Caracladus zamoniensis, Frick, Holger & Muff, Patrick, 2009

Frick, Holger & Muff, Patrick, 2009, Revision of the genus Caracladus with the description of Caracladus zamoniensis spec. nov. (Araneae, Linyphiidae, Erigoninae), Zootaxa 1982, pp. 1-37 : 20-26

Caracladus zamoniensis spec. nov.

( Figs 48–58 )

Caracladus avicula, Lessert 1907: 108 , figs 5–6, ♂ misidentified; Lessert 1910: 160, figs 98–99, ♂ misidentified.

Type material. HOLOTYPE: Switzerland: Grisons: Sur, Alp Flix , Salategnas , 1960 m [46°31'11.00'' N, 9°38'46.00'' E], 1♂ 24.x.2007, litter sieving, close to the edge of a subalpine forest of Norway spruce ( Picea abies ), leg. H. Frick, P. Muff, S. Klopfstein, det. H. Frick ( NMBE Ar6741 ). PARATYPES: Switzerland: Grisons: Sur, Alp Flix , Salategnas , 1960 m [46°31'11.00'' N, 9°38'46.00'' E], 3♂ 4♀ 24.x.2007, litter sieving, close to the edge of a subalpine forest of Norway spruce ( Picea abies ), leg. H. Frick, P. Muff, S. Klopfstein, det. H. Frick ( NMBE AR 6742 ); Sur, Alp Flix , Salategnas , 1960 m [46°31'09.01'' N, 9°38'50.07'' E], 1♀ 17.x.–06.v.2005, pitfall trap, in spruce forest, leg. P. Muff, det. H. Frick ( NMBE Ar6736 ) ( Muff et al. 2007); Sur, Alp Flix , Salategnas , 1960 m [46°31'11.00'' N, 9°38'46.00'' E], 1♂ 19.ix.–16.x.2005, pitfall trap, in spruce forest, leg. P. Muff, det. H. Frick ( NMBE Ar6735 ) ( Muff et al. 2007); Sur, Alp Flix , Salategnas , 1960m [46°31'11.00'' N, 9°38'46.00'' E], 1♀ 27.v.-24.vi.2005, pitfall trap, in spruce forest, leg. P. Muff, det. H. Frick ( MHNG) ( Muff et al. 2007); Sur, Alp Flix, Salategnas, 1960 m [46°31'11.00'' N, 9°38'46.00'' E], 1♂ 21.v.–24.vi.2005, pitfall trap, in spruce forest, leg. P. Muff, det. H. Frick ( MHNG) ( Muff et al. 2007); Sur, Alp Flix, Salategnas, 1960 m [46°31'11.50'' N, 9°38'41.89'' E], 1♂ 17.x.2005 – 06.v.2005, pitfall traps, in spruce forest, leg. P. Muff, det. H. Frick ( NMB 2795b) ( Muff et al. 2007); Sur, Alp Flix, Salategnas, 1960 m [46°31'11.00'' N, 9°38'46.00'' E], 1♀ 17.x.2005 – 06.v.2006, pitfall traps, in spruce forest, leg. P. Muff, det. H. Frick ( NMB 2795a) ( Muff et al. 2007); Sur, Alp Flix, Salategnas, 1960 m [46°31'11.50'' N, 9°38'41.89'' E], 1♂ 17.x.2005 – 06.v.2005, pitfall traps, in spruce forest, leg. P. Muff, det. H. Frick ( SMF) ( Muff et al. 2007); Sur, Alp Flix, Salategnas, 1960 m [46°31'09.24'' N, 9°38'47.74'' E], 1♀ 19.ix.-16.x.2005, pitfall trap, alpine timberline, leg. P. Muff, det. H. Frick ( SMF) ( Muff et al. 2007) .

