Holocnemus reini Huber 2022, comb. nov.
Creators: name: Huber, Bernhard A.
Publisher: name: Zenodo
Year: 2022
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Arachnida
Araneae
Pholcidae
Holocnemus
Holocnemus reini

description: Holocnemus reini (C. Koch, 1873) comb. nov.
Figs 3, 7–8, 59–96
Pholcus reini C. Koch, 1873: 113.
Holocnemus caudatus (misidentification) – Barrientos et al. 2019: 12.
Crossopriza sp. – Hajer & Řeháková 2003: 345–354.
Remarks
While Roewer (1955) considered Pholcus reini a nomen dubium (“nicht zu deuten”), Bonnet (1958) treated it as an available name. Platnick (2000), in his first online catalogue, adopted Roewer’s view, and Pholcus reini has been listed as a nomen dubium ever since (World Spider Catalog 2021). The single male type specimen is apparently lost. It is not in SMF, in contrast to the types of at least one other species collected during the same expedition and published in the same paper (Ocypete fritschi C. Koch, 1873, now Eusparassus fritschi). However, even though C. Koch’s (1873) original description does not provide any figure, it gives some details (size, ventral color pattern) that fit only two pholcid species known to be present in Morocco: Holocnemus pluchei and the species here interpreted to represent Holocnemus reini. Since C. Koch (1873) distinguished between Pholcus barbarus (now H. pluchei) and P. reini, it seems reasonable to assume that C. Koch’s name does not refer to H. pluchei but to the species treated here.
I have not seen the specimens of Barrientos et al. (2019) but since all newly examined Moroccan specimens were H. reini rather than the similar H. caudatus, the specimens of Barrientos et al. (2019) are regarded as misidentified. Vocher specimens of Hajer & Řeháková’s (2003) “ Crossopriza sp. ” were examined (see below).
Diagnosis
Easily distinguished from most known Smeringopinae (except H. caudatus) by deep indentations on sternum (Fig. 81; in males and females; deepest pair between coxae 4), by shape of procursus (Fig. 63; distinctive retrolateral membrane), by shape of genital bulb (Figs 66, 92; 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 (Figs 64, 82–83; rather than one); from H. caudatus only by slightly wider distances between male cheliceral apophyses (70–80% of cheliceral maximum width) and between female epigynal pockets (> 0.5 mm).
Type material
Holotype MOROCCO • ♂; Mtuga plateau (roughly in the area between Essaouira, Marrakesh, and Agadir); 1872; K. von Fritsch and J. Rein leg.; apparently lost.
Material examined
