59 , 61B View Figure 61 ). A sclerotized external atrium is present in a few Mysmenopsis species   (e.g. Fig. 59H View Figure 59 ). On the other hand, in most Mysmeninae   the epigynal area is weakly modified or even absent (i.e. the cuticle in this area is similar to surrounding abdominal cuticle), and it is usually translucent (spermathecae can be observed beneath it; Figs 14C View Figure 14 , 37A View Figure 37 , 52F View Figure 52 , 141I View Figure 141 ). Although seemingly widespread among araneoids ( Levi & Levi, 1962; Millidge, 1984; Scharff & Coddington, 1997; Griswold, 2001), the ventral scape of most mysmenines is unique within symphytognathoids ( Figs 24B View Figure 24 , 29C View Figure 29 , 31G View Figure 31 , 37C View Figure 37 , 42C View Figure 42 , 129E View Figure 129 ).

Internal genitalia

Copulatory ducts: Within Mysmenidae   , the copulatory ducts show varying degrees of sclerotization and width. In Maymena   and most Mysmenopsinae   the copulatory ducts are short, relatively sclerotized, narrow, and of invariable diameter ( Figs 11D View Figure 11 , 37D View Figure 37 , 128A, D, G View Figure 128 , 129G View Figure 129 ). In Trogloneta   the walls of the distal portion of the long copulatory ducts are rather smooth, relatively sclerotized, and uniform in diameter; however, the proximal portion of the ducts of Trogloneta   is highly membranous and has a larger diameter than the distal part ( Figs 64A View Figure 64 , 128F View Figure 128 ). In the aforementioned taxa (i.e. Trogloneta   , Maymena   , and Mysmenopsinae   ), the trajectory of the copulatory ducts is in most cases recognizable. Conversely, the copulatory ducts of Mysmeninae   differ from all other mysmenids (with the exception of a few taxa). The walls of the ducts are extremely membranous, imperceptible under light microscopy, and are of uneven diameter. These irregular membranous ducts follow a convoluted and long trajectory of unclear course. The ducts seem to extend ventrally and anteriorly to the spermathecae, although without a definite pattern ( Figs 18G View Figure 18 , 27D View Figure 27 , 129A, C, E, H View Figure 129 , 130B View Figure 130 ). In some species the ducts can be subtly more sclerotized and definite, and a coiled trajectory can be observed ( Figs 37D View Figure 37 , 129G View Figure 129 ). The increase in the diameter of the proximal portion of the copulatory ducts (i.e. the first half of the ducts from the copulatory openings) has been termed ‘bursae’ by Schütt (2003: character 77), although it is not clear from that study whether the increase in diameter refers to the copulatory ducts or to the membranous atrium (see above).

Within Mysmeninae   , there is a particular turn occurring proximally in the convoluted and membranous copulatory ducts, seemingly originating close to the internal atrium, but immediately before becoming widened and convoluted. This duct turn is characterized by a subtle but consistent sclerotization. This feature occurs in Microdipoena   , and also evolved independently in two clades within Mysmena   and in MYSM-029-MAD ( Figs 129A, B View Figure 129 , 130A View Figure 130 ).

A convoluted trajectory of the copulatory ducts characterizes Mysmenidae   , although the ducts can vary greatly in terms of sclerotization and diameter. Straight ducts evolved independently twice: once in the clade comprising Maymena   and Mysmenopsinae   and once in Mysmena   MYSM-034-MAD. Although rare in the current mysmenid taxon sample, coiled and more distinct copulatory ducts of mysmenines (as in Figs 37D View Figure 37 , 129G View Figure 129 ) appear to be more common than represented here (e.g. Lopardo, Dupérré & Paquin, 2008; Miller et al., 2009; L. Lopardo, A. Janzen, C. Griswold & P. Michalik, unpubl. data). These distinct ducts might represent a plesiomorphic but intermediate condition (i.e. between sclerotized and fully membranous ducts) in the evolution of convoluted and highly membranous copulatory ducts, and might help in elucidating our current interpretation of their trajectory, as well as their identification.