Examined material. Austria: Vorarlberg: Montafon, Garneratal, close to Gaschurn, 1560 m [46°57'56'' N, 10°00'40'' E], 1♂ 19.vii.–29.viii.2000, leg., det. and coll. W. Breuss (Breuss unpubl.). France: Rhône- Alpes: Haute-Savoie, Chamonix, montagne des Posettes (Montroc), 1600 m [45°59'40'' N, 6°56'03'' E], 1♀ 18.viii.1993, spruce forest with some birch trees, ground dwelling, leg., det. and coll. J.-C. Ledoux (Ledoux unpubl.); Vallorcine, entrance to the canyon of Bérard, 1680 m [46°02'30'' N, 6°56'10'' E], 1♂ 17.viii.1993, underbrush of larch trees, in litter, leg., det. and coll. J.-C. Ledoux (Ledoux unpubl.). Provence-Alpes-Côte d’Azur: Alpes-de-Haute-Provence, Banon, ca. 800 m [44°02'16'' N, 5°37'40'' E], 1♂ 11.v.1986, leg. P. Poot, det. and coll. R. Bosmans (Bosmans unpubl.); Hautes-Alpes, Ceillac, ca. 1650 m [44°40'03'' N, 6°46'39'' E], 1♀ 04.viii.1980, leg. P. Poot, det. and coll. R. Bosmans (Bosmans unpubl.). Switzerland: Bern: Axalp, 1550 m [46°43'00'' N, 8°02'20'' E], 1♂ vi., leg. R. de Lessert, det. H. Frick ( MHNG) ( Lessert 1907). Grisons: Sur, Alp Flix, Salategnas, 1960 m [46°31'11.50'' N, 9°38'41.89'' E], 3♂ 17.x.2005 – 06.v.2005, pitfall traps, in spruce forest, leg. P. Muff, det. H. Frick ( NMBE AR 6740) ( Muff et al. 2007); Sur, Alp Flix, Salategnas, 1960 m [46°31'11.00'' N, 9°38'46.00'' E], 1♂ 21.v.–24.vi.2005, pitfall trap, in spruce forest, leg. P. Muff, det. H. Frick (coll. H. Frick, SP _0362) ( Muff et al. 2007); Sur, Alp Flix, Salategnas, 1960 m [46°31'11.00'' N, 9°38'46.00'' E], 1♀ 17.x.2005 – 06.v.2006, pitfall traps, in spruce forest, leg. P. Muff, det. H. Frick (coll. H. Frick, SP _0363) ( Muff et al. 2007); Trins, Mulins, above Purcs, ca. 1800 m [46°50'42.32'' N, 9°21'11.41'' E], 1♂ 2♀ 01.viii.1930, leg. E. Schenkel, det. P. Muff ( NMB 2795f) ( Schenkel 1933); Trins, Mulins, Bargis–Rischiglus–Furca–Flimserstein [46°51'30'' N, 9°17'30'' E], 1♀ 11.viii.1930, alpine zone, leg. E. Schenkel, det. P. Muff ( NMB 2795f) ( Schenkel 1933); Trins, Mulins, Belmont–Bargis, ca. 1550–2000 m [46°51'10'' N, 9°18'40'' E], 1♀ 21.vii.1930, leg. E. Schenkel, det. P. Muff ( NMB 2795f) ( Schenkel 1933); Trins, Mulins, below Alp Mora, ca. 1800 m [46°50'44'' N, 9°21'10'' E], 1♂ 2♀ 11.viii.1931, upper forest part, leg. E. Schenkel, det. P. Muff ( NMB 2795f) ( Schenkel 1933); Trins, Mulins, Si Munt-Uaul Sec, ca. 1200 m [46°50'0'' N, 9°21'10'' E], 1♂ 1♀ 04viii.1930, leg. E. Schenkel, det. P. Muff ( NMB 2795f) ( Schenkel 1933). Nidwalden: Bruniswaldalp close to Altzellen,> 1400 m [46°51'20'' N, 8°23'20'' E], 1♂ 4♀ viii., leg. E. Schenkel, det. P. Muff ( NMB 2795g) ( Schenkel 1923). Ticino: Val Bedretto, Bedretto to Alpe di Folcra, 1400–1800 m [46°30'8'' N, 8°30'59'' E], 1♀ 11.–22.vii.1927 /1928, forest slope on the right valley side, leg. E. Schenkel, det. P. Muff ( NMB 2795e) ( Schenkel 1929). Valais: close to Fiesch, Rafgarten – Ober Titer, 1500 m – 1600 m [46°30'50'' N, 8°18'20'' E], 6♀ 15.vii.1925, leg. E. Schenkel, det. P. Muff ( NMB 2795c) ( Schenkel 1926); Fionnay, 1500 m [46°01'54'' N, 7°18'26'' E], 1♂ 2♀ ix.1906, in moss of spruce forest, leg. R. de Lessert, det. H. Frick ( MHNG) ( Lessert 1907; Thaler 1972); Leukerbad, ca. 1400 m [46°22'30'' N, 7°37'30'' E], 1♂ 4♀ viii.1930, leg. R. de Lessert, det. H. Frick ( MHNG), 1♂ 1♀ viii.1930, leg. R. de Lessert, det. P. Muff ( NMB 2795h) (Lessert 1930); Lötschental, close to Ried, 1500 m – 1600 m [46°24'50'' N, 7°48'20'' E], 1♂ 11♀ vii.1938, leg. E. Schenkel, det. P. Muff ( NMB 2795i) ( Schenkel 1939); Saas-Tal, Saas-Tal below Saas-Fee, Almagell–Saas-Fee, ca. 1600 m [46°06'30'' N, 7°55'40'' E], 1♂ vii./viii., leg. E. Schenkel, det. P. Muff ( NMB 810d) (Schenkel unpubl.) .

