Gollumjapyx smeagol Sendra & Ortuño, 2006

Sendra, Alberto, Ortuño, Vicente M., Moreno, Agustín, Montagud, Sergio & Teruel, Santiago, 2006, Gollumjapyx smeagol gen. n., sp. n., an enigmatic hypogean japygid (Diplura: Japygidae) from the eastern Iberian Peninsula, Zootaxa 1372, pp. 35-52 : 42-49

publication ID	https://doi.org/ 10.5281/zenodo.174943
DOI	https://doi.org/10.5281/zenodo.5677135
persistent identifier	https://treatment.plazi.org/id/0576944C-FFCC-FF99-2F09-4CB80C9EF6AF
treatment provided by	Plazi
scientific name	Gollumjapyx smeagol Sendra & Ortuño
status	sp. nov.

Treatment

Gollumjapyx smeagol Sendra & Ortuño sp. nov.

Holotype: Ψ 23.40 mm (cerci inclusive) nº 0 815, Castellón, Serra d’en Galceran, “Cova del Mas de Gaspar” cave; 26-V-1980; J. Comas & O. Escolà leg.; preserved in ethanol 70º + glycerine and deposited at the Barcelona Zoology Museum ( Spain).

Paratypes: 1 Ψ, Tarragona, Ulldecona, “Avenc Canals” pit; 11-III-2000; F. Fadrique leg. 1 Ψ, Castellón, Cabanes, “Avenc d’en Serenge” pit; 14-III-1982; J. Comas leg. 1 sex?, idem; 12-XI-1995; E. Carabajal leg. 1 ɗ, idem; 19-VII-2003; S. Montagud leg. 2 sex?, idem; 26-II-2006; S. Montagud & A. Sendra leg. 1 ɗ, Castellón, Cabanes, “Avenc Mas de la Cova” pit; 22-VII-1995; E. Carabajal leg. 1 ɗ, Castellón, Coves de Vinromá, “Avenc Mas Nou” pit; 12.III.2005; S. Teruel leg. 1 sex?, Castellón, Sant Mateu, “Cova dels Encenalls” cave; 9-III-2006; J. Comas leg. Preserved in ethanol 70º or 96º and deposited at: Muséum d’Histoire Naturelle Genève, Switzerland; National Museum of Natural Sciences, Madrid, Spain; Barcelona Zoology Museum ( Spain) and Valencia Natural History Museum ( Spain) — Fundación Entomolólogica Torres Sala-.

Total paratypes: 9 specimens; 3 ɗ ɗ, 2 Ψ Ψ and 4 sex?.

Etymology: the specific epithet refers to Gollum’s original name, who, in his epigean origin, was a hobbit named Smeagol .

DNA barcode: A nucleotide sequence of the 5' half of the cytochrome c oxydase subunit I (cox1) has been deposited in Genbank with accession number DQ993154 View Materials .

Body* Cerci** Antennae** Antennomeres** Legs III** sex?, Avenc d’en Serenge broken and - 8.0 5 55 -

not complete

♂, Avenc Mas Nou 13.26 1.50 7.40 53 4.81

♂, Avenc d’en Serenge 12.00 1.17 6.75 54 4.76

* Length body without cerci.

** Measurements taken of the two appendixes from the same specimen, although only the longest is recorded. In the case of two intact antennae from the same specimen, only the antennae with the most antennomeres is recorded.

Description: Long body, maximum and minimum body length (excluding cerci) from 21.20 to 12.00 mm. Other lengths described in Table 1 View TABLE 1 . All three thoracic segments show a remarkable lengthening more obvious in mesothoracic and metathoracic prescutum.

Almost totally unpigmented cuticle. More or less apparent sclerotization of buccal pieces, anterior margin of head, claws and a large portion of urite X, becoming particularly evident in cerci. Abdominal segments clearly show a lateral expansion in the tergi, increasingly apparent from segment I to VII ( Fig. 1 View FIGURE 1 and 2 a View FIGURE 2. a ). Ordinary setae noticeable on head, pronotum and mesonotum, more abundant in larger specimens. These ordinary setae are shorter in the mesonotum, becoming even shorter and sparse in the metanotum and in the abdominal sclerites ( Fig. 4 View FIGURE 4 ).