Spermathecae and accessory glands: Most mysmenids have one sperm-storage compartment in each spermatheca (e.g. Fig. 33A View Figure 33 ). Trogloneta   and some anapids have two pairs of compartments in each spermatheca ( Fig. 128F View Figure 128 ). The spermathecae are usually defined by a thick sclerotized wall ( Figs 129A View Figure 129 , 130B View Figure 130 ), although exceptions occur (e.g. in mysmenine MYSM- 023-MAD and a few anapids; Figs 49A View Figure 49 , 130D View Figure 130 ).

Ovoid spermathecae are plesiomorphic for both symphytognathoids and Mysmenidae   . Nevertheless, the diversity of spermathecal shapes within Mysmenidae   is immense, seemingly following no particular phylogenetic pattern, not even at the genus level. Spermathecae can be ovoid ( Figs 11D View Figure 11 , 18G View Figure 18 , 37D View Figure 37 , 128D View Figure 128 , 129A, G View Figure 129 ), C- or cup-shaped ( Figs 27D,E View Figure 27 , 42D View Figure 42 , 128E View Figure 128 , 130C View Figure 130 ), coiled within the same spermathecal structure ( Figs 33A, B View Figure 33 , 51D View Figure 51 , 129C View Figure 129 , 130B, G View Figure 130 ), tubuliform (as one large tube, sometimes like tracheoles; Figs 5D View Figure 5 , 49A View Figure 49 , 128A, C View Figure 128 , 130D View Figure 130 ), clavate ( Figs 12C View Figure 12 , 128B View Figure 128 ), or even irregular (although consistent within each species), where no particular shape can be defined ( Fig. 129D View Figure 129 ).

An additional paired structure occurs in the internal genitalia of Trogloneta   , the mysmenine MYSM- 029-MAD, Mysmena   MYSM-005-ARG, Microdipoena   , and the anapid Tasmanapis   . This structure resembles either an apodeme or a glandular structure, and appears related to the copulatory ducts or the spermathecae ( Figs 22B View Figure 22 , 27D View Figure 27 , 29A View Figure 29 , 64A View Figure 64 , 129B, E, H View Figure 129 , 130A, F View Figure 130 ). It is better observed by SEM, although it can be distinguished in transparent preparations of the vulva by a higher degree of sclerotization, comparable with that of the spermathecae ( Figs 129B, E, H View Figure 129 , 130A, F View Figure 130 ; see also Brescovit & Lopardo, 2008: fig. 6C– E). Whether these structures are functional glands, muscle attachment points, or perform other functions remains unknown. They are regarded here as accessory glands.

Fertilization ducts: The morphology of fertilization ducts is also variable within the family. Fertilization ducts were identified in Mysmenidae   by discerning the ducts connected to and from the spermathecae, and, with the help of SEM, an attempt was made to follow the trajectory of these ducts. Usually fertilization ducts are located either dorsal to or on the central internal lateral side of the spermathecae. As a convention, when the genital system was mainly composed by membranes, highly developed and convoluted copulatory ducts were first identified, and then fertilization ducts were distinguished by elimination. The degree of sclerotization of the fertilization ducts appears highly homoplastic. Weakly sclerotized fertilization ducts with a distinguishable wall are plesiomorphic for Mysmenidae   (although ambiguously optimized), and occur in Isela   , mysmenine MYSM-023-MAD, and most Mysmena   ( Figs 42D View Figure 42 , 49A View Figure 49 , 128A View Figure 128 , 129G View Figure 129 ); however, membranous, translucent, and almost imperceptible fertilization ducts also occur within the family, in Trogloneta   , most Mysmenopsis   , and all members of Microdipoena   ( Figs 18G View Figure 18 , 27D View Figure 27 , 51D View Figure 51 , 60H View Figure 60 , 64A View Figure 64 , 128F View Figure 128 , 136D View Figure 136 ).

Small, short fertilization ducts providing a direct connection between the spermathecae and the uterus externus (usually in straight fashion and unmodified) are plesiomorphic and widespread within Mysmenidae   ( Figs 42D View Figure 42 , 49A View Figure 49 , 64A View Figure 64 , 128A, F View Figure 128 ). Large, long fertilization ducts, most often longer than the size of the spermathecae, might be provided with modifications or expansions ( Figs 11D View Figure 11 , 18G View Figure 18 , 27D View Figure 27 , 51D View Figure 51 , 60H View Figure 60 , 92F View Figure 92 , 129G View Figure 129 , 136D View Figure 136 ). They occur independently in most Maymena   , most Mysmenopsis   , most species of Microdipoena   , and few Mysmena   .

