Neoscotolemon spinifer (Packard, 1888) comb. rest.

(Figs 21–28, Table 4)

Phalangodes spinifera Packard 1888: 52, pl. 13, figs 2, 2a–c; Banks 1893: 151

Scotolemon spinifera: Banks 1901: 672.

Scotolemon spinigera: Banks 1904: 140 (misspelling).

Neoscotolemon spinifera: Roewer 1912a: 150.

Neoscotolemon spinifer: Roewer 1923: 113.

Rula spinifera: Goodnight & Goodnight 1942b: 13, figs 43–45; 1945: 64.

Stygnomma spinifera spinifera: Goodnight & Goodnight 1951: 9, figs 15–18; 1953: 177, fig. 4; Edgar 1966: 355 (misidentification).

Stygnomma spinifera: Rambla 1969: 391; Duffield et al., 1981: 446, fig. 5; Edgar 1990: 546, fig. 19.44; Peck 1975: 308, 1992: 45 (misidentification?); Acosta et al., 1993: 27; Hounsome 1994: 311, 320 (misidentification?); Peck 1999: 375 (misidentification?).

Stygnomma spiniferum spiniferum: Kury 2003: 236.

Citranus marquesas Goodnight & Goodnight 1942b: 4, figs 7–9 (synonymy established by Goodnight & Goodnight 1951).

Type material. Phalangodes spinifera Packard, 1888: Holotype: major ♂ (MCZ 39047, examined), UNITED STATES OF AMERICA, Florida, Key West or Tortugas [Packard 1888: 53 stated: “The present species [referred to P. spinifera] was collected by us either in Key West or Tortugas, Florida, probably the former locality.”]. Citranus marquesas Goodnight & Goodnight, 1942: Holotype: ♀ (AMNH, examined), UNITED STATES OF AMERICA, Florida, Marquesas Key, under trash palm tree, 23-Jun-1938, George Van Hyning leg. Paratype: ♀ (AMNH, not examined) UNITED STATES OF AMERICA, Florida, Barracuda Key, June 13-Jun-1938, George Van Hyning leg.

Remark. The holotype of Phalangodes spinifera Packard, 1888, was originally designated, incorrectly, as a female specimen (Packard 1988: 52), and the holotype of Citranus marquesas Goodnight & Goodnight, 1942, was originally designated, incorrectly, as a male specimen (Goodnight & Goodnight 1942b: 4).

Other material examined. UNITED STATES OF AMERICA: 1 ♀ (AMNH), Florida, Key Largo, 1-Apr-1957, W.J. Gertsch & R. Forster leg. • 2 ♀ (AMNH), Florida, Miami-Dade, 7 miles west of Florida City, 31-Mar-1957, R. Forster & W.J. Gertsch leg. • 1 ♀ (AMNH), Florida, Florida Keys, Little Duck Key [24.4°; -81.15°], Jean and Wilton Ivie leg. • 1 juvenile (AMNH), Florida, Key Largo [25°; -80°], 30-Jan-1959, H.A. Denmark leg. • 1 ♂ (AMNH), Florida, Miami-Dade. • 1 ♀ (AMNH), Florida, Miami-Dade, Homestead, 11-JUN-1930, N.W. Davis leg. • 2 major ♂ (one photo voucher) 1 ♀ (photo voucher) (AMNH), Florida, Miami-Dade, Homestead, fall 1930, J.B. Tower leg. • 1 ♀ (AMNH), Florida, Miami-Dade, Homestead, Rattlesnake Hannsand [?], 2-Jun-1941. • 1 major ♂ (AMNH), Florida, Miami-Dade, Everglades National Park, Royal Palm Hammock, hardwood hammock, malaiseFIT, 28- Jul / 15-Nov-1985, S. & J. Peck leg. • 1 ♀ (AMNH), Florida, Monroe, Key Largo, Pennekamp State Park, hammock forest leaf litter Ber., 31-Aug-1985, S. & J. Peck leg. • 1 major ♂ (AMNH), Florida, Monroe, Big Pine Key, Watson’s Hammock, hardwood hammock, malaise-FIT, 3- May / 31-Aug-1985, S. & J. Peck leg. • 1 minor ♂ (photo voucher) (AMNH), Florida, Monroe, Big Pine Key, Watson’s Hammock, hammock for malaise-FIT, 3- Jun / 27-Aug-1986, S. & J. Peck, leg. • 1 ♀ (AMNH), Florida, Tamiami Trail, 1-Mar-1936. • 1 major ♂ (AMNH), Florida, Miami-Dade, Everglades National Park, Royal Palm State Park, 28-Dec-1940, A. F. Archer leg. • 1 ♂ (SEM voucher), 2 ♀ (USNM), Florida, Everglades, Rowdy Bend [25.1747°, -80.90428°], 6-Jun-2013, CarBio Team leg.

