Revision of recluse spiders (Araneae: Sicariidae: Loxosceles ) preserved in Dominican amber and a total-evidence phylogeny of Scytodoidea reveal the first fossil Drymusidae

Revision of recluse spiders (Araneae: Sicariidae: Loxosceles ) preserved in Dominican amber and a total-evidence phylogeny of Scytodoidea reveal the first fossil Drymusidae. Arthropod Abstract Recluse or violin spiders in the genus Loxosceles (Scytodoidea: Sicariidae) are a diverse group (~140 extant species) including medically important species and distributed mainly in the Americas, Africa, and the Mediterranean region. In addition, this genus includes three fossil species from Miocene Dominican amber. Here we revise the taxonomy of these fossil species by examining, imaging and re-describing their type specimens. We find that L. defecta Wunderlich, 1988 and L. deformis Wunderlich, 1988 are bona fide members of the genus and report additional characters overlooked in their original descriptions. We further study the holotype of L. aculicaput Wunderlich, 2004 using synchrotron radiation micro-computed tomography to reveal previously unknown morphological details hidden by fissures in the amber. We found several characters inconsistent with Loxosceles but consistent with Drymusa (false violin spiders; Scytodoidea: Drymusidae), such as three claws, well-developed podotarsite, and a broad colulus. This suggests the species is misplaced in Loxosceles . To test this hypothesis, we estimated a total-evidence phylogeny of the superfamily Scytodoidea including extant and fossil taxa, morphological data, traditional molecular markers, and sequences of ultra-conserved elements. The results show unambiguously that L. aculicaput belongs to Drymusa and is a close relative of extant species of the genus inhabiting the Greater Antilles. Therefore, we here transfer this species to Drymusa , establishing a new combination and new family assignment. Drymusa aculicaput comb. nov. represents the first known fossil Drymusidae and shows that crown members of this genus already existed in the Miocene.


