A new, relictual Antilloides from Mexican caves: first mainland record of the genus and revised placement of the fossil Misionella didicostae (Araneae: Filistatidae).
The Mexican fauna associated with caves is a rich one, and includes several species of Filistatidae (Reddell 1981). This medium-sized spider family includes around 150 species in 19 genera (World Spider Catalog 2017), most of which inhabit arid and semi-arid regions of the subtropics; some species are known to be associated to caves (Brescovit et al. 2016a; Magalhaes 2016). The North American fauna includes three genera but is still poorly known, as extensive taxonomic revisions are lacking and only Filistatinella Gertsch & Ivie, 1936 has been recently revised (Magalhaes & Ramirez 2017).
Among filistatid material collected from limestone caves in south-western Coahuila State (Fig. 1), I found some particularly interesting specimens. At first glance, they seemed to belong to yet another undescribed species of Filistatoides F. O. Pickard-Cambridge, 1899. However, a closer look at their fine morphology suggested that they belonged to the recently described genus Antilloides Brescovit, Sanchez-Ruiz & Alayon, 2016, which was previously recorded only from the Antilles. The aims of this paper are thus to describe this new species, to discuss its generic placement, and to document its unique suite of characters and their implications for the taxonomy of New World Filistatidae.
Specimens of the new species and comparative material of Antilloides and Filistatoides (Appendix 1) were borrowed from the collections of the American Museum of Natural History, New York, USA (AMNH; curator L. Prendini), the Field Museum of Natural History, Chicago, USA (FMNH; P. Sierwald), the Instituto Nacional de Biodiversidad, San Jose, Costa Rica (INBIO; C. Viquez) and the Museum of Comparative Zoology, Cambridge, USA (MCZ; G. Giribet).
Measurements and habitus photos were taken using a Leica M165C stereoscopic microscope and Leica Application Suite 3.6. Female genitalia were digested in a pancreatin solution and examined in lactic acid, and the male palp was cleared in clove oil. Genitalia were photographed using an Olympus BH-2 compound microscope and drawn with the aid of a camera lucida. Multi-focus photo stacks were constructed in Helicon Focus 6. The format of descriptions follows Magalhaes & Ramirez (2017). Exact locations for some Mexican caves were taken from Reddell (1981). For scanning electronic microscopy images, specimens were dehydrated in solutions of increasing concentration of ethanol, critical point-dried, mounted on aluminium stubs using copper adhesive tape, gold-paladium sputter-coated, and examined using a Philips FEI XL 30 TMP scanning electron microscope.
RESULTS AND DISCUSSION
The reduced cymbium, feathery setae, lack of tarsal macrosetae and wide cribellum, among other characters, suggest that the specimens from Coahuila belong to Prithinae, one of the two subfamilies of Filistatidae (see Gray 1995; Magalhaes & Ramirez 2017). The spiders are pale in color and have elongate palps and legs (Figs. 2, 3), characters observed in some Filistatoides but also frequently associated with cavedwelling species. However, a closer inspection of the male revealed it had modifications to metatarsus II, with a retrolateral excavation and a pair of macrosetae (Fig. 4). In Filistatidae, these modifications are so far unknown outside of the South American genera Pikelinia Mello-Leitao, 1946 and Misionella Ramirez & Grismado, 1997 (see Ramirez & Grismado 1997). The male palpal morphology is also modified and quite unlike most known Filistatidae species (Figs. 5, 6). The tibia bears a small projection with strong setae (Figs. 5C-E: MS), as in Pikelinia (see Ramirez & Grismado 1997, figs. 25, 27, 33) and Antilloides (see Brescovit et al. 2016b, figs. 11A, 16G). The bulb is small and globose and covered dorsally by an extension of the cymbium (Figs. 5B, D, 6A: Cy); the latter has a dorsal area devoid of setae and is not fused to the bulb. A small unsclerotized outgrowth near the apex of the embolus is probably a reduced paraembolic lamina (Figs. 5A, 6A: PL). The sperm duct has a single coil and the fundus points ventrally and attaches to a very reduced basal bulb sclerite (Figs. 5G, 6A: F, BBS), as in many prithines. The female has a pair of simple receptacles each with a narrowed 'neck', as in Filistatoides and Antilloides (see Brescovit et al. 2016b, figs. 1B, 9F), but also an enlarged interpulmonary fold, a diagnostic character of the latter genus (Figs. 6-8: IF). Interestingly, the receptacles lack glandular pores, which are restricted to the 'neck' region (Figs. 7-8: N). The body is further covered in feathery setae (Fig. 9A), as in Filistatoides, Antilloides, Pikelinia and relatives (see Ramirez & Grismado, 1997, fig. 80; Brescovit et al. 2016a, fig. 10G).
