(Collembola, Sminthuridae): A new cave species from ...

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Disparrhopalites tergestinus (Collembola, Sminthuridae): A new cave species from northeastern Italy a

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Pietro Paolo Fanciulli , Andrea Colla & Romano Dallai

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Dipartimento di Biologia Evolutiva , Università di Siena , Via Aldo Moro 2, Siena, I‐53100, Italy b

Museo civico di Storia Naturale , Piazza Hortis 4, Trieste, I‐34123, Italy

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Dipartimento di Biologia Evolutiva , Università di Siena , Via Aldo Moro 2, Siena, I‐53100, Italy Published online: 28 Jan 2009.

To cite this article: Pietro Paolo Fanciulli , Andrea Colla & Romano Dallai (2005) Disparrhopalites tergestinus (Collembola, Sminthuridae): A new cave species from northeastern Italy, Italian Journal of Zoology, 72:2, 167-173, DOI: 10.1080/11250000509356667 To link to this article: http://dx.doi.org/10.1080/11250000509356667

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Ital. J. Zool., 72: 167-173 (2005)

Disparrhopalites tergestinus (Collembola, Sminthuridae): a new cave species from northeastern Italy PIETRO PAOLO FANCIULLI Dipartimento di Biología Evolutiva, Università di Siena, Via Aldo Moro 2, 1-53100 Siena (Italy)

ANDREA COLLA Museo civico di Storia Naturale, Piazza Hortis 4, I-34123 Trieste (Italy)

ROMANO DALLAI

Downloaded by [183.218.89.154] at 19:33 21 March 2014

Dipartimento di Biologia Evolutiva, Università di Siena, Via Aldo Moro 2, I-53100 Siena (Italy)

INTRODUCTION The genus Disparrhopalites was erected by Stach (1956) to include some specimens from the genus Pararrhopalites with peculiar features concerning the number of eyes, the shape of the foot complex and the dens. The genus includes only one species, D. patrizii, formerly described by Cassagnau & Delamare Deboutteville (1953) as Pararrhopalites patrizii. Disparrhopalites patrizii is considered a troglophilic species and it has been found in several European sites (Fig. 1), as well as on the Canaries and Azores islands (da Gama, 1988); most collecting sites are in fact from caves, catacombs and artificial refuges (Dallai, 1973; Christian, 1998) even though the species can also be found in open habitats (Dallai, 1973; Schieuter, 1985; Ahrens, 1989, Landbau. Diss., Univ. Bonn). Dallai (1970a) discussed the systematic position of the genus Disparrhopalites on the basis of the structure of the cuticle, a character he considered a useful feature for separating Disparrhopalites from the genera Arrhopalites, Sminthurides and Sminthurinus. The differences in cuticle structure between Disparrhopalites and Arrhopalites appear of great interest because they allow us to reject the proposed synonymy of Disparrhopalites with the genus Arrhopalites, as suggested by Richards (1968) supporting an earlier conclusion by Delamare Deboutteville & Bassot (1957). Dallai (1970b) established the close relationship of Disparrhopalites with the genus Gisinurus, and Betsch (1980), and more recently Bretfeld (1999), ultimately placed it within the family Sminthuridae. In the last few years, we had the opportunity to study many collembolan species collected in several Italian caves. In one of these samples a new species of Disparrhopalites, showing clearly different characters from D. patrizii, was found. The aim of the present paper is to describe this new taxon.

MATERIALS AND METHODS

ABSTRACT A new sminthurid Collembola cave species is described from northeastern Italy: Disparrhopalites tergestinus n. sp. The new taxon is characterised by the lack of ocelli, the subdivision of the IV antennal article into 14 sub-articles and the claw without inner teeth. These features allowed the new species to be distinguished from the congeneric D. patrizii, the other species of the genus known so far. The systematic position of the new species and the distribution of the genus are discussed. KEY WORDS: Collembola - Sminthuridae - Disparrhopalites Taxonomy - Cave species. (Received 21 September 2004 - Accepted 14 December 2004)

One specimen (cD of D. tergestinus n. sp. and some of D. patrizii were prepared for scanning electron microscopy observations. Material preserved in 80° ethanol was dehydrated in absolute ethanol, critical-point-dried in a Balzer CPD 030 apparatus, and coated with gold in a Balzers MED 010 sputter coater. Observations were performed with a Philips XL20 scanning electron microscope. Specimens of D. patrizii were from Susegana (Treviso Province) near Castello S. Salvatore (Forte and Vanin Leg).