Introduction
Spiders are a speciose clade of predators containing ~50000 species (WSC 2022), which play a key role in terrestrial food webs (Nyfeller and Birkhofer 2017). Despite their diversity, abundance, and ubiquity, they have delicate bodies, and thus, they only fossilize in exceptional conditions (Selden and Penney 2010). In fact, over three-quarters of the ~1400 species of fossil spiders are preserved as inclusions in amber coming principally from three deposits in northern Myanmar (Late Cretaceous), the Baltic Sea (Eocene), and the Dominican Republic (Miocene) (Dunlop et al. 2020;Magalhaes et al. 2020). While it has been shown that arthropods entrapped in natural resins are only a subset of the total community present in the environment (Solórzano Kraemer et al. 2018), fossils preserved in amber provide an important glimpse into the past diversity of spiders. Miocene amber from the Dominican Republic is the major source of spider fossils in the Neotropical region. Around 170 currently valid species have been named from this deposit (Dunlop et al. 2020), most of them (~120) described in the monographic treatment of this fauna by Wunderlich (1988) (for a historical account on the taxonomy of fossil spiders from Dominican amber see Selden and Penney 2010: 181). Penney and Pérez-Gelabert (2002) compared the fossil fauna with the composition of extant Hispaniolan spiders. While a poor knowledge of extant spiders hampers a more detailed comparison, they showed that many families and genera are shared between the faunas, showing that the deposit represents a typical tropical assemblage. Interestingly, some families and genera are known from the islands only from fossils, prompting the hypothesis that extant members might be present in Hispaniola but remain undiscovered (Penney 1999). This is certainly more probable for specimens described in Holocene copal or Defaunation resin from the Dominican Republic (e.g., Wunderlich 1986) since these resins can have an age of only 60 years BP (conventional radiocarbon age) (Solórzano- Kraemer et al. 2020). For some families, this hypothesis has been confirmed, such as for Filistatidae, initially known from Hispaniola by fossil specimens (Penney 2005) but later showed to include extant Dominican taxa endemic to the island (Brescovit et al. 2016). Some families with extant Hispaniolan taxa have hitherto not been recorded in amber, such as Drymusidae, which includes Drymusa simoni Bryant, 1948from Haiti (see WSC 2022 and unidentified species from the Dominican Republic (Solanlly Carrero, pers. comm.). This spider family has so far no fossil record at all (Dunlop et al. 2020).
Dominican amber yields the only fossils known for a few spider families, such as Sicariidae. Sicariids include six-eyed sand spiders (Sicarius Walckenaer, 1847 andHexophthalma Karsch, 1879) and recluse or violin spiders (Loxosceles Heineken et Lowe, 1832). The family belongs to Synspermiata, a major spider clade that is well-represented in Cretaceous amber (Magalhaes et al. 2020), and molecular clocks estimate that sicariid genera originated in the Cretaceous (Binford et al. 2008; Magalhaes et al. 2019). Despite that, the family has only three known fossils preserved in middle Miocene Dominican amber: Loxosceles defecta Wunderlich, 1988, L. deformis Wunderlich, 1988and L. aculicaput Wunderlich, 2004. Violin spiders comprise ~140 described species distributed mainly in the Americas, Africa, and the Mediterranean region (WSC 2022), ranking among the most speciose spider genera. They are infamous because of their medical importance (Vetter 2008), but are also important models for biogeography (e.g., Binford et al. 2008;Planas and Ribera 2014). A better knowledge of the three Loxosceles fossil species would be desirable since they provide the only suitable calibration point for estimating the dated phylogenetic trees of this family. Because they are only known from their original descriptions (Wunderlich 1988(Wunderlich , 2004, we here strive to revise their morphology and taxonomy.
During a preliminary examination of the holotype of L. aculicaput, we found that some of its characters are inconsistent with a placement in Loxosceles, such as the presence of three tarsal claws and a well-delimited podotarsite (the genus has only two tarsal claws and a poorly delimited podotarsite; see Labarque and Ramírez 2012). These observations suggested that this species was misplaced in Loxosceles. Unfortunately, the holotype is preserved in a piece of amber with large fissures that prevent a clear examination of its ventral side, including the palps, the morphology of which has a high taxonomic value. Recently, the use of X-ray micro-computed tomography (µ-CT) and synchrotron radiation micro-computed tomography (SRµ-CT) has been employed to enhance taxonomic descriptions of fossils (e.g., Penney et al. 2007Penney et al. , 2012Saupe et al. 2012) and to reveal morphological details hidden in the piece, or even in completely opaque amber, allowing a better assessment of the systematic placement of the fossil (e.g., Soriano et al. 2010;Dunlop et al. 2011;Azevedo et al. 2021;Solórzano Kraemer et al. 2011, 2015, 2022. Thus, the use of this technique might help shed light on the phylogenetic placement of L. aculicaput. Recently, systematic advances in spiders have relied heavily upon genomic-scale molecular data (e.g., Kulkarni et al. 2020;Ramírez 2021). While this approach has proved to be a powerful tool for resolving recalcitrant relationships, most of these studies include only molecular data in their phylogenetic matrices. There are legacy morphological matrices that may be used alongside the newly collected molecular data (e.g., Griswold et al. 2005;Labarque and Ramírez 2012;Ramírez 2014). This total evidence approach is useful in testing the placement of taxa for which molecular data is unavailable, such as fossils (e.g., Wood et al. 2013;Wood 2017;Mongiardino Koch et al. 2021;Azevedo et al. 2021;Magalhaes and Ramírez 2022). We thus expect that a phylogenetic analysis including molecular and morphological data is the most straightforward way to test the phylogenetic relationships between the fossils treated here and the extant members of Scytodoidea.
The aims of this contribution are: (1) to revise the taxonomy of Dominican amber Loxosceles, re-illustrate the type specimens and re-assess their morphology; and (2) to test the phylogenetic placement of the fossils using a dataset containing morphological and sequence data for representatives of all extant Scytodoidea families. Amber pieces were re-polished at the SMF using a Phonix Beta polishing machine with grinding paper for metallography, wet and dry: Grip 2500 and 4000. After polishing, pieces were embedded in Araldite 2020® Epoxy resin following Sadowski et al.'s (2021) recommendations.