Spinneret silk gland spigot morphology (refer to characters 304–340)

The spinning organs of Mysmenidae   have been described for a few species, including the kleptoparasitic Isela okuncana ( Griswold, 1985)   , an undescribed Isela   from Cameroon, Mysmenopsis penai   , the Australian Mysmena tasmaniae   and M. leichhardti ( Lopardo & Michalik, 2013)   and Maymena mayana ( Griswold et al., 1998)   . Recently, the gland spigot patterns of Trogloneta cantareira ( Brescovit & Lopardo, 2008)   and several Chinese mysmenids ( Mysmena   , Gaoligonga   , Chanea   , and Maymena, Miller et al., 2009   ) were also described. The data in the aforementioned works suggest that mysmenids have the typical symphytognathoid and higher araneoid silk gland spigot conformation on the anterior lateral and the posterior median spinnerets (ALS and PMS, respectively): few ALS piriform gland spigots, few aciniform gland spigots on posterior (median and lateral) spinnerets, a furrow between major ampullate and piriform fields, and reduced piriform bases. In this study we examined in detail the spinneret gland spigot conformation of 30 mysmenid species. In the following section, the general arrangement of mysmenid spinneret gland spigots is described. Exceptions, singular features, and synapomorphies for the main mysmenid clades are noted below. See Appendix 2 for an explanation of cuticular textures.

In general, the colulus is fleshy and usually relatively large, bearing three or less setae ( Figs 24E View Figure 24 , 33C View Figure 33 , 56D View Figure 56 , 59I View Figure 59 ). On the anterior lateral spinnerets, a glabrous tuberculate intersegmental cuticle occurs ( Figs 6A View Figure 6 , 16B View Figure 16 , 23A, C View Figure 23 , 33F View Figure 33 , 52B View Figure 52 , 61C View Figure 61 ). The major   ampullate gland spigot is accompanied by a nubbin and a tartipore ( Figs 6B View Figure 6 , 23A, C View Figure 23 , 61C View Figure 61 ). The base of the piriform gland spigots is reduced ( Figs 6B View Figure 6 , 52B View Figure 52 , 61C View Figure 61 ), and the cuticle surrounding those piriform gland spigots can be either fingerprint (in Mysmeninae   and most Mysmenopsinae   ) or rugose (in Maymena   , Trogloneta   , and the Isela   representative Kilifina- MYSM-002- KENYA; Fig. 6B View Figure 6 ). Two aciniform, one posterior minor ampullate, and (in females) one cylindrical gland spigot occur on the posterior median spinnerets ( Figs 11F View Figure 11 , 33D View Figure 33 , 58F View Figure 58 ). The minor   ampullate can be accompanied by a tartipore, a tartipore plus a nubbin, or by none of these. The posterior lateral spinnerets bear two cylindrical gland spigots, where both ( Figs 11G, H View Figure 11 , 13F View Figure 13 , 67D View Figure 67 ) or only the posterior spigot is peripheral to the spinning field (the anterior spigot on the edge of the field; Figs 23B View Figure 23 , 37B View Figure 37 , 58H View Figure 58 ), and some aciniform gland spigots. The triad (the assemblage of one flagelliform and two aggregate gland spigots producing sticky silk in araneoids) is present in most female mysmenids ( Figs 11G View Figure 11 , 23B View Figure 23 , 37B View Figure 37 , 52C View Figure 52 , 67D View Figure 67 ), as is usually the case in Araneoidea. Some exceptions occur, however (see below). Both aggregate and flagelliform gland spigots are similar in size ( Figs 13F, G View Figure 13 , 37B View Figure 37 , 52C View Figure 52 ). In most species, the triad on males is vestigial, were remnants of previously functional gland spigots can be observed ( Figs 23E View Figure 23 , 33H View Figure 33 ). In some species, the triad is retained in adult males ( Fig. 13G View Figure 13 ; this change is ambiguously optimized at the base of Mysmenidae   ).

