Holocnemus caudatus ( Dufour, 1820 )

Huber, Bernhard A., 2022, Revisions of Holocnemus and Crossopriza: the spotted-leg clade of Smeringopinae (Araneae, Pholcidae), European Journal of Taxonomy 795 (1), pp. 1-241 : 38-46

Holocnemus caudatus ( Dufour, 1820)

Figs 3 , 97–128

Pholcus caudatus Dufour, 1820: 208 , pl. 76 fig. 2a–b.

Pholcus caudatus – Walckenaer 1837: 654 (description copied from Dufour 1820). — Simon 1866: 123, pl. 2, figs 12–13.

Holocnemus caudatus – Simon 1873: 49. — Franganillo 1925: 35; 1926c: 71. — Wiehle 1933: 244, fig. 2b. — Brignoli 1971a: 84; 1971c: 256 (“ Holocnemus prope caudatus ”). — Melic 1994: 15. — Ribera et al. 2003: 8. — Benhadi-Marín et al. 2013: 75, fig. 2a(1–3). — Breitling 2020: 352, fig. 26a–d.

Dubious records (see Distribution below)

Holocnemus caudatus – Barrientos & Ferrández 1982: 82. — Ferrández et al. 2006: 76.

Misidentifications

Barrientos et al. 2019: 12 (see H. reini ).

Remark

Holocnemus acuminatus Franganillo, 1925 from Spain, Málaga, is currently considered a nomen dubium ( World Spider Catalog 2021). It is probably a synonym of H. caudatus but could also be a senior synonym of H. hispanicus Wiehle, 1933 . The World Spider Catalog (2021) considers Franganillo’s (1925) publication in the Boletín de la Sociedad Entomológica de España as the original publication for this name. However, in that paper, Franganillo writes “ Holocnemus acuminatus Frang. ”, i.e., without adding “sp. nov.” as he did in other occasions, suggesting that the species had been described before. In fact, in the introduction he cites a series of publications from which the listed records are taken. I suspect that H. acuminatus was originally intended to be described in Franganillo’s contribution to the Proceedings volume of the Tercer Congreso Científico Panamericano (Lima, 22 Dec. 1924 – 5 Jan. 1925), under the title “Arácnidos de Andalucía ”. However, that proceedings volume was never published. Thus, Holocnemus acuminatus Franganillo, 1925 is here considered a nomen nudum.

Diagnosis

Easily distinguished from most known Smeringopinae (except H. reini ) by distinct indentations on sternum ( Fig. 112 ; in males and females; deepest pair between coxae 4), by shape of procursus (cf. Figs 62–63 , 117 ; distinctive retrolateral membrane), by shape of genital bulb (cf. Figs 66–68 ; distal bulbal sclerite triangular in prolateral view, with one large and two small rounded processes); from known congeners and representatives of Crossopriza also by 2–3 modified hairs on each male cheliceral apophysis (rather than one; Fig. 115 ); from H. reini only by slightly smaller distances between male cheliceral apophyses (50–60% of cheliceral maximum width) and between female epigynal pockets (<0.4 mm).

Type material

Syntypes SPAIN • Unspecified number; Valencia, Mogente / Moixent (“Moxente”); 38.88° N, 0.76° W; date unknown; L. Dufour leg.; probably lost .

