Collembola (Entognatha) from East Africa

Home , Brachystomella parvula, Entognatha, Friesea, Frieseinae, Hypogastruridae, Neanuridae

Eur. J. Entomol. 95: 217-237, 1998 ISSN 1210-5759

W anda M. WEINER1 and Judith NAJT2

1 Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Slawkowska 17, PL-31016 Krakdw, Poland 2EP 90 du CNRS, Laboratoire d’Entomologie, Muséum National d’Histoire Naturelle, 45, rue Buffon, F-75005 Paris, France

Collembola, Hypogastruridae, Odontellidae, Neanuridae, Onychiuridae, Isotomidae, East Africa, identification keys, new species, redescription

Abstract. Materials of Hypogastruridae, Odontellidae, Neanuridae, Onychiuridae and Isotomidae from East Africa were studied. Several new species are described: Acherontiella kowalskiorum sp. n., Furcu- lanurida grandcolasorum sp. n., Stachorutes dallaii sp. n., and Paleonura cassagnaui sp. n. Friesea vtorovi Tshelnokov, 1977 and Tullbergia kilimanjarica (Delamare Deboutteville, 1953) are redescribed. Stachorutes arlei (Thibaud & Massoud, 1980) is a new combination. Identification keys for Friesea Dalla Torre, 1895 with 2 + 2 eyes and Stachorutes Dallai, 1973 are given.

INTRODUCTION The last systematic account on the Collembola of East Africa was presented by Dehar- veng and Diaz in 1984 with a review of all references concerning this region. The present study is based on the material collected in Tanzania and Kenya.

Abbreviations . ISEA - Institute of Systematics and Evolution of Animals, Polish Academy of Sci ences, Krakdw, Poland; MNHN - Laboratoire d ’Entomologie, Muséum national d’Histoire naturelle, Paris, France.

SYSTEMATIC ACCOUNT Family Hypogastruridae Ceratophysella denticulata Bagnall, 1941

M aterial examined . Tanzania, Ngorongoro Conservation Area, 2,200 m a.s.l., brink of the crater, dry forest near Sopa Lodge, 28.viii.1996, lgt. B. & K. Kowalski, 2 specimens: $ and 6 juv. Geographical distribution . Probably cosmopolitan species.

Xenylla gamae Cardoso, 1967

Acherontiella kowalskiorum sp. n.

D iagnosis . Habitus and buccal cone typical of the genus Acherontiella Absolon, 1913. Labrum with 4/554 setae. Distal labral setae smooth, slightly thicker than other distal se tae. Antenna Ill-organ with two subcylindrical guard sensilla with a very characteristic

217 constitution. Antennal segment IV with 5 globular sensilla of which 2 are large, placed in cavities, and the remaining 3 are smaller. Male with secondary sexual characters on ab dominal sterna IV and V. Tibiotarsi I, II and III with 19, 19 and 18 setae, respectively. D escription . Holotype (male) length: 0.64 mm, paratype length (juvenile female): 0.57 mm. Colour in alcohol white. Tegumental granulation rather strong. Antennae shorter than head (about 3/4 of the length of head). Antennal segment I with 7 setae, antennal segment II with 11 setae. Segments III and IV fused dorsally, the ventral separation well marked. Sensory organ of antennal segment III consisting of two small internal sensilla, two subcylindrical guard sensilla of very characteristic constitution (Deharveng & Diaz, 1984: Fig. 4K) and ventral microsensillum. Antennal segment IV with 5 globular sensilla of which 2 lateroexternal ones are very large, placed in cavities, and the other 3 are smaller; dorsoextemal microsensillum; small subapical organite; and small non-retractile apical vesicle in ventrosubapical position (Fig. 2). Eyes absent. Buccal cone typical of the genus. Labrum with 4/554 setae. Distal labral setae smooth, slightly thicker than other distal setae. Dorsal chaetotaxy as in Fig. 1 with rather short, simple setae, with thin long sensory se tae s, their formula per half tergum: 022/11111. Abdominal tergum VI without anal spines. Thoracic sterna without setae. Ventral abdominal chaetotaxy as in Fig. 3. Ventral tube with 4 + 4 setae. Male without setae a2 on abdominal sternum IV, these are present in fe male. Male with secondary sexual characters on abdominal sterna IV and V: 1 + 1 trans formed setae on abdominal sternum IV (in row p’) and 3 + 3 on abdominal sternum V (in row a) (Fig. 4). Row p’ absent in female. Tibiotarsi I, II and III with 19, 19 and 18 setae respectively, with acuminate distal setae, with setae M in the row B. Femora I, II and III with 12, 11 and 10 setae respectively, tro chanters with 5 setae each, coxae I, II and III with 3, 8 and 7 setae, subcoxae “2” I, II and III with 0, 2 and 2 setae, subcoxae “1” I, II and III with 1, 2 and 3 setae respectively.