Extant material examined for comparison
The following specimens have been examined for the new scorings of the morphological matrix, or to discuss the morphology of scytodoids. We scored Ochyrocera diablo Pérez-González, Rubio et Ramírez, 2016

Light microscopy
The photographs and Z-stacks images of fossil species were taken under a Nikon SMZ25 microscope, using Nikon SHR Plan Apo 0.5× and SHR Plan Apo 2× objectives with a microscope camera Nikon DS-Ri2 and the NIS-Element software (version 4.51.00; www.microscope.healthcare.nikon.com). In some cases, to prevent diffraction caused by irregular surfaces of the amber piece, we placed a coverslip and a drop of water with sugar on top of the piece to flatten the surface to be photographed. Morphological observations on extant specimens were made using Leica M165 C and Leica M125 stereomicroscopes. Pictures were taken with Nikon DXM1200 digital camera mounted on a stereoscopic microscope Nikon SMZ1500 and on a microscope Leica DM4000 M, and with a Leica DFC 500 digital camera mounted on a stereoscopic microscope Leica M165 C or Leica M216. Extended focal range images were composed with the Leica Application Suite version 3.6.0. or Helicon Focus 3.10.3-4.62 (https://www.heliconsoft.com, Ukraine). Pre parations were carefully cleaned using fine brushes and a thin jet of alcohol from a thinned pipette; some setae were removed to expose structures, especially those on legs, palps, spinnerets, and chelicerae.

Scanning electron microscopy
For scanning electron microscope (SEM), all preparations were dehydrated in a series of increasing concentrations of ethanol (80%, 90%, 95%, 100%), and critical-point dried. After drying and brushing, they were mounted on adhesive copper tape (Electron Microscopy Sciences, EMS 77802) affixed to a stub and secured with a conductive paint of colloidal graphite on isopropyl alcohol base (EMS 12660). Prior to SEM examination under a high vacuum with a FEI XL30 TMP or a LEO 1450VP, the structures were sputter-coated with Au-Pd.

Synchrotron radiation microcomputed tomography
The imaging of the holotype of L. aculicaput was performed at the Imaging Beamline -IBL P05 -PETRA III at Deutsches Elektronen Synchrotron (DESY) in Hamburg, operated by the Helmholtz-Zentrum Hereon (Greving et al. 2014;Wilde et al. 2016). The specimen was imaged at a photon energy of 18 keV using a commercial CMOS camera system with an effective pixel size of 1.28 µm.
The sample to detector distance was set to 3 cm. For each tomographic scan, 3601 projections at equal intervals between 0 and π were recorded. Tomographic reconstruction was done by applying transport of intensity phase retrieval approach and using the filtered back-projection algorithm (FBP) implemented in a custom reconstruction pipeline (Moosmann et al. 2014) using Matlab (Math-Works) and the Astra Toolbox (Palenstijn et al. 2011;van Aarle et al. 2015;van Aarle et al. 2016). For processing, raw projections were binned two times, resulting in an effective pixel size of the reconstructed volume of 2.56.
The complete specimen was segmented in three dimensions using region-growing techniques in VGStudioMax (version 3.3.1 www.volumegraphics.com/de, Volume Gra phics, Heidelberg, Germany). We carried out the segmentation and volume rendering of the pedipalps in AMIRA5.4.5 (FEI Visualization Science Group, Burlington, MA, USA). All images were post-processed to adjust contrast and sharpness using Photoshop CS6 (Adobe Inc., San Jose, CA, USA).