MOROCCO – Marrakesh-Safi • 9 ♂♂, 6 ♀♀ (partly used for SEM); near Tilamizene; 31.3500° N, 7.7645° W; 950 m a.s.l.; 11 Sep. 2018; B.A. Huber leg.; near ground; ZFMK Ar 22319 • 2 ♀♀, 1 juv. (in pure ethanol); same collection data as for preceding; ZFMK Mor 72 • 1 ♂, 1 ♀; High Atlas, near Asni; 31.24° N, 7.98° W; 1200 m a.s.l.; 12 May 1977; “PDH” leg.; BMNH • 1 ♂, 2 ♀♀; near Toufliht; 31.4715° N, 7.4332° W; 1465 m a.s.l.; 27 Sep. 2018; B.A. Huber leg.; among rocks near ground above ravine; ZFMK Ar 22320 • 1 ♀ (in pure ethanol); same collection data as for preceding; ZFMK Mor 108 • 1 ♀; near Rakte; 31.033° N, 8.137° W; 1060 m a.s.l.; 12 Sep. 2018; B.A. Huber leg.; among rocks in dry riverbed; ZFMK Ar 22321 • 2 ♂♂, 1 ♀; N of Marrakesh, Oued Tensift; 31.69° N, 8.03° W; 9 Feb. 1996; J. Van Keer leg.; under stones; CJVK • 1 ♀; same locality as for preceding; in and along riverbed; 9 Feb. 1996; R. Bosmans leg.; CRB • 2 ♂♂, 2 ♀♀; near Irhoud (= Ighoud); 31.854° N, 8.873° W; 285 m a.s.l.; 29 Sep. 2018; B.A. Huber leg.; along river and among cacti; ZFMK Ar 22323 • 1 ♂; NE of Tamelelt [Tamallalt], El Kelaa des Sraghna; 32.055° N, 7.400° W; 590 m a.s.l.; 16 Apr. 2012; J. Van Keer leg.; stones in wasteland; CJVK. – Souss-Massa • 1 ♂, 1 ♀; S of Tizi n’Test at R203; 30.850° N, 8.375° W; 1580 m a.s.l.; 12 Sep. 2018; B.A. Huber leg.; among rocks near ground; ZFMK Ar 22322 • 1 juv. (in pure ethanol); same collection data as for preceding; ZFMK Mor 75 • 1 ♂, 1 juv.; 80 km E of Taroudant, Oued Lemdad; 30.64° N, 8.38° W; 18 May 1939; Berland leg.; MNHN Ar 10313 • 1 ♂; N of Agadir, Imouzzer; 30.678° N, 9.483° W; 960 m a.s.l.; 8 Sep. 2014; S. Huber leg.; small cave near waterfall; ZFMK Ar 22326 • 1 ♀; N of Agadir, Imouzzer; 30.678° N, 9.482° W; 960 m a.s.l.; 27 Nov. 2016; S. Huber leg.; olive tree plantation; ZFMK Ar 22327 • 2 ♂♂, 1 ♀; Paradise Valley; 30.588° N, 9.528° W; 305 m a.s.l.; 13 Sep. 2018; B.A. Huber leg.; among rocks near ground; ZFMK Ar 22328 • 2 ♀♀ (in pure ethanol); same collection data as for preceding; ZFMK Mor 77 • 1 ♀; 7 km N of Agadir, Anza; 30.45° N, 9.65° W; 50 m a.s.l.; 3 Feb. 1996; R. Bosmans leg.; stones in Euphorbia vegetation; CRB • 1 ♂; N of Agadir, road Agadir-Alma; 30.486° N, 9.565° W; 440 m a.s.l.; 27 Nov. 2016; S. Huber leg.; ZFMK Ar 22329 • 4 ♀♀, 1 juv.; Agadir, path to Kasbah Hill; 30.430° N, 9.619° W; 110 m a.s.l.; 7 Sep. 2014; S. Huber leg.; ZFMK Ar 22330 • 1 ♂; same locality as for preceding; 60 m a.s.l.; 28 Nov. 2016; S. Huber leg.; ZFMK Ar 22331 • 1 ♀; wadi ~ 5 km NE of Agadir; 30.456° N, 9.610° W; 150 m a.s.l.; 14 Sep. 2014; S. Huber leg.; ZFMK Ar 22332 • 1 ♂, 3 ♀♀; between Gourizim and Tiznit; 29.6690° N, 9.8545° W; 330 m a.s.l.; 14 Sep. 2018; B.A. Huber leg.; among rocks near ground; ZFMK Ar 22333 • 5 ♂♂, 1 ♀; near Jamaa Idaoussemlal; 29.5306° N, 9.2375° W; 1220 m a.s.l.; 15 Sep. 2018; B.A. Huber leg.; along small brook; ZFMK Ar 22334 • 1 ♂, 1 ♀ (in pure ethanol); same collection data as for preceding; ZFMK Mor 84 • 1 ♂; SW of Tafraoute; 29.696° N, 9.010° W; 1025 m a.s.l.; 15 Sep. 2018; B.A. Huber leg.; among rocks and boulders; ZFMK Ar 22335. – Béni Mellal-Khénifra • 4 ♂♂, 1 ♀; N of Demnate; 31.761° N, 7.006° W; 895 m a.s.l.; 26 Sep. 2018; B.A. Huber leg.; among rocks near small cave; ZFMK Ar 22324 • 1 ♂, 1 ♀; Imi n’Ifri; 31.724° N, 6.971° W; 1050 m a.s.l.; 26 Sep. 2018; B.A. Huber leg.; among rocks near river at natural bridge; ZFMK Ar 22325 • 2 ♂♂, 2 ♀♀; 35 km [3.5 km?!] W of Kasba Tadla, Al Kamoun (“ Kamoune ”); 32.56° N, 6.26° W; 21 Dec. 1986; V. and B. Roth leg.; CAS 9027140 • 1 ♂; Oued Zem; 32.86° N, 6.57° W; 750 m a.s.l.; 12 May 1984; R. Bosmans leg.; among stones in Pinus halepensis plantation; CRB • 1 juv.; E of Kasba Tadla, SW Aït-Roadi; 32.62° N, 6.11° W; 560 m a.s.l.; 17 Apr. 2012; R. Bosmans leg.; stones in grassland; CRB • 2 ♂♂, 1 ♀; near El Ksiba, Kehf d’bouba; 32.5165° N, 6.0679° W; 1640 m a.s.l.; 24 Sep. 2018; B.A. Huber leg.; among rocks at cave entrance and near cave; ZFMK Ar 22336 • 1 ♂ (in pure ethanol); same collection data as for preceding; ZFMK Mor 101 • 1 ♂, 3 ♀♀; near Sidi Ben Daoud; 32.5336° N, 6.1282° W; 700 m a.s.l.; 25 Sep. 2018; B.A. Huber leg.; among cacti and in small cave; ZFMK Ar 22337 • 1 ♂, 1 ♀; N of Imilchil, near R317; 32.300° N, 5.660° W; 1585 m a.s.l.; 25 Sep. 2018; B.A. Huber leg.; among rocks near ground in dry riverbed; ZFMK Ar 22338 • 1 ♂; SE of El Ksiba, near R317; 32.486° N, 5.942° W; 1315 m a.s.l.; 25 Sep. 2018; B.A. Huber leg.; among rocks in gorge; ZFMK Ar 22339 • 5 ♂♂, 4 ♀♀; Sources of l’Oum er Rbia; 33.051° N, 5.412° W–33.049° N, 5.411° W; 1270–1330 m a.s.l.; 21 Sep. 2018; B.A. Huber leg.; in small cave and among rocks above waterfall; ZFMK Ar 22342 • 1 ♀ (in pure ethanol); same collection data as for preceding; ZFMK Mor 99. – Drâa-Tafilalet • 1 juv. (in pure ethanol); Timoula; 31.791° N, 5.463° W; 2000 m a.s.l.; 19 Sep. 2018; B.A. Huber leg.; under rocks at roadside; ZFMK Mor 93 • 1 ♂, 1 ♀; E of Tamtetoucht; 31.686° N, 5.521° W; 1775 m a.s.l.; 19 Sep. 2018; B.A. Huber leg.; at rocks near spring; ZFMK Ar 22340 • 1 ♀ (in pure ethanol); same collection data as for preceding; ZFMK Mor 94 • 8 ♂♂, 1 ♀, 7 juvs; N of Tinerhir, Tamtetoucht; 31.68° N, 5.54° W; 6 Dec. 1986; V. and B. Roth leg.; CAS 9027135 • 4 ♂♂, 4 ♀♀; W of Tamtetoucht; 31.659° N, 5.581° W; 1745 m a.s.l.; at rocks; 19 Sep. 2018; B.A. Huber leg.; ZFMK Ar 22341 • 1 ♀ (in pure ethanol); same collection data as for preceding; ZFMK Mor 95 • 1 ♂; SE of Zebzat; 32.625° N, 4.540° W; 1675 m a.s.l.; 