Head. Antennae with 53 to 55 antennomeres, measuring around half of body length ( Table 1 View TABLE 1 ); 13 typical trichobothria on antennomeres IV–VI (1 ventral and 2 dorsal in IV), a clearly distal; all antennomeres generally pilose ( Fig. 3 View FIGURE 3 b); antennomeres II to IV are largest and from antennomere V tapering to penultimate antennomere; pilose patches dense with long, robust setae, arranged in two whorls in proximal antennomeres, tapering to one whorl in distal antennomeres. These long and robust setae are accompanied by many thin setae, clearly shorter, covering almost all of each antennomere. Joining the pilose patch are many long, fine and hooked setae-shaped sensilla and also a few fine and short sensilla ( Fig. 3 View FIGURE 3 a). Placoid sensilla of apical antennomere 14 to 16 in number, in two random groups ( Fig. 3 View FIGURE 3 a). One or two placoid sensilla in the penultimate antennomere in some of the studied specimens. In three of the specimens from “Avenc d’en Serenge” placoid sensilla appear from antennomere XI or XII, ventrally and the next antennomeres show 0, 1 or 2 placoid sensilla ( Fig. 3 View FIGURE 3 b).

Simple vertex setae increase noticeably the larger the body size of the specimen ( Fig. 4 View FIGURE 4 a, b). However, the number and pattern of the large setae does not vary in the specimens under study. A comparison with the pattern of large setae proposed by Pagés (1984) in the case of Indjapyx uvaianus Pagés, 1984 proved complex; nonetheless the setae have been numbered, comparing their pattern of distribution. Out of 40 large setae proposed for I. uvaianus distributed in seven groups (A, S, V, M, I, L y P), only 14 seem to appear in G. smeagol ( Fig. 4 View FIGURE 4 a).

Buccal pieces are consistent with the family: five laminae of the internal lobe of the maxilae with all five laminae pectinate; labial palpi 3.70 times longer than wide at base (423 μm holotype) covered with long setae; submentum with 1+1 large setae (almost equivalent to real macrosetae), postmentum with 1+1 large setae in the internal margin of the labial palpus; admentum with some large setae of which 2+2 are more noticeable; prementum (external lobe) with many setae among which a dozen small short setae. All large setae are longer than the labial palpus.

Thorax. Extremely elongate ( Fig. 1 View FIGURE 1 and 2a View FIGURE 2. a ). Pronotum ( Fig. 4 View FIGURE 4 c): typical 5+5 M and 1+1 M posterior lateral. Ordinary setae abundant, increasing relative to body length. Mesonotum ( Fig. 4 View FIGURE 4 d): prescutum with few setae longer than ordinary (0+1 M in the paratype from “Avenc Canals”); scutum with 5+5 M (M1 and M 2 in more posterior position, M1 even shorter than half of those remaining); with ordinary setae but less abundant than in the pronotum. Metanotum ( Fig. 4 View FIGURE 4 e): Prescutum with 1+1 M or 0+1 M (male from “Avenc Mas de la Cova” lacking M); scutum with 4+4 M (M2 absent; M1 shorter than those remaining); ordinary setae very short and abundant.

Exceptionally long legs. The end of metathoracic pretarsus exceeds half of urite X. Length (units in mm) of the metathoracic leg of the holotype (total length 9.79 mm): coxa (0.95), trochanter (0.60), femur (2.85), tibia (3.06), tarsus (1.92) and pretarsus (0.41). The tibia is the longest segment in the legs. In all specimens examined legs length is either equal or inferior to half the total length of the body ( Table 1 View TABLE 1 ). Femur with short ordinary setae; femur and tibia with dozens of large setae; tarsus with 16–18 spiniform setae scattered in two ventral rows; well developed claws, posterior 1.2 times longer than anterior (holotype).

Abdomen. Tergite I. Prescutum and scutum: 0+0 M. Tergite II: 1+1 M (ma=M). Tergites III–VII ( Fig. 5 View FIGURE 5 a): 5+5 M and 1+ 1 m 3 (ma=M, M1 absent). Tergite VIII ( Fig. 5 View FIGURE 5 a) nearly 2.30 times wider than long, 5+5 M (2+2 anterolateral, 2+2 posterolateral and 1+1 consistent with M5). Tergite IX ( Fig. 5 View FIGURE 5 a) around 3.30 times wider than long, M absent but with 4+4 M in the paratype from “Avenc Canals”.

Urite X ( Fig. 3 View FIGURE 3 c and 5b) nearly 1.60 longer than wide, distinctly marked carinae with subparallel margins, 4+4 M intracarinal (D) (3+3 M lateral and 1+1 M posterior) in both small and large specimens, although larger specimens show 3+3 large setae on both sides preceding posterior macrosetae; 5+5 M carinae (L).

Very short ordinary setae, practically absent from the first tergites and progressively more abundant from tergite V.

Acropygium triangular, slightly protruding, rounded ( Fig. 3 View FIGURE 3 c and 5b).

Tergites I–IV showing posterolateral angles either blunt or slightly rounded; angles found in tergite V are more conspicuous due to a small lobiform projection ( Fig. 5 View FIGURE 5 a); tergites VI and VII show this projection more manifestly, although they are not visibly sclerotized ( Fig. 5 View FIGURE 5 a); tergite VIII maintains this angular lobe but shorter than the two preceding ones; finally, the posterior angle of tergite IX lacks the aforementioned projection ( Fig. 5 View FIGURE 5 a).

Relative lengths of segments VI–X ( Fig. 5 View FIGURE 5 ): 28–28–33–20–100.

Sternite I ( Fig. 6 View FIGURE 6 a): Prescutum 5+5 M; scutum: 22+22 M and other 7–9+7–9 large setae consistent with true macrosetae; sternite totally covered with short ordinary setae, increasing in posteriorly and with longer and more robust setae scattered in two or three rows at the front of the lateral subcoxal organs (between 80 in the male specimen from “Avenc Mas de la Cova” and 60 in the female specimen from “Avenc Canals”). Subcoxal organs appear either invaginated below the posterior margin of sternite I or evaginated outwards, protruding over the margin of the urosternite ( Fig. 6 View FIGURE 6 ).

The rows of setae on the subcoxal organs ( Fig 6 View FIGURE 6 b) represent between 1/5 to 1/3 of interstyle width (0.19–0.27 in females and 0.25–0.57 in males and 0.31 in a specimen of unknown sex from “Avenc d’en Serenge”). About 65 short glandular setae in the male from “Avenc del Mas de la Cova”, set in 1 or 2 rows, GS/st1= 0.14; nearly 80 glandular setae, equally short, in the female from “Avenc Canals”, in 2 rows, GS/st1= 0.13; 25–28 sensory setae, SS/st1= 0.10 in the male and 35–38 sensory setae, SS/st1= 0.10 in the female.

Both visible median glandular organ and central setae-shaped sensilla missing. These sensilla appear in 4–8+ 4–8 in the internal side of the subcoxal organs ( Fig. 6 View FIGURE 6 a).

Urosternites II–VII: 28+28 M, although missing some typical macrosetae (C2, C8, C10). Urosternite VIII: 12+12 M, in the largest specimens and 7+7 M in the smallest. Paratergite VIII: 1 M subposterior.

Elongate styli with sharp sclerotized points: s1/s7= 0.44–0.94; st1/st7= 0.59–0.76; s1/st7= 0.24–0.46; s1/st7= 0.26–0.59.

