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anterolenis -group (new species group) Species group definition. Species of Megalothorax with secondary granulation coverage limited to the abdominal I–IV tergites.

Megalothorax zealanterolenis sp. nov. Figs 6, 7, 8A, B, 9E Material examined. For all following material, name of locality, date of collection and name of collector: New Zealand, South Island, Nelson-Tasman region, Kahurangi National Park, Mt Arthur summit track, 27 January 2017, collected by M. Minor. Holotype. Female on slide (CSCOL _1327), 41.1967°S, 172.7312°E, 1096 m a.s.l., southern beech forest (mostly Fuscospora cliffortioides (Hook.f.) with some Lophozonia menziesii (Hook.f.), in soil with litter and moss Hypnum cupressiforme Hedw., sample MtAb_ T 1-1. Paratypes. Four females and one male on three slides (CSCOL _1326, 1328, 1329), same data as the holotype, sample MtAb_ T1-1. Other material. One female on slide CSCOL _1330, 41.1966°S, 172.7315°E, 1091 m a.s.l., southern beech forest (F. cliffortioides), in soil with litter, sample MtAb_ T 1-4. A juvenile on slide CSCOL _1331, 41.1965°S, 172.7319°E, 1082 m a.s.l., southern beech forest (mostly F. cliffortioides with some F. fusca), in soil with litter, sample MtAb_ T 1-5. A female on slide CSCOL _1332, 41.1962°S, 172.7322°E, 1084 m a.s.l., southern beech forest (mostly F. cliffortioides with some F. fusca), in soil with litter, sample MtAb_ T 1-6. Three juveniles, three males, one female and one adult (sex indet.) on eight slides CSCOL _1333–1340, 41.1923°S, 172.7354°E, 1066 m a.s.l., southern beech forest (F. cliffortioides with some L. menziesii), in soil with litter, sample MtAb_ T 2-1. Four juveniles, two females on three slides CSCOL _1341–1343, 41.1923°S, 172.7354°E, 1066 m a.s.l., southern beech forest (F. cliffortioides with some L. menziesii), in soil with litter under liverwort Bazzania involuta (Mont.) Trevis., sample MtAb_ T 2-2. Two females on a slide CSCOL _1345, 41.1920°S, 172.7359°E, 1077 m a.s.l., southern beech forest (F. cliffortioides with some L. menziesii), in soil with litter, sample MtAb_ T 2-5. Two females on two slides CSCOL _1346, 1347, 41.1901°S, 172.7397°E, 1018 m a.s.l., mixed southern beech forest (F. cliffortioides with some Fuscospora fusca (Hook.f.) and L. menziesii), in soil with litter, sample MtAb_ T 3-1. One female on slide CSCOL _1348, 41.1901°S, 172.7397°E, 1018 m a.s.l., mixed southern beech forest (F. cliffortioides with some F. fusca and L. menziesii), in soil and litter with moss Dicranoloma robustum (Hook.f. & Wilson) Paris, sample MtAb_ T 3-2. One female on slide CSCOL _1349, 41.1902°S, 172.7399°E, 1024 m a.s.l., mixed southern beech forest (F. cliffortioides with some F. fusca), in soil and litter with moss D. robustum, sample MtAb_ T 3-3. Two juveniles, one adult (sex indet.) on three slides (CSCOL _1350–1352), 41.1902°S, 172.7400°E, 1025 m a.s.l., southern beech forest (F. cliffortioides with some F. fusca), in soil and litter with moss D. robustum, sample MtAb_ T 3-4. One male, one female on two slides (CSCOL _1353, 1354), 41.1903°S, 172.7409°E, 1013 m a.s.l., mixed southern beech forest (F. cliffortioides with some F. fusca and L. menziesii), in soil and litter with moss Ptychomnion aciculare (Brid.) Mitt., sample MtAb_ T 3-6. Material deposit. The holotype (CSCOL _1327) and two paratypes (a female and a male, CSCOL _1328, CSCOL _1329) will be deposited at the Museum of New Zealand Te Papa Tongarewa, Wellington, New Zealand. Three female paratypes (CSCOL _1326) will be deposited at the Senckenberg Museum für Naturkunde, Görlitz, Germany. Diagnosis. Whitish in ethanol, labrum a1 chaetae with bifurcate tip, maxillary palp with an internal bifurcate hair, basomedian and basolateral fields of labium each with 1 + 1 chaetae, secondary granulation limited to the abdominal I – IV tergites, on the abdomen s-chaetae s2 elongated and tubular, s3 absent, tenaculum with 4 + 4 teeth, posterior lamellae of the mucro moderately enlarged, internal lamella serrated, external lamella smooth and waved or with weak teeth. Description. General aspect. Habitus and segmentation typical of the genus. Length from labrum to anus: up to 400 µm. Specimens whitish (in 96% ethanol). All typical chaetal types of the genus are accounted for, without any remarkable development. Integument. Secondary granulation restricted to the dorsal part of the abdominal area except Abd. VI, similarly to M. anterolenis sp. nov. Head channels connection with linea ventralis circular, integumentary channels not precisely analysed, apparently limited to the dorsal part of the head. Sensory fields and wax rods. Same observation as in M. anterolenis. sp. nov., see Fig. 6A,B. Head chaetotaxy. Postero-dorsal chaetae without remarkable thickening, stronger than antero-dorsal chaetae. Number of chaetae: 12 + 12 in the postero-dorsal region, 9 + 9 and 2 unpaired in the antero-dorsal region, 2 + 2 in the antero-lateral region (Fig. 6B). Ventrally, 3 + 3 chaetae in the sub-labial region (Fig. 8B). Labium. Basomedian fields of labium with 1 + 1 chaetae, basolateral fields of labium with 1 + 1 chaetae on a small papilla. Labial palps ordinary. Labrum. Same observations as in M. tasmanterolenis. Other mouthparts. Same observations as in M. tasmanterolenis, left mandibula figured in Fig. 6C. Antenna. Ant. I with one chaeta. Ant. II with four chaetae, the anterior one much stronger than the three others (Fig. 6D). Ant. III with 10 chaetae, and five S-chaetae including S1–S4 from the sensory organ and Sb4; S2 and S3 small but clearly protruding from the cupule; without perceptible ornamentation in light microscopy; S1 and S4 appeared well contrasted with perceptible ornamentation rings. Ant. IV with five chaetae (X present), and 11 Schaetae (Sb1–3, Sb 5, Sa 1-5, Sx and Sy; Sa2 notably enlarged and with a clearer appearance than the others common S-chaetae (Sa and Sb); 2 apical and subapical rods (a, sa) present (Fig. 6D). Thoracic and abdominal tergites. As in Fig. 6A, same observations as in M. anterolenis sp. nov., but with schaetae s3 absent and s-chaetae s2 elongated. Pseudopores not studied, 3 + 3 τ-chaetae could be seen (the others to be considered not studied). Abd. VI. Ordinary, with nine chaetae on tergites, a small chaetae on each anal valves. Sternites chaetotaxy not studied. Genital plate. Female with 2 + 2 chaetae, male not studied. Abd. IV sternites. With 2 + 2 usual neosminthuroid chaetae and 2 + 2 ordinary chaetae, the external chaetae much shorter than the internal ones (Fig. 7A). Abdominal appendages. Manubrium with 2 + 2 posterior chaetae. Dens ordinary: basal part of dens with 1 + 1 posterior chaetae, apical part of dens with 1 + 1 posterior chaetae and 7 + 7 small spines. Apex of the five anterior spines tapering to a filament. Mucro elliptical, with wide lamellae; five to seven teeth on the external posterior lamella, internal posterior lamella smooth and waved, sometimes with weak teeth (Figs 7H – J, 9E). Ventral tube with 2 + 2 apical chaetae, retinaculum with 4 + 4 teeth. Legs. Chaetal composition on legs (subcoxa 1, 2, coxa, trochanter, femur and tibiotarsus): leg I, 1, 0, 1 (at least), 2, 8, 12 chaeta(e); leg II, 1, 1, 1, 3, 8, 12 chaeta(e); leg III, 2, 1, 1, 4, 8, 10 chaeta(e). Claws ordinary, as in Fig. 7E, F, G.; with basal lamellae la and lp short. Differential diagnosis. Most similar to M. tasmanterolenis sp. nov., the two species can be distinguished from the shape of s-chaetae s2 on the abdomen. Name etymology. Derived from the name of M. anterolenis sp. nov. combined with the country of discovery (New Zealand). Ecology. The new species was found in soil and litter of montane southern beech forests in the subalpine zone (ca. 1000–1100 m above sea level) of Mt Arthur (1795 m) in the Kahurangi National Park, Nelson-Tasman region of the South Island of New Zealand. The Mt Arthur sites vegetation is dominated by montane forests of southern beech, mostly mountain beech Fuscospora cliffortioides (Hook.f.) Heenan & Smissen, some red beech Fuscospora fusca (Hook.f.) Heenan & Smissen and silver beech Lophozonia menziesii (Hook.f.) Heenan & Smissen, small shrubs and mosses. The soils in sampled plots were cold (7–8 deg Celsius) and wet (21–44% volumetric moisture content) in January (mid-summer). The new species was widespread in samples from Mt.Arthur and locally found in high abundance (mean density 2,800 ind. m-2, max 10,800 ind. m-2). The new species may be geographically restricted, as it was not observed in subalpine beech forests of two other mountains sampled in the same region., Published as part of Schneider, Clément, Minor, Maria A. & D'Haese, Cyrille A., 2023, A new group of species of the genus Megalothorax (Collembola, Neelidae) with Gondwanan distribution, and introducing an open interactive identification key of Megalothorax species, pp. 101-121 in Zootaxa 5228 (2) on pages 110-115, DOI: 10.11646/zootaxa.5228.2.1, http://zenodo.org/record/7532173