Material examined

SPAIN – Valencian Community • 1 ♂; Alicante, Cova del Rull near Val de Ebo ; 38.812° N, 0.177° W; 475 m a.s.l.; 2–3 Jun. 2010; S. Huber and A. Schönhofer leg.; SMF • 6 ♂♂, 7 ♀♀, 1 juv.; Alicante, Elda ; 38.48° N, 0.79° W; 19 Jun. 1971; A. Senglet leg.; MHNG • 3 ♂♂, 3 ♀♀; Valencia, Ayora ; 39.06° N, 1.06° W; 20 Jun. 1971; A. Senglet leg.; MHNG • 4 ♂♂, 3 ♀♀; Valencia, Montroy ; 39.34° N, 0.61° W; 22 Jun. 1971; A. Senglet leg.; MHNG • 8 ♂♂, 9 ♀♀; Valencia, Requena-Chera ; 39.54° N, 1.02° W; 23 Jun. 1971; A. Senglet leg.; MHNG • 1 juv.; Castellón, Cabanes , “Avenc d’en Soria”; 40.15° N, 0.05° E; 2 Jan. 1964; Nebot leg.; CRBA 1424 . – Aragon • 3 ♂♂, 3 ♀♀; Teruel, Mora de Rubielos ; 40.25° N, 0.75° W; 5 Sep. 1971; A. Senglet leg.; MHNG • 12 ♂♂, 13 ♀♀; Teruel, Albarracín ; 40.41° N, 1.44° W; 3 Sep. 1971; A. Senglet leg.; MHNG • 1 ♀, 1 juv.; Teruel, Aguaviva, along Río Bergantes ; 40.82° N, 0.20° W; 500 m a.s.l.; 2 Apr. 1996; R. Bosmans leg.; maquis and Pinus; CRB • 2 ♂♂, 2 ♀♀, 1 juv.; Teruel (the label says “ Lerida ” but Castelnou is not in Lérida ), Castelnou ; 41.23° N, 0.36° W; 8 Jun. 1955; A. Comellini leg.; MHNG . – Castilla-La Mancha • 3 ♂♂, 6 ♀♀; Cuenca, Sta Cruz de Moya ; 39.955° N, 1.255° W; 24 Jun. 1971; A. Senglet leg.; MHNG • 3 ♀♀, 3 juvs; Albacete, S of Tarazona de La Mancha, along Río Júcar ; 39.25° N, 1.91° W; 600 m a.s.l.; 8 Apr. 1997; R. Bosmans leg.; degraded Quercus ilex forest; CRB • 1 ♂, 3 ♀♀, 3 juvs; Albacete , La Gineta; 39.11° N, 2.00° W; 28 Jun. 1971; A. Senglet leg.; MHNG. – Region of Murcia • 2 ♂♂, 5 ♀♀; Caravaca [de la Cruz]; 38.11° N, 1.86° W; 5 Jul. 1971; A. Senglet leg.; MHNG • 5 ♂♂, 9 ♀♀, 1 juv.; Bullas ; 38.05° N, 1.67° W; 4 Jul. 1971; A. Senglet leg.; MHNG • 2 ♂♂, 2 ♀♀ (partly used for SEM); Mazarrón ; 37.60° N, 1.32° W; 1 May 2001; J. Van Keer leg.; CJVK • 1 ♀; Mazarrón, Cueva del Agua ; 37.576° N, 1.219° W; 18 Feb. 1983; C. Ribera leg.; CRBA 2088 • 5 ♂♂, 5 ♀♀; Alberca ; 37.94° N, 1.14° W; 4 May 1930; H. Wiehle leg.; SMF 11442, 11443 , 19430 . – Catalonia • 2 ♀♀; Barcelona, Parque Natural del Garraf ; 41.28° N, 1.84° E; 6 Sep. 2006; R. Bosmans leg.; CRB • 1 ♂; Lleida, Bor, Cova de la Fou de Bor ; 42.339° N, 1.801° E; 28 Sep. 1965; C. Ribera leg.; CRBA 1409 . – Andalusia • 1 ♂, 3 ♀♀; Granada, Puebla de Don Fadrique ; 37.96° N, 2.44° W; 1000–1200 m a.s.l.; 6 Jul. 1971; A. Senglet leg.; MHNG • 1 ♂, 6 ♀♀; Granada, Puerto de La Ragua ; 37.17° N, 3.05° W; 1200 m a.s.l.; 17 Jul. 1971; A. Senglet leg.; MHNG • 1 ♂, 2 ♀♀; Granada, Ugíjar ; 36.96° N, 3.05° W; 15 Jul. 1971; A. Senglet leg.; MHNG • 3 ♂♂, 9 ♀♀; Granada, Las Alpujarras ; 36.98° N, 3.15° W; 1250 m a.s.l.; 9 Jul. 1971; A. Senglet leg.; MHNG • 11 ♂♂, 9 ♀♀ (partly used for SEM); Granada, Trevélez ; 37.00° N, 3.27° W; 14 Jul. 1971; A. Senglet leg.; MHNG • 3 ♂♂, 2 ♀♀; Granada, Sierra Nevada, Alto del Chorrillo ; 37.01° N, 3.31° W; 2100– 2800 m a.s.l.; 11 Jul. 1971; A. Senglet leg.; MHNG • 1 ♀; Granada , La Mamola; 36.75° N, 3.28° W; 50 m a.s.l.; 13 Aug. 1991; R. Bosmans leg.; CRB • 1 ♂; Granada (province or city?); Aug. 1924; M.D. Leonard leg.; AMNH • 8 ♂♂, 14 ♀♀; Granada, Capileira ; 36.96° N, 3.36° W; 1500 m a.s.l.; 10 Jul. 1971; A. Senglet leg.; MHNG • 9 ♂♂, 6 ♀♀; Granada, Pampaneira / Órgiva ; 36.92° N, 3.38° W; 1200 m a.s.l.; 10 Jul. 1971; A. Senglet leg.; MHNG • 2 ♀♀; Granada, Motril ; 36.75° N, 3.52° W; 50 m a.s.l.; 3 Apr. 1987; J. Murphy leg.; garrigue, stones; MMUE Murphy #14441 • 1 ♂, 1 ♀; Granada, Sierra Nevada, Albergue Universitario ; 37.094° N, 3.387° W; 2500 m a.s.l.; date and collector not given; MNHN Ar 10340 • 3 ♂♂, 1 ♀; Granada, Sierra Nevada, Peñones de San Francisco ; 37.10° N, 3.39° W; date not given; H. Janetschek leg.; MNHN Ar 10346 • 3 ♀♀, 1 juv.; Almería, Cabo de Gata ; 36.78° N, 2.24° W; 50 m a.s.l.; 5 Apr. 1996; R. Bosmans leg.; stones in dunes; CRB • 1 ♂, 1 ♀ (in pure ethanol); same locality as for preceding; 36.7389° N, 2.2014° W; Jul.–Aug. 2021; J. Moya-Laraño leg.; subdesert grassland; ZFMK G154 , G155 • 1 ♂; Almería, Sierra de Gádor, Sima de los Locos ; 36.856° N, 2.506° W; 5 Feb. 2000; P. Barranco leg.; CRBA 3620 • 1 ♂, 2 ♀♀; Málaga, Maro ; 36.76° N, 3.84° W; 100 m a.s.l.; 3 Apr. 1987; J. Murphy leg.; stones, litter; MMUE Murphy #14318 • 2 ♂♂; Málaga, Frigiliana ; 36.79° N, 3.89° W; 20 m a.s.l.; 6 Apr. 1987; J. Murphy leg.; stones, litter; MMUE Murphy #14545 • 1 ♀; Málaga, Fuengirola ; 36.55° N, 4.63° W; 26 Jul. 1969; A. Senglet leg.; MHNG • 2 ♂♂, 4 ♀♀, 2 juvs; Málaga, Marbella; 36.52° N, 4.88° W; 12 Aug. 1991; R. Bosmans leg.; stones in Pinus forest ; CRB • 1 ♀; Málaga, San Pedro de Alcántara ; 36.49° N, 4.99° W; 23 Jul. 1969; A. Senglet leg.; MHNG • 14 ♂♂, 21 ♀♀; Málaga, Ronda ; 36.74° N, 5.16° W; 21 Jul. 1969; A. Senglet leg.; MHNG • 1 ♂, 2 ♀♀; same locality as for preceding; 20–23 Jul. 1969; A. Senglet leg.; MMUE Murphy #12790 • 1 ♀; same locality as for preceding; 750 m a.s.l.; 12 Aug. 1991; R. Bosmans leg.; stones; CRB .