Phylogenetic analyses
We augmented the morphological matrix of Scytodoidea families (Labarque and Ramírez 2012) by including the following taxa: Ochyrocera diablo, Althepus maechamensis, Loxosceles defecta, and L. aculicaput. We added the first two to include all extant Scytodoidea families in the sampling, and the latter two to test their phylogenetic position. Loxosceles deformis was not included because the only known specimen is poorly preserved and most characters needed in the matrix are not observable.
To maximize morphological and genetic data overlap, we merged data from closely related species into a single terminal in the outgroup genera Stedocys, Ariadna, Althepus, and Ochyrocera (Table 1) We matched the assembled sequences to a compilation of UCE probes merging the Arachnid probe set (Starrett et al. 2017), and the Spider probe set (Kulkarni et al. 2020). Sequences were aligned using MAFFT v. 7.455 and trimmed with GBLOCKS v. 0.31b (Castresana 2000), with the default parameters implemented in PHYLUCE. The recovery of UCE markers did not work well with the few taxa represented in our analysis, probably because few markers were shared by several species; this small dataset failed to recover the monophyly of well-established groups, such as the genus Loxosceles. We then used a wider sample of 100 short-read archives from 75 species of Synspermiata and outgroups (Table 1), which recovered 1912 markers with a minimum coverage of 10%, and a total length of 510,125 sites. From this data-set, we selected the target species, which were analyzed together with the target-gene markers and morphological data.
The phylogenetic analysis under maximum likelihood was done with IQ-TREE v. 2.1.3 (Minh et al. 2020), selecting models for each target gene and the concatenated UCE markers by Bayesian information criterion with ModelFinder (Kalyaanamoorthy et al. 2017), and estimating support with 1000 rounds of ultrafast bootstrap (Hoang et al. 2018). The models selected were: UCE GTR+F+I+G4, 12S TIM2+F+G4, 16S TIM2+F+I+G4, 18S TNe+R3, 28S TIM3+F+R3, COI TIM3+F+I+G4, H3 K2P+R2. We removed the invariant characters of the morphological partition and replaced polymorphic scor- ings with missing entries. Morphological data were analyzed afterwards as two separate partitions, one for 74 unordered characters, treated with the Mk model, and the other for 5 ordered characters, with the Ordered model; both partitions were corrected for the absence of invariant characters with the Asc model. The analysis under maximum parsimony was done with TNT v. 1.5 (Goloboff and Catalano 2016), under equal weights; since 100 out of 100 heuristic searches of 1 RAS followed by TBR branch swapping reached the same tree and length, it is likely that the optimal tree was found. We estimated branch support with 1000 rounds of bootstrap. Synapomorphies were calculated with TNT.

Data resources
The supplementary figures and data underpinning the analyses reported in this paper are deposited in the Zenodo repository at https://zenodo.org/record/6954956 under www.doi.org/10.5281/zenodo.6954956.

Imaging results
We present new photographs under light microscopy of the holotypes of Loxosceles defecta (Fig. 1), L. deformis ( Fig. 2) and L. aculicaput (Fig. 3). This revealed some details that had been overlooked in the original descriptions, such as a third tarsal claw in L. aculicaput (Fig. 3E-F), or the modified legs, bifid cheliceral lamina and stridulatory files in the chelicerae of L. defecta (Fig. 1D-F).
The phylogenetic significance of these observations is discussed below (see 4.2.). In the case of L. aculicaput, a large fissure in the amber prevented observation of the ventral aspects of the specimen, and thus only the dorsal side could be imaged (Fig. 3), hampering a detailed examination of some taxonomically important structures, such as the palps (Fig. 3G).
After scanning the amber piece containing the holotype of L. aculicaput, we rendered a 3D reconstruction of the specimen (Figs 4, 5) that revealed additional details of key ventral structures of the spider, such as a notched labium (Fig. 4C, inset), a broad colulus (Fig. 5A, B) and the shape of both pedipalps (Fig. 5D-J). It also allowed better visualization of the spinnerets, which are disposed of in a compact group (Fig. 5B); the ALS are about the same length as the PLS (Figs 3D, 5C). The resolution did not resolve some of the smaller structures, such as the number of tarsal claws (although these can be seen in Fig.  3E, F), the presence and number of cheliceral teeth, or the presence of a double row of teeth in the first and second prolateral claws of the legs. The 3D rendering of the genitalia shows some aberrant structures (marked with an asterisk in Fig. 5) that we interpret as artifacts (perhaps thin layers of air surrounding the specimen) since each is only present in a single palp and similar structures are lacking in related species (see Fig. 7).