Megalothorax anterolenis sp. nov. Figs. 1–3, 8A–D. Material examined. Holotype. Female on slide (CSCOL _104), Chile, X Región de Los Lagos, Llanquihue, Parque Nacional Alerce Andino, 2011.xi.23, 41.5838°S, 72.5698°W, dwelling among epiphytic plants on a living tree, sample CHL002. Paratypes. Four females on a single slide (CSCOL _105), same data as the holotype; two females and two males on two slides (CSCOL _111, 113), same data as the holotype but sampled on another tree: 41.5836°S, 72.5692°W, sample CHL004. Other material. One female, one juvenile on two slides (CSCOL _190, 191), Chile, X Región de Los Lagos, Llanquihue, Parque Nacional Alerce Andino, 2011.xi.24, 41.5882°S, 72.5808°W, among mosses on dead wood, sample CHL015. One female on slide (CSCOL _125), Chile, X Región de Los Lagos, Llanquihue, Parque Nacional Alerce Andino, 2011.xi.26, 41.5787°S, 72.5599°W, among mosses and epiphytic plants naturally fallen from the host tree, sample CHL028. Three females on slide (CSCOL _213, 214, 221), Chile, Región de Aysén, Sendero Laguna Los Pumas, 2011.xii.04, 44.2354°S, 72.5195°W, among mosses growing on the base of a tree, sample CHL218. All material collected by C. Schneider. Material deposit. The holotype and two paratypes (CSCOL _104, CSCOL _111) will be deposited at the Museo Nacional de Historia Natural de Chile, Santiago, Chile. Four paratypes (CSCOL _105) will be deposited at the Muséum National d’Histoire Naturelle, Paris, France. Rest of the material will be deposited at the Senckenberg Museum für Naturkunde, Görlitz, Germany. Obtained molecular data and genbank accession number. One specimen from sample CHL002 (same a holotype): 28S rDNA: OP942371 (808bp). One specimen from sample CHL004, COI: OP933760 (658bp), 16S rDNA: OP942368 (454bp), 28S rDNA: OP942372 (808bp). One specimen from sample CHL218, COI: OP933759 (658bp), 16S rDNA: OP942369 (468bp), 28S rDNA: OP942373 (784bp). Vouchers could not be recovered. Diagnosis. Colour varying from whitish to red (in 96% ethanol), labrum a1 chaetae ordinary with acuminate tip, maxillary palp with an internal bifurcate hair, basomedian and basolateral fields of labium each with 1 + 1 chaetae, secondary granulation limited to the abdominal I–IV tergites, s-chaetae s3 present on the abdomen, tenaculum with 3 + 3 teeth, posterior lamellae of the mucro moderately enlarged and finely serrated. Description. General aspect. Habitus and segmentation typical of the genus. Length from labrum to anus: up to 450µm. Specimens whitish to red (in 96% ethanol). All typical chaetal types of the genus are accounted for, without any remarkable development. Integument. The ordinary secondary granulation is limited to the posterior half of the trunk (roughly corresponding to the abdominal region), absent on the head, the thoracic region and Abd. VI (Fig. 1A). Anteriorly on the abdomen, dorso-median line also without secondary granulation until the terminal secondary granule T (same as in Fig. 9A). Primary granulation pattern distinctly visible dorsally on the head and the thorax: primary grains stronger and hexagons surface larger than on the secondary granulated part of the abdomen (same as in Fig. 9A). On the clypeal area, a speckled pattern of hexagons with enlarged primary grain yielding superficial resemblance to ordinary secondary granulation (same as in Fig. 9B). Integumentary channels moderately developed on the head (Fig. 1B, C). On each side of the head, the basal channel splits successively. At each split, the external branch is always terminal (never splitting). The first external branch extending to the antero-dorsal region of the head, then up to four branches extending to the latero-posterior part of the head. Finally, the two most posterior branches reaching the dorso-posterior part of the head. Channels connection with linea ventralis from circular (mostly observed, Fig. 1C) to almost crossed (seen in one specimen, Fig. 1D). Channels absent on trunk. Sensory fields and wax rods. Ordinary distribution of sensory fields and wax rods secretory crypts: 2 + 2 wrc on head, 12 + 12 wrc on body; including the ones associated with the 6 + 6 sensory fields (Fig. 1A, B). Sensory fields include the swollen inner chaetae in the usual distribution (from sf1–6: 0, 1, 3, 2, 2, 1). The swollen inner chaetae are all short and flame-shaped, some with a tendency toward the T-shape (Fig. 1A). Level of separation of wrc5 and 6 from sf5 cannot be quantified with the amount of secondary granules (non applicable). Head chaetotaxy. Postero-dorsal chaetae without remarkable thickening, only subtly stronger than antero-dorsal chaetae (Fig. 1B). Number of chaetae: 12 + 12 in the postero-dorsal region, 8 + 8 and 2 unpaired in the antero-dorsal region, 2 + 2 in the antero-lateral region (Fig. 1B, 8A). Ventrally, 3 + 3 chaetae in the sub-labial region (Fig. 1C, 8B). Diagram of head chaetotaxy provided in Fig. 8A, B. Labium. Basomedian fields of labium with 1 + 1 chaetae, basolateral fields of labium with 1 + 1 chaetae on a small papilla (Fig. 1C). Labial palps ordinary. Labrum. Chaetae a1 and a2 thicker than chaetae m0-2; a1 shorter than a2; m0-2 smooth; a1 and a2 acuminate, with inward curvature, each with one external, basal tooth (Fig. 2A, B). A median, very fine tooth can sometimes be perceived on a2 (Fig. 2B). Labral anterior process apparently as in M. minimus, with a continued transversal crest separating a1-2 from m0-2. Ridge of the labrum with at least two small teeth (Fig. 2B). Other mouthparts. Oral fold with 2 + 2 chaetae (Fig. 1B). Maxilla outer lobe with two chaetae (apical and basal) and with a strong bifurcate hair in subapical internal position (Figs 1B, 2C). Sub-lobal plate without hair, but with a small lobe near the anterior ridge (Fig. 2C). Mandibula with the ordinary asymmetry: one strong basal tooth on the left mandibula (Fig. 2D), missing on the other side. Maxilla with a well developed apical lamella (Fig. 2E). Antenna. Ant. I with one chaeta (Fig. 2F). Ant. II with four chaetae, the anterior one stronger than the three others (Fig. 2F). Ant. III with eight chaetae and five S-chaetae including S1–S4 from the sensory organ and Sb4; S2 and S3 small but clearly protruding from the cupule; without perceptible ornamentation in light microscopy (Fig. 2F, G). Ant. IV with five chaetae (X present) and 11 S-chaetae (Sb1–3, Sb5, Sa1–5, Sx and Sy); Sa2 is notably enlarged in regard to the others common S-chaetae Sa and Sb (as in Fig. 8C); apical and subapical rods (a, sa) present (Fig. 2F, G). Diagram of antennal chaetotaxy provided in Fig. 8C. Thoracic tergites. Th. II with 12 + 12 ordinary chaetae, 1 + 1 s-chaetae s1 and three τ-chaetae (only the bases of τ-chaetae could be perceived using light microscopy); chaetae a5, a6 present and chaeta a7 missing; the laterodorsal τ-chaeta is far from chaeta p4, in the lateral direction (Fig. 1A). Th. III with 10 + 10 ordinary chaetae, 6 + 6 wrc, 4 + 4 τ-chaetae and 2 + 2 pseudopores (ps); a5 sensibly bigger than a6, wrc 2 distant from p4 (Fig. 1A). Diagram of chaetotaxy provided in Fig. 8D. Abd. I–V tergites. With 22 + 22 ordinary chaetae, 2 + 2 globular s-chaetae (s 2, s 3) s3 notably smaller than s2, 2 + 2 τ-chaetae, 1 + 1 pseudopores and 2 + 2 wrc; chaetae ζ1 and η4 present (Fig. 1A). Abd. VI. Tergite with 4 + 4 and 1 unpaired chaetae. Each anal valves with one small chaeta. Sternite with 10 + 10 chaetae (Fig. 2 H). Genital plate. Ordinary: female with 2 + 2 chaetae (Fig. 2 H). Males present, but the genital plate could not be studied in detail. Abd. IV sternites. With 2 + 2 usual neosminthuroid chaetae, usually with 2 + 2 ordinary chaetae and 1 + 1 small lobes but anomalous asymmetry observed in one specimen (Fig. 2 H). Abdominal appendages. Manubrium with 2 + 2 posterior chaetae (Fig. 2 H). Dens ordinary: basal part of dens with 1 + 1 posterior chaetae, apical part of dens with 1 + 1 posterior chaetae and 7 + 7 small spines (Fig. 2 H, I). Mucro elliptical, with moderately wide lamellae, with a dozen of teeth on each posterior lamellae (Fig. 2 J). Ventral tube with 2 + 2 apical chaetae, retinaculum with 3 + 3 teeth. Legs. Chaetal composition on each legs subcoxa 1, 2, coxa, trochanter, femur and tibiotarsus: Leg I, 1, 0, 1, 2, 8, 12 chaeta(e) (Fig. 3A); leg II, 1, 1, 1, 3, 8, 12 chaeta(e) (Fig. 3B); leg III, 2, 1, 1, 4, 8, 10 chaeta(e) (Fig. 3C). Claws ordinary, as in Fig. 3A, C., with short external basal lamellae. Differential diagnosis. Megalothorax anterolenis sp. nov. shares some uncommon chaetotaxic similarities with M. granulosus: only 1 + 1 chaetae on the basomedian field of labium, and presence of chaetae a5 and a6 on Th. II and chaetae ζ1 and η4 on the abdomen. However M. anterolenis sp. nov. and M. granulosus are clearly distinct on many aspects, such as the shape of the connection of the head integumentary channels (circular vs crossed), the secondary granulation (ordinary vs coarse), the antennal chaetotaxy and the mucro lamellae (toothed vs smooth). In general, the repartition of the secondary granulation clearly distinguish Megalothorax anterolenis sp. nov. from any species of the incertus and minimus -group (absent on head and anterior part of trunk vs full dorsal coverage). Name etymology. Combination of the Latin anterior and lçnis (smooth)., Published as part of Schneider, Clément, Minor, Maria A. & D'Haese, Cyrille A., 2023, A new group of species of the genus Megalothorax (Collembola, Neelidae) with Gondwanan distribution, and introducing an open interactive identification key of Megalothorax species, pp. 101-121 in Zootaxa 5228 (2) on pages 104-107, DOI: 10.11646/zootaxa.5228.2.1, http://zenodo.org/record/7532173