Diagnosis. C. zamoniensis spec. nov. is most similar to C. avicula but differs in the shape of the male and female genitalia and the shape of the male cephalic lobe.

Males: Cephalic lobe of C. zamoniensis spec. nov. more robust than in C. avicula : the neck-like prolongation of C. zamoniensis spec. nov. is of equal diameter directly below and above the eye-field (AME, ALE, PLE) ( Fig. 54 ) but much thinner below the eye-field in C. avicula ( Fig. 23 ); distance between sulcus and AME is below 0.11 mm in C. zamoniensis spec. nov. ( Fig. 53 ) and above 0.12 mm in C. avicula ( Fig. 22 ); sulcus cup-like in C. zamoniensis spec. nov. and channel-like in C. avicula . Embolus of C. zamoniensis spec. nov. short, broad and robust basally, thin and U-shaped distally ( Figs 49, 50 ); C. avicula with long, straight and whip-like embolus that narrows constantly towards the end ( Figs 18, 19 ). C. zamoniensis spec. nov. tibia I proximally bent and dorsally with glabrous area on the proximal half ( Fig. 55 ) and no macroseta, in C. avicula with one dorsal macroseta in small glabrous field ( Fig. 24 ).

Females: Epigyne of C. zamoniensis spec. nov. with two anterior pouches formed by the ventral and dorsal plate, anterior borders highly sclerotised ( Fig. 56 ). Pouches in C. avicula much larger and less sclerotised ( Fig. 26 ). C. zamoniensis spec. nov. with ventrally visible square dorsal plate, sclerotised parts of the vulva visible in transparency through ventral and dorsal plate defining a bright hourglass-like form centrally ( Fig. 56 ). C. avicula with rectangular dorsal plate without sclerotised parts visible in transparency through dorsal plate but lateral to it ( Fig. 26 ). Vulva of C. zamoniensis spec. nov. without copulatory duct, those of C. avicula with. Vulva of C. zamoniensis spec. nov. simple with hook-like sclerotised pouch borders, originating anterior and mesal to the receptacula ( Figs 57, 58 ), in C. avicula shapes more complex ( Figs 27, 28 ).

Description. Male (Holotype, NMBE Ar 6741): Total length 2.18 mm. Cephalothorax: honey brown (138 U); reticulated; broad oval; 0.85 mm long without cephalic lobe ( Fig. 54 ), 1.22 mm long with cephalic lobe ( Fig. 54 ); 0.65 mm wide. Cephalic lobe: honey brown (138 U); shaft with few long hairs ( Fig. 52 ); shaft constantly thick, at thinnest part below the eye-field 0.10 mm wide laterally, 0.11 mm wide dorsally ( Figs 52, 54 ); tip of lobe laterally flattened with many short, stout and few long, slender hairs anterior to the PME ( Figs 52, 54 ); sulcus 0.08 mm below AME ( Fig. 53 ). Eyes: PME topmost on the cephalic lobe; AME projecting forward, lateral eyes besides the AME; one long macroseta projecting forward between AME ( Fig. 54 ). Clypeus: directed obliquely backwards. Sternum: very fine brown (469 U) pigmentation on yellow (124 U) ground, dark brown (469 U) on the margins; 0.47 mm long; 0.51 mm wide; shield-shaped. Chelicerae: yellow (124 U); promargin with 5 teeth; retromargin with 5 denticles; stridulatory striae very dense and fine. Legs: yellow to light brown (120 U); formula 4-1-2-3; tibia I proximally bent and dorsal with glabrous area from proximal to more than half its length ( Fig. 55 ), tibia III–IV with one dorsal proximal macroseta (0-0-1-1); metatarsi I–III with one trichobothrium, Tm I: 0.54 mm, metatarsus IV without trichobothria. Pedipalp: patella two times longer than broad, tibia retrolateral with expansion (round glabrous area, Fig. 51 ), one retrolateral and one prolateral trichobothrium ( Fig. 51 ); paracymbium a simple clasp; tegulum distal with short and long papillae on protegulum ( Fig. 48 ); suprategular apophysis semi-circular; marginal suprategular apophysis rather small, emerging close to the tip; distal suprategular apophysis robust, highly sclerotised ( Figs 49, 50 ); column broad; embolic membrane slender; radix simple without any processes other than the elongated radical tailpiece and the embolus; embolus strongly sclerotised, twisted; broad at the base; very thin, curved tip ( Fig. 50 ). Abdomen: dark olive green-brown (125 U); booklung covers very light brown (467 U); scaly.