20 Sep. 2018; B.A. Huber leg.; among rocks outside of cave; ZFMK Ar 22343 • 1 ♂, 1 ♀; near Tizi n’Tairhemt; 32.593° N, 4.526° W; 1855 m a.s.l.; 20 Sep. 2018; B.A. Huber leg.; among rocks in pine forest; ZFMK Ar 22344 • 1 juv. (in pure ethanol); same collection data as for preceding; ZFMK Mor 97. – Rabat-Salé- Kénitra • 1 ♂, 2 juvs; 2 km N of Sidi Allal El Bahraoui, Forêt de la Maâmora; 34.03° N, 6.55° W; 8 Feb. 1996; R. Bosmans leg.; litter in Quercus suber forest; CRB • 1 ♂, 2 ♀♀; Oued Beht; 33.88° N, 5.93° W; 7 Feb. 1996; J. Van Keer leg.; Pinus plantation, stones; CJVK • 1 ♀; same locality as for preceding; 7 Feb. 1996; R. Bosmans leg.; CRB. – Fès-Meknès • 2 ♂♂, 2 ♀♀; Sefrou; 33.83° N, 4.83° W; 25 Dec. 1986; V. and B. Roth; travertine falls; CAS 9027126 • 2 ♂♂, 3 ♀♀; NE of Bab Boudir; 34.0895° N, 4.1080° W; 1400 m a.s.l.; 23 Sep. 2018; B.A. Huber leg.; among rocks; ZFMK Ar 22345 • 1 ♂, 1 ♀; SW of Bab Boudir; 34.0618° N, 4.1317° W; 1570 m a.s.l.; 23 Sep. 2018; B.A. Huber leg.; among rocks; ZFMK Ar 22346 • 1 ♂, 1 ♀; Grotte Kef el Ghar (= Rhar); 34.4788° N, 4.2766° W; 600 m a.s.l.; 22 Sep. 2018; B.A. Huber leg.; among rocks outside of cave; ZFMK Ar 22347. – Tanger-Tetouan-Al Hoceima • 1 ♂, 1 ♀; El Araich (“ Larache ”), near Lixus; 35.20° N, 6.11° W; Sep. 1997; J. Hajer leg.; edge of phosphate mining pit; ZFMK Ar 22348 • 6 ♂♂, 6 ♀♀ (vouchers of Hajer & Řeháková 2003); same collection data as for preceding; ZFMK Ar 22349. – Laâyoune-Sakia El Hamra • 2 ♂♂, 2 ♀♀, 1 juv.; Laayoune, 40 km S of Tarfaya; 27.70° N, 12.95° W; 11 Feb. 2007; R. Bosmans leg.; stony desert, stones along road; CRB.
ALGERIA – Sidi bel Abbès • 1 ♀; Mezaourou; 34.81° N, 0.62° W; 900 m a.s.l.; 23 May 1990; R. Bosmans leg.; clearing in Pinus halepensis forest, under stones; CRB. – Blida • 2 ♂♂, 3 ♀♀; Atlas Blidéen, W of Chréa, Pic. E Abdelkader; 36.42° N, 2.86° E; 1520 m a.s.l.; 20 Jun. 1987 – 9 May 1988; R. Bosmans leg.; open Cedrus forest with Berberis; CRB. – M’Sila • 1 ♀, 1 juv.; Djebel Maadid towards south; 35.80° N, 4.79° E; 1350 m a.s.l.; 29 Apr. 1988; R. Bosmans leg.; along river, under rocks; CRB • 2 ♂♂; M’Sila, Oultem; 35.12° N, 4.38° E; 600 m a.s.l.; 2 Nov. 1987; R. Bosmans leg.; under rocks in steppe; CRB. – Sétif • 1 ♀ (only cleared epigynum); N of Magra, along Oued Nakhar; 35.727° N, 5.161° E; 850 m a.s.l.; 2 Nov. 1988; R. Bosmans leg.; litter; CRB. – Batna • 1 ♀; S of Tazoult Lambese, road to Forêt de S’gag; 35.45° N, 6.25° E; 1800 m a.s.l.; 16 Oct. 1987; R. Bosmans leg.; stones along dry rivulet in grassland; CRB. – Biskra • 1 ♂, 2 ♀♀ (without abdomens), 19 juvs; Biskra; 34.84° N, 5.73° E; date and collector unknown; Simon collection #12283; MNHN Ar 10325.