The phylogenetic relationships among New World filistatid genera, and the limits of some of the genera themselves, are still largely untested or unknown. Existing studies have a limited sampling of taxa and/or characters (Gray 1995; Ramirez & Grismado 1997; Magalhaes & Ramirez 2017) and do not include some of the characteristics mentioned in the previous paragraph. However, the polarity of some of these characters can be inferred by comparison with other filistatid genera, some of which are becoming better-known due to recent redescriptions and revisions of several taxa worldwide (e.g., Zonstein & Marusik 2015, 2016; Brescovit et al. 2016a, b; Magalhaes 2016; Legittimo et al. 2017; Magalhaes & Ramirez 2017). From these revisions, it can be inferred that some characters are of particular interest for placing the new species. For example, the presence of modified setae on a small projection of the male palpal tibia combined with a modified metatarsus II in the male have hitherto been observed only in Pikelinia (Ramirez & Grismado 1997) - a genus that belongs to a clade (along with Misionella and Lihuelistata Ramirez & Grismado, 1997) where the cymbium is fused to the bulb (Ramirez & Grismado 1997). Thus, if the new species belongs to Pikelinia, the presence of an unfused cymbium would represent at least one instance of homoplasy. On the other hand, the new species shows a particular shape of the cymbium: large, anteriorly convex, covering the bulb dorsally, and with an area devoid of setae. We have recently argued (Magalhaes & Ramirez 2017) that an anteriorly concave cymbium (the "horseshoe-shaped cymbium" that Lehtinen (1967) suggested to be diagnostic of Pritha Lehtinen, 1967) is widespread among prithine genera, including Pikelinia and Filistatinella, and is probably a synapomorphy of the subfamily. Thus, an anteriorly projected, convex cymbium covering the bulb is a rare character among the Prithinae, to my knowledge present only in Antilloides (see Brescovit et al. 2016b, figs. 11A-C). The anteriorly elongate interpulmonary fold (termed "uterus externus" by Brescovit et al. 2016b) is also a diagnostic feature of some (though not all) Antilloides species. Finally, the new species lacks pores in the receptacles of the spermathecae; these are restricted to the 'necks'. My own observations of Antilloides females (including A. abeli Brescovit, Sanchez-Ruiz & Alayon, 2016, A. mesoliticus Brescovit, Sanchez-Ruiz & Alayon, 2016, and a morphospecies similar to A. cubitas Brescovit, Sanchez-Ruiz & Alayon, 2016) show that at least these species have the glandular pores restricted to the 'necks' of the spermathecae; this character is unknown to me in other filistatid genera.
Taken together, these observations suggest that this species belongs to Antilloides, the first to be described from mainland North America, as all previously known species are restricted to the Antilles. It is likely that the genus had a wider distribution in the past, and now survives on the Caribbean islands and as a relict in Mexican caves in dry areas. In addition, I report a novel putative synapomorphy for Antilloides (or at least a group of species within the genus): the glandular pores restricted to the 'neck' of the spermathecae. The presence of a modified metatarsus II in the new species - the first to be described outside of Pikelinia and Misionella - suggests that Antilloides is closely related to these genera, or that metatarsal modifications evolved independently several times within Prithinae. Both hypotheses should be tested within a phylogenetic framework that includes a wide sample of taxa and characters.