Female (Paratype, NMBE Ar 6742): Total length 1.81 mm. Cephalothorax: honey brown (138 U); reticulated; 0.89 mm long; 0.65 mm wide. Eyes: posterior row slightly procurved; anterior row straight. Sternum: very fine brown (469 U) pigmentation on yellow (124 U) ground, dark brown (469 U) on the margins; 0.46 mm long; 0.46 mm wide; shield-shaped. Chelicerae: honey brown (138 U); promargin with 5 large teeth; retromargin with 5 denticles; stridulatory striae very fine and dense. Legs: yellow (122 U); formula 4-1-2-3; tibia I–IV with one dorsal proximal macroseta (1-1-1-1); metatarsi I–III with one trichobothrium, Tm I: 0.52 mm, metatarsus IV without trichobothria. Epigyne: simple with hook-like sclerotised pouch borders, originating anteriorly and mesally to the receptacula ( Figs. 57, 58 ); dorsal plate square, fully visible in ventral view; sclerotised parts of vulva visible in transparency through ventral and dorsal plate, defining a bright hourglass-like form centrally ( Fig. 56 ). Vulva: without copulatory duct; receptacula globular, incoming dorsally. Abdomen: dorsal olive green-brown (119 U), ventral darker (147 U).

Variation. The measurements are based on all type material (10♂ 9♀) plus specimens from the NMB (810i: 1♂ 2♀) and the MHNG (Axalp: 1♂; Fionnay: 1♂ 1♀).

Males (n=13, means in brackets): The coloration is variable. Total length 1.91–2.18 mm (2.09 mm). Cephalothorax: 0.73–0.86 mm (0.82 mm) long without cephalic lobe, 1.10–1.23 mm (1.18 mm) long with cephalic lobe; 0.61–0.69 mm (0.65 mm) wide. Cephalic lobe: at thinnest part below the eye-field 0.10–0.13 mm (0.11 mm) wide laterally, 0.09–0.11 mm (0.11 mm) wide dorsally; sulcus 0.07–0.11 mm (0.08 mm) below AME ( Fig. 53 ). Legs: Tm I: 0.50–0.59 mm (0.54 mm).

Females (n=12, means in brackets): The colorations are variable. Total length 1.62–2.00 mm (1.82 mm). Cephalothorax: 0.75–0.89 mm (0.82 mm) long; 0.60–0.65 mm (0.62 mm) wide. Legs: Tm I: 0.48–0.60 mm (0.53 mm).

Distribution. Endemic to the Alps, occurring in the Western- and Central Alps in France, Switzerland and Austria ( Fig. 59 ). The Eastern distribution border seems to be in Western Austria. Checking of specimens of C. avicula collected west of Vorarlberg ( Austria) revealed no misidentifications.