T ype material . Holotype8 in ISEA, paratype juvenile 9 in MNHN. T ype locality . Kenya, near Massai Mara National Reserve, savanna, fissure in granite rock with moss and liverworts, 25.viii.1996, lgt. B. & K. Kowalski, 1 8 and 1 specimen juvenile. Etymology . This species is cordially dedicated to Polish paleontologists, Barbara Rzebik-Kowalska and Kazimierz Kowalski, who collected for us the material from Kenya and Tanzania.

D iscussion . The new species shares two characters (constitution of guard sensilla in an tenna Ill-organ and the number and type of sensilla on the antennal segment IV) with two species: Acherontiella Candida (Delamare Deboutteville, 1952 sensu Deharveng & Diaz, 1984) and A. colotlipana Palacios-Vargas & Thibaud, 1985. The latter species possesses 18, 18 and 17 setae on the tibiotarsi I, II and III, and 1, 2 and 2 setae on the subcoxae “2”, while the other two species have 19, 19 and 18 setae on the tibiotarsi and 1, 2 and 3 setae on the subcoxae “2”. A. kowalskiorum sp. n. is very close to A. Candida. They differ by the presence of setae ml on the abdominal sternum IV in A. Candida, absent in the new spe cies; setae m’ 1 are present in both species. Males of both species differ by the type of the secondary sexual characters. In A. kowalskiorum sp. n. the transformed setae are arranged on the posterior part of the abdominal sternum IV (1 + 1 setae) and on the abdominal ster num V in front of genital plate (3 + 3 setae). In A. Candida, they are situated only on the genital plate (2 + 2 setae). R emarks . Acherontiella Candida was briefly described by Delamare Deboutteville (1952) in the genus Xenyllina Delamare Deboutteville, 1948 from Ivory Coast and

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Figs 1-4. Acherontiella kowalskiorum sp. n. 1 - dorsal chaetotaxy (scale 0.1 mm); 2 - antennal seg ments III and IV dorsally (scale 0.01 mm); 3 - abdominal sterna II-V (scale 0.1 mm); 4 - male secondary sexual characters (scale 0.01 mm).

219 redescribed by Deharveng & Diaz (1984). The type [according to Fig. 4 on page 63 in De- lamare Deboutteville (1952), it was a juvenile specimen] was lost, and we therefore use the redescription based of material from Mt. Kenya (Deharveng & Diaz, 1984) for comparison.

Family Odontellidae Afrodontella septemlobata (Salmon, 1954)

M aterial examined . Tanzania, Ngorongoro Conservation Area, 2,200 m a.s.l., brink of the crater, dry forest near Sopa Lodge, 28.viii.1996, lgt. B. & K. Kowalski, 1 specimen juv.; Kenya, near Massai Mara National Reserve, savanna, fissure in granite rock with moss and liverworts, 25.viii.1996, lgt. B. & K. Kowalski, 1 9. Geographical distribution . Described from Ruwenzori in Uganda and found by Deharveng (1981) on Mt. Kenya and Aberdare in Kenya. This is the first record of this species from Tanzania.

Family Neanuridae Subfamily Frieseinae Friesea vtorovi Tshelnokov, 1977

D iagnosis . Habitus and buccal cone typical for genus Friesea Dalla Torre, 1985. 2 + 2 eyes present. Some stronger and feebly serrated setae on abdominal terga V and VI, for mula of sensory setae s per half tergum: 022/11111. Abdominal tergum VI without anal spines. Thoracic sternum without setae. Vestigial furca reduced to finely granulated small area with 4 small microchaetae. Tibiotarsi I, II and III with 18, 18 and 17 setae respec tively, with acuminate distal setae, without setae M. Redescription . Holotype (juvenile male) length 0.61 mm, length of other specimens: 0.53 (juv.) - 0.73 (adult male) and 0.89 (female) mm. Colour in alcohol very ligth gray, ocular plate blue-black. Tegumental granulation rather strong. Antennae shorter than head (about 2/3 of the length of head). Antennal segment I with 7 setae, II with 12 setae. Seg ments III and IV fused dorsally, ventral separation well marked. Sensory organ of antennal segment III consisting of two small internal sensilla bent in the same direction, two subcylindrical guard sensilla (ventral longer than dorsal), and ventral microsensillum. Anten nal segment IV with 6 distinct subcylindrical sensilla and dorsoextemal microsensillum; subapical organite present; apical vesicle simple (Figs 6 and 7). Eyes 2 + 2. Buccal cone typical for the genus. Mandible with 7 teeth. Dorsal chaetotaxy as in Fig. 5 with quite short simple setae, some stronger and feebly serrated setae on abdominal terga IV-VI, with thin long sensory setae s, their formula per half tergum: 022/11111. Abdominal tergum VI without anal spines. Thoracic sterna with out setae. Ventral abdominal chaetotaxy as in Fig. 10. Ventral tube with 4 + 4 setae. Ves tigial furca reduced to finely granulated small area with 4 small microchaetae. Tibiotarsi I, II and III with 18, 18 and 17 setae respectively, with acuminate distal setae, without setae M. Femora I, II and III with 12, 11 and 10 setae respectively, trochanters with 5 setae each, coxae I, II and III with 3, 7 (8) and 7 setae respectively, subcoxae “2” I, II and III with 0, 2 and 2 setae respectively, subcoxae “1” I, II and III with 1, 2 and 2 setae respectively. Claw without teeth, empodial appendage absent (Fig. 9).