TAXONOMIC ACCOUNT

Disparrhopalites tergestinus n.sp. Material

One holotype ($) and three paratypes (2$ and lcf) of D. tergestinus n. sp. were found in the cave named "Grot-


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Fig. 1 - Distribution map of Disparrhopalites patrizii (open points) and D. tergestinus n. sp (triangle).

ta dei Partigiani" located near Basovizza (Province of Trieste), along the road connecting Basovizza to Gropada (no. 3477 V.G. = 689R; G.C. N45°39'12.5", E13O51'23,7", 373 m a.s.L, at 11 °C) (A. Colla Leg.). Holotype and one paratype are conserved in the collection of R. Dallai at the University of Siena Department of Evolutionary Biolo-

gy; one paratype is conserved in the collection of Andrea Colla at the Trieste Natural History Museum. Etymology

The specific name is from the ancient name for Trieste, Tergeste, where the species was found. Description

Fig. 2 - Scanning electron microscopy of the cuticle of D. tergestinus n. sp. (A-B) and D. patrizii (C-D). Both species show a ground plan constituted by primary granules arranged as hexagonal meshes (A, C); the dorsal surface of the large abdomen is covered by secondary granules derived from the fusion of the primary ones (B, D).

Size (measurements and ratios from the holotype): habitus of the species as in Figure 7A; length of body 880 pm; diagonal of head 495 um. Body pigmentation absent. Epicuticle of the head and both sides of the great abdomen, as well as on the ventral body region, antennae, legs and furca with primary granules arranged in hexagons (Figs 2A, 3A, C, 4A-D). On the contrary, the dorsal surface of the great abdomen is covered by secondary granules derived from the fusion and growth of some primary granules; small areas with only primary granules are visible intermingled with others provided with the more complex structures (Fig. 2B). Total length of the antenna 2.140 mm; ratio between length of antennae and head diagonal = 4.3. Relative lengths of antennal articles I : II : III : IV = 1 : 1.7 : 5.4 : 19.7. Ant. I and II with 7 and 15 setae, respectively (Fig. 5A). Ant. Ill (Fig. 5A) provided with only thin long setae and with an apical sensory organ consisting of two sensory rods protected by a cuticular pocket (Figs 4A-B, 5A, 6B). Ant. IV formed by 14 subarticles (Fig. 5B); the first one has nine setae; the second one three basal setae and an apical whorl of nine

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DISPARRHOPAUTES TERGESTINUS: A NEW COLLEMBOLAN SPECIES

er two (I: II: 111= 1 : 0.85 : 1.08) without tenent hairs (Fig. 9C); claw elongated with a non adherent thin dorsal tunica (Figs 4C-D, 6D), with neither inner nor outer teeth and with no small setae. The third claw is more slender than the other two. The empodium of all claws with evident lamellae (Fig. 4C-D) without teeth and terminal filament. Chaetotaxy of dens as in Figure 9D. On the ventral (posterior) side 3, 2, 2, 1...1 setae. Dens : muero = 2.73- Muero : anal appendage = 2.15. Muero with the inner and outer edges serrated. Setae missing. Retinaculum with three teeth and two setae on the central body (Fig. 6F). Ventral tube provided with filament showing papillae on the surface (Fig. 6C).


Justification

Fig. 3 - Scanning electron microscopy of the head and the ocular plate of D. tergestinus n. sp (A, C) and D. patrizii (B, D). Note the lack of ocelli in the ocular plate of D. tergestinus, while D. patrizii shows eight ocelli.

setae; the third one has both a basal and an apical whorl with six and nine setae, respectively. Also the fourth sub-article has two whorls, the basal with four setae and the apical with nine setae. Each sub-article from the fifth to the thirteenth has a whorl usually containing 10 setae, two of them being represented by thin sensilla (arrows in Fig. 5B). The last sub-article carries several setae, short sensilla and an apical vesicle (Figs 5B, 6A). Head without eyes (Fig. 3A, C). Head chaetotaxy as in Figure 7B with a single setae in the frontal area, ventral side with setae only in the distal region, without spiny setae (Fig. 7C). Labrum as in Figure 6E. Chaetotaxy of the great abdomen as in Figure 7D, provided with three trichobotria A, B, C, arranged according to the inverted pattern (Richards, 1968) without neosminthuroid setae. Chaetotaxy of the small abdomen of the female and male as in Figure 8A and 8B, respectively. Trichobotrium D present. Circumanal setae long; appendices anales long, thick, setalike. The male has about 20 short setae around the genital opening (Fig. 8C). Trochanter of the third leg with 5 setae, a spinelike seta and an oval organ (Fig. 9A). Femur with 16 setae as in Figure 9B; third tibiotarsus longer than the oth-