Finally, the finding of this new Antilloides species with a modified metatarsus II in males calls for a reappraisal of the only fossil Filistatidae known to date, Misionella didicostae Penney, 2005. This fossil comes from amber deposits from the Dominican Republic (where at least two extant species of Antilloides can still be found) and date from the Miocene. Penney (2005a) placed the fossil species in Misionella based on the modified metatarsus II combined with the absence of a tibial apophysis. However, the new extant species here described also combines these characters. Furthermore, in the specimen PRC A-10-255 of M. didicostae, the cymbium is convex and covers almost half of the bulb; setae can be observed along its margin (Fig. 12D, arrow). This indicates M. didicostae also belongs in Antilloides, and a new combination is here proposed.
Family Filistatidae Ausserer, 1867
Genus Antilloides Brescovit, Sanchez-Ruiz & Alayon, 2016 Antilloides Brescovit, Sanchez-Ruiz & Alayon, 2016: 413.
Type species.--Antilloides abeli Brescovit, Sanchez-Ruiz & Alayon, 2016, by original designation.
Diagnosis.--Antilloides males can be recognized by the combination of a retrolateral projection bearing thick setae on the palpal tibia (Figs. 5C, D, F) and a convex, anteriorly drawn cymbium partly covering the bulb dorsally and bearing a glabrous dorsal area (Fig. 5B, D); the metatarsus II may or may not present modifications such as a retrolateral excavation and macrosetae (Fig. 4). Antilloides females are similar to Filistatoides in having a single pair of spermathecae, usually with a narrow 'neck' between the base and the receptacle (Fig. 7); they differ from Filistatoides by having at least one of the two following characters: a large, anteriorly drawn interpulmonary fold covering the spermathecae (Figs. 6D, 8A) (short, not covering the spermathecae in A. cubitas, A. zozo Brescovit, Sanchez-Ruiz & Alayon, 2016 and A. mesoliticus) or glandular pores restricted to the 'neck' and absent from the receptacles (Figs. 7, 8E) (pores present on the receptacles of A. haitises Brescovit, Sanchez-Ruiz & Alayon, 2016).
Antilloides chupacabras sp. nov.
Filistatoides sp. Reddell, 1981: 25.
Type material.--Holotype male. MEXICO: Coahuila: Sierra de Mayran, Cueva de Yeso, [102.8[degrees]W, 25.5[degrees]N], 24 July 1965, J. Fish & J. Reddell (AMNH, IFM-1302).
Paratypes. MEXICO: Coahuila: 6 [female], 1 juvenile, La Cuchilla, Sierra de Mayran, Cueva de Empalme, 24 February 1966, W. Bell & J. Reddell (AMNH IFM-1303, 1304); subadult [female], 50 km E. of Torreon, September 1964, B. Russell (AMNH); 2 [female], Sierra de Mayran, 0.25 miles W. of Cueva de Yeso, Cueva de Los Grillos, 24 July 1965, J. Reddell & J. Fish (AMNH); subadult [male], Sierra San Lorenzo, 4 km E. of Coyote, Cueva del Vapor, [103.15[degrees]W, 25.65[degrees]N], 9 June 1980, J. Reddell (AMNH); 1 [female], Cueva del Guano (AMNH); 1 [female], 4.5 km E. of Coyote, Cueva Granjero (AMNH).
Note.--The labels associated with the specimens have no geographical coordinates, or exact locations other than the cave names. For instance, the label of the holotype reads only "Cueva de Yeso, Coah., MEX. / 7-24-65. J Fish, J. Reddell". I have traced their approximate placement using the list of Mexican caves provided by Reddell (1981; see caves listed under Sierra de Mayran Region, p. 294; see also map on page 18, pinpointing the location of Sierra de Mayran). Reddel collected most of the specimens himself, and was the first to notice that this was an undescribed species, although he listed it under Filistatoides (Reddell 1981).