Phylogenetic analyses
The target-gene and UCE datasets produced similar trees (Figs S1, S2), with higher support in the last case, as expected. The combination of all sequence data (Fig. S3) produced a tree with all the groups in agreement with the UCE analysis. The morphological dataset produced trees with the same resolution for Sicariidae, Scytodidae, Periegopidae, and Drymusidae, but discordant with the molecular data in the more basal splits, also differing between maximum likelihood (ML) and maximum parsimony (MP) analyses (Fig. S4). The ML and MP analyses of the complete dataset produced very similar results, only differing in three polytomies of the MP tree, that were resolved in the ML tree ( Fig. 6; Fig. S5). Because these resolved branches in the ML tree had bootstrap below 75% and no molecular or morphological synapomorphies, we base our subsequent analyses on the phylogeny with those branches collapsed (Fig. 6). The morphological matrix, DNA alignments, and the total evidence tree found under ML are available as Supplementary material 2, 3, and 4, respectively.
The total evidence analysis (Fig. 6) recovered a monophyletic Scytodoidea including Ochyroceratidae, Psilo-dercidae, and Scytodidae as three successive splits in the base of the superfamily. Sicariidae is recovered including Sicariinae (Hexophthalma + Sicarius) and two extant species of Loxosceles grouped with the fossil L. defecta. This family is recovered as sister to Periegopidae + Drymusidae. Within Drymusidae, the South African Izithunzi Labarque, Pérez-González and Griswold, 2018 is recovered as the sister group to a clade containing American Drymusa Simon, 1892 and Loxosceles aculicaput. This latter species is recovered in a clade of Antillean species (D. armasi Alayón, 1981 and D. spectata Alayón, 1981 from Cuba).
Sicariidae is supported by five unambiguous morphological apomorphies, only one of which is observable in the fossils (third tarsal claw absent; Fig. 1G

Taxonomy
All three species are redescribed and re-diagnosed below. Based on the phylogenetic results, we propose the transfer of Loxosceles aculicaput from Sicariidae to Drymusa in Drymusidae, resulting in Drymusa aculicaput (Wunderlich, 2004) comb. nov. and new family assignment.

Relationships among Scytodoidea families
The phylogenetic results we obtained are consistent with those of the most recent phylogeny with a broad sampling of Synspermiata families (Ramírez et al. 2021; ultra-conserved elements), although that study lacked Ochyroceratidae in its sampling. Our findings partially contradict the results found with morphology by Labarque and Ramírez (2012), who recovered Scytodidae (rather than Sicariidae) as closer to Periegopidae + Drymusidae; because our dataset includes a morphological matrix almost identical to Labarque and Ramírez's, it seems that the signal of the massive number of molecular markers (1912 markers) overrides that of morphological data (79 variant characters). Our results also contrast with those of Labarque et al. (2018; Sanger sequences) and Li et al. (2020;transcriptomes), who obtained Ochyroceratidae as sister to Scytodidae. It seems the phylogenetic position of ochyroceratids and psilodercids is far from settled (e.g., see the low bootstrap below Althepus maechamensis in the parsimony analysis, Fig. S5), although the evidence that they are indeed members of Scytodoidea is mounting. Discussing synapomorphies for the extended Scytodoidea (including Ochyroceratidae and Psilodercidae) is difficult since the morphology of both families needs further study. We recovered a single morphological synapomorphy for Scytodoidea (including Ochyroceratidae and Psilodercidae), the fused 3 rd abdominal entapophyses. This might be an artifact of our taxon sampling, since some members of the Pholcoidea (the sister group of Scytodoidea) present this character state as well; also, it is open to interpretation, since the entapophyses are vestigial in Ochyroceratidae, Psilodercidae and Pholcoidea.
The following node (Psilodercidae + the remaining Scytodoidea families) has a single synapomorphy: the simple (as opposed to branched) lateral tracheae in the posterior respiratory system.

Phylogenetic affinities of fossil species
We tested the position of Loxosceles defecta in the phylogeny of Scytodoidea and found it to be a true member of Loxosceles, supported in our analysis by the bifid cheliceral lamina (Fig. 1D). Other characters are also consistent with this genus, such as the absence of a third tarsal claw (Fig.  1G). Loxosceles is divided into several species groups (see Gertsch 1967, Gertsch andEnnik 1983) and we strived to place the fossil into one of those groups by comparing its morphology to that of extant species (L. simillima, L. hirsuta, L. laeta, L. rufescens, L. deserta, L. cubana, and L. caribbaea, each belonging to a different clade; see Binford et al. 2008, Magalhaes et al. 2019. The cheliceral stridulatory files of L. defecta consist of relatively deep and well-delimited ridges (Fig. 1D) when compared to the shallow scales that had been documented for L. rufescens (see Labarque and Ramírez 2012: fig. 16E), and thus this was a potentially useful character. However, examination of several other Loxosceles revealed that deep ridges are the most common state in this genus, and only L. rufescens (among the species examined) presents shallow scales. Another potential synapomorphic state is the prolaterally expanded cymbium (Fig. 1I, arrow). Gertsch and Ennik (1983: 280) noted the "tarsus broadly lobed on the prolateral side" as diagnostic of the L. reclusa species group.
Confirming this, we only observed this character state in L. reclusa, L. cubana, L. taino, L. caribbaea and L. defecta, suggesting it might be a synapomorphy of the reclusa group and indicating the fossil species belongs here (the prolateral shape of the cymbium is not observable in L. deformis, see Fig. 2D-F). Finally, L. defecta shows a few unusual features: the first tibia is sinuous, distally incrassate, and bears macrosetae (Fig. 1E-F fig. 34), but it apparently belongs to the distantly related amazonica group, although its phylogenetic position remains untested. Thus, it might be that this character state evolved independently in L. carinhanha and L. defecta. Regarding the strong macrosetae, the only species examined by us whose tibia bears strong macrosetae is L. cubana (Fig. 8). Loxosceles caribbaea and L. taino (examined by us) and other members of the reclusa group from the mainland (A. Valdez-Mondragón, pers. comm.) lack such tibial macrosetae, although a few species present strong macrosetae in the first femur. Thus, the strong macrosetae in the tibia may be a synapomorphy uniting L. defecta with L. cubana, and indicate the fossil could be more closely related to the Antillean species of the reclusa species group. We could not include Loxosceles deformis in the phylogeny, since the only known individual is poorly preserved and details of the legs, chelicerae, and spinnerets cannot be observed. However, the genitalia is similar to that of L. defecta, and it has the flattened, ribbon-shaped embolus that is common in the Loxosceles reclusa species group (see Gertsch and Ennik 1983). It also lacks the third claw and has an elongate palpal femur that is synapomorphic of Loxosceles (see Magalhaes et al. 2017a). Thus, we conclude that it is a true member of the genus, most likely in the Loxosceles reclusa species group.
We here transfer Loxosceles aculicaput to Drymusa. This species presents three tarsal claws (Fig. 3F), a well-developed podotarsite (Fig. 3F), relatively short anterior lateral spinnerets (Fig. 3D), and the spinnerets are in a compact group (Fig. 4B). All of these characters dispute a placement in Loxosceles, which presents only two tarsal claws in a poorly developed podotarsite (Labarque and Ramírez 2012: fig. 5A, B), relatively long anterior spinnerets, and a diastema between the anterior and posterior spinnerets (Magalhaes et al. 2017b: fig. 1). Our phylogenetic results (Fig. 6) suggest a placement in Drymusa instead, particularly close to extant Antillean species, based on three characters: the broad colulus (Figs 5A, 7I), the femoral macrosetae (Figs 3C, 7A) and the notched labium (Figs 4C, 7B). Interestingly, two of these characters were revealed by imaging the piece with SRµCT, highlighting the usefulness of this technique in the study of fossils. The genitalia of Drymusa aculicaput comb. nov. are here studied in detail for the first time ( Fig. 5D-J). They are similar to other species of Drymusa, including the Antillean representatives, in the short, incrassate tibia and the cymbium with a small apical extension (see Fig. 7D-F), but are clearly different from the extant Hispaniolan species Drymusa simoni (see 5.2.1). Penney (1999) observed that extant, endemic species of Drymusa can be found today in Hispaniola and that fossils would shed light on the timing of arrival of this group to the islands. The discovery that Drymusa aculicaput comb. nov. belongs in this genus indicates the genus arrived at Hispaniola during (or before) the Miocene. It is also a crown Drymusa, showing the genus had diversified by this epoch, and thus that it is an ancient group. It is not unlikely that the separation between the African Izithunzi and the American Drymusa pre-dates the separation of these continents, as is the case of Sicarius and Loxosceles (Binford et al. 2008;Magalhaes et al. 2019). A more complete phylogeny of drymusids, building on previous efforts by Labarque et al. (2018), will be useful to put this hypothesis to test, and will certainly benefit from the new morphological data on D. aculicaput comb. nov. gathered here.

Conclusions
Our phylogenetic and taxonomic revision of spider taxa preserved in Dominican amber and previously assigned to Loxosceles allows us to conclude: (1) Loxosceles defecta and L. deformis are bona fide members of this genus, and L. defecta can be further placed in the L. reclusa species group based on the prolaterally expanded palpal cymbium; this species also presents an incrassate first tibia bearing macrosetae. Tibial macrosetae are also present in the extant species Loxosceles cubana, suggesting that L. defecta may be closely related to extant taxa from the Antilles.
(2) Loxosceles aculicaput presents three claws and shares a broad colulus, femoral macrosetae, and a notched labium with extant members of Drymusa inhabiting the Antilles, and thus was misplaced in Loxosceles; we here propose the new combination Drymusa aculicaput, representing the first known fossil of Drymusidae. This extends the stratigraphic range of crown Drymusa to the Miocene and indicates these spiders had already reached the islands by then.   Preservation. Both spiders are preserved in the same reddish yellow amber piece. Both individuals are partly incomplete and preserved in moderate condition. In both spiders the abdomen is severely shrunk. The holotype has both palps (right palp slightly deformed, with a bubble trapped in the tibia), while the paratype has none. The holotype is missing right leg I, right leg IV and parts of right legs II and III; left leg I is detached from the body and preserved to the left of the animal. The paratype has all the legs on the right side but they are distinctly deformed and covered by emulsion; only part of femur IV remains in the left side. Syninclusions: a large leg of an entelegyne spider, a leaf, and a Collembola.

Diagnosis.
Loxosceles defecta can be diagnosed from the other Loxosceles in Dominican amber (L. deformis), as well as from extant Antillean species, by the sinuous, distally incrassate first tibia bearing macrosetae (Fig. 1E, F); the palp has a gently curved, flattened embolus that is ~1.5 times as long as the globose base of the bulb and is less flattened than that of L. deformis.
Description. Male holotype (SMF-Be 970a). Structure: Carapace slightly wider than long, narrowed anteriorly, particularly hirsute in the cephalic region, with a well-developed fovea. Leg I with sinuous tibia, distally incrassate, with ~5 visible macrosetae in the retrolateral face of the incrassate portion (Fig. 1E); third claw absent. Palps (Fig. 1H Preservation. The spider is completely and well-preserved in an orange piece of amber. The abdomen is slightly shrunken and shriveled but no other structures are obviously deformed. The amber piece has large fissures in the portion ventral to the spider. The piece has been embedded in artificial resin. There are no syninclusions other than some pieces of unidentifiable plants and detritus. Diagnosis. Drymusa aculicaput comb. nov. can be distinguished from other scytodoids preserved in Dominican amber, as well as from extant congeners, by the palp with short, incrassate tibia and by the bulb with a thin, straight, needle-like embolus (Fig. 5E, G, H, J).

Competing interests
The authors have declared that no competing interests exist.

Acknowledgements
We are indebted to curators and curatorial staff from the collections holding the material studied in this revision. We thank Alejandro Val-