ANDORRA • 3 ♂♂, 1 ♀; “Andorre”, no further locality data; 42.51° N, 1.53° E; date and collector unknown; MNHN Ar 10328 .

Redescription

Male ( Granada, Trevélez, MHNG)

MEASUREMENTS. Total length 5.5, carapace width 1.6. Distance PME–PME 110 µm; diameter PME 110 × 130 µm; distance PME–ALE 50 µm; diameter AME 70 µm; distance AME–AME 25 µm. Leg 1: 33.4 (9.7 + 0.8 + 8.8 + 12.1 + 2.0), tibia 2: 6.3, tibia 3: 4.7, tibia 4: 5.7; tibia 1 L/d: 49; femora 1–4 diameters: 0.32, 0.24, 0.21, 0.22.

COLOR (in ethanol). Carapace ochre-yellow; ocular area, carapace pit and posterior median area light brown; sternum brown, with dark brown radial marks; legs ochre-yellow, without darker rings, with black lines on femora and (few) on tibiae; abdomen ochre-gray, with some dark marks dorsally and laterally; ventrally with distinct black median band, partly disrupted, with three parallel longitudinal marks behind gonopore.

BODY. Habitus very similar to H. reini (cf. Fig. 7 ). Ocular area slightly raised. Deep thoracic pit and pair of shallow furrows diverging from pit toward posterior margin. Clypeus unmodified but rim more sclerotized than in female. Sternum wider than long (1.05/0.85), with distinct indentations between consecutive leg coxae and between coxae 4. Abdomen strongly elongated beyond spinnerets, dorsoposteriorly pointed. Gonopore with six epiandrous spigots ( Fig. 118 ); ALS with one widened spigot and one pointed spigot, PMS with two pointed spigots ( Fig. 125 ).