Etymology.--The 'chupacabras' is the name of a mythological creature from an urban legend that is widespread in Latin America; it holds that this strange, reptilian creature roams around rural areas and feeds by sucking the blood of livestock. The new species is given this name because it has a strange morphological appearance, feeds by sucking the fluids of its prey, and is the only known species to have been separated from its closest relatives in the Antilles. The specific epithet is to be treated as a noun in apposition.
Diagnosis.--Individuals of this species are pale-colored, with reduced pigmentation, and both sexes have more elongate legs than any other Antilloides (femur-carapace length ratios of 3.02 and 2.24 in A. chupacabras sp. nov. males and females, respectively; at most 2.3 and 1.4 in other Antilloides males and females, respectively; Figs. 2-3). Males are easily distinguished from other extant species of Antilloides by the metatarsus II bearing two macrosetae on the retro-lateral face (Fig. 4) and by the long palp with a short bulb (Fig. 5). Females can be distinguished by the deep furrow bearing setae in the genital region (Fig. 3D), and by the internal genitalia, which have slender, straight 'necks' leading to globose receptacles (Figs. 6B-D, 7, 8).
Description.--Male holotype (Figs. 2-4, 6A). Coloration pale yellowish cream except where noted (Fig. 2). Carapace with dark coloration around eyes and a median elongate spot dorsally. Chelicerae orange. Legs pale yellowish cream, except for light orange legs I and II. Abdomen dorsum light cream, with faint brown chevron markings posteriorly. Anterior margin of the carapace unmodified (Fig. 2D). Sternum oval, narrowest posteriorly, sigilla not visible. Total length 5.03. Carapace length 2.30; width 1.85. Clypeus length 0.57. Eye diameters and interdistances: anterior median eye (AME) 0.11; posterior median eye (PME) 0.12; anterior lateral eye (ALE) 0.15; posterior lateral eye (PLE) 0.13; AME-AME 0.03; PME-PME 0.15. Sternum length 1.26; width 1.00. Palp: femur (fe) length 1.67, height 0.25; tibia (ti) length 1.15, height 0.20. Leg I: fe 6.95; patella (pa) 0.85; missing from tibia-tarsus. II: fe 4.50; pa 0.83; ti 5.48; metatarsus (mt) 3.94; tarsus (ta) 1.48. III: fe 3.51, pa 0.81, ti 3.58, mt 3.85, ta 1.49. IV: fe 4.19, pa 0.85, ti 4.03, mt 4.85, ta 2.57. Abdomen: length 2.88; width 1.65. Leg II modified: metatarsus II with a shallow retrolateral apical excavation, a strong retrolateral condyle, and two macrosetae, one more proximal, ventral and parallel to the metatarsus, and the other more distal, dorsal, and pointing dorso-apically (Fig. 4). Leg macrosetae otherwise absent. Palp (Figs. 5, 6A): femur and tibia unusually slender; cymbium convex, covering at least half of the bulb, unfused to bulb (Figs. 5C, D); bulb short and globose, bent slightly upwards distally (Figs. 5G, 6A); sperm duct N-shaped, proximally with a ventral bend (Fig. 6A); paraembolic lamina a small triangular projection situated prolateral to the embolus (Fig. 6A); embolus short. The specimen is in relatively poor condition, with most setae lost, both legs I missing from the tibiae, femora I broken in the midline, left leg III and right leg IV disarticulated from the tibiae, and the left palp dissected.