Comparative diagnosis. Major males of Neoscotolemon spinifer differs from males of all other species of Neoscotolemon (except from major males of N. tancahensis) by the presence of enlarged pointed setiferous tubercles on the lateral regions of free tergite III (Figs 22A; 23A; 24A). Males of Neoscotolemon spinifer are easily distinguished from N. armasi by the absence of a pseudochela in the enlarged tarsus of the pedipalp (Figs 22B, C; 23D, E; 25A, B vs. Fig. 8E). Additionally, N. spinifer can be separated from N. pictipes by the presence of a medial long pointed setiferous tubercle on free tergites I–III (Figs 22A, C; 23A, B; 24A, E vs. Figs 1A; 2A, D). Another distinguished feature is that Neoscotolemon spinifer has scattered granules on the dorsal surface of the pedipalp tibia (Figs 23D, E; 25B), whereas N. pictipes has a smooth pedipalp tibia (Fig. 3A, B); this characteristic is also useful for differentiating N. spinifer from its most morphologically similar species, N. tancahensis, which lacks the long, pointed medial setiferous tubercles on each free tergite exhibited by N. spinifer (Figs 22A, C; 23A, B; 24A, E vs. Figs 29A, C; 30A, B; 31A, D).

Redescription. Major male (holotype, AMNH; 1 male AMNH; 1 male USNM). Body measurements: Total body length 3.14, carapace length 1.00, scutum magnum length 2.46, carapace maximum width 1.51, abdominal scutum maximum width 2.20. Appendage measurements in Table 4.

Dorsum: Outline slightly hourglass-shaped with an Eta (η) shape, with a constriction posterior to eyes level (Figs 22A; 23A; 24A). Carapace granulated, wider than long; frontal hump not well marked; anterior border slightly convex, each lateral corner with three small conical tubercles (Figs 23A; 24A). Cheliceral sockets not marked (Fig. 24A). Eyes separated, slightly posterior to the medial region of the carapace and located at the base of a poorly defined ocularium; ocularium with a wide base and apically armed with a long spiniform apophysis slanted forward, with small setiferous granules on the base (Fig. 24E, F) and a row of small setiferous granules on the posterior region (Fig. 23B); ocularium extends from the posterior to just before the frontal hump (Figs 22C; 23B; 24E, F). Abdominal scutum in lateral view convex (Figs 22C, 23B, 24E). Sulcus I deep and well-marked, in dorsal view curved to the posterior body region (Figs 22A, C; 23A, B; 24A, E, F). Mesotergal areas coarsely granulated and not well defined (Figs 23A, B; 24A, E). Mesotergal areas I–II with small medial conical setiferous granules; mesotergal areas III–IV with a row of small conical setiferous granules, with medial granules slightly longer than lateral ones (Figs 23A, B; 24A, E). Mesotergal area V with a posterior row of conical setiferous granules, the medial granule slightly longer (Figs 23A, B; 24A, E). Lateral borders with two rows of granules, the inner row consisting of setiferous granules (Fig. 24A, E). Ozopore with an oval, narrow, and elongated orifice with a descending channel extending toward the posterior region (Fig. 24E). Free tergites granulated; free tergites I–II with a posterior row of conical setiferous granules, a medial long spiniferous tubercle, and smaller lateral tubercles; medial tubercle of free tergite II longer than tubercle of free tergite I; free tergite III with a medial row of short setiferous tubercles, the medial setiferous tubercle slightly longer, and the lateral region with long setiferous tubercles (Figs 23A, B; 24A, E).

Venter: Coxae I–IV with setae and small granules (Fig. 24B); coxa I with setiferous granules; anteroposterior borders of coxa III with a row of strong granules connecting with coxae II and IV, respectively (Fig. 24B); posterior border of the spiracular area and free sternites I–V with a row of setiferous granules (Figs 23C; 24B); anal operculum with several conspicuous setiferous granules and setiferous tubercles (Figs 23B, C; 24D, E). Spiracles not concealed (Figs 23C; 24B, C).