Megalothorax rubidus Salmon, 1946 Material examined. A single slide labeled as “Dominion Museum. N.Z. 536. Slide 3/1823, Deep red alive, Paratype. Gertrude Cirque, Homer in moss and lichen on forest floor. Coll. J. T. Salmon. 28/12/1944. Mnt KOH & P. V. A. Det. J. T. Salmon 1946 ”. New elements of description. Fine secondary granulation present dorsally on the head, thorax and abdomen. Labrum a1 chaetae acuminate. Basomedian field of labium with 1 + 1 chaetae. Ventrally on head, with 1 + 1 external post-labial chaetae much larger than the 2 + 2 median post-labial chaetae. Differential diagnosis. Megalothorax rubidus is similar to the three new species and to M. granulosus from the reduced chaetotaxy of the basomedian field of labium. The presence of the secondary granulation on the head and thorax clearly sets it apart from the three new species group. It can be distinguished from M. granulosus by its fine granulation (coarse in M. granulosus). Furthermore, its external post-labial chaetotaxy is so far unique among Megalothorax species. Comments. The specimen poor preservation prevents a complete chaetotaxic analysis, redescription from new material would be desirable. Due to the slide preservation, it was not possible to observe the whole extension of the secondary granulation but could confirm that it is at least partially present dorsally on the head, the thorax and the abdomen., Published as part of Schneider, Clément, Minor, Maria A. & D'Haese, Cyrille A., 2023, A new group of species of the genus Megalothorax (Collembola, Neelidae) with Gondwanan distribution, and introducing an open interactive identification key of Megalothorax species, pp. 101-121 in Zootaxa 5228 (2) on page 116, DOI: 10.11646/zootaxa.5228.2.1, http://zenodo.org/record/7532173, {"references":["Salmon, J. T. (1946) Collembola - Symphypleona from the Homer district. Dominion Museum Records in Entomology, 1 (4), 27 - 61."]}