The spinnerets of mysmenids differ from that of all other families represented in this data set by the presence of a lobe on the intersegmental groove of the ALS ( Figs 23A, C, G, H View Figure 23 , 52B View Figure 52 ). This lobe is also present within Anapidae   , although with high homoplasy. Mysmenids also differ from other families in the separation of the major   ampullate and piriform fields by a subtle (shallow) groove ( Figs 13C View Figure 13 , 23C View Figure 23 , 33F View Figure 33 , 52B View Figure 52 , 61C View Figure 61 , 64C View Figure 64 ), where the connection between both fields is distinctly evident proximally within the ALS segment (ambiguously occurring in Theridiosomatidae   ). Finally by the characteristic shape of a seta on the major   ampullate field, with either one or two rows of long ‘branches’ ( Figs 6A View Figure 6 , 23F View Figure 23 , 33F View Figure 33 , 52B View Figure 52 , 61C View Figure 61 ).

Trogloneta   is the only symphytognathoid so far examined with minute but distinguishable colulus ( Figs 64D View Figure 64 , 67F View Figure 67 , 68C View Figure 68 ; as opposed to the remnant colulus of Patu   , see character 317). An additional anterior discrete cluster of cuticular protuberances of unknown function also occurs in the ALS of Trogloneta   ( Figs 64C View Figure 64 , 66F View Figure 66 ). Other attributes occurring in Trogloneta   , although not exclusive to this genus, include: a rugose cuticle on the piriform field on ALS ( Fig. 64C View Figure 64 ); minor ampullate gland spigot accompanied solely by a tartipore on PMS; both PLS cylindrical gland spigots equally large and larger than the flagelliform gland spigot ( Fig. 67D View Figure 67 ); and triad spigots retained in adult males (at least in T. granulum   ).

In Maymena   , the shape of the seta on the major   ampullate field has a distinct single row of long ‘branches’ ( Figs 11E View Figure 11 , 13C View Figure 13 , 16B View Figure 16 ); in other mysmenids, two rows occur. The PMS minor ampullate gland spigot is accompanied by a nubbin and a tartipore ( Fig. 11F View Figure 11 ). The following features occur independently in both Maymena   and Mysmeninae   , and were not observed in any other taxa examined in this data set: an anteri- or distinctly flat spatulate modified seta on PLS ( Figs 11G,H View Figure 11 , 13E View Figure 13 ) and aciniform gland spigots of different shape in both the posterior spinnerets (median and lateral, Figs 11F, G View Figure 11 , 13F, G View Figure 13 ; see character 304). Other attributes occurring in Maymena   , although not exclusive of the genus, include: rugose cuticle on piriform field on ALS ( Figs 11E View Figure 11 , 16B View Figure 16 ), and both PLS cylindrical gland spigots equally large and larger than the flagelliform gland spigot ( Figs 11G, H View Figure 11 , 13F View Figure 13 ). Triad spigots are retained in adult males of M. mayana   ( Fig. 13G View Figure 13 ), but this retention appears autapomorphic for this species rather than a generic condition, given that the triad is vestigial in at least M. rica   .

In Mysmenopsinae   the adult male aggregate gland spigots are absent, and those of the females are distinctly absent as well (see below). Other spinneret features of this subfamily include: a fingerprint cuticle on ALS piriform field; PMS minor ampullate gland spigot accompanied by neither nubbin nor tartipore ( Figs 6C, F View Figure 6 , 58F View Figure 58 , 61D View Figure 61 ); and both PLS cylindrical gland spigots slim as other gland spigots (not larger) and subequal to flagelliform gland spigot ( Fig. 58H View Figure 58 ). In Mysmenopsis   , the colulus bears four or more setae ( Figs 56D View Figure 56 , 59I View Figure 59 ). Although both aggregate gland spigots are absent in males and females, the flagelliform gland spigot in the representatives of Mysmenopsis   studied has been retained, and seems to be functional in both sexes ( Fig. 58G, H View Figure 58 ). In Isela   both flagelliform and aggregate gland spigots are distinctly absent in both sexes ( Fig. 6D, G View Figure 6 ).