Habitat. C. zamoniensis spec. nov. occurs in the litter layer of Norway spruce ( Picea abies ) forests at the alpine timberline. Most sampling sites were inside the forest with no direct sunlight under branches of Norway spruce. The collection site and its surroundings were sampled intensively in two previous studies ( Frick et al. 2006; Frick et al. 2007; Muff et al. 2007). We found no specimens of C. zamoniensis spec. nov. around stand alone trees in the dwarf-shrub heath with a similar microclimate as the closed forests. C. zamoniensis spec. nov. seems to avoid the open land. We only found two specimens in more open areas in the dwarf-shrub heath close to the subalpine forest. C. zamoniensis spec. nov. was never collected together with C. avicula in the same pitfall trap but already in pitfall traps about 20 m away from C. avicula . The locus typicus is approximately 50 m away from the alpine timberline in the subalpine deciduous forest. We found the type specimens in litter under snow close to the tree trunk under a Norway spruce ( Fig. 47 ) at 1960 m a.s.l. Other specimens were found between 1400–2000 m in litter and moss of spruce forests (e.g. Schenkel 1939). One record was much lower at app. 800 m in France (Bosmans pers. comm.).

Phenology. This species seems to be eurychronous. All records of other authors at altitudes from 1400 m to 1800 m were between July and September. However, at the type locality (1960 m) specimens were exclusively found between September and June. This corresponds with the time between the first snow fall and the beginning of the snow free time.

Etymology. The cephalic lobe of the male is morphologically very similar to the noses of the so called dwarf pirates and other imaginary figures from “Zamonia”. Zamonia is a continent inhabited by freaky creatures in the novel “The 13 ½ Lives of Captain Bluebear” by the German writer Walter Moers (2000). Translated, the species name means “ Caracladus from Zamonia”.

Remarks. C. zamoniensis spec. nov. lacks a copulatory duct. The insertion of sperm is assumed to take place through a space between the ventral and the dorsal plates which are supposed to be pressed apart during copulation.

The specimens that Lessert (1907, 1910) shows have been evaluated by H.F. The figures of males in Lessert (1907: figs 5, 6) and reprinted in Lessert (1910: figs 98, 99) show C. zamoniensis spec. nov. and not C.

avicula . The female mentioned in Lessert (1907: fig. 7) and Lessert (1910: fig. 100) shows Diplocentria bidentata ( Emerton, 1882) ( Thaler 1972) .

The specimen that was pictured by Pesarini (1996: figs 9–10) was not available to the authors. A definite assignment to either C. avicula or C. zamoniensis spec. nov. is not possible. However, his records are referred to as C. avicula in the distribution map ( Fig. 59 ) and the list of records.

The remaining pictures so far named as C. avicula in Heimer and Nentwig (1991: figs 350.1–350.5), Millidge (1977: fig. 162), Simon (1884: figs 408, 409 and fig. 8 on plate 27) and Thaler (1969: figs 16–21, 1972: figs 7–11) are correctly assigned to C. avicula .

Figures

FIGURE 47. Type locality of C. zamoniensis spec. nov .. Type specimens found in sieved Norway spruce litter, collected close to the tree trunk under approximately 5 cm of snow on 24 October 2007.

FIGURE 59. Distribution map of the European Caracladus species. C. avicula (L. Koch, 1869) (white circles); C. leberti (Roewer, 1942) (grey asterisks) and C. zamoniensis spec. nov. (black circles).

FIGURES 48 – 55. Caracladus zamoniensis spec. nov .. Male: palp retrolateral (48), prolateral (49), embolic division prolateral (50); palpal tibia dorsal (51); cephalothorax dorsal (52), frontal (53) and lateral (54); right leg tibia I retrolateral (55). Scale: 200 Μm. Specimens: Paratypes, NMBE Ar 6742 (palp), Ar 6735 (cephalothorax).

FIGURES 56 – 58. Caracladus zamoniensis spec. nov .. Female: epigyne ventral (55), vulva ventral (56), vulva dorsal (57). Transections through sclerotised walls marked in grey. Scale: 100 Μm. Specimens: Paratype, NMBE Ar 6736.

FIGURES 17 – 24. Caracladus avicula (L. Koch, 1869). Male: palp retrolateral (17), prolateral (18), embolic division prolateral (19); palpal tibia dorsal (20); cephalothorax dorsal (21), frontal (22) and lateral (23); right leg tibia I retrolateral (24). Scale: 200 Μm.

FIGURES 25 – 28. Caracladus avicula (L. Koch, 1869). Female: tracheal system dorsal (25); epigyne ventral (26), vulva ventral (27), vulva dorsal (28). Transections through sclerotised walls marked in grey. Scale: 100 Μm.