Megalothorax tasmanterolenis sp. nov. Figs 4, 5, 8A, B, 9A–D Material examined. Holotype. Female on slide (CSCOL _98), Australia, Tasmania, Ida Bay, 26.iii.2015, 43.5070°S, 146.8755°E, mosses on the ground and rotten branches, sample number TAS001d. Paratype. Female on slide (CSCOL _99), same data as the holotype. All material collected by C. D’Haese. Material deposit. The holotype will be deposited at the Senckenberg Museum für Naturkunde, Görlitz, Germany. Paratype will be deposited at the Muséum National d’Histoire Naturelle, Paris, France. Obtained molecular data. One specimen from TAS001d (same as holotype), 28S rDNA: OP942370 (806bp), 16S rDNA: OP942367 (468bp). Voucher could not be recovered. Diagnosis. Yellow-orange in ethanol, labrum a1 chaetae with bifurcate tip, maxillary palp with an internal bifurcate hair, basomedian and basolateral fields of labium each with 1 + 1 chaetae, secondary granulation limited to the abdominal I – IV tergites, s-chaeta s2 globular, s3 absent on the abdomen, tenaculum with 4 + 4 teeth, posterior lamellae of the mucro moderately enlarged, internal lamella serrated, external lamella irregular. Description. General aspect. Habitus and segmentation typical of the genus. Length from labrum to anus: up to 500µm. Specimens whitish (in 96% ethanol). All typical chaetal types of the genus are accounted for, without any remarkable development. Integument. Distribution of the secondary granulation identical to M. anterolenis sp. nov. (Fig. 4A); the speckle of enlarged primary granules on the clypeal area is present but subtle on our available specimens. Integumentary channels moderately developed on the head (Fig. 4A, B). On each side of the head, the basal channel splits nine times successively. First the two external branches extend to the antero-dorsal region of the head, then up to five branches extend to the latero-posterior part of the head, finally the five most posterior branches reaching the dorsoposterior part of the head. Head channels connection with linea ventralis circular. Channels absent on trunk. Sensory fields and wax rods. Same observation as in M. anterolenis sp. nov. (also see Fig. 4A, B). Head chaetotaxy. Postero-dorsal chaetae without remarkable thickening, only subtly stronger than antero-dorsal chaetae. Number of chaetae: 12 + 12 in the postero-dorsal region, 9 + 9 and 2 unpaired in the antero-dorsal region, 2 + 2 in the antero-lateral region (Fig. 4B). Ventrally, 3 + 3 chaetae in the sub-labial region. Diagram of head chaetotaxy provided in Fig. 8A, B. Labium. Basomedian fields of labium with 1 + 1 chaetae, basolateral fields of labium with 1 + 1 chaetae on a small papilla. Labial palps ordinary. Labrum. Chaetae a1 and a2 thicker than chaetae m0-2; a1 shorter than a2; m0-2 smooth (Fig. 4D); apex of a1 with bifurcate tip (Fig. 4C); apex of a2 acuminate, with inward curvature, subtly serrated (Fig. 4C). Organisation of the labral anterior process not studied in detail. Two small teeth near the ridge of the labrum (Fig. 4C). Other mouthparts. Oral fold with 2 + 2 chaetae (Fig. 4E). Maxilla outer lobe with two chaetae (apical and basal) and with a strong bifurcate hair in subapical internal position (Fig. 4E). Sub-lobal plate without hair, with two lobes on the anterior ridge (Fig. 4E). Mandibula ordinary (Fig. 4F). Maxilla with a well developed apical lamella (Fig. 4G). Antenna. As in Fig. 5A, same observations as M. anterolenis sp. nov., but with 9 to 10 normal chaetae on Ant. III (either 4 or 5 chaetae in the apical whorl) and Sb6 above S2, S3. Sa2 with a swollen aspect in comparison to the other Sa and Sb chaetae. Thoracic and abdominal tergites. As in Fig. 4A, same observations as in M. anterolenis sp. nov., but with schaetae s3 absent. Pseudopores not studied. Abd. VI. Ordinary, with nine chaetae on tergites, a small chaetae on each anal valves, and with 10 + 10 chaetae on sternites. Genital plate. Female with 2 + 2 chaetae, male not seen. Abd. IV sternites. With 2 + 2 usual neosminthuroid chaetae and 2 + 2 ordinary chaetae, the internal chaetae longer than the external ones. Abdominal appendages. Manubrium with 2 + 2 posterior chaetae (Fig. 5E). Dens ordinary (Fig. 5E): basal part of dens with 1 + 1 posterior chaetae, apical part of dens with 1 + 1 posterior chaetae and 7 + 7 small spines; basal spines with blunt apex, apical spines with filamentous apex. Mucro elliptical, with moderately wide lamellae; internal posterior lamella with around seven teeth (but could not precisely studied), external lamella without hardened teeth, but with an irregular aspect (Fig. 5E). Ventral tube with 2 + 2 apical chaetae, retinaculum with 4 + 4 teeth. Legs. Chaetal composition on legs (subcoxa 1, 2, coxa, trochanter, femur and tibiotarsus): leg I, 1, 0, 1, 2, 8, 12 chaeta(e); leg II, 1, 1, 1, 3, 8, 12 chaeta(e); leg III, 2, 1, 1, 4, 8, 10 chaeta(e). Claws ordinary, as in Fig. 5B–D; with external basal lamellae short. Differential diagnosis. Very similar to M. anterolenis sp. nov., but differs from the latter by the shape of the labral chaetae a1, the morphology of posterior lamellae of the mucro, the number of teeth on the retinaculum, and dorsally on the forehead by the additional pair of chaetae and the additional integumentary channel branch. After much scrutinization, we concluded on the absence of s3 on the abdomen (present in M. anterolenis sp. nov.). Name etymology. Derived from the name of M. anterolenis sp. nov. combined with the region of discovery (Tasmania)., Published as part of Schneider, Clément, Minor, Maria A. & D'Haese, Cyrille A., 2023, A new group of species of the genus Megalothorax (Collembola, Neelidae) with Gondwanan distribution, and introducing an open interactive identification key of Megalothorax species, pp. 101-121 in Zootaxa 5228 (2) on pages 108-110, DOI: 10.11646/zootaxa.5228.2.1, http://zenodo.org/record/7532173

International audience; Longtemps considérés comme des Insectes et placés avec les Protoures, les Diploures et les ex Thysanoures au sein d’un groupe artificiel (paraphylétique) appelé « Aptérygotes », les Collemboles constituent aujourd’hui une classe au même titre que les Insectes. Ce sont les plus anciens Hexapodes connus. Ils occupent, avec les Acariens, une place prépondérante au sein des peuplements édaphiques, mais également dans de nombreux milieux terrestres, hypogés et épigés, du niveau du sol à celui de la canopée. Géographiquement, ils sont présents sur tous les continents, sous toutes les latitudes et à toutes les altitudes.