Finally, the subfamily Mysmeninae   shares with Maymena   the anterior distinctly flat spatulate modified seta on PLS ( Figs 23B, E View Figure 23 , 33G, H View Figure 33 , 37B View Figure 37 , 52C View Figure 52 ) and aciniform gland spigots of different shape in both posterior spinnerets (median and lateral, Figs 23D View Figure 23 , 33D View Figure 33 , 37B View Figure 37 ; see character 304). These two features evolved independently in the two clades. Other features occurring in mysmenines include a fingerprint cuticle on ALS piriform field ( Figs 23C View Figure 23 , 52B View Figure 52 ); PMS minor ampullate gland spigot accompanied solely by a tartipore ( Figs 19F View Figure 19 , 23D View Figure 23 , 33D View Figure 33 ); both PLS cylindrical gland spigots slim as other gland spigots (not larger) and subequal to flagelliform gland spigot ( Figs 23B View Figure 23 , 37B, E View Figure 37 , 52C View Figure 52 ); and vestigial triad in males ( Fig. 23E View Figure 23 ), independently functional in Mysmena   MYSM-005-ARG and Mysmena tasmaniae   .

Other morphological features of Mysmenidae  

Clasping spines

The males of all mysmenids except Maymena mayana   have a prolateral metatarsal clasping spine (macroseta) on leg I (e.g. Fig. 34C View Figure 34 ). The phylogenetic hypothesis from the total-evidence analysis agrees with the morphological hypothesis from this study and with previous studies that have suggested this macroseta as a synapomorphy (or diagnostic) for the family ( Thaler, 1975; Platnick & Shadab, 1978; Brignoli, 1980; Griswold, 1985; Wunderlich, 1995; Griswold et al., 1998; Schütt, 2003). Although some members of Anapidae   have a clasping structure on the first legs, these structures are not spines (macrosetae) but spurs (short and stout seta), and can occur in both sexes. The clasping spines of mysmenids are sexually dimorphic, occurring in males, are unique for the family, and might be involved in mating behaviour ( Schütt, 2003). The widespread condition is a medial and straight metatarsal clasping spine ( Figs 34C View Figure 34 , 42H View Figure 42 , 45H View Figure 45 , 65C View Figure 65 ), but the spine can be basal (e.g. Maymena   ; Figs 16G View Figure 16 , 141K View Figure 141 ), apical (few Mysmenopsis   and mysmenines species; Figs 50H View Figure 50 , 59B View Figure 59 , 143C View Figure 143 , 144A View Figure 144 ), twisted ( Isela   and Microdipoena   ; Figs 3B View Figure 3 , 8B, C View Figure 8 , 26C View Figure 26 , 140E, F View Figure 140 , 141K, L View Figure 141 , 142N View Figure 142 ), or strongly curved proximally and accompanied by adjacent strong setae (most Mysmenopsis   ; Figs 54D View Figure 54 , 57I View Figure 57 , 59B View Figure 59 ). Additional apical tibial clasping spines can also be found in some mysmenids, as is the case of one tibial clasping spine occurring in all Maymena species   (except M. mayana   ) and in Mysmenopsinae   ( Figs 3A View Figure 3 , 8D View Figure 8 , 16G View Figure 16 , 54C View Figure 54 , 59B View Figure 59 , 62E, F View Figure 62 , 140E, J, K View Figure 140 ); or two clasping spines in Mysmenopsis dipluramigo   ( Fig. 140H View Figure 140 ) and Microdipoena   ( Figs 26C View Figure 26 , 27I View Figure 27 , 57F, G View Figure 57 , 141L, O View Figure 141 ).

Femoral spot or other femoral structures

The femoral spot has been previously suggested as diagnostic or synapomorphic for the family ( Thaler, 1975; Platnick & Shadab, 1978; Brignoli, 1980; Griswold, 1985; Wunderlich, 1995; Griswold et al., 1998; Schütt, 2003), as this cuticular structure is unique to mysmenids. The results of our study corroborate this hypothesis. The adult females of all mysmenid species have either a sclerotized spot ( Figs 34A View Figure 34 , 39D View Figure 39 , 140G View Figure 140 , 141C View Figure 141 , 143N View Figure 143 ) or a cuticular projection ( Fig. 57A, E View Figure 57 ) on the apical ventral surface of at least femur I, although a few exceptions occur (e.g. Mysmena   MYSM-005-ARG and a few Mysmenopsis species   ). The femoral structure of most female mysmenids occurs in both femora I and II (symplesiomorphic in this data set, Figs 141C View Figure 141 , 142B View Figure 142 ). The occurrence of this feature only on femur I is convergent in Trogloneta   , Mysmenopsis   (when present), and a number of times within Mysmena   ( Fig. 140G View Figure 140 ). The femoral sclerotization (i.e. spot) was first described by Marples (1955) for Tamasesia   (= Mysmena   ). Its function remains unknown. The absence of pores in its surface indicates that we should rule out a glandular function. It has been suggested that the spot could be a ‘... functionless remnant of an unknown structure, because it shows a great variability in size and shape among specimens and can be differently pronounced on the two first femora of a single specimen. It is too large to be the socket of a former spine and apart from this there is no space for a large spine so close to the patella...’ ( Schütt, 2003: 143). Based on the results of the combined analysis, the spot originates at the node of Mysmenidae   , becomes a femoral projection in Mysmenopsis   , and is lost distally in some species. Whether the femoral spot actually has a function (e.g. a behavioural function), or is just a remnant of a functional structure, remains an unsolved puzzle.