CHELICERAE. As in Figs 97–98 , with pair of low proximal frontal humps and pair of frontal lateral apophyses, each with two large modified cone-shaped hairs ( Fig. 115 ); distance between tips of modified hairs: 320 µm; several ‘regular’ frontal hairs with small but distinct processes at basis; lateral stridulatory ridges very fine ( Fig. 113 ; distances between ridges proximally ~1.5 µm, distally ~4 µm), barely visible in dissecting microscope.

PALPS. In general apparently indistinguishable from H. reini (cf. Figs 59–61 ); coxa with rounded retrolateral hump; trochanter barely modified; femur slightly curved towards dorsal, distally widened, with rounded ventral protrusion, proximally with prolateral stridulatory pick ( Fig. 116 ), with barely visible retrolateral transversal line, without retrolateral proximal process; femur-patella joints shifted toward prolateral side; tibia very large compared to femur, tibia-tarsus joints shifted toward retrolateral side; tarsus without macrotrichia; tarsal organ capsulate ( Fig. 121 ); procursus (cf. Figs 62–63 ) straight, few hairs slightly curved upwards; proximally on prolateral side with strong hump free of hairs, procursus tip with distinctive flat membranous element retrolaterally (arrow in Fig. 117 ) covering proximal part of ventral sclerite in retrolateral view, with dorsal transversal flap; genital bulb (cf. Figs 66–68 ) with simple basal sclerite connected to distal (main) sclerite, sperm duct opening in membranous area prolaterally at basis of distal sclerite; distal sclerite with one large apophysis (fold) and two smaller apophyses on prolateral side; with indistinct retrolateral parallel ridges.

LEGS. Femur 1 with single row of ~23 ventral spines; without curved hairs; few vertical hairs ( Figs 119– 120 ); retrolateral trichobothrium of tibia 1 at 4%; prolateral trichobothrium absent on tibia 1, present on other leg tibiae; tarsal pseudosegments indistinct except 2–3 distally; tarsal organs capsulate, with round or slightly irregular rim ( Figs 122–123 ).

Male (variation)

Tibia 1 in 82 males: 6.8–9.9 (mean 8.5). Chelicerae maximum width (N = 43): 0.46–0.62 (mean 0.55); procursus length (N = 46): 0.86–1.00 (mean 0.92). Coloration of fresh specimens very similar to H. reini (cf. Figs 7–8 ), with distinct pattern on abdomen. Ventral dark band on abdomen variably distinct, section behind gonopore with 2–4 longitudinal bands.

Female

In general similar to male (cf. Fig. 8 ) but without spines on legs. Without stridulatory apparatus between carapace and abdomen; with fine cheliceral stridulatory ridges as in male ( Fig. 114 ; distances between ridges 3 µm). Tibia 1 in 103 females: 5.1–8.8 (mean 6.9). Epigynum as in Figs 101–106 and 127–128 , main epigynal plate triangular to semicircular, weakly protruding; only posteriorly laterally strongly sclerotized; with pair of variably distinct pockets (distance between pockets 0.32–0.35); internal sclerotized arc and median round structure variably visible in uncleared specimens; posterior epigynal plate large but simple; pair of low but distinct elevations in front of epigynum. Internal genitalia ( Figs 99–100 , 107–110 ) with triangular pore plates converging anteriorly, dorsal arc simple and slender, ventral arc laterally strongly widened, medially strong, with simple median pouch.

Natural history

The biology of this common Spanish spider has apparently been little studied. According to Wiehle (1933) it occupies similar habitats as H. hispanicus (i.e., wall niches, rock cavities, etc.). The domed web was described as having a surprisingly regular mesh; the size of the dome is similar to H. hispanicus (i.e., approximately 35–40 cm), but the silk puffs are not as close together as in H. hispanicus . Females lay up to 50 eggs per egg-sac and produce up to three egg-sacs per season ( Wiehle 1933).

Distribution

Widely distributed in southern and eastern Spain ( Fig. 3 ), where it occurs from sea level up to 2500 m a.s.l. (Albergue Universitario). I have not seen any specimens from central-western Spain (Castile and León, Extremadura, and Madrid), and consider all previous records from this region (Fernández Galiano 1910 in Ferrández et al. 2006; Barrientos & Ferrández 1982; Ferrández et al. 2006) dubious. I also follow Brignoli (1971a) in considering Simon’s (1873) statement about the presence of H. caudatus in Sicily as dubious to highly improbable.