The author regrets at the beginning of the sub-section "Sex pheromones" (page 428 of the published version) within the section "The use of pheromones, allomones and kairomones by Collembola"

International audience; The present knowledge on chemical communication in springtails (Collembola), one of the two most abundant invertebrate groups living in soil and environments in tight contact with soil (e.g. plant litter, moss), is reviewed here. Chemical communication in an environment where light is absent or dimmed becomes a prominent driver of trophic and non-trophic interactions between soil organisms at a time when better knowledge on the biological determinants of soil communities is required. Like insects and many other arthropods, collembolan individuals of the same population intercommunicate by pheromones, which allow them signalling a risk or clustering in places favourable for feeding, mating, moulting and ovipositing. Olfaction is also used to select preferred food and mates. Researches so far conducted allowed discerning common trends in the role and chemical composition of odour blends used by Collembola. However, much more needs to be done before reaching straightforward conclusions about chemical communication issues at evolutionary and community levels, making this domain even more rewarding.

International audience; Natural recolonization of disturbed environments is a key issue for the maintenance of biodiversity, especially in a context of habitats threatened by exotic species. Recolonization by collembola of habitats destroyed by lava flows has never been documented. These soil-dwelling arthropods provide an essential basis for the functioning of terrestrial ecosystems. Our objective was to answer the following questions: Which habitats (forests, crops) serve as sources for collembola recolonization? Do exotic plant species influence this recolonization? Which traits allow rapid recolonization? Our study was performed in the Reunion island, where several habitats have been destroyed by lava flows and where 3500 introduced plant species threaten the functioning of ecosystems. Collembola were collected from soil, litter, herbaceous layer, moss and tree bark sampled from different habitats (lava flows, forests and sugarcane fields). They were identified at the species or morphospecies level. Their functional traits were measured. For the first time collembolan species were inventoried in forests, crops and lava flows on Reunion Island: 56 species and morpho-species belonging to 12 families were observed. Species richness was increased in crops and forests compared to lava flows indicating that recolonization was incomplete. Our results show that the recolonization of habitats destroyed by lava flows depends on many factors that differ according to the considered micro-habitat (vertical stratum). Colonization was a function of the percentage of exotic plants, the distance to the nearest forest and the age of lava flow, in herbaceous layer, soil and litter, respectively. Dominant functional traits observed in lava flows were those belonging to edaphic species. The rapidity with which edaphic species colonized lava flows, added to the observation of many of these species on tree trunks, suggests that they could have benefited from air transport for settlement in lava flows.

International audience; I-Principe de l'analyse à trois éléments. L'analyse à trois éléments ("three-taxon analysis" ou "three-item analysis" en anglais, abrégée 3ia dans ce qui suit) est une méthode phylogénétique qui s'applique aux taxons et aux aires biogéographiques. Elle se justifie par l'interprétation suivante des principes de base de la théorie cladistique :  La 3ia est conçue comme une méthode d'argumentation des relations de parenté entre taxons. Elle permet de contrôler de façon pertinente les idées de parenté émises par un systématicien et s'exempte de celles qui seraient seulement proposées par l'application d'un algorithme.  Les caractères sont considérés comme des arguments en faveur d'une hypothèse phylogénétique. Ils ne sont pas de simples données ou de simples observations : ce sont des hypothèses d'homologie formellement structurées, c'est-à-dire des hypothèses sur l'identité due à la parenté.  Le but de l'analyse 3ia est de contrôler si l'hypothèse phylogénétique proposée par le systématicien peut être argumentée à partir des hypothèses d'homologie explicitées sur chacun des caractères. L'analyse à trois éléments représente donc une forme originale de penser et de pratiquer la phylogénétique et la biogéographie. Les caractères sont considérés comme des arguments phylogénétiques (Hennig 1966 : 90) que le systématicien fournit pour défendre une hypothèse phylogénétique. Les caractères dérivent directement d'hypothèses d'homologie, la notion d'homologie utilisée ici se rapprochant de celle donnée par Richard Owen (1843 : 373). Ce dernier ne définit pas l'homologie mais un homologue. Un homologue constitue un même organe chez différents organismes, sous toute la diversité de ses formes et de ses fonctions. Peut-on dire que le bras chez l'humain est la même chose que la patte antérieure d'un chien ? Il est difficile de répondre par l'affirmative tout autant que par la négative. En revanche, dire que ces structures sont en même temps la même tout en étant différentes est plus satisfaisant. L'assertion prend du sens lorsqu'on affirme que le bras humain ressemble davantage à une patte avant de chien qu'à une aile de pigeon, c'est-à-dire lorsque l'on introduit un troisième terme. Les homologues représentent donc des classes d'équivalence de parties d'organismes qui établissent des relations d'identité relative à d'autres classes d'équivalence (Prin 2016 : 432). Ainsi, l'ensemble « bras humain » et l'ensemble « pattes antérieures de chien » peuvent être réunis dans une classe d'équivalence plus inclusive, un nouvel homologue qui exclut l'ensemble « aile de pigeon ». Les trois membres (de l'homme, du chien et du pigeon) sont à leur tour réunis dans une nouvelle classe de degré d'équivalence, celle des membres antérieurs de tétrapodes (figure 1). FIGURE 1. Hypothèse d'homologie sous forme de classes de degré d'équivalence du membre antérieur de tétrapodes.

International audience; Les arbres évolutifs, ou phylogénies, racontent une histoire, l’histoire des êtres vivants, de leur morphologie et de leurs gènes, mais aussi, l’histoire des langues, des textes, des faits culturels ou même des idées. Ces arbres sont avant tout des hypothèses sur les liens de parentés qui exigent réflexions à la fois sur les faits et sur les méthodes. Ce livre définit les concepts fondamentaux de la reconstruction phylogénétique. Il explique la nature et la diversité des approches pratiquées depuis leurs balbutiements au XIXe siècle jusqu’à nos jours. Il insiste, de manière pédagogique et aussi objectivement que possible, sur leurs performances et leurs limites, en fonction de la nature des données étudiées.Cet ouvrage constitue une version largement augmentée de La Reconstruction phylogénétique. Concepts et méthodes publié en 1993 dans la collection « Biologique théorique » des éditions Masson. Il tient donc compte des méthodes et débats qui ont marqué l’évolution rapide de la discipline ces vingt-cinq dernières années.Il s’adresse aux étudiants des 1er, 2e et 3e cycles et aux chercheurs non spécialistes de phylogénie mais désireux de connaître l’état actuel de la question.