Chelicerae: Basichelicerite unarmed, with an elongated and slightly marked bulla (Figs 23F, G; 25C, D). Cheliceral hand with sparse setae and small frontal setiferous granules (Figs 23F–H; 25C–E). Movable finger with a proximal lamina with sub-square teeth, followed by a medial conical tooth and a distal lamina with sub-square teeth (Figs 23I; 25E); fixed finger with a large medial conical tooth (Figs 23I; 25E).

Pedipalps: Coxa elongated (i.e., remarkably longer than trochanter), with one small dorsomesal protuberance and one small medial setiferous granule on the ventral surface (Figs 24 A, B). Trochanter rounded, with three dorsal, one mesal setiferous pointed tubercles, and two ventral setiferous conical tubercles (Figs 23D, E; 24B; 25A, B). Femur dorsally convex, ventrally armed with a row of six small ectal setiferous pointed tubercles, the fifth distal tubercle longest (Figs 23E; 25B); ventroproximally with two large spines, fused at the base (Figs 23D, E; 25A, B); ventromesal surface with a medial spine followed by one setiferous pointed tubercle (Figs 23D; 25A). Patella short, with scattered dorsal granules, and ventrodistally with one mesal spine and one small ectal setiferous pointed tubercle (Figs 23D, E; 25A, B). Tibia with dorsal granules; ventromesally with three spines, increasing in size from proximal to distal (Figs 23D; 25A); ventroectally with one small proximal setiferous pointed tubercle, followed by one spine, one setiferous pointed tubercle, and two spines fused at the base, the longest spine featuring a socket with an apical square-shaped projection (Figs 23E; 25B); ventral surface with scattered small granules (Fig. 25A). Tarsus remarkably elongated, incrassate, and ventrally flattened (Figs 22C; 23D, E; 25A, B); ventromesally with one proximal setiferous pointed tubercle, followed by a row of five spines (Figs 23D, 25A); ventroectally with three spines interspersed with two setiferous pointed tubercles (Figs 23E). Claw remarkably short, robust, and triangular (Figs 22C; 23D, E; 25A, B).

Legs: Coxae II and IV with setiferous granules on dorsolateral surface (Fig 24A, E). Trochanters I–IV covered by setiferous granules (Fig. 24A, B). Femurs I–IV with longitudinal rows of ventral conical setiferous tubercles (Fig. 22B, C). Metatarsus III swollen at calcaneus region, in lateral view with a rectangular shape (Figs 26A, C; 28F); calcaneus extends from the medial region of the metatarsus (Figs 26A, C; 28F), ventrally with trichomes and some lateral sensilla chaetica (Fig. 26A, C, E); apical region of calcaneus with a high concentration of acuminate trichomes densely covering numerous aggregated pores (glandular openings) (Fig. 26A–D). Tarsi III–IV without scopula and modified spatulate setae (Fig. 26F). Tarsal formula: 4(2):8(3):5:5.

Color (specimen preserved in 80% ethanol): General body appearance yellowish-brown; appendages light yellowish-brown (Fig. 22A–C); coloration clearer at the level of cheliceral insertion, creating a false appearance of a marked cheliceral socket (Fig. 22A). Coxae I–IV, free sternites V and anal operculum light yellowish-brown; free sternites I–IV dark yellowish-brown (Fig. 22B).

Genitalia: General shape of penis tubular, with a strong distal constriction and widening apically to a blunt, rectangular tip; boundary not well defined between pars basalis and pars distalis (Fig. 27A, C, E, G). Pars distalis with a ventral plate ending in a calyx (Fig. 27B, F); calyx dorsally open with two thin laminar projections (wings) (Fig. 27B, F); dorsally, pars distalis with a medial deep neckline (Fig. 27B, F). Pars distalis armed with two groups of macrosetae arranged bilaterally: a basal row of four pairs (B1–B4) extending along the edge, from the dorsal neckline to the ventrolateral region (Fig. 27B, D, F, H), and an apical row (A1–A3) located on the ventrolateral region of the calyx (Fig. 27D, H, I). Capsula externa with follis invaginated and not visible in resting position (Fig. 27B, F). Capsula interna with two laminar conductors that are arrow-shaped apically (i.e., medially pointed and with two lateral projections) (Fig. 27 B, D, F, H); conductors flank a shorter, pointed, laminar stylus (Fig. 27 B, D, F, H–I). When the penis is expanded and the capsula interna is everted it is possible to see that the conductors are basally fused between them and with the stylus, forming an integrated structure (Fig. 27B, D).