All femoral structures are absent in juvenile mysmenids (which argues against the remnant hypothesis), although they can sometimes be perceived in subadult stages. The femoral projection occurs only in females of some species of Mysmenopsis   . The femoral spot, however, can also occur on males ( Fig. 21A View Figure 21 ). Male femoral spots evolved ambiguously in Trogloneta   and Maymena   , independently in Microdipoena   [excluding its basal species M. (= Mysmenella   ) samoensis   ], and in two instances within Mysmena   . In contrast with females, most mysmenid males have the femoral spot (when present) only on femur I ( Fig. 141O View Figure 141 ), although in two cases the spot occurs in both femora I and II ( Maymena ambita   and Mysmena tasmaniae   ; Fig. 140M View Figure 140 ).

Prolateral row of modified setae in the male

first leg tarsi

First observed and described as a ventral row of modified setae by Thaler (1975, 1995) for males of Trogloneta granulum   and Mysmeniola spinifera   (respectively), this prolateral row of modified setae (see below) is an ambiguously optimized synapomorphy for Mysmenidae   . As is the case of most morphological features within Mysmenidae   , and even though this row of modified setae occurs in all examined species, the particular details of this feature differ among clades. The modified setae are usually shorter than the surrounding tarsal setae, and can be slimmer and curved or stouter and straight. These modified setae can also be distributed along the entire length of the tarsus or just on the distal half. In Mysmeninae   , the setae are slimmer and the row occupies the distal half of the tarsus (except in Mysmeniola   , see below; Figs 26A View Figure 26 , 34D View Figure 34 , 45I View Figure 45 , 50F View Figure 50 ). In Maymena rica   and in Mysmeniola spinifera   the setae are also slimmer but the row is distributed all along the segment ( Fig. 16H View Figure 16 ). Trogloneta   and Isela   both have stout setae distributed along the tarsus ( Figs 8F View Figure 8 , 65D View Figure 65 , 68A View Figure 68 ). And lastly, Mysmenopsis   also have stout setae but, as in Mysmeninae   , the row occupies the distal half of the tarsus ( Figs 54G View Figure 54 , 59D View Figure 59 ). Although the function of this row of modified setae remains unknown, the fact that these setae are found only in the males suggests that it may be involved in mating behaviour.

Other morphological features of Mysmenids

Size of the tarsal organ opening on leg I: A large opening of the tarsal organ (i.e. subequal or larger than setal sockets) evolved independently in Mysmenidae   and Anapidae   (see Figs 26E View Figure 26 , 39C View Figure 39 , 48F View Figure 48 , 50D View Figure 50 , 73B View Figure 73 , 83E View Figure 83 , 95I View Figure 95 , 101E View Figure 101 , respectively). An opening distinctly smaller than setal sockets is the plesiomorphic condition in symphytognathoids. Within Mysmenidae   , the tarsal organ opening becomes secondarily small in Mysmenopsinae   , with a reversal in Mysmenopsis palpalis   ( Figs 3F View Figure 3 , 54F View Figure 54 , 62H View Figure 62 ).

Distinctly thick distal promarginal curved seta on chelicerae: Most taxa in this data set have a particularly distinct curved seta located distally at the promarginal edge of chelicerae, near the fang base. This seta differs from surrounding promarginal setae by its thickness and/or serration. Although a distinctly curved seta can also occur on the cheliceral retromargin of some araneomorph spiders (see Griswold et al., 2005, character 34), retromarginal setae of the taxon sample examined in this study do not differ from surrounding setae (e.g. Fig. 7J View Figure 7 ). Within symphytognathoids, the distinctly curved promarginal seta is lost in Synaphridae   and most Symphytognathidae   ( Figs 108E View Figure 108 , 118E View Figure 118 , 122A View Figure 122 ; the optimization of this character under parsimony is ambiguous). All examined mysmenid representatives have a uniquely thicker distal promarginal curved seta ( Figs 19E View Figure 19 , 25E View Figure 25 , 38H View Figure 38 , 48B View Figure 48 ), except Maymena   . In some mysmenids this seta is equally serrated as surrounding seta, which is the plesiomorphic condition for the family; however, the thicker seta is strongly serrated on one side independently in Mysmeninae   and most Mysmenopsinae   ( Figs 19E View Figure 19 , 38H View Figure 38 , 42E View Figure 42 , 48B View Figure 48 ).

Intermediate sternum posterior margin: Pointed and truncate sternal margins are quite distinct in the taxa represented here, although the systematic value of this character has been questioned because of imprecision in shape definition, reliability of observation, homoplasy, and possible influence of overall body proportions on sternum shape ( Coddington, 1986a; Griswold et al., 1998; Schütt, 2003). A truncate sternum is a synapomorphy of symphytognathoids, occurring in all families except Mysmenidae   . In Mysmenidae   , an intermediate condition between pointed and truncate posterior sternal margin is consistently found ( Figs 2C View Figure 2 , 7C View Figure 7 , 25B View Figure 25 , 35C View Figure 35 , 46A View Figure 46 , 59G View Figure 59 , 140B View Figure 140 , 143B, I View Figure 143 ); however, an ambiguously optimized reversal to pointed posterior sternum occurs in Maymena   ( Fig. 141F View Figure 141 ), and two independent instances of truncate sternum occur within mysmenines ( Fig. 143O View Figure 143 ).

Sparse imbricate cuticular pattern on carapace border: Almost all mysmenids here examined have a typical cuticle pattern on the carapace lateral edges that was not observed in other taxa. It consists of slender (i.e. not prominent) ridges running mostly parallel with each other and with the edge of the carapace, delimiting elongated scales ( Fig. 50A View Figure 50 ). A smooth cuticle is widespread in both outgroup taxa and symphytognathoids ( Figs 118A, B View Figure 118 , 121B, E View Figure 121 ). Within Mysmenidae   , a smooth cuticle is secondarily and independently present in Maymena mayana   and the mysmenine MYSM-019-MAD.

Cheliceral fang furrow denticles: Minute denticles in the cheliceral fang furrow ( Figs 7G View Figure 7 , 15H View Figure 15 , 48B View Figure 48 ) occur in almost all mysmenids studied (e.g. Forster, 1959; Brignoli, 1974; Thaler, 1975; absent in Microdipoena jobi   ; Platnick & Shadab, 1978; Griswold et al., 1998; Schütt, 2003), and have been proposed as synapomorphic for the family ( Platnick & Shadab, 1978). Although not unique for symphytognathoids (cheliceral denticles have been reported at least in nesticids, Wiehle, 1963; uloborids, Peters, 1982; and araneids and nephilids, Hormiga, Eberhard & Coddington, 1995), similar denticles also occur in Theridiosomatidae   and some anapids ( Coddington, 1986a; see also Schütt, 2003). Within symphytognathoids, denticles are an ambiguously optimized synapomorphy for both Theridiosomatidae   and Mysmenidae   .

Anterior median eyes on protruded area: Another particular feature shared between mysmenids and theridiosomatids is the arrangement of the anterior median eyes. Character optimization under parsimony for this feature is ambiguous. Both sexes of mysmenids (except Trogloneta   , see below) and theridiosomatids have a depression around the anterior median eyes defining a protruded area ( Figs 15B, D View Figure 15 , 25C View Figure 25 , 46D, E View Figure 46 , 59C View Figure 59 ). This area is clearly protruded, not just smoothly raised from the rest of the carapace, and can be best observed with SEM and in frontal view. In Trogloneta   , males have all eyes in a tubercle or a narrow elevation of the ocular area ( Figs 63G, H View Figure 63 , 66A View Figure 66 ).