Additional file 1: Description of Electrosminthuridia helibionta sp. nov.

International audience

D'Haese, Cyrille A., Stevens, Mark I., Weiner, Wanda M. (2017): Friesea najtae n. sp. (Collembola, Neanuridae, Frieseinae) from southern Western Australia. Zoosystema 39 (1): 21-29, DOI: 10.5252/z2017n1a3, URL: http://dx.doi.org/10.5252/z2017n1a3

Genus Friesea Dalla Torre, 1895 Friesea Dalla Torre, 1895: 14. Polyacanthella Schäffer, 1897: 15. TYPE SPECIES. — Friesea mirabilis (Tullberg, 1871)., Published as part of D'Haese, Cyrille A., Stevens, Mark I. & Weiner, Wanda M., 2017, Friesea najtae n. sp. (Collembola, Neanuridae, Frieseinae) from southern Western Australia, pp. 21-29 in Zoosystema 39 (1) on page 22, DOI: 10.5252/z2017n1a3, http://zenodo.org/record/4578463

Zon, Serge Déméango, Bedos, Anne, D'Haese, Cyrille A. (2015): Phylogeny of the genus Willemia (Collembola: Hypogastruridae) and biogeography of the W. buddenbrocki - group with description of a new species from Ivory Coast (western Africa). Zootaxa 3980 (2): 230-240, DOI: http://dx.doi.org/10.11646/zootaxa.3980.2.4

Willemia tondoh sp. nov. Figs 1���6 Type material. Holotype female (IV-VJW 2 L 1, LZBA-UFHB) and four paratypes females (VC02P 2, LZBA- UFHB; VC01S 7, MNHN-EA 012013; VJ 12 L 1, MNHN-EA 012014 and VC01L 8, MNHN-EA 012015). Type locality and sampling dates. Ivory Coast, Taabo, Zougoussi terroir near Lamto reserve. Holotype (sample VJW 2 L 1): 29 October 2012, litter in old fallow, 06�� 15 ��� 28 ���N and 05��01��� 42.1 ���W. Two paratypes (sample VC01L 8 and VC01S 7): 13 November 2012, in cocoa culture, 06�� 15 ��� 43.6 ���N and 05��01��� 29.8 ���W. One paratype (sample VJ 12 L 1): 27 October 2012, in mixed cultures, 06�� 15 ��� 25.9 ���N and 05��01��� 36 ���W. One paratype (sample VC02P 2): 15 November 2012, in cocoa culture, 06�� 15 ��� 30 ���N and 05��01��� 38.7 ���W. Description. Body length: holotype 400 ��m, paratypes 400���540 ��m. Color in alcohol white, body with short acuminate ordinary chaetae, some slightly longer. Sensory chaetae a little thicker and longer than ordinary chaetae. Tegumental granulation fine and regular. Antennae somewhat shorter than head���s diagonal. Ant. I and Ant. II with 7 and 11 chaetae respectively. Ant. III and IV only hardly separated. Sensorial organ of Ant. III consisting of a large integumentary fold hiding 2 internal s -microchaetae, 2 long guard chaetae, 2 long sensory rods, and one ventral microsensillum (Fig. 4). Ant. IV with a small globular apical vesicle, s-chaetae comprising: S 2 /d and S 9 /e 1 absent or not differentiated from ordinary chaetae; S 1 / i 1 and S 8 /e 2 subcylindrical or hardly differentiated from ordinary chaetae; S 4 / i 2 and S 7 /e 3 globular in cavity covered by a tegumental fold. External part of the cavity with microsensillum. Subapical organite present. Postantennal organ with 9 vesicles (Figs 1 & 3), no eyes. Chaetae a0 and c 1 absent on head (Fig. 1). Prelabrum/labrum with 2 /5,3,4 chaetae. Three pairs of labial chaetae. Head ventral chaetotaxy as in Fig. 2. Dorsal body chaetotaxy presented in Fig. 1. Some variation and asymmetries present. Chaetae s per half tergum formula: 22 / 11111 generally, in m 7 and p 4 position on thorax II and III terga, in p 4 position on abdominal terga I to IV and p 2 position on Abd. V i.e. ordinary chaeta p 2 absent. Thorax II and III with reduced chaetotaxy: m 3 likely to be missing (Fig. 1). Abd. II and III with a 2, but without m-row. Abdomen IV without m 1, m 2, m 3, or m��� 3 or p 5. Tibiotarsal chaetotaxy I, II and III with 11 chaetae each. Unguis with a small empodial appendage and without teeth (Fig. 6). Ventral tube with 4 + 4 chaetae. Ventral abdominal chaetotaxy as in Fig. 5. Furcula absent. Female genital plate with 2 chaetae. Anal valves with 2 hr chaetae and without e and z chaetae (Fig. 5). Two anal spines well developed. Derivatio nominis. The new species is cordially dedicated to J��r��me E. Tondoh, Professor of soil ecology at Nangui Abrogoua University (Abidjan, Ivory Coast), who is the initiator of Collembola studies within the research unit on soil fauna. The noun is in apposition (article 31.1 of the International Code of Zoological Nomenclature). Distribution. It has to be noted that W. brevispina belongs to the buddenbrocki -group (D'Haese & Weiner 1998) and has been originally described from South America but it has been mentioned from Gambia (Murphy 1965). At that time two other species only were known in the group: W. nadchatrami Yosii, 1959 (Continental South-East Asian and Malaysian regions) and W. buddenbrocki H��ther, 1959 (widely distributed). The specimen collected by Murphy has possibly been identified as W. brevispina because its general morphology was compatible with its description, at that time. Since Gambia and Ivory Coast are located in western Africa, it is highly probable the species mentioned by Murphy is actually a new undescribed species closely related to if not W. tondoh sp. nov., Published as part of Zon, Serge D��m��ango, Bedos, Anne & D'Haese, Cyrille A., 2015, Phylogeny of the genus Willemia (Collembola: Hypogastruridae) and biogeography of the W. buddenbrocki - group with description of a new species from Ivory Coast (western Africa), pp. 230-240 in Zootaxa 3980 (2) on pages 231-233, DOI: 10.11646/zootaxa.3980.2.4, http://zenodo.org/record/240011, {"references":["D'Haese, C. A. & Weiner, W. M. (1998) A review of Willemia buddenbrocki - group (Collembola, Poduromorpha, Hypogastruridae) with cladistic analysis. Journal of Natural History, 32 (7), 969 - 986. http: // dx. doi. org / 10.1080 / 00222939800770501","Murphy, D. H. (1965) Collembola Poduromorpha from the Gambia (West Africa). Journal of Zoology, 146, 388 - 411. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.1965. tb 05216. x","Yosii, R. (1959) Studies on the Collembolan fauna of Malay and Singapore. With special reference to the genera: Lobella, Lepidocyrtus and Callyntrura. Contributions from the Biological Laboratory, Kyoto University, 10, 1 - 65.","Huther, W. (1959) Willemia buddenbrocki n. sp., und zur chaetotaxie von W. persimilis Bonet, 1945 (Collembola). Senckenbergiana biologica, 40, 173 - 177."]}

Willemia tondoh sp. nov. Figs 1–6 Type material. Holotype female (IV-VJW 2 L 1, LZBA-UFHB) and four paratypes females (VC02P 2, LZBA- UFHB; VC01S 7, MNHN-EA 012013; VJ 12 L 1, MNHN-EA 012014 and VC01L 8, MNHN-EA 012015). Type locality and sampling dates. Ivory Coast, Taabo, Zougoussi terroir near Lamto reserve. Holotype (sample VJW 2 L 1): 29 October 2012, litter in old fallow, 06° 15 ’ 28 ”N and 05°01’ 42.1 ”W. Two paratypes (sample VC01L 8 and VC01S 7): 13 November 2012, in cocoa culture, 06° 15 ’ 43.6 ”N and 05°01’ 29.8 ”W. One paratype (sample VJ 12 L 1): 27 October 2012, in mixed cultures, 06° 15 ’ 25.9 ”N and 05°01’ 36 ”W. One paratype (sample VC02P 2): 15 November 2012, in cocoa culture, 06° 15 ’ 30 ”N and 05°01’ 38.7 ”W. Description. Body length: holotype 400 µm, paratypes 400–540 µm. Color in alcohol white, body with short acuminate ordinary chaetae, some slightly longer. Sensory chaetae a little thicker and longer than ordinary chaetae. Tegumental granulation fine and regular. Antennae somewhat shorter than head’s diagonal. Ant. I and Ant. II with 7 and 11 chaetae respectively. Ant. III and IV only hardly separated. Sensorial organ of Ant. III consisting of a large integumentary fold hiding 2 internal s -microchaetae, 2 long guard chaetae, 2 long sensory rods, and one ventral microsensillum (Fig. 4). Ant. IV with a small globular apical vesicle, s-chaetae comprising: S 2 /d and S 9 /e 1 absent or not differentiated from ordinary chaetae; S 1 / i 1 and S 8 /e 2 subcylindrical or hardly differentiated from ordinary chaetae; S 4 / i 2 and S 7 /e 3 globular in cavity covered by a tegumental fold. External part of the cavity with microsensillum. Subapical organite present. Postantennal organ with 9 vesicles (Figs 1 & 3), no eyes. Chaetae a0 and c 1 absent on head (Fig. 1). Prelabrum/labrum with 2 /5,3,4 chaetae. Three pairs of labial chaetae. Head ventral chaetotaxy as in Fig. 2. Dorsal body chaetotaxy presented in Fig. 1. Some variation and asymmetries present. Chaetae s per half tergum formula: 22 / 11111 generally, in m 7 and p 4 position on thorax II and III terga, in p 4 position on abdominal terga I to IV and p 2 position on Abd. V i.e. ordinary chaeta p 2 absent. Thorax II and III with reduced chaetotaxy: m 3 likely to be missing (Fig. 1). Abd. II and III with a 2, but without m-row. Abdomen IV without m 1, m 2, m 3, or m’ 3 or p 5. Tibiotarsal chaetotaxy I, II and III with 11 chaetae each. Unguis with a small empodial appendage and without teeth (Fig. 6). Ventral tube with 4 + 4 chaetae. Ventral abdominal chaetotaxy as in Fig. 5. Furcula absent. Female genital plate with 2 chaetae. Anal valves with 2 hr chaetae and without e and z chaetae (Fig. 5). Two anal spines well developed. Derivatio nominis. The new species is cordially dedicated to Jérôme E. Tondoh, Professor of soil ecology at Nangui Abrogoua University (Abidjan, Ivory Coast), who is the initiator of Collembola studies within the research unit on soil fauna. The noun is in apposition (article 31.1 of the International Code of Zoological Nomenclature). Distribution. It has to be noted that W. brevispina belongs to the buddenbrocki -group (D'Haese & Weiner 1998) and has been originally described from South America but it has been mentioned from Gambia (Murphy 1965). At that time two other species only were known in the group: W. nadchatrami Yosii, 1959 (Continental South-East Asian and Malaysian regions) and W. buddenbrocki Hüther, 1959 (widely distributed). The specimen collected by Murphy has possibly been identified as W. brevispina because its general morphology was compatible with its description, at that time. Since Gambia and Ivory Coast are located in western Africa, it is highly probable the species mentioned by Murphy is actually a new undescribed species closely related to if not W. tondoh sp. nov.

Antarctica contains some of the most challenging environmental conditions on the planet due to freezing temperatures, prolonged winters and lack of liquid water. Whereas 99.7% of Antarctica is permanently covered by ice and snow, some coastal areas and mountain ridges have remained ice-free and are able to sustain populations of microinvertebrates. Tardigrades are one of the more dominant groups of microfauna in soil and limno-terrestrial habitats, but little is known of their diversity and distribution across Antarctica. Here, we examine tardigrades sampled from across an extensive region of continental Antarctica, and analyse and compare their partial mitochondrial cytochrome c oxidase subunit 1 (COI) gene sequences with those from the Antarctic Peninsula, maritime and sub-Antarctica, Tierra del Fuego and other worldwide locations in order to recognise operational taxonomic units (OTUs). From 439 new tardigrade COI sequences, we identified 98 unique haplotypes (85 from Antarctica) belonging to Acutuncus, Diphascon, Echiniscus, Macrobiotus, Milnesium and unidentified Parachela. Operational taxonomic units were delimited by Poisson tree processes and general mixed Yule coalescent methods, resulting in 58 and 55 putative species, respectively. Most tardigrades appear to be locally endemic (i.e. restricted to a single geographic region), but some (e.g. Acutuncus antarcticus (Richters, 1904)) are widespread across continental Antarctica. Our molecular results reveal: (i) greater diversity than has previously been appreciated with distinct OTUs that potentially represent undescribed species, and (ii) a lack of connectivity between most OTUs from continental Antarctica and those from other Antarctic geographical zones.

International audience; Amber is nearly ubiquitous in lignites from Otago and Southland and cornmon throughout New Zealand; however, no amber inclusions have been reported previously. We studied amber from 22 Cretaceous to Miocene sites in southem New Zealand, recovering inclusions at three localities: Cosy Dell (late Oligocene), Roxburgh (early Miocene), and Hyde (?early Miocene). Preparation of New Zealand amber to expose inclusions for study under incident and transmitted light is challenging and time-consuming, with most samples brittle and opaque. Thus, we stabilize and clear the amber lumps using epoxy preparation under vacuum before grinding and imaging under light microscopy. To date we have recovered 63 arthropods, as weIl as plant remains, fungi, and nematodes. Arachnids include diverse mites (Mesostigmata, Oribatida, Astigmata, and Prostigmata), a variety of spiders and web remains with prey, and a pseudoscorpion. Sorne Collembola were identified as belonging to the family Entomobryidae (Entomobryomorpha). Insects include members of the families Dermestidae (Coleoptera), Mymaridae and Scelionidae (Hymenoptera), Veliidae (Heteroptera), Ceratopogonidae (Forcipomyia) and Mycetophilidae (Diptera), as well as Psocoptera, and Lepidoptera. The most abundant fungi in New Zealand amber are hyphomycetes similar to the genus Casparyotorula from European Palaeogene ambers and we discovered similar fungi growing on resin of the extant Agathis australis, the iconic New Zealand Kauri. Furthermore, specimens of the genus Metacapnodium (Capnodiales) represent the first Southem Hemisphere fossil sooty moulds; saprophytic ascomycetes with brown hyphae, often forming extensive subicula on living plant surfaces. These fungi are ubiquitous and diverse in New Zealand today. Many of these new amber fossils represent groups with an otherwise poor fossil record for the entire Southem Hemisphere. The systematic and ecological diversity of the inclusions highlights the potential of New Zealand amber for reconstructing past terres trial ecosystems of Zealandia, one of the biogeographically crucial former Gondwanan landmasses.