Female paratype (AMNH, IFM-1303). Coloration as in male except for more yellowish legs and chelicerae (Fig. 3). Anterior margin of the carapace unmodified, eye apodemes present (Fig. 9A). Sternum oval, narrowest posteriorly, sigilla not visible, but detected in another specimen examined under SEM (Fig. 9D). Total length 7.16. Carapace length 2.83; width 2.03. Clypeus length 0.72. Eye diameters and interdistances: AME 0.06; PME 0.10; ALE 0.13; PLE 0.13; AME-AME 0.06; PME-PME 0.17. Sternum length 1.40; width 1.09. Palp: fe length 2.31, height 0.47; ti length 1.65, height 0.34. Leg I: fe 6.35; pa 0.95; ti 8.88; mt 8.07; ta 2.93. II: fe 4.19; pa 0.83; ti 4.83; mt 4.09; ta 1.54. III: fe 3.33, pa 0.91, ti 3.20, mt 3.48, ta 1.49. IV: fe 3.91, pa 0.93, ti 3.70, mt 4.07, ta 1.76. Abdomen: length 4.47; width 2.65. Chelicerae with a posterior row of chemosensory setae, and gland with projected openings (Fig. 10B). Palp and leg macrosetae absent. Tarsal organ capsulate (Fig. 10C), metatarsus stopper narrowed (Fig. 10E). Calamistrum three-rowed (Fig. 10D), missing most setae. Cribellum very wide, with narrow spinning fields (Fig. 11 A), posterior median spinnerets with spatulate setae (Fig. 11D), posterior lateral spinnerets with several aciniform gland spigots (Fig. 11E). Epigastric furrow unsclerotized, with a fold delimiting a depression bearing many setae (Fig. 3D). The specimen is in relatively good condition, but most setae are missing.
Female paratype (AMNH, IFM-1304). Interpulmonary fold enlarged anteriorly, covering the spermathecae, with ragged anterior margin, and laterally subsquarish (Figs. 6D, 7A, 8); with a single pair of receptacles, each connected to the uterus externus through a long and thin spermathecal 'neck' covered by glandular pores; receptacles globose, without glandular pores, set close to each other (Figs. 7B, 8E).
Variation.--Females (n = 5): total length 6.55-7.16 (mean 6.84), carapace length 2.43-3.01 (2.72), femur I length 5.54-6.70 (6.14), tibia I length 7.62-9.22 (8.62), femur/carapace length ratio 2.15-2.41 (2.26). The receptacles in the female genitalia can be rounded (Fig. 7B) to oval (Fig. 7D); subadult females have simpler, smaller genitalia (Fig. 6C).
Natural history.--All of the specimens of A. chupacabras I examined come from caves. They are much more lightly coloured than other species of Antilloides, and also have much longer legs (see Diagnosis). It is tempting to suspect that these aspects represent troglomorphisms, but the lack of epigean records might be a consequence of undercollecting outside the caves. More studies on the region should reveal whether this species also occurs outside caves or is a cave obligate, but for now this species should be considered a potential troglophile.
Reddell (1981: 25) stated that the caves in the Sierra de Mayran are formed by a solution of gypsum beds in Cretaceous limestone, stand at an elevation of ca. 1150 m above sea level, and are very dry, with a fauna typical of Mexican arid areas.
Antilloides didicostae (Penney, 2005), comb. nov. (Fig. 12)
Misionella didicostae Penney, 2005a: 95, figs. 1-5. Male holotype deposited in the Museo del Ambar Dominicano, not examined; Penney 2005b, fig. 1.
Note.--I could not examine the holotype of this species directly, but have examined photos by G. Poinar of a second specimen (Oregon State University, PRC A-10-255), examined and identified by Penney (2005b) shortly after his original description. The palpal morphology (Fig. 12D) fits with the original description. The new combination is here proposed because the examined specimen has a convex cymbium covering the bulb, as do other Antilloides. The two characters originally used for placing the species in Misionella (the modified second metatarsus and lack of tibial apophysis) are also shared with Antilloides chupacabras sp. nov.