Minor male (AMNH). Body measurements: Total body length 2.40, carapace length 0.90, scutum magnum length 2.21, carapace maximum width 1.34, abdominal scutum maximum width 2.05. Appendage measurements in Table 4. Minor male differentiated from major male by the absence of lateral setiferous tubercles on free tergite III and shorter medial setiferous tubercle; medial setiferous tubercles on free tergites I–II shorter, contrasting with the long setiferous tubercles present in major male (Fig. 28D, E vs. Fig. 28G, H). Additionally, minor male has a shorter pedipalp with smaller spines; trochanter of pedipalp with small setiferous tubercles, resembling female; femur of pedipalp with the mesal spine closer to the distal setiferous tubercle; tarsus not elongated and enlarged, with a proximal setiferous tubercle but with four spines, instead of the five observed in major males; claw is remarkably long, thin, and pointed, similar to that of female (Fig. 28E vs. Fig. 28B, H). Metatarsus III similar to major male (Fig. 28F vs. 28I). Tarsal formula 4(2):8–9(3):5:5.

Female (AMNH). Body measurements: Total body length 2.61, carapace length 0.85, scutum magnum length 1.28, maximum carapace width 1.32, abdominal scutum maximum width 2.06. Appendage measurements in Table 4. Resembles minor and major males in the armature of scutum magnum, but differs from major male by the absence of long lateral setiferous tubercle tubercle on free tergite III (Figs 21A; 28A, B vs. Figs 21B; 28G, H); medial setiferous tubercle of free tergite I–III shorter than in major male (Fig. 28A, B); pedipalp remarkably shorter than in major male and with smaller spines; trochanter without dorsal setiferous tubercles and only one small setiferous granule; femur of pedipalp with the mesal spine and distal tubercle closer than in major male; tarsus not elongated, with four spines and without proximal setiferous tubercle; claw elongated, thin and pointed as in minor male (Fig. 28B vs. Fig. 28E, H). Female differs from minor and major males by having metatarsus III not swollen, lacking aggregated pores and associated setae, and without the deep intrusion of the astragalum by the calcaneus (Fig. 28C vs. 28F, I). Tarsal formula 4(2):7(3)–8(3):5:5.

Geographical distribution. United States of America: Florida: Key West or Tortugas (Packard 1888); Biscayne Bay (Banks 1904); Everglades National Park—Royal Palm State Park, Homestead, Marquesas Key, Barracuda Key (Goodnight & Goodnight 1942b); Everglades National Park (Duffield et al., 1981). New records: Key Largo, 7 miles west of Florida City, Little Duck Key, Key Largo—Pennekamp State Park, Big Pine Key—Watson’s Hammock, Tamiami Trail (Fig. 39).

Unconfirmed records. Cayman Islands: Little Cayman (Hounsome 1994); probably another species of Neoscotolemon, potentially a new species (Fig. 39).

Doubtful records. Jamaica: Clarendon, Jackson Bay Cave; Saint Ann, Dairy Bull Cave and Ken Connell Hole (Peck 1992), probably a misidentification of a Jamaican Samoidae species (Akdalima jamaicana Šilhavý, 1979, Reventula amabilis Šilhavý, 1979, or another related species) (Fig. 39).

Spurious records. United States of America, Ohio, Clear Creek; Hocking County; Rockbridge; Sugar Creek (Walker, 1928: 157, fig. 11). There are no reliable records of Neoscotolemon spinifer outside peninsular Florida and adjacent Keys. The record by Walker is surely a misidentification, probably of Erebomaster acanthinus (Crosby & Bishop 1924) (Cladonychiidae) that commonly occurs in southern Ohio (W. Shear 2010 pers. comm.).

Natural history. Duffield et al. (1981) stated that: “The opilionids [N. spinifer] were collected in Everglades National Park, Florida, in November 1977 and March 1978, from solitary retreats on the undersides of coral rocks. Also, specimens have been collected (in August 1985) in Hammock forest leaf litter.” See “Other material examined” above.