International audience; The adequacy and utility of behavioural characters in phylogenetics is widely acknowledged, especially for stereotyped behaviours. However, the most common behaviours are not stereotyped, and these are usually seen as inappropriate or more difficult to analyze in a phylogenetic context. A few methods have been proposed to deal with such data, although they have never been tested on samples larger than six species, which limits their evolutionary interest. In the present study, we perform behavioural observations on 13 cockroach species and derive behavioural phylogenetic characters with the successive event-pairing method. We combine these characters with morphological and molecular data (approximately 6800 bp) in a phylogenetic study of 41 species. We then reconstruct ancestral states of the behavioural data to study evolution of social behaviour in these insects with regard to their social systems (i.e. solitary, gregarious, and subsocial) and diversity of habitat choice. We report for the first time that nonstereotyped behavioural data are adequate for phylogenetic analyses: they are no more homoplastic than traditional data, and support several phylogenetic relationships that we discuss. From an evolutionary perspective, we show that the solitary species Thanatophyllum akinetum does not display original behavioural interactions, suggesting phylogenetic inertia of interactive behaviours despite a radical change in social structure. Conversely, the subsocial species Parasphaeria boleiriana shows original behavioural interactions, which could result from its peculiar social system or habitat. We conclude that phylogenetic approaches in studies of behaviour are useful for deciphering evolution of behaviour and discriminating between its different modalities, even for nonstereotyped characters.(c) 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111, 58-77.

Willemia zeppelini sp. nov. Figs 1���5 Type material. Holotype (female) and 2 paratypes (females). Type locality. Brazil, Joao Pessoa, Beach of Tambau, central region, Joao Pessoa, PB. 17.vi. 2010. Other localities. Beach of Camboinha 3, Cabedelo, PB. 19.vi. 2010; 3 specimens, 1 male, 2 females (MNHN- EA010008 to MNHN-EA010010). Description. Body length: holotype 300 ��m (MNHN-EA010005), paratypes 270���330 ��m (MNHN-EA010006 to MNHN-EA010007). Color in alcohol white, body with short acuminate ordinary chaetae, some slightly longer. Sensory chaetae a little thicker and longer than ordinary chaetae. Tegumental granulation fine and regular. Antennae somewhat shorter than head���s diagonal (40 ��m; and 50 ��m respectively). Ant. I and Ant. II with 7 and 11 chaetae respectively (Fig. 2). Ant. III and IV only hardly separated. Sensorial organ of Ant. III consisting of a large integumentary fold hiding 2 internal s-microchatae, 2 long guard chaetae, 2 long sensory rods with a dark axis and lighter expansions on both sides, and one ventral microsensillum (Fig. 1). Ant. IV with a large globular apical bulb, microsensillum and subapical organite present. Ant. IV s-chaeta comprising of S 2 / d and S 9 / e 1 absent or not differentiated form ordinary chaetae, S 1 / i 1 and S 8 / e 2 subcylindrical or hardly differentiated from ordinary chaetae, and S 4 / i 2 and S 7 / e 3 enlarged with the shape of a slightly flattened pear. Some variation is observed with S 1, S 8, S 4 and S 7 being in a continuum from subcylindrical hardly differentiated from ordinary chaetae to flattened pear-shaped (Figs 1���2). Postantennal organ with 4 to 5 tubercles (Fig. 3), no eyes. Chaetae a 0 and c 1 absent on the head (Fig. 3). Prelabrum/labrum with 2 /2,5,4 chaetae. Three pairs of labial chaetae. Dorsal body chaetotaxy presented in Fig. 3. The specimens observed present some variation and asymmetries. Chaetae s per half tergum formula: 22 / 11111 generally, in m 7 and p 4 position on thorax II and III terga, in p 4 position on abdominal terga I to IV and p 2 position on Abd. V i.e. chaeta p 2 absent; some specimens have p 2 present (i.e. chaeta s in p 3 position). Thorax II and III with reduced chaetotaxy: a 1, a 2 or m 3 seem to be missing (Fig. 3). Abd. II and III with a 2, but without m -row. Abdomen IV without m 1, m 2, m 3, or m��� 3 or p 5. No anal spine. Ventral tube with 4 + 4 chaetae. Ventral abdominal chaetotaxy as in Fig. 4. Furcula absent. Female genital plate with 2 eugenital chaetae; male genital plate with 8 eugenital chaetae. Anal valves with one hr chaeta and without e and z chaetae (Fig. 4). Tibiotarsal chaetotaxy I, II and III with 12 chaetae each. Unguis with a small empodial appendage and without teeth (Fig. 5). Derivatio nominis. The new species is cordially dedicated to the brazilian collembologist Professor Douglas Zeppelini from Universidade Estadual da Paraiba in Joao Pessoa, who participated to the collection of this new species. Distribution. Phylogenetically, W. zeppelini fits among central american/neotropical species (W. subbulbosa, persimilis -group and W. brevispina) which makes a lot of sense biogeographically. The only other Willemia species recorded from Brazil is W. brevispina H��ther, 1962, found in littoral sand near Rio coast (Thibaud & Palacios-Vargas 1999) and it is one of the closest to the new W. zeppelini on the phylogeny (Fig. 6)., Published as part of D'Haese, Cyrille A. & Thibaud, Jean-Marc, 2011, Description and phylogenetic position of a new Willemia species (Collembola: Hypogastruridae) from the littoral coast of Brazil, pp. 33-40 in Zootaxa 2932 on page 34, DOI: 10.5281/zenodo.202784, {"references":["Huther, W. (1962) Beitrag zur gattung Willemia Borner (Collembola). Beitrage zur Entomologie, 12, 511 - 526.","Thibaud, J. - M. & Palacios-Vargas, J. G. (1999) Brazilian Collembola from littoral sand with descriptions of Austrogastrura gen. n. and Isotomodes carioca sp. n. Revue francaise d'Entomologie, 21, 25 - 31."]}

D'Haese, Cyrille A., Thibaud, Jean-Marc (2011): Description and phylogenetic position of a new Willemia species (Collembola: Hypogastruridae) from the littoral coast of Brazil. Zootaxa 2932: 33-40, DOI: 10.5281/zenodo.202784