I am indebted to L. Prendini, N. Platnick and L. Sorkin (AMNH), P. Sierwald and J. Boone (FMNH), C. Viquez (INBIO) and G. Giribet and L. Leibensperger (MCZ) for making specimens available for this study. Peter Sprouse helped with finding the exact location of some Mexican caves. George Poinar kindly photographed the Misionella didicostae specimen and authorized use of his pictures. I thank D. Penney for suggesting some references and pointing out the existence of the second specimen of M. didicostae. Fabian Tricarico helped with the acquisition of SEM images. Earlier versions of the manuscript have been improved by commentaries from M.J. Ramirez and A. Perez-Gonzalez, and by a careful revision by A. Sanchez-Ruiz, an anonymous reviewer and the editor, M. Rix. I have been supported by a CONICET doctoral fellowship and by grants from CONICET (2012-0943) and FONCyT (2011-1007) to Martin J. Ramirez.
Brescovit, A.D., I.L.F. Magalhaes & I. Cizauskas. 2016a. Three new species of Misionella from northern Brazil (Araneae, Haplogynae, Filistatidae). ZooKeys 589:71-96.
Brescovit, A.D., A. Sanchez-Ruiz & G. Alayon. 2016b. The Filistatidae in the Caribbean region, with a description of the new genus Antilloides, revision of the genus Filistatoides F. O. P. Cambridge and notes on Kukulcania Lehtinen (Arachnida, Araneae). Zootaxa 4136:401-432.
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Ivan L. F. Magalhaes: Division Aracnologia, Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"-CONICET, Av. Angel Gallardo 470, C1405DJR, Buenos Aires, Argentina. E-mail: firstname.lastname@example.org
Manuscript received 29 May 2017, revised 18 August 2017.
Material examined.--The following comparative material of Antilloides and Filistatoides was examined for this study:
Antilloides abeli Brescovit, Sanchez-Ruiz & Alayon, 2016. CUBA: Pinar del Rio: 5 [female], Vifiales, San Vincente, cave entrance, [83.7166[degrees]W, 22.6166[degrees]N], 9 January 1954, L. Ross (FMNH 2857656).
Antilloides mesoliticus Brescovit, Sanchez-Ruiz & Alayon, 2016. CUBA: Pinar del Rio: 5 [female], San Diego de Los Bafios, Cueva de los Indios, [83.36558[degrees]W, 22.64661[degrees]N] (AMNH IFM-1310).
Antilloides morphotype 'IFMsp114' (cf. A. cubitas Brescovit, Sanchez-Ruiz & Alayon, 2016). BRITISH VIRGIN ISLANDS: Virgin Gorda: 3 [female], Baths & Devil's Bay, [64.44336[degrees]W, 18.42663[degrees]N], 25 June 1966, Island Project Staff (AMNH IFM-1285).
Antilloides morphotype 'IFMsp151'. FRENCH WEST INDIES: Saint Barthelemy: 3 [female], [62.83333[degrees]W, 17.9[degrees]N], 2008, K. Questel (MCZ 79820).
Antilloides morphotype 'IFMsp153'. BRITISH VIRGIN ISLANDS: West Dog Seal. 1 [female], [64.43467[degrees]W, 18.50665[degrees]N], Island Project Staff (AMNH IFM-1309).
Filistatoides insignis (O. Pickard-Cambridge, 1896). GUATEMALA: Tijax: 3 [female], [89.26903[degrees]W, 15.37521[degrees]N], C. Viquez (INBIO IFM-1189).
Filistatoides morphotype 'IFMsp23'. MEXICO: Nuevo Leon: 1 [male], 9 [female], 5 juveniles, near Mt. El Candela, Gruta del Carrizal, in darkness, [100.57037[degrees]W, 26.77644[degrees]N], B. Russell (AMNH IFM-1282).
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|Author:||Magalhaes, Ivan L. F.|
|Publication:||The Journal of Arachnology|
|Date:||May 1, 2018|
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