TUNISIA – Siliana • 1 ♂, 1 ♀; E of Makhtar [Mactaris]; 35.85° N, 9.22° E; 900 m a.s.l.; 23 Jan. 1995; R. Bosmans leg.; stones along oued; CRB. – Sidi Bouzid • 1 ♂, 1 ♀; Djebel Bou Hedma National Park; 34.5° N, 9.6° E; 25–29 Mar. 2001; U. Moldrzyk leg.; CRB. – Gafsa • 1 ♀; near Bou Omrane; 34.35° N, 9.11° E; 26 Mar. 2006; A. López and C. Ribera leg.; CRBA 1315. – Kebili • 2 ♂♂, 9 ♀♀, 8 juvs; “Nefzana” [presumably Nefzaoua]; 33.5° N, 9.0° E; date and collector unknown; Simon collection #4890; MNHN Ar 10338.
Redescription
Male (Morocco, near Tilamizene, ZFMK Ar 22319)
MEASUREMENTS. Total length 6.7, carapace width 1.75. Distance PME–PME 120 µm; diameter PME 120 × 140 µm; distance PME–ALE 50 µm; diameter AME 100 µm; distance AME–AME 25 µm. Leg 1: 38.4 (11.7 + 0.8 + 10.0 + 13.6 + 2.3), tibia 2: 6.9, tibia 3: 5.5, tibia 4: 6.5; tibia 1 L/d: 47; femora 1–4 diameters: 0.31, 0.25, 0.22, 0.23.
COLOR (in ethanol). Carapace ochre-yellow; ocular area posteriorly with light brown V-mark; sternum dark brown; legs ochre-yellow, femora and tibiae with light tips and indistinct darker subdistal rings, with black lines on femora and (few) on tibiae, very few also on metatarsi; abdomen with distinct pattern of dark marks and whitish marks dorsally and laterally; ventrally with distinct black median band, partly disrupted, behind gonopore divided by two thin longitudinal light lines.
BODY. Habitus as in 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 (1.10/0.85), with distinct indentations between consecutive leg coxae and between coxae 4 (cf. Fig. 81). Abdomen strongly elongated beyond spinnerets, dorso-posteriorly pointed. Gonopore with seven epiandrous spigots (Fig. 93); ALS with one widened spigot and one pointed spigot (Fig. 94).
CHELICERAE. As in Figs 64–65, with pair of low proximal frontal humps and pair of frontal lateral apophyses, each with two large modified cone-shaped hairs; distance between tips of modified hairs: 520 µm; several ‘regular’ frontal hairs with small but distinct processes at basis; lateral stridulatory ridges very fine (Fig. 84; distances between ridges: 2–3 µm), barely visible in dissecting microscope.
PALPS. As is 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. 86), 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; procursus (Figs 62–63) straight, few hairs slightly curved upwards; tarsal organ capsulate (Fig. 87); proximally on prolateral side with strong hump free of hairs, procursus tip with distinctive flat membranous element retrolaterally (arrow in Fig. 63) covering proximal part of ventral sclerite, with dorsal transversal flap and complex retrolateral elements (Figs 89–91); genital bulb (Figs 66–68) with simple basal sclerite connected to distal (main) sclerite, sperm duct opening in membranous area prolaterally at basis of distal sclerite (Fig. 92); distal sclerite with one large apophysis (fold) and two smaller apophyses on prolateral side; with indistinct retrolateral parallel ridges (Fig. 68).
LEGS. Femur 1 with single row of ~24 ventral spines; without curved hairs; few vertical hairs; retrolateral trichobothrium of tibia 1 at 3%; prolateral trichobothrium absent on tibia 1, present on other leg tibiae; tarsal pseudosegments indistinct and irregular except 4–5 distally.
Male (variation)
Tibia 1 in 73 males: 7.6–12.8 (mean 10.6). Chelicerae maximum width (N = 57): 0.56–0.80 (mean 0.67); distance between tips of cheliceral apophyses: 0.41–0.58; cheliceral apophyses sometimes with 3 modified hairs, sometimes asymmetric (2 and 3 hairs, respectively; Figs 82–83). Procursus length (N = 58): 0.82–1.16 (mean 1.03). Coloration of fresh specimens slightly variable, usually with dark median band on carapace. Posterior elongation of abdomen variably long, ~0.5–1.0 × distance pedicel– spinnerets. Gonopore with 6–8 epiandrous spigots.
Female
In general similar to male (Fig. 8) but without spines on legs; often with slightly longer dorsal abdominal elongation. Without stridulatory apparatus between carapace and abdomen; with fine cheliceral stridulatory ridges as in male (Fig. 85; distances between ridges ~3 µm). Tibia 1 in 53 females: 6.4– 12.5 (mean 9.2). Epigynum as in Figs 71–76 and 96, main epigynal plate triangular to trapezoidal, weakly protruding; only posteriorly laterally strongly sclerotized; with pair of variably distinct pockets (distance between pockets 0.5–0.6); 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 69–70, 77–80) with triangular pore plates converging anteriorly, dorsal arc simple and slender, ventral arc laterally strongly widened, medially strong, with simple median pouch.
Natural history
This species lives in a variety of habitats, ranging from shady pine forests and river valleys to very dry arid slopes. Most specimens were found in sheltered spaces among and under rocks. The large webs (diameter ~ 30 cm) were often exposed to direct sunlight, but during the day the spiders rested on the shady rock surface. On some occasions, the species was present outside of caves while the caves were occupied by a different species (e.g., SE of Zebzat: Maghreba aurouxi gen. nov.; Grotte Kef el Ghar: Micropholcus sp.). On several occasions, H. reini was found on rocks very close to undescribed representatives of Micropholcus, occupying much the same microhabitat (Paradise Valley; Imi n’Ifri; near Ksiba). Near Sidi Ben Daoud, the spiders were extremely abundant in a very dense field of cacti. The domed webs transformed to distinct funnels that led into the barely accessible space among the cacti. High abundances were also reported by Hajer & Řeháková (2003), who observed “communities of up to several hundred individuals” (vouchers of their “ Crossopriza sp. ” were reexamined; see above).
Hajer & Řeháková (2003) also described web construction and the production of silk puffs, which were in principle identical to those of H. pluchei (see above). Unlike, H. pluchei, however, mated females built a more open spotted dome (rather than a sphere) within the main web and left this dome for attacking prey. Also, females tolerated the presence of males inside the dome and copulated repeatedly while still holding the egg sac. After the second molt of the spiderlings, females destroyed the dome and the spiderlings dispersed (Hajer & Řeháková 2003).
Distribution
The species is widely distributed in northwestern Africa, ranging from Morocco to Tunisia (Fig. 3). The MNHN has a female with a dubious label saying “ Espagne ”, while the original label is no longer readable.
description: Published as part of Huber, Bernhard A., 2022, Revisions of Holocnemus and Crossopriza: the spotted-leg clade of Smeringopinae (Araneae, Pholcidae), pp. 1-241 in European Journal of Taxonomy 795 (1) on pages 29-38, DOI: 10.5852/ejt.2022.795.1663, http://zenodo.org/record/6299150