Figures

Fig. 3. Known distributions of Holocnemus reini (C. Koch, 1873) comb. nov. and H. caudatus (Dufour, 1820). Light blue: dubious records; see Distribution section for details. Abbreviation: TL = type localities (the type locality of H. reini is not exactly known; see text for details).

Figs 97–100. Holocnemus caudatus (Dufour, 1820); male and female from Spain, Valencia, Montroy (MHNG). 97–98. Male chelicerae, frontal and lateral views. 99–100. Cleared female genitalia, ventral and dorsal views; arrows: pockets. Scale bars = 0.3 mm.

Figs 101–106. Holocnemus caudatus (Dufour, 1820); female abdomens and epigyna, ventral views. 101–102. Spain, Albacete, S of Tarazona de La Mancha (CRB). 103–104. Spain, Granada, Trevélez (MHNG). 105–106. Spain, Granada, Motril (MMUE Murphy #14441). Scale bars = 0.5 mm.

Figs 107–110. Holocnemus caudatus (Dufour, 1820). 107–109. Female from Spain, Valencia, Montroy (MHNG); internal genitalia, ventral view, regular dorsal view, and dorsal view with dorsal arc tilted backwards. 110. Female from Spain, Albacete, S of Tarazona de La Mancha (CRB); internal genitalia, dorsal view with dorsal arc tilted backwards. Abbreviations: da = dorsal arc; va = ventral arc. Scale bars = 0.5 mm.

Figs 111–118. Holocnemus caudatus (Dufour, 1820); male and female from Spain, Granada, Trevélez (MHNG). 111. Female ocular area. 112. Male sternum; arrows: indentations. 113–114. Male and female cheliceral stridulatory files. 115. Modified hairs on male cheliceral apophysis. 116. Stridulatory pick on male palpal femur. 117. Right procursus tip, prolateral view; arrow: retrolateral membrane. 118. Male gonopore. Abbreviation: vs = ventral sclerite. Scale bars: 111–112, 117–118 = 100 µm; 113–114 = 20 µm; 115–116 = 10 µm.

Figs 119–128. Holocnemus caudatus (Dufour, 1820); male and female from Spain, Granada, Trevélez (MHNG). 119. ‘Short vertical hair’ on male metatarsus 4. 120. Tip of hair from preceding figure. 121. Male palpal tarsal organ. 122–123. Tarsal organs on male tarsi 1 and 4. 124. Female palpal tarsal organ. 125–126. Male and female spinnerets. 127–128. Epigyna and anterior epigynal humps (arrows). Scale bars: 119, 121–125 = 10 µm; 120 = 1 µm; 126 = 20 µm; 127–128 = 100 µm.

Figs 62–70. Holocnemus reini (C. Koch, 1873) comb. nov.; male from Morocco, near Tilamizene (ZFMK Ar 22319), female from Morocco, Al Kamoun (CAS 9027140). 62–63. Left procursus, prolateral and retrolateral views; arrow: retrolateral membrane. 64–65. Male chelicerae, frontal and lateral views. 66–68. Left genital bulb, prolateral, dorsal, and retrolateral views; arrow: sperm duct opening. 69–70. Cleared female genitalia, ventral and dorsal views; arrows: pockets. Abbreviations: bs = basal sclerite; ds = distal sclerite; vs = ventral sclerite. Scale bars: 62–63, 66–68 = 0.3 mm; 64–65, 69–70 = 0.5 mm.

Figs 5–12. Live specimens and webs. 5–6. Holocnemus pluchei (Scopoli, 1763); male and female from Morocco, Imi n’Ifri. 7–8. Holocnemus reini (C. Koch, 1873) comb. nov.; male from Morocco, near Tilamizene; female from Morocco, W of Tamtetoucht. 9–10. Maghreba amezyan gen. et sp. nov., male and female from Morocco, between Lakhssas and Bouizakarne. 11–12. Maghreba amezyan gen. et sp. nov., pair in spotted web, and detail of web with silk puffs, SW of Ida Ougnidif.

Figs 59–61. Holocnemus reini (C. Koch, 1873) comb. nov.; male from Morocco, near Tilamizene (ZFMK Ar 22319); left palp, prolateral, dorsal, and retrolateral views. Scale bar = 0.5 mm.