The new species belongs to Sminthuridae because of the presence of one pair of bothriotrichia D, the presence of normal setae and the lack of spines on the head and the body and of neosminthuroid setae. It lacks postantennal seta and shows a trochanter III posteriorly with one spine-like seta. Dens anteriorly with nine setae, muero with small teeth and without seta. The cuticle structure consists of primary granules on the head and appendages and secondary granules on the large abdomen. However, D. tergestinus n. sp. exhibits some important and peculiar characters. Contrary to D. patrizii, it lacks eye-patches and has a slender claw without inner teeth but provided with a tapering tunica not adherent to the claw; moreover, its antennae are longer than in D. patrizii; Ant. IV shows 14 sub-articles. These characters indicate that the new species is a specialized troglobiont; moreover, the presence of an elongated and narrow claw also suggests that it is well adapted to wet surfaces as it occurs on stalagmites. The genus Disparrhopalites now includes two species: Disparrhopalites patrizii - troglophilic, with 8 + 8 eyes, Ant IV with 12 sub-articles and Illrd claw with teeth and tunica. Disparrhopalites tergestinus - troglobiont, without eyes, Ant. IV with 14 sub-articles and Illrd claw elongated without teeth, and with slender tunica.

CONCLUSIVE REMARKS The Partigiani cave is located in the famous karstic area of the Trieste Province, NE Italy; this area is characterized by fossiliferous limestone dating to the Cretaceus period, from Lower Turonian (about 91 Mya) to Upper Senonian (Cucchi et al, 1989). The area was exposed to strong karstic phenomena that led to the formation of many wide and deep dolines, furrowed fields, many hollows and caves with large and deep shafts. The Partigiani cave, where the new species was found, is a small cave that consists of some rooms connected to each other. It opens to the outside through a small hidden opening near the road connecting Basovizza to Gropada and rather close to the famous Pluton

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Fig. 4 - Scanning electron microscopy of D. tergestinus n. sp. (A-D) and D. patrizii (E-F). The sensorial organ of Ant. Ill of D. tergestinus is formed by two sensorial rods inside a cuticular pocket (A, B). The claw of D. tergestinus n. sp. does not show any inner teeth (C, D), while that of D. patrizii (E, F) has numerous inner teeth.


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mites and on the surface of a small dripping basin. Together with them, other smaller but similar Collembola (probably young specimens of the new taxon) were observed, but not collected. In the dripping basin, together with the specimens of D. tergestinus n. sp., there was also a Palpigradi species (Eukoenenia sp.) that was probably plundering them. The air humidity of the cave is close to saturation almost throughout the year, but the new species appears only during the wettest period, when the surface of many stalagmites is covered by a film of water. During the dry months, unidentified but clearly different Collembola are abundant; they were not observed when the new species was collected. Christiansen (1961, 1965), Deharveng & Leclerc (1989), Vannier & Thibaud (1984) and Thibaud (1994) stated that the troglobitic Collembola show many morphological and physiological modifications compared with the epigeic forms. These modifications regard the total absence of pigmentation and eyes, and the elongation of the appendages and claws, while the body size appears larger than the most closely related epigeic species; the new species shows all these troglobitic features. Disparrhopalites patrizii, the other species of the genus, has frequently been found inside caves, but it has also been collected in open habitats and in caves whose large openings allow the light to reach the bottom; this species can also be found in artificial refuges such as Fig. 5 - Disparrhopalites tergestinus n. sp. A, antennal articles I, II and III. B, ant. IV showing 14 sub-articles. Arrows indicate the two sensorial setae usually present in each whorl.

abyss (no 23 V.G.), well known by local speleologists because of its vertical shaft 115 meters deep (Fig. 1). By means of a narrow passage, the cave opens into a first room whose ceiling is reached by the roots of some trees; the exchange of air appears quite limited, therefore the cave has remarkable environmental stability, with only small variations during the year, practically absent in the innermost part of the cave. The cave has already been studied for its fauna (Gasparo, 1995) and several troglobitic species were found. Among them the pseudoscorpion Troglocbthonius doratodactylus Helversen, the isopod crustacean Titanethes dahli Verhoeff and the land gastropod Zospeum spelaeum Rossmaessler. In addition, the cave also hosts many troglophilic species such as the coleopterans Laemostenus (Antisphodrus) cavicola cavicola (Schaum) (Carabidae), the orthopteran Troglophilus neglectus Krauss, the spider Meta menardi (Latreille) and some diplopods of the genus Brachydesmus probably belonging to the species B. subterraneus Heller. The 10 Mm coleopteran Leptinus testaceus (Mueller) (Leptnidae) is associated with the presence of small mammals as Myo- Fig. 6 - Disparrhopalites tergestinus n. sp. A, apical sub-article of xus glis (Linnaeus) (Myoxidae): both were observed in the Ant. IV. B, sensorial organ of Ant. III. C, ventral tube showing two filaments covered by numerous papillae. D, shape of the the first room of the cave. The four individuals of D. the third claw. E, shape of the labrum. F, retinacle with three teeth tergestinus n. sp. were observed at the top of the stalag- and two setae on the corpus.

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100 (im

Fig. 7 - Disparrhopalites tergestinus n. sp. A, habitus of the species. B, dorsal and C, ventral chaetotaxy of the head. D, chaetotaxy of the large abdomen showing the three trichobotria A, B and C; D is the trichobotrium on the small abdomen.

catacombs (Christian, 1998); Dallai (1973) found some individuals in an artificial repair for boats along the coast of the island Salina of Eolean Archipelago, Sicily. These last findings seem to confirm that D. patrizii is a troglophilic species and this agrees with the assumption that troglophilic species usually have a wider geographic distribution than troglobitic ones (Christiansen, 1982; Stomp, 1986; da Gama, 1990. Delamare Deboutteville & Bassot (1957) suggested a very ancient distribution for D. patrizii, but Dallai (1970a), considering the life style of the species, its larger distribution and the lack of morphological modifications in the specimens from different Mediterranean populations, hypothesized a recent adaptation to cave environment, possibly from the Quaternary. The climate changes during the Quaternary could have forced many coUembolans to seek refuge in caves where the environmental conditions are more stable and suitable for their survival. The adaptation to the cave environment became a permanent condition for some species, like D. tergestinus n.sp., that developed the morphological and physiological properties to survive as troglobitic species. The different level of adaptation between the two species could simply be a consequence of a different environmental selection or reveals a more ancient colonization of caves by the new taxon.

Fig. 8 - Disparrhopalites tergestinus n. sp. A, lateral view of the small abdomen of a female. B, frontal view of the small abdomen of a male. C, higher magnification of the male genital opening.

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Fig. 9 - Disparrhopalites tergestinus n. sp. A, trochanteral organ. B, chaetotaxy of the femur. C., chaetotaxy of the tibiotarsus III. D, chaetotaxy of the dens and shape of the muero.

Dallai R., 1970b - Ricerche sui Collemboli. XIV. Le Alpi Apuane. Lav. Soc. Ital. Biogeogr., 1: 433-482. Dallai R., 1973 - Ricerche sui Collemboli. XVII. Le isole Eolie. Lav. Soc. Ital. Biogeogr., 3: 481-590. Deharveng L., Leclerc P., 1989 - Recherches sur les faunes cavernicoles d'Asie du sud-est. Mém. Biospéol., 16: 91-110. Delamare Deboutteville C., Bassot J. M., 1957 - Collemboles symphypléones de Madeire et remarques biogéographiques. Vie Milieu, 8: 76-86. da Gama M., 1988 - Colêmbolos das Canarias (Insectos, Apterigotas). In: Actas III Congr. Ibér. Entomol., pp. 73-79. da Gama M., 1991 - Collemboles (Insectes, Aptérygotes) cavernicoles de la France et de la Suisse. Mém. Biospéol., 18: 171-183. Gasparo F., 1995 - La fauna delle grotte e delle acque carsiche sotterranee della Venezia Giulia, stato delle ricerche e checklist delle specie cavernicole. Atti Mem. Comm. Grotte "E. Boegan", 32: 17-42. Richards W. R., 1968 - Generic classification, evolution and bio-

geography of the Sminthuridae of the world (Collembola). Mem. Entomol. Soc. Canada, 53: 1-54. Schleuter M., 1985 - Zur Kenntnis der Collembolenfauna des Naturparkes Kottenforst-Ville: das Artenspektrum. Decheniana 138: 149-156. Stach J., 1956 - The apterygotan fauna of Poland in relation to the world fauna of this group of Insects. Family: Sminthuridae. Polska Akad. Nauk Inst. Zool., Krakow. Stomp N., 1986 - Le groupe de Pseudosinella vandeli (Insectes, Collemboles) dans les Alpes Orientales. In: R. Dallai (ed.), Second international seminar on Apterygota. University of Siena, Siena, pp. 85-91. Thibaud J. M., 1994 - For a biological and ecological classification of cavernicolous Collembola (Hexapoda). Mém. Biospéol., 21: 147-149. Vannier G., Thibaud J. M., 1984 - Conséquences de la vie cavernicole sur l'écophysiologie et la biologie de l'insecte Collembole Tomocerus catalanus Denis. Mém. Biospéol., 11: 221-231.