220 Figs 5-10: Friesea vtorovi Tshelnokov, 1977. 5 - dorsal chaetotaxy (scale 0.1 mm); 6 - antennal seg ment III and IV dorsally (scale 0.01 mm); 7 - antennal segment III and IV ventrally (scale 0.01 mm); 8 - vestigial furca (scale 0.01 mm); 9 - tibiotarsus II with claw (scale 0.01 mm); 10 - abdominal sterna IV-V (scale 0.1 mm).

221 T ype material . Holotype juvenile 6 and two paratypes in the Department of Entomology, State Uni versity of Sankt Petersburg (Russia). T ype locality . Democratic Republic of Congo (formerly Zaire), Kahuzi-Bega National Park, Mitumba Mts, forest with bamboos on the slope of Kahuzi Mt., 1,800 m a.s.l., litter with soil, 15.ix.1975, lgt. P.P. Vtorov & N.N. Drozdov. Other material . Tanzania, Ngorongoro Conservation Area, 2,200 m a.s.l., brink of the crater, dry for est near Sopa Lodge, 28.viii.1996, lgt. B. & K. Kowalski, 4 specimens. Kenya, near Massai Mara Na tional Reserve, savanna, fissure in granite rock with moss and liverworts, 25.viii.1996, lgt. B. & K. Kowalski, 1 specimen juv.

D iscussion . Among the twelve species of Friesea with 2 + 2 eyes, Friesea vtorovi Tshelnokov, 1977 and F. neptunia Greenslade & Deharveng, 1997 are distinguished by the absence of the furca and anal spines. The remaining species with 2 + 2 eyes invariably possess either the furca or anal spines, or both:F. microphthalma Deharveng & Bedos, 1991 (3 anal spines, furca present), F. jaliscoensis Palacios-Vargas & Mejia, 1988 (3 anal spines, furca absent), F. gemioculata Loksa, 1964 (3 anal spines, furca present), F. kardosia (Wray, 1952) (8 or more anal spines, trilobed apical vesicle, furca absent; in our opin ion, the structures described by Wray, 1952 as anal spines should rather be regarded as spiniform setae), F. judithae Palacios-Vargas, 1986 (6 spiniform setae, furca present), F. stachi Kseneman, 1936 (3 anal spines, furca present), F. deharvengi Izarra, 1980 (6 feebly serrated anal spines, furca present), F. arnei Weiner & Najt, 1985 (4 spiniform setae, furca present), F. alaskella Fjellberg, 1985 (6 anal spines, furca present), F. wilkeyi Christiansen & Bellinger, 1974 (2 anal spines, furca present), F. troglophila Cassagnau, 1958 (6 anal spines, furca absent). F. vtorovi possesses the paurochaetotic type of chaetotaxy with short setae, whileF. neptunia shows the plurichaetotic type with very long macrochaetae and fairly long mesochaetae. Subcylindrical sensilla on antennal segment IV are rather short and subequal in F. vtorovi and quite long and different in F. neptunia. The latter species possesses a trilobate apical vesicle, while the apical vesicle in F. vtorovi is simple. Remarks . In the description of F. vtorovi, the median line is incorrectly drawn (Tshel nokov, 1977: Fig. 3). Setae placed near this line are in fact setae c2 and p2. The holotype shows some asymmetry in the chaetotaxy in abdominal sternum IV.

Key to the species ofFriesea Dalla Torre, 1895 with 2 + 2 eyes

1 Furca present (stage 2-4 after Cassagnau, 1958) ...... 2 - Furca absent (stage 5 after Cassagnau, 1958) ...... 9 2 Mucro present, hook-shaped, fused with dens ...... 3 - Mucro absent, dens with 1-3 setae ...... 4 3 Abdominal tergum VI with two anal spines, tenaculum with 1 + 1 teeth; U S A ...... F. wilkeyi Christiansen & Bellinger, 1974 - Abdominal tergum VI with three anal spines, tenaculum with 2 + 2 teeth; Thailand ...... F. microphthalma Deharveng & Bedos, 1991 4 Abdominal tergum VI with three anal spines ...... 5 - Abdominal tergum VI with four or more anal spines or spiniform setae ...... 6 5 Tenaculum with 2 + 2 teeth, dens with two setae (stage 4), color whitish, 1 + 1 anterior eyes and 1 + 1 posterior eyes with dark-blue pigment; Ukraine ...... F. stachi Kseneman, 1936 - Tenaculum with 3 + 3 teeth, dens with one distal seta, color white-yellow, 2 + 2 anterior eyes without pigment; Hungary ...... F. geminioculata, Loksa, 1964

222 6 Abdomen with 6 anal spines or spiniform setae, setae m present on abdominal tergum IV ...... 7 - Abdominal tergum VI with 4 spiniform setae, setae m absent on abdominal tergum IV; Korea ...... F. amei Weiner & Najt, 1985 7 Abdominal tergum VI with 6 anal spiniform setae, setae m absent on thoracic terga II and III ...... 8 - Abdomen with 6 anal spines, setae m4 present on thoracic terga II and III, abdominal tergum III with setae m2, m3, without setae a2 and a4; USA ...... F. alaskella Fjellberg, 1985 8 Abdominal terga I—III with setae m2, m3, without setae a2 and a4, body without blue or gray pigmen tation (colour white-yellow), eyes with dark-blue pigment; F ran ce...... F. deharvengi Izarra, 1980 - Abdominal terga I—III without setae m, a2 and a4, body with blue-gray pigmentation, eyes with blue pigment; Mexico ...... F.judithae Palacios-Vargas, 1986 9 Abdominal tergum VI with anal spines or spiniform setae ...... 10 - Abdominal tergum VI without anal spines or spiniform s e ta e ...... 12 10 Abdominal tergum VI with six or more anal spines or spiniform setae, furcal remnant with 2 + 2 microchaetae in one row inF. troglophila (character state unknown in F. kardosia) ...... 11 - Abdominal tergum VI with three anal spines, furcal remnant with 3 + 3 microchaetae; Mexico ...... F. jaliscoensis Palacios-Vargas & Mejia, 1988 11 Abdominal tergum VI with six anal spines, furcal remnant with 2 + 2 microchetae in one row, 2 + 2 anterior eyes; France ...... F. troglophila Cassagnau, 1958 - Abdominal tergum VI with eight spiniform setae, 1 + 1 anterior eyes and 1 + 1 posterior eyes; USA ...... F. kardosia (Wray, 1952) 12 Body with very long macrochaetae and fairly long mesochaetae, apical vesicle trilobate; Australia ...... F. neptunio Greenslade and Deharveng, 1997 - Body with short simple setae, apical vesicle simple; Democratic Republic of Congo, Tanzania ...... F. vtorovi Tshelnokov, 1977

Subfamily Pseudachorutinae Aethiopella pedifalx (Salmon, 1956)

M aterial examined , Tanzania, Ngorongoro Conservation Area, 2,200 m a.s.l., brink of the crater, dry forest near Sopa Lodge, 28.viii. 1996, lgt. B. & K. Kowalski, 1 specimen juv. Geographical distribution . Described from Democratic Republic of Congo (Belgian Congo).

Cephalachorutes sp.

M aterial examined . Tanzania, East Usambara Mts, Amani, 1,000 m a.s.l., evergreen tropical forest, v.1995, lgt. L. & P. Grandcolas, 1 specimen juv. Remarks . Probably a new species (with anterior part of the head, meso- and metathorax, abdominal segments I-VI and legs dark gray-blue, the posterior part of head and prothorax white), but we do not have enough material to describe it.

Furculanurida grandcolasorum sp. n.

D iagnosis . Habitus and buccal cone typical of the genus Furculanurida Massoud, 1967. Postantennal organ with 9-10 vesicles arranged eliptically. 5 + 5 eyes present. Very short ordinary setae, formula of sensory setae s per half tergum: 022/11111. Thoracic sterna without setae. Furca well developed with 5 setae on each dens. Mucro straight with apex slightly hooked. Tibiotarsi I, II and III with 19, 19 and 18 setae respectively, with microchaeta M in row B. D escription . Holotype (male) length 0.75 mm, paratypes (females) length 0.82 mm and 0.93 mm. Colour in alcohol gray-blue, ocular plate blue-black. One specimen represents another form (Fig. 17) with antennae, anterior part of the head, thoracic segments II—III,

223 Figs 11-16. Furculanurida grandcolasorum sp. n. 11 - dorsal chaetotaxy (scale 0.1 mm), postantennal organ and eyes (scale 0.01 mm); 12 - antennal segment III and IV dorsally (scale 0.01 mm); 13 - mandi ble (scale 0.01 mm); 14 - maxilla (scale 0.01 mm); 15 - dens with mucro (scale 0.01 mm); 16 - tibiotarsus III with claw (scale 0.01 mm). abdominal segments I-VI and legs dark gray-blue, the posterior part of head and prothorax white. Ventral side of entire head white, only labium gray-blue. Ocular plate dark. Tegu mental granulation rather strong. Antennae shorter than head (about 3/4 of the length of head). Antennal segment I with 7 setae, antennal segment II with 11 setae. Segments III

224 and IV fused dorsally, ventral separation well marked. Sensory organ of antennal segment III consisting of two small straight internal sensilla in lateroventral position, two subcylindrical guard sen silla, both dorsolateral in comparison with microsensilla; a ventral microsensillum present. Antennal segment IV with long simple setae, with 6 distinct subcylindri cal sensilla; dorsoextemal microsensillum absent, truncated subapical organite pre sent; apical vesicle trilobate (Fig. 12). Postantennal organ (Fig. 11) almost twice larger than ocellus, bearing 9-10 vesicles arranged elliptically. Eyes 5 + 5. Buccal cone typical of the genus. Mandi ble with 6 teeth (Fig. 13), maxilla with two lamellae (Fig. 14). Labral chaetotaxy 4/352. Dorsal chaetotaxy as in Fig. 11 with very short simple setae, with long sensory setae s, their formula per half tergum: 022/11111. Thoracic sterna without setae. Ventral tube with 3 + 3 setae. Fig. 17. Furculanurida grandcolasorum sp. n. - Furca well developed with 5 setae on bicolour form. each dens (Fig. 15). Muero straight with apex slightly hooked. Ratio muero : dens = 1 : 1.5. Tenaculum with 3 + 3 teeth. Tibiotarsi I, II and III with 19, 19 and 18 setae respectively, with acuminate distal setae, with setae M (= microchaeta) between setae B4 and B5 present. Femora I, II and III with 13, 12 and 11 setae respectively, trochanters with 6 setae each, coxae I, II and III with 3, 5 and 7 setae respectively, subcoxae “2” I, II and III with 0, 2 and 2 setae respectively, sub coxae “ 1” I, II and III without setae. Claw with inner tooth in basal 1/4 length of its inner edge (Fig. 16). Empodial appendage absent. T ype material. Holotypeó in MNHN, paratypes 1 9 in MNHN and 1 9 in ISEA. T ype locality. Tanzania, East Usambara Mts, Amani, 1,000 m a.s.l., evergreen tropical forest, v.1995, Igt. L. & P. Grandcolas. Etymology. This species is cordially dedicated to our friends, French entomologists Laure Desutter- Grandcolas and Philippe Grandcolas, who collected the material for us in Tanzania.

D iscussion . The new species is closest to Furculanurida belemensis Arlé & Rufino, 1976 with 5 + 5 eyes, described from the Amazonian basin. The two species differ by shape of postantennal organ (oval in the new species and circular inF. belemensis), by the number of vesicles in postantennal organ (9-10 in the new species and 8-9 in F. belemen sis), number of setae on dens (5 in the new species and 6 in F. belemensis) and by the ratio muero : dens (1 : 1.75 in the new species and 1 : 2.15 in F. belemensis). Thibaud & Massoud (1980) described Furculanurida arlei with 5 + 5 eyes from Mo rocco. This species belongs, however, toStachorutes Dallai, 1973 (see below).

225 Pseudachorutes ouatilouensis Najt & Weiner, 1997

M aterial examined . Tanzania, Zanzibar, Jozani Forest, tropical humid forest, 3.ix.l996, lgt. B. & K. Kowalski, 3 specimens. Geographical distribution . Described from New Caledonia. This is the first record of this species from Tanzania.

R emarks . Having made our description (Najt & Weiner, 1997) ofP. ouatilouensis, we obtained further material from New Caledonia and can complete the description. P. ouatilouensis possesses a minute inner tooth on the claw, small apical denticles on labium and a large finely granulated area on the ventral side of the dens. Specimens from Tanza nia present these characters more distinctly than the New Caledonian specimens. All the specimens have femora I, II and III with 12, 11 and 10 setae respectively, trochanters with 5 setae each, coxae I, II and III with 3, 7(6) and 7(6) setae respectively, subcoxae “2” I, II and III with 0, 2 and 2 setae, and subcoxae “1” I, II and III with 1, 2 and 2 setae respectively.

Stachorutes dallaii sp. n.

D iagnosis . Habitus typical of the genus Stachorutes Dallai, 1973. Antennal segment IV with large simple apical vesicle, ventrally with one basal lanceolate sensillum. Postanten- nal organ bearing 9-10 vesicles arranged in a circle. Eyes 2 + 2. Buccal cone typical for the genus. Mandible with 2 teeth, maxilla with two lamellae. Furca reduced, with 5 setae on each dens and stump-shaped mucro. Tenaculum with 2 + 2 teeth. Tibiotarsi I, II and III with 18, 19 and 18 setae respectively. Claw without inner tooth. D escription . Holotype (male) length 0.58 mm, paratype (juvenile) length 0.38 mm. Col our in alcohol white-gray, eyes dark. Tegumental granulation rather strong. Antennae shorter than head (about 3/4 of the length of head). Antennal segment I with 7 setae, antennal segment II with 12 setae. Segments III and IV fused dorsally, ventral sepa ration well marked. Sensory organ of antennal segment III consisting of two small internal sensilla bent in the same direction, two subcylindrical, subequal guard sensilla, ventral microsensillum present. Antennal segment IV with simple setae, dorsally with 6 distinct subcylindrical sensilla, dorsoextemal microsensillum and subapical organite present; large simple apical vesicle (Fig. 19); ventrally with one basal lanceolate sensillum, without sen sory rasp (Fig. 20). Postantennal organ twice larger than ocellus, bearing 9-10 vesicles arranged in a circle. Eyes 2 + 2. Buccal cone typical of the genus. Mandible with 2 teeth, maxilla with two lamellae (Fig. 22). Labium as in Fig. 21, without seta E and with 1 + 1 apical denticles. Dorsal chaetotaxy as in Fig. 18, with short ordinary setae, with rather long sensory setae s; their formula per half tergum: 022/22111. Head without setae aO and oc3. Thoracic sterna without setae. Ventral tube with 4 + 4 setae. Ventral chaetotaxy as in Fig. 25. Furca reduced, with 5 setae on each dens, mucro stump-shaped (Fig. 24). Ratio mucro : dens : claw III = 1 : 10.5 : 8.5. Tenaculum with 2 + 2 teeth. Tibiotarsi I, II and III with 18 (17 on one leg), 19 and 18 setae, respectively, with acuminate distal setae, with setae M almost in row B; tibiotarsi I and III without setae B1 and B3, tibiotarsus II without seta B3. Femora I, II and III with 13, 12 and 11 setae respec tively, trochanters with 6 setae each, coxae I, II and III with 3, 6 and 6 setae respectively,

226 /

Figs 18-25. Stachorutes dallaii sp. n. 18 - dorsal chaetotaxy (scale 0.1 mm); 19 - antennal segment III and IV dorsally (scale 0.01 mm); 20 - antennal segment III and IV ventrally (scale 0.01 mm); 21 - labium (scale 0.01 mm); 22 - mandible and maxilla (scale 0.01 mm); 23 - tibiotarsus III with claw (scale 0.01 mm); 24 - furca (scale 0.01 mm); 25 - abdominal sterna II-VI (scale 0.1 mm).

227 subcoxae “2” I, II and III with 0, 3 and 3 setae respectively, subcoxae “1” I, II and III with 1, 2 and 2 setae. Claw without inner tooth (Fig. 23). Empodial appendage absent.

T ype material . Holotype6 in ISEA, paratype juvenile in MNHN. Type locality . Tanzania, Ngorongoro Conservation Area, 2,200m a.s.l., brink of the crater, dry forest near Sopa Lodge, 28.viii.1996, lgt. B. & K. Kowalski. Etymology . This species is cordially dedicated to our friend Romano Dallai, a prominent Italian collembologist.

D iscussion . Until now, only one species of genusStachorutes Dallai, 1973 with 2 + 2 eyes was described: Stachorutes dematteisi Dallai, 1973 from Italy. The new species dif fers by presence of 5 setae on the dens (4 in S. dematteisi), of a stump-shaped mucro (ab sent in S. dematteisi), and by 9-10 vesicles in the postantennal organ (4-7 in S. dematteisi).

Key to the species ofStachorutes Dallai, 1973

1 Antennal segment IV with flame-shaped sensilla, 2 + 2 eyes; Poland ...... S. sphagnophilus Slawska, 1996 - Antennal segment IV with subcylindrical sensilla ...... 2 2 2 + 2 eyes present ...... 3 - 5 + 5 eyes present ...... 4 3 Mucro present, postantennal organ with 9-10 simple vesicles, dens with 5 setae; Tanzania ...... S. dallaii sp. n. - Mucro absent, postantennal organ with 4-7 vesicles, each dens with 4 setae; Italy ...... S. dematteisi Dallai, 1973 4 Mucro very well separated from the dens, setae pi on head present or ab se n t ...... 5 - No distinct border between mucro and dens, setae aO and cl on the head absent, setae pi present . . 6 5 Dens with 6 setae each, mandible with 3 teeth, head with setae aO, pi and cl; USA ...... S. navajellus Fjellberg, 1984 - Dens with 5 setae each, mandible with 2 teeth, head with setae c 1, without setae aO and p 1; Morocco ...... S. arlei (Thibaud & Massoud, 1980) comb. nov. 6 Short buccal cone, simple apical vesicle, seta a2 on thoracic tergum II p resen t...... 7 - Elongated buccal cone, trilobate apical vesicle, seta a2 on thoracic tergum II absent; France ...... S. longirostris Deharveng & Lienhard, 1983 7 Postantennal organ with 8 vesicles, dens with 5 setae, manubrium with seta aO, without setae al, seta p4 on head present; Spain ...... S. valdeaibarensis Arbea & Jordana, 1991 - Postantennal organ with 9-10 vesicles, dens with 6 setae, manubrium with setae al, without seta aO, seta p4 on head absent; Switzerland ...... S. sherae Deharveng & Lienhard, 1983

Remarks . Slawska (1996) included Stachorutes riebi Barra, 1991 to the key to the spe cies of Stachorutes, but it does not belong to this genus, as well as Stachorutes jizuensis Tamura, 1997 described from southwest China (Yunnan Province). In 1983 Deharveng & Liehnard suggested that Micranurida ashrafi Yosii, 1966 described from Nepal, could be another species of the genusStachorutes. At present we do not have enough evidence to include it into this genus. Stachorutes arlei (Thibaud & Massoud, 1980) comb. n. is hereby transferred from Fur- culanurida Massoud, 1967 to Stachorutes Dallai, 1973 sensu Deharveng & Lienhard, 1983. The species possesses one microsensillum on antennal segment IV, mandible with only two teeth, and a reduced furca with small mucro (ratio mucro : dens =1:3).

228 Figs 26-32. Paleonura cassagnaui sp. n. 26 - dorsal chaetotaxy (scale 0.1 mm); 27 - antennal seg ment III and IV dorsally (scale 0.01 mm); 28 - apex of labrum ventrally (scale 0.01 mm); 29 - mandible and maxilla (scale 0.01 mm); 30 - labium (scale 0.01 mm); 31 - dorsolateral tubercle (abdominal tergum V; scale 0.01 mm); 32 - abdominal sterna IV-VI (scale 0.1 mm).

Subfamily Neanurinae Paleonura cassagnaui sp. n.

D iagnosis . Head with two setae oc, setae A, B, C, D, F and G (seta E absent). Thoracic terga II and III with two setae Di, abdominal tergum IV with only one seta De (seta s).

229 D escription . Holotype (male) length 0.53 mm, paratype (female) length about 1.2 mm. Colour in alcohol white, eyes without pigmentation. Tubercles not developed. Abdominal segment VI not bilobate (Fig. 26). Antennae shorter than head (about 3/4 of the length of head). Antennal segment I with 7 setae, antennal segment II with 11 setae. Segments III and IV fused dorsally, ventral sepa ration well marked. Sensory organ of antennal segment III consisting of two small internal sensilla bent in the same direction, two subcylindrical guard sensilla (ventral slightly longer than dorsal), ventral microsensillum present. Antennal segment IV with simple se tae (some of them with blunt apex), dorsally with 8 distinct subcylindrical sensilla, with out microsensillum, subapical organite present; without apical vesicle, but with a swollen smooth area (Fig. 27). Eyes 2 + 2. Buccal cone typical for the genus. Mandible with two teeth, maxilla unilamellated (Fig. 29). Labrum rather long, rounded at apex (Fig. 28). Labium without seta B (Fig. 31). Cephalic chaetotaxy as in Fig. 26 and Table 1.

T able 1. Palaeonura cassagnaui sp. n., cephalic chaetotaxy.

Group of setae Tubercle Number of setae Type of setae Setae Cl - 4 M F me G Af (+) 8 M B me C,D mi A Oc (+) 2 M Oca mi Ocp

Di, De - 4 me Dil, Del mi Di2, De2 DL - 4 me DL6, DL5 mi DL3

L - 4 M acumin. L4 me LI, L2, L3 So - 5 me So2, 3, 4, 5, 6

Postcephalic chaetotaxy as in Table 2, Figs 26 and 32. Dorsal chaetotaxy with ordinary setae of three types: thin microchaetae (smooth and acuminate), mesochaetae of two types (acuminate and rounded at apex), and rather short macrochaetae, slightly serrated on ab dominal terga IV-VI, covered with weak sheath rounded at apex (Fig. 31). Sensory setae s longer than the closest meso- and macrochaetae: their formula per half tergum: 022/11111. Thoracic sterna without setae. Ventral tube with 4 + 4 setae. Tibiotarsi I, II and III with 18, 18 and 17 setae respectively, distal setae acuminate. Seta M absent. Femora I, II and III with 13, 12 and 11 setae respectively, trochanters with 6 setae each, coxae I, II and III with 7 setae each, subcoxae “2” I, II and III with 0, 1 and 1 setae respectively, subcoxae “1” I, II and III (as above) with 0, 1 and 1 setae. Claw without teeth. Empodial appendage absent.

T ype material . Holotype d in MNHN, paratype 2 in ISEA.

230 T ype locality . Tanzania, East Usambara Mts, Amani, 1,000 m a.s.l., evergreen tropical forest, v.1995, lgt. L. & P. Grandcolas. Etymology . This species is cordially dedicated to Paul Cassagnau, a French collembologist and spe cialist in Neanurinae. D iscussion . The new species is closest toPaleonura dejeani Cassagnau, 1996 from Bu rundi and Democratic Republic of Congo, toP. anosyennica Cassagnau, 1996 from Mada gascar, and to P. monochaeta Deharveng & Bedos, 1993 from Thailand. All these species possess two setae Oc and setae A, B, C, D, F and G on head (seta E absent). P. cassagnaui sp. n. differs from the other species by number of setae Di on thoracic terga II and III (2 setae in the new species, 3 setae in all others) and by presence on abdominal tergum IV of only one seta De (seta s), while there are two setae and seta s inP. anosyennica, one seta and seta s in P. monochaeta, 4 setae and seta s of De + Dl in P. dejeani.

Table 2. Palaeonura cassagnaui sp. n., postcephalic chaetotaxy.

Thoracic terga Di De Dl L = Sex 1/pi I 1 2 1 - II 2 2 + s 2 + s + ms 3 III 3 2 + s 2 + s 3

Abdominal terga Di De Dl L = Scxl/pl I 2 1 + s 2 2 II 2 1 + s 2 2 III 2 1 + s 2 2 IV 2 s 2 4 V 2 - 3 + s ------VI 7 Abdominal sterna Ve Ag/An Furca VL L’ II 4 - - - - III 2 - 2 + 2 1 - IV 6 - - 4 - V 1 3 + 3 - ‘ 1 1 VI 12(11 in $) 1 + 1 - - -

Family Onychiuridae Subfamily Tullbergiinae Tullbergia kilimanjarica (Delamare Deboutteville, 1953)

Mesaphorura kilimanjarica Del am are Deboutteville, 1953. Tullbergia (Mesaphorura) kilimanjarica: Paclt, 1959: 38.

D iagnosis . Body elongated. Formula of pseudocelli per half tergum: 11/111/11121. An tennal segment IV with small apical vesicle, five sensilla a-e, three dorsal sensory clubs and two small sensory rods protected by integumental fold in antenna Ill-organ. Postan- tennal organ 3.5^1 times longer than diameter of pseudocellus, elongated, with simple vesicles in two regular rows. Thoracic tergum II and III with setae m3, m4 and m5. Redescription . Lectotype (preadult male) length 0.95 mm, length of other specimens: 0.44 mm for I stage, 0.91 (juvenile female) - 1.24 (female) mm. White in alcohol.

231 Granulation on the head and on the lateral parts of thoracic terga slightly coarse, on the two last abdominal terga coarse. Pseudocelli present per half tergum: 11/111/11121. Antennal segment IV with five thickened sensilla a-e, one microsensillum, subapical organite and small apical vesicle. Sensory organ of antennal segment III with four guard setae, two small sensory rods protected by integumental fold, three thick, cylindrical sen sory clubs, and thick long bent sensory club on ventral side (Fig. 36). Antennal segments II and I with 11 and 7 setae respectively. Elongated postantennal organ 3.5^1 times longer than the diameter of nearest pseudocellus, with 48-57 (52 in the holotype) simple vesicles in two rows (Fig. 34). Chaetotaxy as in Figs 33 and 37, with macrochaetae and microchaetae well differenti ated. Dorsal chaetotaxy as in Fig. 33 and Table 3; remark: anomalies in chaetotaxy of lectotype. Sensory setae (s) very weakly marked on thoracic terga II—III and on abdominal terga IV-V, pleurae II—III. Microsensilla (ms) on thoracic terga II and III present. Seta pi on head sligthly longer (1.2 times) than p2. On abdominal tergum VI one pair of yellow anal spines on distinct papillae, seta aO 1.7 times longer than pO.

Table 3. Tullbergia kilimanjarica, formula of dorsal chaetotaxy per half tergum. th l th II th III abd I abd II abd III abd IV abd V a - 5(1) 5(1) 6 6 6 5(5) 3(8) m - 5(2) 5(2) - 1(4) 1(4) 2(6) 2(9) p 4 5(3) 5(3) 6 6 6 6(7) 2(10) sbc/pl 2 3 3 2 3 3 5 2 1 - a4 absent; 2 - m l, m3, m5 and m6-seta s present, microsensilla present; 3 - p6 absent; 4 - m4 pre sent; 5 - a6 absent; 6 - m4 and m6 present; 7 - p5-seta s; 8 - a3 absent; 9 - m5 and m6 present; 10 - p2, p4 - seta s, present.

Subcoxae “2” I, II, and III with 2, 3, and 3 setae respectively, pleura of abdominal seg ments I-V with 2, 3, 3, 5, and 2 setae. Anterior setae on abdominal pleura II and III are sensory setaes. Thoracic sterna II and III with 1 + 1 setae each. Ventral abdominal chaetotaxy as in Fig. 37. