Louis Deharveng

Sopark Jantarit

Anne Bedos

Revisiting Lepidonella Yosii (Collembola: Paronellidae): character overview, checklist of world species and reassessment of Pseudoparonella doveri Carpenter

Keywords: Taxonomy, checklist, chaetal types, pseudopores, cave fauna

 [START_REF] Carpenter | XIX. Fauna of the Batu caves, Selangor[END_REF]described a remarkable new species of Paronellidae from the Batu Caves in Selangor (Malaysia), Pseudoparonella doveri, characterized in its original description by elongate appendages and claws, and lacking eyes (in fact, it has 3 + 3 small eyes, see below). It was at that time the most troglomorphic species of springtails known from Southeast Asia. He assigned it to the genus Pseudoparonella [START_REF] Handschin | Beiträge zur Collembolenfauna der Sundainseln[END_REF] on the basis of its short bidentate mucro. [START_REF] Yoshii | Paronellid Collembola of Sabah[END_REF][START_REF] Yoshii | Studies on Paronellid Collembola of East Asia[END_REF] did not accept this assignment, and considered the species as incertae sedis. [START_REF] Deharveng | Cave Collembola of South-East Asia[END_REF][START_REF] Deharveng | A new troglomorphic Collembola from Thailand: Troglopedetes fredstonei, new species (Collembola: Paronellidae)[END_REF] placed the species in Troglopedetes Absolon, 1907 following the redefinition of the genus by [START_REF] Palacios-Vargas | Taxonomia y Biogeographia de Troglopedetes (Collembola: Paronellidae) en America, con enfasis en las especies cavernicolas[END_REF]. [START_REF] Mitra | Chaetotaxy, Phylogeny and Biogeography of Paronellinae (Collembola: Entomobryidae). Records of the Zoological Survey of India[END_REF][START_REF] Mitra | Status of Microparonella Carpenter, 1916 (Collembola: Entomobryidae: Paronellinae)[END_REF] assigned P. doveri to the genus Microparonella [START_REF] Carpenter | The Apterygota of the Seychelles[END_REF], which he had redefined. Microparonella was shown to be a synonym of Troglopedetes to which its type species clearly belongs [START_REF] Deharveng | Lepidonella lecongkieti n.sp., premier Collembole cavernicole du Vietnam (Collembola, Paronellidae)[END_REF], but these authors did not reassess the taxonomic position of P. doveri in this context, which remains therefore uncertain.

With the current acceleration of researches on cave Collembola in Southeast Asia, it is increasingly important for biogeographic knowledge of the region to better place this species taxonomically, which remains the most troglomorphic of Southeast Asia Collembola. We had the opportunity to examine a few specimens of Pseudoparonella doveri from the Batu Caves given by Max Moseley and Liz Price, and a paralectotype of the species from the Natural History Museum (NHM) in London. Comparing our specimens to the original description of Paronella appendiculata (Schött, 1917), type species of the genus Pseudoparonella [START_REF] Ellis | An annotated list of the generic names of Collembola (Insecta) and their type species[END_REF], led us to the conclusion that "Pseudoparonella doveri" is not a member of the genus Pseudoparonella, in agreement with the authors cited above. The species is neither a Troglopedetes as currently understood, as its *Corresponding author. Email: dehar.louis@wanadoo.fr fourth antennal segment is not subdivided. Its inclusion in the genus Microparonella has to be abandoned as the genus was found to be based on a species of Troglopedetes [START_REF] Deharveng | Lepidonella lecongkieti n.sp., premier Collembole cavernicole du Vietnam (Collembola, Paronellidae)[END_REF]. The short mucro and presence of only two trichobothria on Abd. IV exclude doveri from Cyphoderopsis Carpenter, 1917or Trogolaphysa Mills, 1938 as well [START_REF] Thibaud | Collemboles (Insecta) de l'Équateur IV. Paronellidae avec révision de quatre genres[END_REF][START_REF] Jantarit | The genus Cyphoderopsis (Collembola: Paronellidae) in Thailand and a faunal transition at the Isthmus of Kra in Troglopedetinae[END_REF][START_REF] Soto-Adames | Comparative analysis of the dorsal chaetotaxy of Troglopedetes, Trogolaphysa and Campylothorax supports the synonymization of tribes Paronellini and Troglopedetini (Collembola: Paronellidae)[END_REF]). Conversely, it matches the most recent definition of the genus Lepidonella [START_REF] Yosii | On some Collembola of Afghanistan, India and Ceylon, collected by the Kuphe-expedition[END_REF] [START_REF] Deharveng | Lepidonella lecongkieti n.sp., premier Collembole cavernicole du Vietnam (Collembola, Paronellidae)[END_REF][START_REF] Soto-Adames | Dorsal chaetotaxy of neotropical species supports a basal position for the genus Lepidonella among scaled Paronellidae (Collembola, Entomobryoidea)[END_REF][START_REF] Bellini | The Neotropical genera of Paronellinae (Collembola, Entomobryoidea, Paronellidae) with description of two new species and redescription of Campylothorax mitrai[END_REF], except for its reduced eye number, a well-known adaptive character among cave-obligate species.

In this paper, several new or rarely used taxonomical characters are introduced for the taxonomy of Lepidonella and Paronellidae. The genus Lepidonella is rediagnosed, and a list of its species is given. Pseudoparonella doveri is transferred to the genus Lepidonella, and redescribed in detail. Its affinities with L. lecongkieti [START_REF] Deharveng | Lepidonella lecongkieti n.sp., premier Collembole cavernicole du Vietnam (Collembola, Paronellidae)[END_REF] from southern Vietnam are emphasized.

Material and methods

The examined specimens include a slide of a paralectotype borrowed from the London NHM and six specimens collected in the Batu Caves by Max Moseley and Liz Price. These last specimens in alcohol were examined and photographed under a Leica (Wetzlar, Germany) MZ16 stereomicroscope, then mounted on slides in Marc Andre II after clearing in lactic acid. Morphological characters were observed and drawn using a Leica DMLB microscope with Differential Interference Contrast (DIC). Photographs were taken with a Jenoptik (Jena, Germany) ProgRes C10+ camera mounted on the microscope. Figures including photos were improved with Photoshop Elements v9 (Adobe Inc., San José, USA).

Abbreviations used in the descriptions

Abd. = abdominal segment; Ant. = antennal segment; mac = macrochaeta; mes = mesochaeta; mic = microchaeta; ms or s-microchaeta = microsensillum auct.; omma = ommatidium, -ia; psp = pseudopore; Scxa = furcal anterior subcoxa; sens or s-chaeta = sensillum or sensory seta auct.; tric = trichobothrium; Th. = thoracic segment; Tita = tibiotarsus.

Nomenclature

We follow [START_REF] Yoshii | Lepidocyrtid Collembola of Sabah[END_REF] for the nomenclature of various morphological structures, [START_REF] Zhang | Cave-dwelling Coecobrya from southern China with a survey of clypeal chaetae in Entomobryoidea (Collembola)[END_REF] for clypeal chaetotaxy and [START_REF] Fjellberg | The Labial Palp in Collembola[END_REF] for labial palp. We use the notation of [START_REF] Gisin | Espèces nouvelles et lignées évolutives de Pseudosinella endogés[END_REF] for labium basis, with upper case letter for ciliated chaetae, lower case letter for smooth chaetae; we use brackets for mic.

Regarding chaetal categories, the term S-chaeta [START_REF] Deharveng | Morphologie évolutive des Collemboles Neanurinae, en particulier de la lignée néanurienne[END_REF]) is preferred to the term sensillum, considering that a chaeta is best defined as the external part of a sensillum, which is a composite organ. The term chaeta is used instead of seta for coherence with the widely used term "chaetotaxy", and macrochaeta, microchaeta instead of macroseta, microseta for etymological coherence (chaeta is a Greek root, like macro and micro, while seta is a Latin root). For dorsal chaetotaxy, we rely (but see below) on the nomenclatures of [START_REF] Szeptycki | Chaetotaxy of the Entomobryidae and its phylogenetical significance[END_REF], [START_REF] Mitra | Chaetotaxy, Phylogeny and Biogeography of Paronellinae (Collembola: Entomobryidae). Records of the Zoological Survey of India[END_REF][START_REF] Soto-Adames | Two new species and descriptive notes for five Pseudosinella species (Hexapoda: Collembola: Entomobryidae) from West Virginian (USA) Caves[END_REF][START_REF] Soto-Adames | Dorsal chaetotaxy of neotropical species supports a basal position for the genus Lepidonella among scaled Paronellidae (Collembola, Entomobryoidea)[END_REF]. We used most characters listed in [START_REF] Soto-Adames | Comparative analysis of the dorsal chaetotaxy of Troglopedetes, Trogolaphysa and Campylothorax supports the synonymization of tribes Paronellini and Troglopedetini (Collembola: Paronellidae)[END_REF], with some additional ones. The formula of dorsal macrochaetae and trichobothria are given by half-tergite from head to Abd. IV; mac along peri-antennal area, anterior rows of mac on Th. II, lateral mac of Abd. IV and mac of Abd. V-VI are not included in the formula. S-chaetae formulae are given separately for ms and for s, by half-tergite from Th. II to Abd. V.

Most chaetae had fallen during clearing and mounting processes. Sockets of scales can often be distinguished from those of chaetae on slide specimens of Entomobryoidea, but this was not that easy on the few available specimens of L. doveri, and we were not able to reconstruct reliably the dorsal chaetotaxic pattern of mesochaetae as thoroughly analyzed by Soto-Adames & [START_REF] Soto-Adames | Dorsal chaetotaxy of neotropical species supports a basal position for the genus Lepidonella among scaled Paronellidae (Collembola, Entomobryoidea)[END_REF].

Results

Morphological characters

Below we list and comment on some important morphological characters for the taxonomy of Lepidonella. Several are newly introduced characters, and a few are overlooked features drawn from various papers of Yoshii (mostly, 1981Yoshii (mostly, , 1982)).

Pseudopores

Figure 2. Pseudopores were first discovered by [START_REF] Gisin | Collemboles d'Europe[END_REF] on tergites and coxae I and II of Entomobryidae, then distally on manubrium [START_REF] Gisin | Collemboles d'Europe. VI[END_REF]. Later on, they were observed in other locations in Isotomidae [START_REF] Deharveng | Note sur un type d'organites tégumentaires originaux rencontré chez les Isotomidae (Collembola)[END_REF], Poduromorpha [START_REF] Deharveng | Morphologie évolutive des Collemboles Neanurinae, en particulier de la lignée néanurienne[END_REF], Entomobryidae [START_REF] Lukic | The puzzling distribution of Heteromurus (Verhoeffiella) absoloni Kseneman, 1938 (Collembola: Entomobryidae: Heteromurinae) resolved: detailed redescription of the nominal species and description of a new species from Catalonia (Spain)[END_REF] and Tomoceridae [START_REF] Yu | A peculiar cave species of Tomocerus (Collembola, Tomoceridae, Tomocerinae) from Vietnam, with a discussion of the postantennal organ and prelabral chaetae in Tomocerinae[END_REF]. In fact, they are also present in other locations not mentioned in the literature, as listed below for Lepidonella doveri:

(1) Apically on antennal segments II and III, probably firstly mentioned on Ant. III of Paronella vivieni [START_REF] Barra | Collemboles du Gabon[END_REF][START_REF] Barra | Collemboles du Gabon[END_REF]. [START_REF] Christiansen | The cave collembola of Mexico[END_REF] call them lenticular organs.

Bellini & Cipola (2017) mention a "discrete shallow subapical organite on Ant. IV" that we have not observed in other Entomobryoidea so far, which is perhaps a pseudopore. (2) Externally to antennal basis on head in the periantennal area (initially mentioned in Neanuridae, [START_REF] Deharveng | Morphologie évolutive des Collemboles Neanurinae, en particulier de la lignée néanurienne[END_REF]).

(3) Externally on coxae I, II and III [initially mentioned by [START_REF] Hüther | Beiträge zur Kenntnis der Collembolenfauna des Sudans[END_REF] as "chitinringe", and by [START_REF] Gisin | Collemboles d'Europe[END_REF]]. (4) On manubrium between its basis and the genital plate, probably those observed in Coecobrya by [START_REF] Zhang | Three new species of Coecobrya (Collembola: Entomobryidae) from southern and northwest China[END_REF] and [START_REF] Zhang | Species delimitation in the morphologically conserved Coecobrya (Collembola: Entomobryidae): A case study integrating morphology and molecular traits to advance current taxonomy[END_REF], as a group of 2-3 in L. doveri.

(5) Dorsodistally on manubrium (initially mentioned in [START_REF] Gisin | Collemboles d'Europe. VI[END_REF]. ( 6) Basally on dentes (new location, observed in many Entomobryidae). ( 7) Laterally on Abd IV along anterior subcoxa margin, already mentioned in [START_REF] Mateos | Towards understanding Lepidocyrtus Bourlet, 1839 (Collembola, Entomobryidae) I: diagnosis of the subgenus Setogaster, new records and redescriptions of species[END_REF] in several Lepidocyrtinae. (8) Dorsally, 1 + 1 on each tergite from Th. II to Abd.

IV (present in all Entomobryoidea, all Tomoceridae, and probably Isotomidae).

The line of pseudopores located behind the posterior line of chaetae on Abd. IV tergite in Troglopedetes Absolon, 1907[START_REF] Deharveng | A new troglomorphic Collembola from Thailand: Troglopedetes fredstonei, new species (Collembola: Paronellidae)[END_REF], Cyphoderopsis Carpenter, 1917[START_REF] Jantarit | The genus Cyphoderopsis (Collembola: Paronellidae) in Thailand and a faunal transition at the Isthmus of Kra in Troglopedetinae[END_REF]), Trogolaphysa Mills, 1938[START_REF] Soto-Adames | The dorsal chaetotaxy of Trogolaphysa (Collembola, Paronellidae), with descriptions of two new species from caves in Belize[END_REF] and Cyphoderus Nicolet, 1842 [START_REF] Jantarit | Cyphoderus (Cyphoderidae) as a major component of collembolan cave fauna in Thailand, with description of two new species[END_REF] has not been observed in L. doveri and in L. lecongkieti and is not reported in other species of the genus [START_REF] Zeppelini | Chaetotaxy of Neotropical Cyphoderus caetetus sp. nov. with comments on the taxonomic position of Cyphoderinae within Paronellidae (Collembola, Entomobryoidea)[END_REF]. The presence of these pseudopores is a strong putative synapomorphy of (Troglopedetinae + Cyphoderidae), that separates them from other Paronellidae and Entomobryidae.

In five other locations where pseudopores may be present in some Entomobryoidea, they were not detected in Lepidonella, due to unfavorable preparations or real absence: (i) anterointernally on the peri-antennal area of head (figured in Jantarit et al. 2014, p. 7); (ii) apically on Ant. I [START_REF] Lukic | The puzzling distribution of Heteromurus (Verhoeffiella) absoloni Kseneman, 1938 (Collembola: Entomobryidae: Heteromurinae) resolved: detailed redescription of the nominal species and description of a new species from Catalonia (Spain)[END_REF]; (iii) on thoracic sternites (observed in Lepidocyrtus sp., unpublished); (iv) between tenaculum and ventral tube (observed in Lepidocyrtus sp. and Cyphoderus sp., unpublished); and (v) on the central part of Abd. IV tergite in Lepidocyrtus curvicollis Bourlet, 1839, as figured by [START_REF] Mateos | The European Lepidocyrtus Bourlet, 1839 (Collembola: Entomobryidae)[END_REF]. A large number of pseudopores are at least present on the manubrium of Tomoceridae [START_REF] Yu | A peculiar cave species of Tomocerus (Collembola, Tomoceridae, Tomocerinae) from Vietnam, with a discussion of the postantennal organ and prelabral chaetae in Tomocerinae[END_REF], that have never been observed in Lepidonella, nor in other Entomobryomorpha. Evidence is therefore increasing that these tiny pseudopores have diversified distribution patterns that provide (or may provide) powerful characters at all taxonomic levels.

Eyes arrangement. When the complete set of eyes (8 + 8) is present, they may be arranged as two main different patterns in Entomobryoidea: either in two parallel lines with omma G anterior to or at level of F (most atmobiotic Paronellidae s. l.), or in a compact group, with G posterior to F. This last pattern is the only one reported in Lepidonella and known in soil Paronellidae (Paronella spp., Trogolaphysa spp.).

Ventral complex of labrum. Figure 3D. The presence of two slightly asymmetrical multi-toothed combs as described here for L. doveri has been already reported in a number of Entomobryoidea, for instance by [START_REF] Rusek | Zweiter Beitrag zur Kenntnis der Collembola (Apterygota) Chinas[END_REF] in Willowsia nigromaculata (Lubbock, 1873) and in Cyphoderus asiaticus Yosii, 1959;[START_REF] Palacios-Vargas | Taxonomia y Biogeographia de Troglopedetes (Collembola: Paronellidae) en America, con enfasis en las especies cavernicolas[END_REF] in Trogolaphysa spp.; by [START_REF] Yoshii | Lepidocyrtid Collembola of Sabah[END_REF] in Acrocyrtus heterolepis (Yosii, 1959); [START_REF] Gruia | Collemboles Arthropléones de Cuba récoltés par les expéditions cubano-roumaines en 1969-1973[END_REF] in Pseudosinella (Plumosinella) kenchristianseni [START_REF] Gruia | Collemboles Arthropléones de Cuba récoltés par les expéditions cubano-roumaines en 1969-1973[END_REF][START_REF] Deharveng | Collemboles cavernicoles. V/Une nouvelle Pseudosinella du gouffre de la Pierre St Martin (France: Pyrénées-Atlantiques)[END_REF] in Pseudosinella cabidochei [START_REF] Deharveng | Collemboles cavernicoles. V/Une nouvelle Pseudosinella du gouffre de la Pierre St Martin (France: Pyrénées-Atlantiques)[END_REF][START_REF] Jantarit | Cyphoderus (Cyphoderidae) as a major component of collembolan cave fauna in Thailand, with description of two new species[END_REF] in Cyphoderus spp. These combs are also present in the Hypogastruridae Triacanthella madiba [START_REF] Janion | A new species and first record of the genus Triacanthella Schäffer, 1897 (Collembola, Poduromorpha, Hypogastruridae) for Africa[END_REF][START_REF] Janion | A new species and first record of the genus Triacanthella Schäffer, 1897 (Collembola, Poduromorpha, Hypogastruridae) for Africa[END_REF] and in several Isotomidae [START_REF] Fjellberg | Revision of the European species of the Isotoma olivacea group (Collembola, Isotomidae)[END_REF]. They are associated in many (all?) species with a pair of thin, sinuous, unequal tubules (that we propose to name tongues), located side by side, which are however not mentioned in most cited cases. They show low inter-specific and inter-generic variations among Entomobryoidea. Conversely, they are more or less reduced in Isotomidae where a distal brush, never found in Entomobryoidea, tends to develop between the combs and the distal edge of the labrum [START_REF] Fjellberg | Revision of the European species of the Isotoma olivacea group (Collembola, Isotomidae)[END_REF][START_REF] Potapov | Synopses on Palaearctic Collembola. Volume 3. Isotomidae[END_REF]; in the few studied Tomoceridae where this labral part is described, only the distal brush is clearly seen [START_REF] Lukic | A new relictual and highly troglomorphic species of Tomoceridae (Collembola) from a deep Croatian cave[END_REF].

Labium basis. Figures 3A,4A,B. The succession of ciliated/smooth chaetae along the proximal edge of labium basis is widely used to separate species of Entomobryoidea following [START_REF] Gisin | Espèces nouvelles et lignées évolutives de Pseudosinella endogés[END_REF]. In L. doveri, on 4 specimens examined, chaeta M2 was asymmetrical in one specimen (ciliated/smooth), and some variability has often been reported in chaetae M of various species in the literature. This character should therefore be used cautiously as a diagnostic character. Chaeta L2 is similar in size to L1 in L. doveri, as well as in most species of Entomobryoidea, whereas it is reduced to a minute mic in Troglopedetes [START_REF] Deharveng | A new troglomorphic Collembola from Thailand: Troglopedetes fredstonei, new species (Collembola: Paronellidae)[END_REF], Cyphoderopsis [START_REF] Jantarit | The genus Cyphoderopsis (Collembola: Paronellidae) in Thailand and a faunal transition at the Isthmus of Kra in Troglopedetinae[END_REF]Campylothorax (Mari Mutt, 1987). Labium basis chaetotaxy may therefore provide interesting generic or supra-specific characters in Paronellidae s. l., though information is still lacking for a large number of species.

Maxillary outer lobe. Figure 3C. The number of sublobal hairs is known in only 5 species of Lepidonella. In the two cave species L. doveri and L. lecongkieti, there are 4 hairs, of which one is much smaller than the others. In L. nigrofasciata (Handschin, 1928) and L. oudemansi [START_REF] Yoshii | Studies on Paronellid Collembola of East Asia[END_REF], there are 3 subequal hairs, but the fourth minute additional hair observed in the above species may have been overlooked. The American species L. zeppelinii Soto-Adames & Bellini, 2015 departs from the previous four Asian species in having only two sublobal hairs (Soto-Adames & Bellini 2015), a number frequent among other Paronellidae s. l. Conversely, the presence of 4 sublobal hairs including one much smaller than others is retrieved in Entomobryidae like Entomobrya nivalis (Linnaeus, 1758) after [START_REF] Fjellberg | The Collembola of Fennoscandia and Denmark. Part II: Entomobryomorpha and Symphypleona[END_REF], Pseudosinella argentea Folsom, 1902 after Soto-Adames (2010), or in Verhoeffiella absoloni Kseneman, 1938after Lukic et al. (2015). [START_REF] Soto-Adames | Dorsal chaetotaxy of neotropical species supports a basal position for the genus Lepidonella among scaled Paronellidae (Collembola, Entomobryoidea)[END_REF] for the South American species L. zeppelinii but is unknown in other species of the genus. Only macrochaetotaxy and S-chaetotaxy are described here for L. doveri, as the state of available slides were too poor to allow a detailed description of their chaetotaxy.

Complete dorsal chaetotaxy. Dorsal chaetotaxy has been analyzed in detail by

Femoral organ. Femoral organ is a group of modified chaetae, thick, subcylindrical and rather short, located on the femur of leg III. It is the main diagnostic character which separates Lepidonella (without femoral organ) from Bromacanthus Schött, 1925 (with femoral organ) according to [START_REF] Mitra | On Lepidonella Yosii, 1960 and Bromacanthus Schött, 1925 complex (Collembola: Entomobryidae: Paronellinae)[END_REF]. In L. nigrofasciata, [START_REF] Yoshii | Studies on Paronellid Collembola of East Asia[END_REF] figures a rather dense group of chaetae in the place of the femoral organ, that he did not consider to represent the femoral organ itself, as chaetae were not morphologically modified. Femoral organ is absent in L. doveri and in L. lecongkieti (unpublished data), supporting their inclusion in Lepidonella rather than in Bromacanthus.

Filamentous organ. The presence of filamentous organs laterally on Abd. III has been observed in several poorly delimited genera of Lepidocyrtinae (Yoshii 1982, p. 34;[START_REF] Yoshii | Notes on the Collembolan fauna of Indonesia and its vicinities. I. Miscellaneous notes, with special references to Seirini and Lepidocyrtini[END_REF][START_REF] Mitra | Fixation of the concept of Paronella Schötte, 1893 (Collembola: Entomobryidae)[END_REF][START_REF] Mateos | Towards understanding Lepidocyrtus Bourlet, 1839 (Collembola, Entomobryidae) I: diagnosis of the subgenus Setogaster, new records and redescriptions of species[END_REF]. Surprisingly, these organs are also present in L. doveri and in L. lecongkieti, at the same place as in the species of "Ascocyrtus (Cinctocyrtus)" [START_REF] Yoshii | Notes on the Collembolan fauna of Indonesia and its vicinities. I. Miscellaneous notes, with special references to Seirini and Lepidocyrtini[END_REF]cited in Yoshii & Suhardjono (1992). In L. lecongkieti, they are constituted by a tuft of very long chaetae (up to 450 μm) inserted laterally on Abd. III. In L. doveri, only the sockets of these chaetae were visible. In both species, the filamentous organ was seen in a few specimens only.

Coxal complex. Figure 8G,H,I. The coxal pattern of mac and psp seems to be rather stable across many Entomobryoidea, and that of Lepidonella doveri is quite similar to those described for Heteromurtrella puertoricensis [START_REF]Heteromurtrella, a new tropical subgenus of Heteromurus with descriptions of two new species (Insecta: Collembola: Entomobryidae)[END_REF][START_REF]Heteromurtrella, a new tropical subgenus of Heteromurus with descriptions of two new species (Insecta: Collembola: Entomobryidae)[END_REF], Lepidocyrtus flexicollis Gisin, 1965[START_REF] Mateos | The European Lepidocyrtus Bourlet, 1839 (Collembola: Entomobryidae)[END_REF], and Verhoeffiella absoloni Kseneman, 1938 [START_REF] Lukic | The puzzling distribution of Heteromurus (Verhoeffiella) absoloni Kseneman, 1938 (Collembola: Entomobryidae: Heteromurinae) resolved: detailed redescription of the nominal species and description of a new species from Catalonia (Spain)[END_REF] for instance.

Claw complex. Figure 8D,E,F. In most Lepidonella, the unguiculus is clearly truncated, with four exceptions: L. doveri and L. lecongkieti, which are the only cave species in the genus, and L. subcarpenteri Denis, 1948 and L. zeppelinii. This last species is the only representative of the genus in America and departs by several characters from other Lepidonella. A re-examination of L. subcarpenteri is needed, in order to re-evaluate the taxonomic value and evolutionary significance of unguiculus morphology.

Ventral tube morphology. Ventral tube is long, slightly longer than head in L. lecongkieti, L. doveri and undescribed species of Southeast Asia, and relatively longer in cave than in soil species. Its apical part is a large exsertile vesicle sensu [START_REF] Willem | Recherches sur les Collemboles et les Thysanoures[END_REF] [= terminal tubule in [START_REF] Yoshii | Paronellid Collembola of Sabah[END_REF]], a structure often well developed in Paronellidae s. l. It is mentioned and figured under the name "protrusible sac" by [START_REF] Carpenter | The Apterygota of the Seychelles[END_REF] in the description of the genus Microparonella and of Lepidonella flava [START_REF] Carpenter | The Apterygota of the Seychelles[END_REF]. The protrusible structure is smooth, subdivided in two paired parts and oriented obliquely forward, towards the mouth, in the examined species of Lepidonella preserved in ethanol, with anterior diverticule reduced to a short swelling.

Multiplets on dens. Figure 10B,C. The arrangement of chaetae in multiplets is well known on Th. II of many Entomobryoidea [multiplets sensu [START_REF] Szeptycki | Chaetotaxy of the Entomobryidae and its phylogenetical significance[END_REF]]. In L. doveri and in L. lecongkieti scales are arranged in a similar way on dentes [START_REF] Deharveng | Lepidonella lecongkieti n.sp., premier Collembole cavernicole du Vietnam (Collembola, Paronellidae)[END_REF], as it is also observed on the dens of Cyphoderopsis spp. [START_REF] Jantarit | The genus Cyphoderopsis (Collembola: Paronellidae) in Thailand and a faunal transition at the Isthmus of Kra in Troglopedetinae[END_REF]. This pattern, though less obvious, also exists in some Lepidocyrtinae, like Pseudosinella (Plumosinella) kenchristianseni [START_REF] Gruia | Collemboles Arthropléones de Cuba récoltés par les expéditions cubano-roumaines en 1969-1973[END_REF].

Lateral organ of Abd. V-VI. The lateral organ of [START_REF] Yoshii | On some Collembola of New Caledonia, with notes on the "Colour Pattern Species[END_REF] is a conspicuous invaginate paired structure located on each side at the joint between Abd. V and Abd. VI of several Entomobryidae. It is absent in Lepidonella and has not been reported so far among Paronellidae s. l., but was observed in Callyntrura sp. (unpublished).

Taxonomy

Family Paronellidae Börner, 1906 (sensu [START_REF] Absolon | Troglopedetini. Vergleichende Studie über eine altertümliche höhlenbewohnende Kollembolengruppe aus den dinarischen Karstgebieten. Bericht über eine naturwissenschaftliche forschungsreise und biospeologische Erforschung der Insel Brac (Brazza) in Dalmatien[END_REF] Subfamily Bromacanthinae [START_REF] Mitra | Chaetotaxy, Phylogeny and Biogeography of Paronellinae (Collembola: Entomobryidae). Records of the Zoological Survey of India[END_REF](sensu Bromacanthini Mitra, 1993) Genus Lepidonella [START_REF] Yosii | On some Collembola of Afghanistan, India and Ceylon, collected by the Kuphe-expedition[END_REF] Type species: Lepidonella tokiokai Yosii, 1960 (Bougainville in Solomon Islands)

Diagnosis of the genus (*: character known in the type species) Similar to Lepidocyrtinae in body shape, at least for smallest species. Medium to large size (1-2.7 mm)*. Head perpendicular to body in specimens stored in alcohol*. Body curvature well marked at level of the middle or posterior part of Th. III, Th. II not protruded*. Dark pigment present as patches on the body (soil species)*, or body unpigmented or very lightly and uniformly pigmented (cave species). Flexed-cross mac absent. Scales present on the body*, present dorsally on Ant. I*, on Ant. I-II or absent, and present on furca at least ventrally. Scales finely and uniformly sculptured, hyaline to slightly beige, rounded to oval. Omma 3 + 3 to 8 + 8*. Extraocular structure sensu [START_REF] Mitra | An accessory ocular structure in two genera of Paronellinae (Collembola: Entomobryidae)[END_REF] absent. Labrum granulated area with anterior U-shape indentation*. Maxillary outer lobe with 3 or 4 sublobal hairs (2 in L. zeppelinii). Antennae shorter (surface species) or longer (cave species) than body; first segment shorter than head. Fourth antennal segment neither subdivided nor annulated, without apical bulb* (except in L. marimuti). Metafemoral organ absent*. Trochanteral organ well developed*. Posterior row of mes present on head*. Macrochaetotaxy very reduced on head and body*. Trichobothrial pattern as 1/0,0/0,2,3,2. Trichobothrial complexes well differentiated, with short and strongly enlarged mes. Filamentous organ present in L. doveri and L. lecongkieti, unknown in other species. Posterior line of pseudopores absent on Abd. IV. Lateral organ absent at the suture of Abd. V-VI. Manubrial spines absent. Two longitudinal rows of numerous spines on dentes (23-110)*. Distal lobe and enlarged distal chaetae on dens absent. Mucro plump, short, with 3-4 teeth*, apically inserted on dens.

Note

The type species of the genus, Lepidonella tokiokai, was described from Solomon Islands on a single specimen. It has never been revised since its original description, which is globally accurate. The state of many of the characters subsequently introduced, e.g. pseudopore distribution, is however unknown in this species as well as in most other species of Lepidonella. Compared to previous definitions of the genus [START_REF] Carpenter | The Apterygota of the Seychelles[END_REF][START_REF] Deharveng | Lepidonella lecongkieti n.sp., premier Collembole cavernicole du Vietnam (Collembola, Paronellidae)[END_REF][START_REF] Mitra | On Lepidonella Yosii, 1960 and Bromacanthus Schött, 1925 complex (Collembola: Entomobryidae: Paronellinae)[END_REF][START_REF] Soto-Adames | Dorsal chaetotaxy of neotropical species supports a basal position for the genus Lepidonella among scaled Paronellidae (Collembola, Entomobryoidea)[END_REF], the main changes in the current diagnosis concerns eye number that may be as few as 3 + 3, pigmentation which may vary from large dark patches to uniformly white, and antenna length which may vary from shorter to longer than body. Several characters newly introduced need to be checked on a larger number of species. Dorsal chaetotaxy of Lepidonella is "complete or almost complete" according to Soto-Adames & Bellini (2015, p. 31). However, this observation concerns only two species, of which one is incertae sedis (L. marimuti from Antillas), and the other one, L. zeppelinii from Brazil, is the only other species known from America which departs from its congeners by the reduced chaetotaxy of its maxillary outer lobe. No information on dorsal chaetotaxy is available for Lepidonella of Southeast Asia and Australasia.

List of species of Lepidonella in the current definition of the genus

Complete references for the current species of Lepidonella as well as those which have been previously considered as Lepidonella but are now placed in other genera are drawn from the unpublished lists of Bellinger et al. (1996Bellinger et al. ( -2017)). We selected here the most significant citations, i.e. new combinations, or additional distribution data. All citations from the works of [START_REF] Deharveng | Lepidonella lecongkieti n.sp., premier Collembole cavernicole du Vietnam (Collembola, Paronellidae)[END_REF], [START_REF] Mitra | On Lepidonella Yosii, 1960 and Bromacanthus Schött, 1925 complex (Collembola: Entomobryidae: Paronellinae)[END_REF] and Soto-Adames & [START_REF] Soto-Adames | Dorsal chaetotaxy of neotropical species supports a basal position for the genus Lepidonella among scaled Paronellidae (Collembola, Entomobryoidea)[END_REF] are included. Later references include citations giving additional locations or new combinations, and those in the recent review of the genus by Soto-Adames & [START_REF] Soto-Adames | Dorsal chaetotaxy of neotropical species supports a basal position for the genus Lepidonella among scaled Paronellidae (Collembola, Entomobryoidea)[END_REF]. Complete reports of species citations anterior to 2007 or 2015, depending on species, may be found on the web (Bellinger et al. 1996(Bellinger et al. -2017)).

Lepidonella annulicornis [START_REF] Oudemans | Apterygota des Indischen Archipels. In Weber's Ergeb[END_REF] The Oudemans paper is dated 1890-1891. We therefore quote it as 1891. The report from India by [START_REF] Denis | Collemboles de Costa Rica avec une contribution au species de l'ordre (Deuxième note)[END_REF] is probably a mistranslation of "Ost Indies" (nowadays Indonesia) from the title of the paper of [START_REF] Oudemans | Apterygota des Indischen Archipels. In Weber's Ergeb[END_REF].

The record of the species from "Am. insulae" (American insulae) by von Dalla Torre ( 1895) is a geographical misplacement, as the species has never been recorded in America.

The record of the species from India by [START_REF] Denis | Collemboles de Costa Rica avec une contribution au species de l'ordre (Deuxième note)[END_REF] is also a geographical misplacement: the author probably misunderstood the title of Oudemans (1891) paper as referring to India, having overlooked that the species was described from Java.

Lepidonella ceylonica (Yosii, 1966) Microparonella ceylonica This species was overlooked in the lists of [START_REF] Deharveng | Lepidonella lecongkieti n.sp., premier Collembole cavernicole du Vietnam (Collembola, Paronellidae)[END_REF] and Soto-Adames & [START_REF] Soto-Adames | Dorsal chaetotaxy of neotropical species supports a basal position for the genus Lepidonella among scaled Paronellidae (Collembola, Entomobryoidea)[END_REF].

Lepidonella subcarpenteri [START_REF] Denis | Collemboles d'Indochine. Récoltes de M.C. Dawydoff[END_REF] Paronella subcarpenteri The incorporation of the genus Lepidonella into Microparonella by [START_REF] Yoshii | Paronellid Collembola of Sabah[END_REF] implied the transfer of its type species, L. tokiokai, to Microparonella, though the species itself was not cited in the paper. and Pseudosinella, notoriously known to be phyletically heterogeneous). On these grounds, L. marimuti is considered here as incertae sedis. Description or redescription of more Southeast Asia species of the genus would be necessary to validate these conclusions.

Lepidonella zeppelinii

Species transferred to Trogolaphysa

Because of the presence of three trichobothria on Abd. IV, two species placed in Lepidonella by [START_REF] Deharveng | Lepidonella lecongkieti n.sp., premier Collembole cavernicole du Vietnam (Collembola, Paronellidae)[END_REF] were transferred to the genus Trogolaphysa by Soto-Adames & [START_REF] Soto-Adames | Dorsal chaetotaxy of neotropical species supports a basal position for the genus Lepidonella among scaled Paronellidae (Collembola, Entomobryoidea)[END_REF].

Trogolaphysa monomaculata [START_REF] Barra | Collemboles du Gabon[END_REF] Paronella Redescription of Lepidonella doveri [START_REF] Carpenter | XIX. Fauna of the Batu caves, Selangor[END_REF] Type locality. Malaysia: Selangor: Gombak: Batu Caves: Dark Cave (X = 101.684132°; Y = 3.237908°; alt = 85 m).

Type material. The species was described based on several specimens according to [START_REF] Carpenter | XIX. Fauna of the Batu caves, Selangor[END_REF]

Description.

Habitus. Figure 1A,C. Length: 2.3-2.7 mm. Color: pale yellow according to original description; specimens in ethanol whitish with very small, almost inconspicuous microdots of bluish grey pigment especially behind eye spots; 2 or 3 small black separated eye patches on each side with 3 omma (Figure 1D). Extraocular structure sensu [START_REF] Mitra | An accessory ocular structure in two genera of Paronellinae (Collembola: Entomobryidae)[END_REF] absent. Slender, with elongate legs, furca and antennae. In lateral view, head down perpendicularly to the body. Body strongly bent at the level of Th. II but not humped, straight backwards or slightly arched at the level of Th. III-Abd. II (Figure 1A,C). Th. II more than 1.5 as long as Th. III; Abd. IV about 4 times longer than Abd. III (Figure 1A).

Pseudopores. Figure 2. Pseudopores present as round flat disks, larger than mac sockets on various parts of the body: antennae, head, tergites, coxae, manubrium and dens. On antennae, psp located ventroapically between the tip of antennal segments and the chaetae of the apical row, or just below apical row of chaetae (absent or not detected on Ant. I, 1 on Ant. II and 2 on Ant. III). On head, 1 or 2 psp located externally on peri-antennal area, difficult to observe. On tergites, 1 + 1 psp close to the axis from Th. II to Abd. IV, and a file of 4 conspicuous psp on Abd. IV along the suture with the anterior furcal subcoxa. Coxae I, II and III with 2-(3), 3-(4) and 1-2 psp respectively, located close to longitudinal rows of chaetae described below. On manubrium, 3 dorsobasal psp close to genital plate and 2 + 2 apical ones. Dens dorsally with a line of 4 psp on the proximal third.

Chaetae. More detail later in the description of antennae, tergites, legs and furca. Ordinary chaetae differentiated into ciliated to serrated mac and mes, some enlarged and modified (in trichobothrial complexes), smooth mes (distally on Tita), smooth pointed mic (on Ant. I, on pretarsus and on anal valves), and weakly ciliated mic (on Tita and femora). Mac ill-differentiated from large mes on Abd. IV. S-chaetae of various morphologies. Dense cover of overlapping scales on head and body. Scales present on Ant. I-II, on manubrium and dentes, absent on legs and ventral tube. Scales roundish or oval, never pointed, of variable size, hyaline, adorned with many minute spicules in irregular lines. Two types of S-chaetae on tergites, like in other Entomobryoidea Trichobothrial pattern: 1/0,0/0,2,3,2 by half tergite.

Mouthparts. Figures 3,4. Clypeal area (Figure 3F) with 12 chaetae (and an extra-chaeta asymmetrically in the examined specimen), not matching exactly any of the patterns described in [START_REF] Zhang | Cave-dwelling Coecobrya from southern China with a survey of clypeal chaetae in Entomobryoidea (Collembola)[END_REF], but close to that of L. nigrofasciata as illustrated by Yoshii (1983: 9). Distal border of the apical non-granulated area of the labrum with a relatively narrow median intrusion into the granulated area dorsally (Figure 4C); apical edge not adorned with spines. Ventrodistal complex of labrum well differentiated, asymmetrical, with 2 distal combs of 11-12 and 8-9 teeth and an axial pair of sinuous tubules (Figure 3D). Labral formula 4/5,5,4; prelabral chaetae bent and ciliated, labial chaetae smooth and acuminate; labral chaetae of the distal row slightly shorter than those of the median and proximal row (30 vs 39 and 46 μm), their socket almost fused to the integument. Maxillary outer lobe with 1 papillate chaeta, 1 basal chaeta and 4 sublobal hairs of which the upper one is 4 times shorter than the others (Figure 3C).

Labium (Figures 3A,4A,B). Chaetae of labial basis as M1M2(R)EL1L2 or M1m2(R)EL1L2. Chaetae M1 and E subequal, ciliated; m2 or M2 as long as M1, smooth or ciliated (asymmetrically in one specimen); R reduced to a short and thick ciliated spine; L1 and L2 ciliated, other chaetae of the submentum smooth and acuminate, of similar size. Labial palp similar to that described by [START_REF] Fjellberg | The Labial Palp in Collembola[END_REF] for Troglopedetes sp., with strong papillate chaetae. Number of guards for each major papillate chaetae as: h (2), A (0), B (5), C (0), D (4) and E (4). Lateral process subcylindrical, not reaching the apex of papilla E. Five smooth proximal chaetae.

Mandible head strong, asymmetrical (left one with 4 teeth, right one with 5 teeth); molar plate with one strong pointed basal tooth, and 4-( 5) very strong inner distal teeth, identical in both mandibles.

Maxilla head (Figures 3B,4D,E,F) with a 3-toothed claw and several stout ciliated lamellae; lamella 2 large and broad, lamella 3 well developed; several other lamellae present, but homologies are uncertain; a thin elongate structure, arising from the basis of the maxilla head, reaches claw basis; close to it is a spiny structure 10 μm long (Figures 3B,4E,F).

Hypopharynx with many long, thin or thick pointed teeth.

Antennae. Figures 1A,5. Antennae longer than the body, about 3.7 times as long as cephalic diagonal. Lengths of antennal segments I to IV in μm: 284/568/ 625/852 (N = 1), similar to the values of the original drawing of [START_REF] Carpenter | XIX. Fauna of the Batu caves, Selangor[END_REF] (273/466/602/943). Antennal segments entire, not sub-segmented or annulated; in at least one specimen, antennae asymmetrical, one normal with 4 segments, the other one shortened with 3 segments (Figure 1A). A large cuticular notch devoid of chaetae ventrodistally on Ant. I and II, without (Ant. I) or with 1 pseudopore (Ant. II), not differentiated on Ant. III. Ant. III with at least one ventrodistal psp. Ant. IV. not subdivided, without apical bulb; dorsoexternal subapical organite relatively large (3.5 × 1 μm to 4 × 1.3 μm) swollen, oval, slightly enlarged medially, inserted dorsally at 35-45 μm from the apex (Figure 5G).

Antennal scales subequal, small (7-9 × 4-5 μm), oval, present dorsally on Ant. I (Figure 5B) and Ant. II, absent on Ant. III-IV.

Other antennal chaetae of various types (Figure 5A), arranged in a complex pattern rather similar to those described in Cyphoderopsis [START_REF] Jantarit | The genus Cyphoderopsis (Collembola: Paronellidae) in Thailand and a faunal transition at the Isthmus of Kra in Troglopedetinae[END_REF]) and in Cyphoderus [START_REF] Jantarit | Cyphoderus (Cyphoderidae) as a major component of collembolan cave fauna in Thailand, with description of two new species[END_REF]).

• Ciliated mac, very long (108-120 μm), flexuous, thinner than ciliated mes, tapering, relatively hyaline.

A few basally on Ant. II, absent on Ant. I, III and IV. IV), straight, almost perpendicular to the integument contrary to other chaetae, often curved at tip, smooth, thin, acuminate, more refractive than other chaetae. Present internodistally on 2/3 on Ant. III (Figure 5D) and all along Ant. IV, especially numerous in its distal part (Figure 5G); absent on Ant. I and Ant. II. • Swollen sens, rather short (7.5-11.5 μm), straight, smooth, ribbed, rounded apically, hyaline. Absent on Ant. I and not seen on Ant. II; two located externodistally, close together, in AIIIO of Ant. III (Figure 5F); absent on Ant. IV. • Short sens (6-14 μm), curved or straight, smooth, slender, thickened basally or subcylindrical, hyaline. Located ventrally on Ant. I and Ant. II (Figure 5C); present in distal 2/3 on Ant. III (Figure 5D,E) and on Ant. IV except in its distal part (Figure 5G).

• microsens (3-4 μm), smooth, thick, subcylindrical, hyaline. Located in AIIIO, internally to swollen s-chaetae (Figure 5F), absent elsewhere.

Head. Figures 1D, 3E, F. On head, mac limited to antennal lines (8-9 mac on each side); dorsal mes short (12-13 μm), subequal, indistinctly serrated, pointed, symmetrically arranged (Figure 1D). Pattern of mes incompletely visible on a single specimen; on anterior part of the head, mes arranged similarly to those of L. zeppelinii, but with additional mes near the antennal line, and fewer along the suture line; those of the posterior row slightly stronger. All postlabial chaetae distinctly to strongly ciliated, pointed rather than acuminate distally, scattered among a cover of large oval scales (45×40 to 60×50 μm) (Figure 3E). Dorsal scales smaller than ventral ones (20×14 to 28×22 μm).

Cephalic trichobothria short, close to and posteroexternal to ocular area.

Tergites. Figures 6, 7. Tergite chaetae of six types as usual in the group: ordinary chaetae of various size, ciliated or serrated, modified mes of trichobothrial complexes, short S-chaetae (S and ms), S-like chaetae, trichobothria and scales. Dense clothing of round to oval scales (15-40 μm long), mixed with sparse, short, slightly serrated, subequal mes (9-15 μm), symmetrically arranged. State of material examined did not allow to reconstruct accurately the detailed pattern of dorsal mes on tergites.

Trichobothrial complexes of Abd. II, III and IV well differentiated (Figure 6C,D,E), comprising of short, thick, often triangular modified mes, strongly ciliated unilaterally. Trichobothrial complex of Abd. II with 3 modified mes anterior to internal tric and, 2 modified mes anterior to external tric (possibly more). Trichobothrial complex of Abd. III with 3 modified mes anterior to internal tric and 7 close to lateral tric. Trichobothrial complex of Abd. IV with 4 modified chaetae associated to each of the two trichobothria.

S-chaetotaxy (Figure 6A-D) formulae from Th. II to Abd. V: ms as 1,0/1,0,1,0,0; S as 1,?1/0,1,1,n,?. On Abd. IV, a short posterolateral sens and numerous longer S-like or Schaetae. On Abd. V, S-chaetae not distinctly seen. On Abd. II-III and posteriorly on Abd. IV, S and ms are associated to mes in a tandem configuration (Figures 6A,C,D,7B).

Tergite chaetotaxy with few mac differentiated (Figure 6A). Th. II collar as a row of small mac directed forward, associated with irregularly arranged mes behind them (Figure 7A), and 2-3 irregular rows of shorter mac and mes anterolaterally. Anterolateral sensorial complex of Th. II present, comprising an opaque pointed ms posterior to and side by side with a slightly longer, thinner, hyaline sens. No posterolateral sens detected. No mac detected. Most mes fallen, their sockets small and difficult to differentiate from scale sockets in our specimens.

Th. III devoid of mac, with more than 14 + 14 mes (2 + 2 posterointernal between the axis and the pseudopores, 8 + 8 posterior, 4 + 4 anterior and several lateral); S and ms not seen.

Abd. I devoid of mac, with at least than 6 + 6 mes (of which 1 + 1 between the axis and the pseudopores), and 1 + 1 lateral ms.

Abd. II devoid of mac, with 2 + 2 trichobothria, about 8 + 8 non-modified mes, 5 + 5 modified mes and 1 + 1 S near internal trichobothria (Figure 6C).

Abd. III with 3 + 3 trichobothria, 3 + 3 lateral mac (am6, pm6 and p6), close and external to lateral trichobothria; more than 8 + 8 non-modified mes, 10 + 10 modified mes; 1 + 1 external ms and 1 + 1 internal S, both in tandem with mes (Figures 6D,7B). Presence of a filamentous organ on each side of Abd. III, constituted of about 30 thin and presumably very long mac, but all fallen on our slides.

Abd. IV with 2 + 2 posterointernal mac (B5 and B6), several (6-10) + (6-10) lateral mac ill-differentiated from large mes, more than 20 + 20 thin, slightly serrated S-like mes (18-25 μm long, some of them possibly smooth S), and 9 + 9 long (about 70 μm), thin, serrated mes in the posterior row; at least 1 + 1 s-chaetae posterolaterally. No pseudopore behind the posterior line of chaetae, 4 pseudopores along the suture Abd. IV -Scxa (Figure 6A). Chaetotaxy of Abd. V-VI not examined.

Legs. Figure 8. Legs long; Tita of leg III as long as head diagonal, Tita of legs I and II slightly shorter. Legs devoid of scales, covered with ordinary ciliated chaetae of various length, ranging from mic to mac. Coxa of leg I with 2 proximal psp and 6 chaetae posteriorly; coxa of leg II with 11-14 chaetae in anterior row, 9-11 chaetae in posterior row and 3-4 proximal psp in between; coxa of leg III with 4-5 + 8-12 chaetae in anterior row, and 2 proximal psp posteriorly (Figure 8G-I). Trochanteral organ of leg III with 17-21 smooth, straight, unequal spiny chaetae. Femur with a long ventrobasal mac (140-165 μm), rather thin, 50-60% as long as femur dorsal length; 4-5 shorter mac (60-100 µm); 5-6 ciliated mic in the proximal half (10-15 μm long); and a dense cover of mes, 30-100 μm long (Figure 8A). Tita with strong, slightly sinuous, finely ciliated ventral mes, including 1-2 basal ones longer and thicker than others (70-80 μm versus 40-60 μm); dorsally, dense cover of thinner ciliate-serrated mes, and a dorsal row of about 5-11 ciliated mic (10-15 μm long) (Figure 8A,B,C). The distal whorl of Tita with 9-10 subequal ciliated mes, irregularly arranged, and a thin, acuminate, smooth dorsoapical tenent hair, slightly shorter (85%) than inner edge of claw (35 μm) and than surrounding ciliated mes (Figure 8E, F); ventrodistal smooth chaeta of Tita III thick, erected, pointed, rather short (27 μm, N = 1).

Pretarsal mic minute (2.5-3 μm). Unguis slender and long (50-60 μm long, 10.5-12 μm wide at basis), with (N = 3) or without (N = 2) an unpaired inner tooth at about 72% of inner edge; the claw of the examined paratype had even 2 strong inner teeth, probably a teratological case; a pair of very small, subequal, basal teeth present at 1/8 of inner side of claw (Figure 8E, F); a pair of minute teeth dorsobasally in two specimens, but may be merely the ledge between the smooth and the primary granule areas at the basis of the claw (Figure 8D). Unguiculus about 3/5 as long as inner edge of claw, slightly swollen basointernally, blunt apically, devoid of inner tooth, not truncated, with a minute, often inconspicuous outer tooth at 1/3 of its length (Figure 8E,F).

Ventral tube. Figure 6F. VT four times longer than wide. Lateral flaps with 12-13 + 12-13 chaetae: 6-7 + 6-7 smooth chaetae distally (17-34 μm), including an external one distinctly longer (48 μm), and 6 + 6 more hyaline, subequal, ciliated mes proximally (about 20 μm). Anteriorly, 14 + 14 large chaetae, all ciliated-serrated, including 2 + 2 very large mac side by side distally (Figure 6F). Posteriorly, about 35 mes, all serrated, arranged roughly symmetrically, and 1 + 1 smooth, straight, distal mac close together. The two exsertile vesicles are broad, smooth and short, turned forwards, and devoid of anterior diverticule.

Furca complex. Figures 9, 10. Tenaculum with 4 large teeth of decreasing size from the basal to the distal one of each ramus, on a prominent, irregular body, with a posterobasal strong serrated chaeta bent distally. Mucrodens 1.15-1.30 times longer than manubrium (Figure 10A). Furcal chaetae of 3 types: scales, ciliated chaetae and spines. Inside the manubrium, two thin, straight longitudinal structures running on 3/4 of manubrium length from its apex. Manubrium ventrally with a dense cover of rounded scales (25-35 × 20-23 μm), more elongated and relatively narrow distally (up to 70 × 25-30 μm), and 2 + 2 thick ciliated centroapical mes (Figure 9A); scale ornamentation constituted of short spicules arranged in more or less regular lines. Manubrium dorsally covered by subequal ciliated and rather thin chaetae, with approximate symmetrical arrangement and with a streak bearing less or no chaetae medially; a dorsolateral line of 12-13 strong, straight, finely ciliated mac (90-110 μm) (Figure 9B); distal area with 4 + 4 ciliated mes and 2 + 2 psp. Presence of 2-3 psp between manubrium basis and genital plate (Figure 2G,H).

Dens slightly and progressively tapering, ventrally entirely and densely covered by oval, more or less fusiform scales (Figure 10B), dorsolaterally with 2 rows of spines throughout and some ciliated chaetae dorsally between these rows (Figure 9D). Scales and spines not uniform in size on the whole ventral side of the furca. Dental scales small to medium-size (8×4.5 to 14×8 μm), oval-fusiform, rounded apically, densely and regularly covered by minute spinules arranged in lines. Lateral scales and distal scales longer than ventral ones (up to 30×10 μm).

Scales grouped in multiplets made of 3-6 chaetae on the distal 2/3 of the dens (Figure 10B,C), more irregularly arranged on the 1/3 proximal (Figure 10D). In the multiplets, scales increase in size from the distal to the proximal ones.

Dental spines thick, pointed, small to medium, the shortest ones often swollen proximally, in two longitudinal rows. Dorsointernal row with 84-105 spines, dorsoexternal row with 50-65 spines (Figure 9C). Spines serrated to almost smooth (Figure 10C), with small (from 10-15 μm) and long (35-40 μm in external row, about 20 μm in internal row) spines interspersed along each row; in the proximal part of the external row, spines are replaced by thick, strong, straight, spiny chaetae (65-80 μm) (Figure 9C).

Thinner, more flexible chaetae present dorsally between the two rows of spines, very variable in size (mes to mac, 20 to 95 μm). In the 1/3 dorsoproximal side of dens, a file of 4-( 5) pseudopores (Figure 9C).

Mucro separated from dens, short (subequal or shorter than distal mac and about 20 times shorter than dens, Figure 9C), sclerotized, triangular, with 4 teeth, an apical one rather strong, two anterodistal and a minute proximal tooth (Figure 9E). Genital plate. Female genital plate with 2 + 2 smooth mic. Male genital plate with 3 + 3 genital mic and 28-30 smooth, short, acuminate circumgenital mes, without modified chaetae.

Discussion

Lepidonella species exhibit two different habitus. Epigean species are usually characterized by the presence of pigment patches on the body (unclear for L. flava), including often darkly pigmented subcoxae. Cave species are characterized by larger size, longer appendages, and absence of pigmented patches except in eyes (body, head and appendages either unpigmented or with a very light uniform coloration). Whether these two morphologies reflect classical adaptations to surface versus cave habitats or correspond to different lineages is unclear, because morphological information is lacking for most soil species. As presently circumscribed, the genus Lepidonella is widely distributed in Southeast Asia sensu lato, with numerous undescribed forms from New Guinea to southern China (unpublished data). Only two species are present in America, of L. marimuti which is strongly divergent from other species of the genus.

Authors have repeatedly underlined the striking similarity of soil Paronellidae s. l. with soil Lepidocyrtinae, expressed in the curvature of their body, their relatively small size and short appendages, the morphology of their scales and their oligochaetosis [START_REF]Puerto Rican species of Paronellidae (Insecta: Collembola)[END_REF][START_REF] Yoshii | Paronellid Collembola from caves of Central and South America collected by P. Strinati[END_REF]). Mitra's redefinition of the genus Lepidonella in 2003 states that some of its species can be "Lepidocyrtiform" sensu [START_REF] Denis | Catalogue des Entomobryens Siraeformes et Lepidocyrtiformes[END_REF]. Soto-Adames & Bellini (2015) more recently confirmed the great chaetotaxic similarity with Lepidocyrtinae of the two American species they assign to Lepidonella, as well as the basal position of Lepidonella among Paronellidae s. l. A puzzling observation, even if not conclusive given the low phylogenetic signal brought by COI, is that barcode distance trees often place side by side Lepidonella and Lepidocyrtinae species (unpublished). In spite of these similarities, mucrodens morphology always allows unambiguous assignment of any species to its family, Paronellidae s. l. or Entomobryidae. It cannot be ruled out therefore that the resemblance between Lepidonella and Lepidocyrtinae are the result of selective pressure of their edaphic habitat on one hand (small size, short appendages), and the share of plesiomorphic characters (eye arrangement, trichobothrial pattern) on the other hand. Molecular approaches will be necessary to test these competing hypotheses. [START_REF] Carpenter | XIX. Fauna of the Batu caves, Selangor[END_REF] mentions in the original description of Pseudoparonella doveri that eyes are "wanting". In the type specimen we examined, two pigment patches are visible on ocular area, a small posterior one and a larger anterior one, and two or three in all the specimens subsequently collected that we examined (Figure 1A,C,D).

Lepidonella doveri shares with L. lecongkieti from Vietnam a typical troglomorphic facies classically linked to subterranean life (Figure 1B): reduced eyes, depigmentation (no color pattern, only very diffuse pigment dots), elongate antennae, legs and claws. The two species are easily recognized from all other Lepidonella by this set of characters alone. Among other, non-adaptive characters, several are potentially discriminant but remain undocumented in most species of the genus, in particular its type species, L. tokiokai. It is the case of macrochaetotaxy or sublobal hairs of the maxillary outer lobe, where a tiny fourth chaeta present in L. lecongkieti (overlooked in [START_REF] Deharveng | Lepidonella lecongkieti n.sp., premier Collembole cavernicole du Vietnam (Collembola, Paronellidae)[END_REF]) and L. doveri has not been reported in the three other species of the genus where maxillary outer lobe has been described.

Several character states considered as adaptive to cave life differ between the two species. Two are more "advanced" in L. doveri: eye number (3 + 3 versus 6(8) + 6(8)), and tibiotarsal tenent hairs (acuminate versus clavate). Other troglomorphic characters (depigmentation, elongate appendages and claws) are almost identical. Inner teeth on claw tend to disappear in cave adapted Entomobryoidea [START_REF] Christiansen | Convergence and parallelism in cave Entomobryinae[END_REF][START_REF] Deharveng | Cave Collembola of South-East Asia[END_REF]). This character is present in L. lecongkieti and present or absent in L. doveri.

The striking similarity between L. doveri from Batu Caves (Malay Peninsula) and L. lecongkieti from southern Vietnam caves raises a biogeographical problem. They are only 850 km apart in a straight line but separated by the large gulf of Thailand. Intermediate regions along the coast of this sea have many caves where Lepidonella have not been recorded so far. Western Cambodia and Southeastern regions of Thailand are poorly known in this respect, but several caves of northern peninsular Thailand and Eastern Thailand were significantly sampled [START_REF] Deharveng | 7 -Le sud: provinces de Ranong, Chumphon, Surat Thani et Phangnga[END_REF][START_REF] Leclerc | 6 -Le sud: provinces de Kanchanaburi, Ratchaburi, Phetchaburi et Prachuap Khiri Khan[END_REF][START_REF] Jantarit | An annotated checklist of the Collembolan fauna of Thailand[END_REF] and did not host cave Lepidonella. An independent adaptation of surface species of Lepidonella to cave life in the two regions cannot be ruled out. However, it is not suggested by the high similarity between L. doveri and L. lecongkieti compared to their important differences with surface species of Southeast Asia, though most of these differences concern troglomorphic adaptations. This disjunct distribution could also result from a vicariance event following the last separation of the Sunda Shelf in South-East Asia during the mid-Holocene (c.11,000 years ago), after a long period of direct land connection [START_REF] Voris | Maps of Pleistocene sea levels in Southeast Asia: shorelines, river systems and time durations[END_REF][START_REF] Woodruff | Neogene marine transgressions, palaeogeography and biogeographic transitions on the Thai-Malay Peninsula[END_REF][START_REF] Sathiamurthy | Maps of Holocene sea level transgression and submerged lakes on the Sunda Shelf[END_REF]. However, the large Chao Phraya river and its continuation southwards during this period already represented a major barrier between Malay Peninsula and Southwestern Vietnam [START_REF] Voris | Maps of Pleistocene sea levels in Southeast Asia: shorelines, river systems and time durations[END_REF][START_REF] Sathiamurthy | Maps of Holocene sea level transgression and submerged lakes on the Sunda Shelf[END_REF]. Further investigations on the limits of cave Lepidonella west of Southwestern Vietnam and north of the Malay Peninsula, as well as phylogeographic reconstructions will be necessary to understand the pattern.

  (more details in Zhang & Deharveng 2015): ms (S-chaetae type 1 in Jantarit et al. 2013; or microsensillum in Soto-Adames et al. 2014) short, opaque, pointed and S (S-chaetae type 2 in Jantarit et al. 2013; lateral sensillum in Soto-Adames et al. 2014) slightly longer, thinner, rod-like, hyaline. Dorsal macrochaetae: 0/0,0/0,0,(3),2

Figure 1 .

 1 Figure 1. Lepidonella, photos (A, B, C, stereomicroscope; D, microscope). A, habitus of L. doveri (asymmetrical antennae); B, habitus of L. lecongkieti; C, anterior part of L. doveri; D, anterior part of head and ocular patches of L. doveri, with recognized sockets marked in black. Scales: 1 mm (A, B); 0.5 mm (C); 0.1 mm (D).

Figure 2 .

 2 Figure 2. Lepidonella doveri, photos (microscope). Pseudopores on different body parts. A, Ant. II, ventroapically; B, Th. III; C, Abd. III; D, two pseudopores of the lateral group on Abd. IV; E, Coxa 1; F, Coxa 2; G, H, between manubrial basis and genital plate. Scales: 25 μm (E, F, G); 10 μm (A, B, C, D, H).

•

  Ciliated mes, short to long (15 to 75 μm), curved, thick basally, tapering, relatively hyaline. Numerous on all antennal segments, mostly ventrally on Ant. I, small and subequal on Ant. IV (Figure 5F, G). • Mic (10-20 μm), straight, smooth, ± spiny. Located ventrobasally (3) and dorsobasally (3) on Ant. I (Figure 5B); dorsobasally (3) on Ant. II; not seen on Ant. III; absent on Ant. IV. • Finely serrated mes (23-26 μm), curved, distinctly thinner than rugose mes, subcylindrical, relatively hyaline. Present on Ant. IV except in its distal part, absent on Ant. I, II and III • Rugose mes, rather short (10-18 μm), straight or weakly curved, thick, subcylindrical, blunt apically, hyaline. Present on Ant. II, on Ant. III and on Ant. IV except in its distal part (Figure 5G); absent on Ant. I.

Figure 3 .

 3 Figure 3. Lepidonella doveri. Mouthparts and ventral side of head. A, labium basis; B, maxilla, with its two special structures; C, maxillary outer lobe; D, ventrodistal part of labrum; E, ventral chaetotaxy of head; F, clypeus. Legend: filled circles, chaeta sockets; U symbols, scale sockets. Scales: 50 μm (E); 20 μm (A, F); 10 μm (B, C, D).

Figure 4 .

 4 Figure 4. Lepidonella doveri, photos (microscope). Mouthparts. A, B, labium triangle, with m2 smooth (A) or M2 ciliated (B); C, distal part of labrum with its non-granulated intrusion; D, E, F, maxilla head at three focus. Legend: x, y, special structures of the maxilla head; 2, 3, lamellae 2 and 3 of the maxilla head. Scales: 20 μm (B); 10 μm (A, C, D, E, F).

Figure 5 .

 5 Figure 5. Lepidonella doveri. Antennae. A, types of chaetae (M, mac; m, mes; μ, mic; f, fine, serrated; r, rugose; S, long sens; e, erected; w, swollen; s, short sens; ms, micro-sens; see text for description); B, dorsobasal mic, one scale and scale sockets of right Ant. I; C, ventroexternal chaetotaxy of the proximal half of right Ant. I; D, E, externodistal part of left Ant. III, ventrally (D) and dorsally (E); F, AIIIO, left antenna; G, distal part of Ant. IV with subapical organite. Legend: r, rugose type chaetae (D); U shapes, scale sockets (B); empty circles, chaeta sockets; struck circle, pseudopore (D); 1 to 6, S-chaeta numbering of AIIIO. Scales: 30 μm (C); 25 μm (D, E, G); 20 μm (A, B, F).

Figure 6 .

 6 Figure 6. Lepidonella doveri. Chaetotaxy of tergites and of ventral tube. A, dorsal chaetotaxy. B, enlargement of the anterolateral S-chaetae of Th. II; C, D, E, enlargement of trichobothrial complex of Abd. II, Abd. III, Abd. IV. F, ventral tube, anterior side. Legend: inverted U, scale sockets (C, D, E); small black dots, mes sockets (A, D); larger black dots with numbering, mac sockets (A, D); empty circles, chaetae sockets of ventral tube (F); large blue dots with p, pseudopores (A); struck circles, pseudopores (C, D); black field triangle, modified mes of trichobothrial complex (C, D, E); large circles with central dot and T, trichobothria sockets (C, D, E); blue cross, trichobothria position (A); ms, pointed s-microchaeta; s, hyaline S-chaeta). Scales: 100 μm (A); 50 μm (E); 25 μm (D); 20 μm (B, C, D).

Figure 7 .

 7 Figure 7. Lepidonella doveri, photos (microscope). Tergites. A, Th. II, anterior line of mac with associated mes; B, Abd. III, internal trichobothrial complex including a tandem (mes+ s). Legend: m, 3 sockets of modified mes; p, pseudopore; s, S-chaeta; T, trichobothria. Scales: 15 μm (A, B).

Figure 8 .

 8 Figure 8. Lepidonella doveri. Legs. A, femur and proximal part of Tita (leg I); B, tibiotarsus (leg II); C, two mac and a mes on ventral side of Tita I; D, claw II in dorsal view; E, Claw I; F, claw III (other specimen); G, coxa I; H, coxa II; I, coxa III; J, orientation of G, H, I. Legend: empty circles, chaeta sockets; struck circles, pseudopores (G, H, I). Scales: 50 μm (A, B, G, H, I); 15 μm (C, D, E, F).

Figure 9 .

 9 Figure 9. Lepidonella doveri. Furca. A, ventrodistal chaetae of manubrium; B, diagrammatic section of manubrium; C, rows of spines on dentes; D, diagrammatic section of dens; E, Mucro. Legend: U shapes, scale sockets (A); black filled circles, spine sockets of different size (C); struck circles, pseudopores (C); empty circles, chaeta sockets (E). Scales: 100 μm (C); 50 μm (B); 25 μm (D); 10 μm (A, E).

Figure 10 .

 10 Figure 10. Lepidonella doveri. Furca, photos (A, stereomicroscope; B, C, D, microscope). A, furca in lateral view; B, scales arranged in multiplets on ventrodistal side of dens; C, dental spines and scale multiplets on distal part of dens; D, ventroproximal scales on dens, not arranged in multiplets. Scales: 25 μm (A, D); 10 μm (B, C).

  Paronella seychellesiensis was proposed as a replacement name for flava by[START_REF] Salmon | An index to the Collembola[END_REF] as a result of the synonymy of Microparonella with Paronella Schött, 1893, which already included the species Paronella flava Carpenter, 1916 described from Myanmar. However, the transfer of P. seychellesiensis (= M. flava) into the genus Lepidonella led to the decision being reversed and P. seychellesiensis being synonymized with Lepidonella flava[START_REF] Deharveng | Lepidonella lecongkieti n.sp., premier Collembole cavernicole du Vietnam (Collembola, Paronellidae)[END_REF].

	Lepidonella duodecimoculata in Deharveng & Bedos 1995: 22
	Yosii 1966b: 378 (Sri Lanka) Lepidonella ceylonica in Mitra 1993 (n. comb.): 86; Deharveng & Bedos 1995: 22 (erroneously given as n. comb.); Mitra 2003: 80 (erroneously given as n. comb.); Soto-Adames & Bellini 2015: 337 Lepidonella doveri (Carpenter, 1933) Pseudoparonella doveri Carpenter 1933: 220 (Malaysia); Absolon & Kseneman 1942: 23; Moseley 2004: 81 Troglopedetes doveri in Deharveng 1987: 167 as n. comb.; Deharveng 1988: 97; Fanciulli et al. 2003: 203 ?Microparonella doveri in Yoshii 1981: 41; Yoshii 1983: 11 (as incertae sedis) Microparonella doveri in Mitra 1993: 54 (n. comb., without Lepidonella oudemansi in Deharveng & Bedos 1995: 22 (n. comb.); Mitra 2003: 80 (erroneously given as n. comb.); Soto-Adames & Bellini 2015: 337 The color pattern of L. oudemansi recalls that of L. incerta. Soto-Adames & Bellini (2015: 337) consider on this basis that the two species might be synonyms. Lepidonella papuana (Womersley, 1937) Pseudoparonella papuanus Womersley 1937: 208 (Papua New Guinea) Lepidonella papuanus in Mitra 2003: 82 (holotype examined, n. comb.) The assignation of this species to Lepidonella by Mitra (2003) was overlooked in Deharveng & Bedos (1995) and Soto-Adames & Bellini (2015). Microparonella nigrofasciata in Yoshii 1983: 8 (n. comb.Microparonella oudemansi Yoshii 1983: 7 (Malaysia) (n. comb.); Soto-Adames & Bellini 2015: 337 Lepidonella flava (Carpenter, 1916) Microparonella flava Carpenter 1916: 44 (Seychelles) Syn. Paronella seychellesiensis Salmon 1964: 545 (replacement name) Microparonella seychellensis in Yoshii 1981: 38 (misspelling for seychellesiensis Salmon 1964) Lepidonella flava in Deharveng & Bedos 1995: 22 (n. comb.); Soto-Adames & Bellini 2015: 337 Lepidonella incerta (Handschin, 1925) Microparonella kei Yoshii & Suhardjono 1992: 47 (Kei Island, lndonesia) Lepidonella kei in Deharveng & Bedos 1995: 22 (n. comb.); Soto-Adames & Bellini 2015: 337 Lepidonella lecongkieti Deharveng & Bedos, 1995 Lepidonella lecongkieti Deharveng & Bedos 1995: 23 (Vietnam) Lepidonella lecongkieti in Soto-Adames & Bellini 2015: 337 Lepidonella nigrofasciata (Handschin, 1928) Pseudoparonella nigrofasciata Handschin 1928: 262 (Java) Handschinella nigrofasciata in Yosii 1966a: 50 (n. comb.) Bromacanthus nigrofasciatus in Yoshii 1981: 41 (incertae sedis) Lepidonella picta (Schäffer, 1898)

justification);

Mitra 2002: 70 (n. comb., with justification)

Lepidonella duodecimoculata

(Prabhoo, 1971)

Microparonella duodecimoculata Prabhoo 1971: 43 (India) Pseudoparonella incerta Handschin 1925: 255 (Sumatra) Microparonella incerta in Yoshii 1983: 11 (n. comb.) Lepidonella incerta in Deharveng & Bedos 1995: 22 (n. comb.); Mitra 2003: 80 (erroneously given as n. comb.); Soto-Adames & Bellini 2015: 337

According to Soto-Adames &

Bellini (2015, p. 337)

, "the original species description is very brief, and the color pattern described suggests that L. incerta may be a senior synonym of L. oudemansi."

Lepidonella kei

[START_REF] Yoshii | Notes on the Collembolan fauna of Indonesia and its vicinities. II: collembola of Irian Jaya and Maluku islands[END_REF]

) ) Lepidonella nigrofasciata in

Deharveng & Bedos 1995: 22 (n. comb.)

; Mitra 2003: 80 (erroneously given as n. comb.); Soto-Adames & Bellini 2015: 337

Lepidonella oudemansi

[START_REF] Yoshii | Studies on Paronellid Collembola of East Asia[END_REF]

Paronella picta Schäffer 1898: 410 (Papua New Guinea: New Britain) Lepidonella picta in

Mitra 1992: 212 (n. comb.)

;

Mitra 2003: 74, 80

(Solomon Islands) (erroneously given as n. comb.)

  [START_REF] Soto-Adames | Dorsal chaetotaxy of neotropical species supports a basal position for the genus Lepidonella among scaled Paronellidae (Collembola, Entomobryoidea)[END_REF] The presence of two sublobal hairs on the maxillary palp(Soto-Adames & Bellini 2015, p. 333) separates this American species from the four Asian species in which this character is known.Yoshii (1983, p. 11) placed Pseudoparonella incerta Handschin in the genus Microparonella. Hence, the species described as Microparonella incerta by Mari-Mutt (1987) was a homonym and should have been renamed.[START_REF] Deharveng | Lepidonella lecongkieti n.sp., premier Collembole cavernicole du Vietnam (Collembola, Paronellidae)[END_REF], having shown that Microparonella was a synonym of Lepidonella, placed incerta Handschin in this last genus, but they mentioned that there was a doubt (underlined below) about the assignation of incerta Mari-Mutt to

	Incertae sedis
	Lepidonella marimuti Soto-Adames & Bellini, 2015
	Microparonella incerta Mari-Mutt 1987: 404 (Puerto Rico), nec
	Handschin 1925: 255 (Sumatra).
	Lepidonella marimuti Soto-Adames & Bellini 2015: 337 (US
	Virgin Islands, Dominican Republic) (Replacement name)
	?Microparonella incerta in Deharveng & Bedos 1995: 22 (incertae
	sedis)
	Lepidonella zeppelinii Soto-Adames & Bellini 2015: 332
	(Brazil).

Lepidonella: "Microparonella incerta Mari Mutt, 1987 (Puerto Rico), s'il s'agit bien d'un Lepidonella, devient un homonyme junior de L. incerta

[START_REF] Handschin | Beiträge zur Collembolenfauna der Sundainseln[END_REF]

". Actually, as stressed by Soto-Adames & Bellini (2015), L. marimuti is very isolated within Lepidonella, by its apical bulb on Ant. IV and its tridentate mucro, but also by the presence of several ciliated labral chaetae, and hardly modified accessory chaetae of its trichobothrial complexes. Apical bulb on Ant. IV has a high taxonomic value given its distribution among suprageneric taxa in most Entomobryoidea (except in Lepidocyrtus s. l.

  Lepidonella vivieni inDeharveng & Bedos 1995: 22 (n. comb.) Trogolaphysa vivieni in Soto-Adames& Bellini 2015: 337 (n. comb.)

	(Microparonella) monomaculata Barra 1969: 212 (Gabon)
	Lepidonella monomaculata in Deharveng & Bedos 1995: 22 (n.
	comb.)
	Trogolaphysa monomaculata in Soto-Adames & Bellini 2015:
	337 (n. comb.)
	Trogolaphysa vivieni (Barra, 1969)

Paronella (Microparonella) vivieni

Barra 1969: 207 (Gabon)

Acknowledgments

We are grateful to Max Moseley and Liz Price for having supplied to us fresh material of the species, Paul Brown (curator) from the Natural History Museum of London for the loan of a type specimen and David Ouvrard for having facilitated the loan.


Gabriel Nève
email: neve@up.univ-mrs.fr

Michael C Singer

Protandry and postandry in two related butterflies: conflicting evidence about sex-specific trade-offs between adult size and emergence time

Keywords: Protandry, postandry, size, Euphydryas aurinia, Euphydryas editha

Natural selection acting on timing of metamorphosis can be sex-specific, resulting in differences in timing between males and females. Insects with discrete generations frequently show protandry: males usually mature before females. Both Euphydryas editha and E. aurinia butterflies followed this trend. The present study was motivated by the unusual observation of consistent postandry in addition to protandry. In a single E. editha population observed over 20 years the emergence period of males was longer than that of females, both the first and last emerging individuals being males. Variance of timing among individual E. editha larvae is imposed by spatial patchiness of the snowmelt that releases them from winter diapause. If individual larvae released late from diapause were to compensate for their lateness by shortening their development times, they would be small at maturity. If such compensation were only partial, they would be both late and small. Size and timing would become associated. If females were more prone to such partial compensation than males, the observations of postandry could be explained and the prediction made that any tendency for late individuals to be small should be stronger in females than in males. This was the case: in one year late males were the same size as early males, in a second year they were larger. Late females were significantly smaller than early females in both years. In E. aurinia, results were opposite both to theoretical prediction and to the observations from E. editha: although the male emergence period was longer than

that of females exactly as in E. editha, late males were smaller than early ones, while late females were not small. The data from E. editha support the hypothesis of a sex-specific trade-off between size and emergence time, the data from E. aurinia do not.

Introduction

In animals which metamorphose, a juvenile that has reached a size where transition to the adult stage is possible can choose between commencing metamorphosis or delaying it and gathering further resources as a larva. When larvae are more susceptible than adults to predation, such delay incurs a cost in terms of prolonged larval exposure to predators [START_REF] Benard | Predator-induced phenotypic plasticity in organisms with complex life histories[END_REF][START_REF] Vonesh | Opposite shifts in size at metamorphosis in response to larval and metamorph predators[END_REF][START_REF] Stearns | The Evolution of Life Histories[END_REF]. In such cases individuals programmed to delay metamorphosis suffer greater exposure to predators as larvae but, if they survive, have increased body size, conferring higher fecundity to females and competitive advantage to males [START_REF] Berger | What keeps insects small? -Size dependent predation on two species of butterfly larvae[END_REF]. The opposing selective forces associated with predation and reproduction then result in a trade-off between maximizing size, hence fecundity (e.g. [START_REF] Taylor | Effects of size at metamorphosis on stone fly fecundity, longevity, and reproductive success[END_REF], and minimizing predation [START_REF] Forrest | Insect size tactics and developmental strategies[END_REF][START_REF] Candolin | Size-dependent selection on arrival times in sticklebacks: why small males arrive first[END_REF].

In most temperate-zone butterfly species emergence is protandrous (males emerge before females) and males are smaller than females [START_REF] Wiklund | Sexual size dimorphism in relation to female polygamy and protandry in butterflies: a comparative study of Swedish Pieridae and Satyridae[END_REF], through shorter male larval developmental and pupal times (Fischer and Fiedler, 2001a). Selection for protandry arises because males that emerge before peak female emergence encounter more virgin females than later males [START_REF] Iwasa | Emergence patterns in male butterflies: a hypothesis and a test[END_REF]. In a north temperate butterfly community (Minnesota), species with discrete generations tended to be more sexually dimorphic (females larger than males) than species with overlapping generations or than those which refrained from mating till after an adult diapause [START_REF] Singer | Sexual selection for small size in male butterflies[END_REF]. This difference between species with discrete and overlapping generations implies some influence of sexual selection for protandry on size dimorphism. These bodies of evidence suggest that in univoltine butterfly species the selection pressures on size and phenology are sexdependent [START_REF] Wiklund | Sexual size dimorphism in relation to female polygamy and protandry in butterflies: a comparative study of Swedish Pieridae and Satyridae[END_REF].

Size and phenology may differ between the sexes even in a constant environment if each sex were programmed to grow for a particular length of time to a particular size, with males normally being the smaller and earlier sex. However, in the presence of environmental variation, sex-specific plastic responses could complement and alter this relationship. Here we use field data from two congeneric butterflies species (genus Euphydryas), to test ideas about how such sex-specific responses might interact with the trade-offs between size and emergence time that have been predicted [START_REF] Singer | Sexual selection for small size in male butterflies[END_REF][START_REF] Zonneveld | Being big or emerging early? Polyandry and the trade-off between size and emergence time in male butterflies[END_REF].

Materials and methods

Our study populations of Euphydryas aurinia and E. editha are univoltine (Lepidoptera, Nymphalidae) which diapause as partly-grown larvae. In spring, larvae resume feeding, and produce a single, discrete generation of adults which fly for about a month between May and July, with precise timing of depending on the local weather conditions. Most individuals tend to stay in their natal habitat patch, but occasional movements between patches occur.

Euphydryas editha (Boisduval) Size At Rabbit Meadow, Tulare County, California, E. editha individuals were captured and measured at the beginning and at the end of the period of adult emergence. This was done in both 1995 and 1996. Each individual was measured, marked and released to prevent measuring the same insect twice. Altogether, 139 and 215 individuals were measured in 1995 and 1996 respectively. Comparisons of wing lengths of early-emerging and lateemerging insects were performed using standard t-tests.

Phenology

We noted the sex of the last freshly-emerged individual E. editha (with no visible scale loss) seen at Rabbit Meadow in each year from 1983 and to 2006, omitting the years 1984, 1987, 1990 and 2003. In each year observations were made daily. Because of the possibility that the fresh males observed at the end of the emergence season were immigrants, in one year (1986) ten small (5 m square) patches were searched intensively to record the sex of the last teneral insect (that could not yet fly). This was done in a year of extremely high insect density [START_REF] Thomas | Catastrophic extinction of population sources in a butterfly metapopulation[END_REF], in which an average of around 20 teneral insects in each 5 m square patch were found.

Euphydryas aurinia (Rottemburg)

Our study site was at Deiffelt, Luxembourg province, Belgium. A Mark-Release-Recapture survey of E. aurinia was performed daily from 27 May to 4 July 1995, except on rainy days. On each of the 23 sunny days, the whole site was visited and a 20 m grid was systematically searched. Each captured individual was individually marked with a number on the underside of the left hindwing. Upon capture, the length of the forewing was measured and the wear of the wings was assessed on a 4 point scale, from fresh to very worn. All individuals were released at the site of capture immediately after marking. A total of 209 males and 118 females were captured, of which 152 and 86 respectively had been first captured when fresh or undamaged (wing wear classes 1 and 2).

As most males emerge before females do, a model incorporating both sex and capture date as possible explanatory variables would have violated the assumption of independence between variables. The data for each sex were thus analyzed separately. Correlations between two variables (size and day), for each sex, were computed using Spearman's  rank correlation coefficient. Log transformations did not normalize the data. All statistical analyses were done using R (R Development Core Team, 2004), except daily population size and births which were estimated using Jolly-Seber model with the POPAN5 program [START_REF] Arnason | POPAN-5: A data maintenance and analysis system for markrecapture data[END_REF][START_REF] Arnason | Using POPAN-5 to analyse banding data[END_REF]. As the site was isolated from other populations of the species, all new individuals caught on the site had emerged locally. The mean day of emergence was calculated using the mean of all individual birth dates.

Results

Euphydryas editha

Wing size distribution did not differ significantly from Normal distribution for each time/sex combination (Shapiro-Wilk test, smallest P=0.16). In both years, females were significantly smaller at the end of the emergence season, whereas there was a trend the other way, males being bigger at the end of the season than at the beginning, in 1995, and there was no trend for males in 1996 (Table 1).

There was a strong and significant tendency for first and last insects to emerge to be males. The last freshly-emerged insect was male in 18 of the 20 years and female in one year. In the remaining year the last two freshly-emerged insects were a male and a female, observed on the same day. The observed ratio of 18:2 is significant at P=0.0004. (two-sided binomial test, R Development Core Team, 2004). We missed the first few days of the emergence season in all but 9 years; in 8 of those 9 years the first insect observed was also a male; in the last year of observation, 2006, the first E. editha seen was female. The ratio 8:1 is also significantly different from the null expectation of 50:50 (P=0.039).

Euphydryas aurinia

There was a strongly protandrous pattern of emergence (Fig. 1), with the mean day of emergence of males on day 20.8 (day 21 is 16 June) and of females on day 29.4 (24 June). The male emergence period lasted for longer (sd of emergence date=9.6) than that of females (sd of emergence date=4.2). The last fresh individuals newly captured were 2 males on day 43, while the last fresh females were on the two previous days (with one individual each time). The total population sizes over the entire flight period were estimated at 324 males and 204 females.

Neither days of capture for each sex, nor wing length follow a normal distribution (all four Shapiro-Wilk normality test, being highly significant P<0.001). For males the size decreased from the beginning to the end of the season (Fig. 2, Spearman's =-0.206, n=152, P =0.011), while the trend for females was for increasing size with time, albeit not significantly (Spearman's =-0.205, n=86, P =0.058).

Discussion

In many temperate-zone arthropods the developmental stage of all individuals in the same generation is synchronized at diapause, so that variation in timing of birth in summer has lost its influence when development re-starts in the following springtime. Furthermore, generations are generally discrete, such that adults from one generation are dead by the time the next generation of adults appears. In the absence of variation in speed of development (which does exist, [START_REF] Nylin | Plasticity in life-history traits[END_REF][START_REF] Fischer | Physiological costs of growing fast: does accelerated growth reduce pay-off in adult fitness?[END_REF], insects programmed to be larger will take longer to develop and emerge later [START_REF] Zonneveld | Being big or emerging early? Polyandry and the trade-off between size and emergence time in male butterflies[END_REF]. Conversely, insects programmed to emerge early may need to be small, in which case natural selection acting on timing of maturity will affect evolution of adult size. Evolution of sex-specific timing may generate sexual dimorphism in size [START_REF] Singer | Sexual selection for small size in male butterflies[END_REF].

These genetic constraints generate the expectation of a phenotypic covariance between size and timing: early insects should be smaller, late ones larger. However, this expected relationship can be reversed by plastic responses to environmental variation [START_REF] Higgins | The interaction of season length and development time alters size at maturity[END_REF] and these plastic responses may themselves be sex-specific (Fischer and Fiedler 2001b).

Why might the sexes respond differently to environmental stress? The collective works of [START_REF] Wiklund | Sexual selection for large male size in a polyandrous butterfly: the effect of body size on male versus female reproductive success in Pieris napi[END_REF] and of [START_REF] Leimar | Unpredictable food and sexual size dimorphism in insects[END_REF] give a relevant example. In the butterfly Pieris napi, male-donated nuptial gifts were large, so that females could potentially increase their mating frequency to compensate any reduction of fecundity, such as that conferred by small body size. These authors suggested that size should be less important to females than to males. This prediction was supported by the experimental finding that females yielded size more readily than males when larvae were fed a low quality diet. This experiment showed clearly that the sizes achieved by adult males and females can respond differently to the same environmentally-imposed variation.

Our own study began with a set of observations that both the first and the last butterflies to emerge in our study population of E.editha were almost always males. This dataset now extends to 20 years, in only one of which the latest freshly-emerged insect observed was female. The male emergence season was longer that that of females, contrary to the hypothesis of approximately equal emergence time for males and females used in [START_REF] Fagerström | Why do males emerge before females ? Protandry as a mating strategy in male and female butterflies[END_REF] model. The last males to emerge appeared when they had no opportunity to encounter virgin females. This did not mean that a late male could not mate, as approximately 10% of female E. editha mate twice [START_REF] Ehrlich | Reproductive strategies in the butterflies. I. Mating frequency, plugging, and egg number[END_REF]) but a late male's opportunities to mate were much reduced, as was the expected number of offspring that it could father if it did mate. The consistent appearance of late males is especially paradoxical in E. editha, because its larvae can respond to perceived adversity by repeated diapause. A male larva that found itself late could re-enter diapause and try again next year, starting from a larger initial size.

Why might females emerge over a shorter period than males? If size were more important to male E.editha than to females, as is the case in P. napi, then males that suffer environmentally-imposed time delays should be less likely than equally-delayed females to yield size in an effort to maintain phenology. This would provide a satisfying explanation of both the postandry that occurs in the field and the difference between the sexes in the size trend over time. The necessary environmental variation to fuel this effect is provided by patchy snowmelt. E.editha larvae diapause under snow and commence feeding almost immediately after snowmelt. Within a single site, snow patches melt at different times, creating phenological differences of 1-3 weeks among individual larvae. Delayed larvae have the option to re-enter diapause and complete development in the following year but not all of them do so. Those delayed larvae that continue development could be viewed as having two very different options. They could maintain predetermined size regardless of phenology or predetermined phenology regardless of size. Neither of these extreme cases would generate a phenotypic association between time and size. However, any intermediate response between the two extremes would do so. For example, a larva that was delayed ten days by snowmelt would have the option of emerging with only seven days' delay but reduced in size by three days' growth. If larvae in general did this, then late-emerging adults would be smaller than early ones. This is the relationship that we see between size and phenology of female E. editha. In contrast, delayed male E. editha evidently were not reduced in size. The male emergence period was extended relative to that of females, as both the first and last individuals to emerge were typically males.

The entire dataset from E. editha is consistent with the hypothesis that females delayed by late snowmelt traded size against time, while delayed males did not. Why should females do this? We might expect to discover some reason why size is less important to females than to males, or some reason why timing is less important to males than to females. We have observed no such phenomena. It is not likely that females can compensate for small size by mating frequently as in P. napi, since nuptial gifts in E. editha are very small [START_REF] Boggs | Checkerspot Reproductive Biology[END_REF]. Neither is it true at this site that strong selection on timing of oviposition arises from seasonal effects on offspring mortality, since survival on the traditional host at Rabbit Meadow was identical for early, mid-season and late eggs [START_REF] Boughton | Empirical evidence for complex source-sink dynamics with alternative states in a butterfly metapopulation[END_REF]. In sum, our current knowledge of E. editha biology merely allows us to discount some obvious possible sources of selection that would shorten the female emergence season but identifies no clear source of such selection.

In our other study species, E. aurinia, the most likely source of environmental variation is not time of breaking diapause but shortage of food for large larvae that have defoliated their hosts and must either search for new hosts or pupate at the size they have already achieved, especially as individuals of the E. aurinia's foodplant, Succisa pratensis, often grow singly on this site. [START_REF] Boggs | Larval food limitation in butterflies: effects on adult resource allocation and fitness[END_REF] showed that a lowering of larval resources reduces adult mass, forewing length and fitness. This effect could render late individuals small by the same type of trade-off we suggested for E. editha. Like E. editha, the E.aurinia population that we studied showed a difference between the sexes in the association between adult size and time of emergence. However, this difference was in the opposite direction from that in E. editha. Late-emerging males were small, late-emerging females were not. By the argument we have developed here, we expect the sex that sacrifices size when it is delayed to have the shorter emergence period. This is true for E. editha but not for E. aurinia. At present we lack a sensible explanation for this difference. In this as in any comparison involving only two species and two sites, the question remains open as to whether the observed difference is due to site or to species difference or both [START_REF] Garland | Why not to do two-species comparative studies[END_REF].

Any difference between E. aurinia and E. editha in the experience of environmental variation by larvae should apply equally to males and females, so it would not account for the difference between the species that we observed. Any comprehensive explanation should involve differences between the species in the relative importance of size and timing to male and female fitness. Investigation of these differences should be a fruitful avenue to understanding how genetic and environmental variation interact to generate correlations among important life-history traits in natural habitats.

Figure 1 .

 1 Figure 1. Emergence curves for E. aurinia at Deiffelt in 1995, as the estimates of numbers of new individuals in the population at each capture session, according to using Jolly-Seber model. Vertical bars indicate s.e. of estimates, when available. Hollow symbols indicate the numbers of freshly emerged individuals having actually been captured. Day 1 is 27 May 1995.

Figure 2 .

 2 Figure 2. Forewing lengths of E. aurinia males (top) and females (bottom) captured at Deiffelt, against the days on which first captured in 1995 (Day 1 is 26 May) . Only wingwear classes 1 and 2 (out of 4) are considered. Multiple points are plotted with the numbers of radius bars corresponding to the numbers of observations. Lines depict running mean wing length for individuals captured during three consecutive trapping sessions.

Table 1 .

 1 Forewing size of Euphydryas editha specimens, according to sex and time in 1995 and 1996. All t tests were between data from time 1 and time 2 per year and sex.

		N	Mean	t	P
	1995 Female wing length
	Time 1	25	21.80 mm
	Time 2	49	20.89 mm	4.40	0.0001
	1995 Male wing length
	Time 1	27	18.93 mm
	Time 2	38	19.37 mm	-2.17	0.034
	1996 Female wing length
	Time 1	66	21.69 mm
	Time 2	45	21.06 mm	3.16	0.002
	1996 Male wing length
	Time 1	67	19.37 mm
	Time 2	37	19.56 mm	-1.19	0.237

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Acknowledgements

Philippe Goffart, David Boughton and Chris Singer helped gather the data. We thank Philippe Lebrun and Michel Baguette (University of Louvain) and the Ministère de la Région Wallonne for support to the E. aurinia study.