Male genital papillae could not be examined in the largest specimen from “Avenc Mas de la Cova”; however, genital appendices are wholly membranous and cylindrical-conic. No ventral glandular fossae have been found in males.

Cerci. ( Fig. 3 View FIGURE 3 c, d and 5b). Large, at first slightly curved, becoming more curved towards the extreme, pointed. Length ranges from 3.25 mm in holotype to 1.17 mm in the smallest specimen; the relative size of the cercus increases larger in larger specimens ( Table 1 View TABLE 1 ). In all specimens the length of the right cercus is slightly inferior to that of the left (0.95 to 0.97). Cerci slightly longer than the normally uncovered part of urite X in smaller specimens, but shorter in the larger specimens. Both cerci heavily sclerotized, in particular both external and internal margins and also in the dorsal carinae projecting from the acetabula, running the length of the right cercus and halfway on the left.

Right cercus. Triangular tooth, very strong, distinctly proximal, r d= 0.40–0.65 (ΨΨ), 0.57–0.58 (ɗɗ); width of cercus at tooth level equals that of base; predental margin with 2 to 4 strong tubercles and very prominent in larger specimens, wherein 1 or 2 appear to be bitubercles; in smaller specimens tubercles are simple and smaller. Postdental arista is slightly concave, surrounded by tiny rounded denticles, tapering in size extremely even to the point of disappearance.

Left cercus. Sharp postmedian tooth, r g= 1.14–1.60 (Ψ), 1.33–1.51 (ɗ); width of cercus at tooth level slightly larger or equal to half the base of the appendix; concave predental arista with 8–12/11–15 sharp/rounded tubercles; postdental arista slightly concave with very small denticles or showing a smooth surface in the smallest specimens.

Chaetotaxy: in dorsal view, the inside margin of both cerci has anterolateral M and a large setae, at tooth level, in the right cercus. The outside margins of both cerci support a dozen large setae each.

Comments. During this study, our colleague at the University of Murcia, José Luis Lencina, gave us a large japygid specimen (body length 22.6 mm, including cerci), displaying troglobiomorphic features. Both the cercal armature and the lengthening of thorax and legs in this specimen show a strong resemblance to Gollumjapyx smeagol ; its poor state of conservation, however, made it impossible to conduct a thorough study that could have confirmed or denied its inclusion in the species described in the present article.

Here follow the collection data: 1 Ψ, Comunidad de Murcia, Murcia, Jumilla, “Cueva del Pozo”; 8-XII-1992; Grupo espeleológico Hinneni leg. (deposited at the Valencia Natural History Museum, Spain). Pozo cave (Jumilla, Murcia, Spain) is a fossil cave, more than 1 km in length and some 50 metres deep, located at an altitude of 498 metres, open in limestone and dolomites from the upper cretaceous at the Rajica de En Medio range, in the southern Prebetic mountain range. The specimen was collected 200 metres into the entrance, on damp clayey soil (despite low humidity), at a mean temperature of 26ºC, higher than expected according to the annual average for this locality (unpublished data, forwarded by the Hinneni Group).

Distribution, habitat and biology. To date Gollumjapyx smeagol gen. n. sp. n. has been found in six caves scattered along the limestone relief of “The Coastal Ranges”. This geomorphological region defined by Garay (1995), comprises the low altitude coastal ranges (400–600 metres average height) from the Oropesa range (Castellón, Spain) to the Montsià range (Tarragona, Spain). It displays a series of particular palaeogeographic and stratigraphic traits which have seemingly allowed it to remain a refugium area throughout the Cenozoic era for a host of species, presently recognised to be relics or paleo-endemics ( Ortuño et al., 2005).

The northernmost locality is the “Avenc Canals” in the Montsià range, a small cave with stalagmite formations. The specimen was retrieved at the base of the entry shaft, in a formation 20–40 cm from a debris base made up of dirt and vegetable detritus (F. Fadrique com. pers.).

In the “la Vall d’Àngel” range, towards the south, there is “Cova dels Encenalls” cave; it hosts a considerable catalogue of troglobite elements, including Zuphiini carabid Ildobates neboti Español, 1965 and Anillini carabids Speleotyphlus jusmeti (Español, 1971) and Iberanillus vinyasi Español, 1971 ; Dysderidae Araneae Speleoharpactea levantina Ribera, 1982 ; Neobisidae Pseudoscorpiones Acantocreagris relicta Mahnert, 1977; and Campodeidae Diplura Paratachycampa hispanica Bareth & Condé, 1981 . This cave is, for the greater part of its 75 m width and only 10 m depth, covered in stalagmite floors, with small gours, generally flooded. The specimen was found wandering on the stony soil of a side gallery (J. Comas com. pers.).

“Avenc del Mas Nou” pit lies on the southern foothills of the “Serra d’en Galceran” range; access to its 28 m drop is gained through a narrow gap. The cave extends for 150 m and reaches a depth of 40 m. It is covered with stalagmite formations as well as clay deposits, all in an extremely humid environment. The specimen was collected crawling on the damp formations at over 30 m depth. Also spotted in that same area were many specimens of two detrivorous species of Campodeidae Diplura : Paratachycampa hispanica and Campodea maestrazgoensis Sendra & Escolà, 2004 . In this same mountain range there is a small cave, less than 70 m of practically horizontal extension, known as “Cova del Mas de Gaspar” or “Cova Santa”, where the first specimen of this new taxon was discovered — which has been used as holotype in the description. Gollumjapyx smeagol shares this cave with other troglobite elements such as Leiodidae Coleoptera Anillochlamys cullelli Lagar, 1978 and Campodeidae Diplura Campodea (Campodea) maestrazgoensis .

The two southernmost localities are two pits in close proximity, the “Avenc d’En Serenge” and the “Avenc Mas de la Cova”. Both are located in limestone elevations near the town of Cabanes, in Ferradura mountain. The “Avenc d’en Serenge”, as well as being the typical locality of Ildobates neboti , is home to a great variety of troglobite forms with great interest from a palaeo- and biogeographical point of view: Neobisidae Pseudoscorpiones Troglobisium racovitzai (Ellingsen, 1912) ; Campodeidae Diplura Paratachycampa hispanica ; Anillini carabid Speleotyphlus aurouxi (Español, 1966) and Pselaphidae Tychobythinus escolai (Besuchet, 1974) . Five of the specimens of Gollumjapyx smeagol come from this cave which extends for 110 m and 34 m depth. Four of the aforementioned specimens were collected in the deepest and dampest areas of the cave, totally covered by calcareous formations, at more than 20 m depth. In the case of the “Avenc del Mas de la Cova” pit no precise data are available about the way the specimen was located. The cave runs for more than 141 m and reaches a depth of 43 m. From the base of the first shaft at 17 m depth, the cave remains very damp, with an abundance of stalagmitic formations and dripstones.

Gollumjapyx smeagol has been found in mesovoid deep substratum, in caves of average extension and little depth, excavated in limestone from the Lower Cretaceous, in altitudes between 300–500 metres. Ground temperature ranges from 14ºC to 17ºC, always in humid areas. It is therefore likely to be stenoterm and stenohygrobious, although not strictly, which would presumably allow it to survive in mesovoid shallow substratum (MSS).

It is well known that the most common feeding habit in Japygidae is to predate on smaller invertebrates. It has been stated that japygids refuse to feed on dead organisms ( Pagés, 1967). Even though feeding in Gollumjapyx smeagol has not been observed, a thorough examination of the digestive content has been performed. Two specimens from the “Avenc d’en Serenge” showed fragments of Acari (legs and sclerites) inside the digestive tract. A third specimen from the same cave showed fragments of Speleotyphlus aurouxi (5 antennomeres, a leg, part of the ring of the genital segment and the whole aedeagus- median lobe and parameres still articulated) ( Fig. 2 View FIGURE 2. a b).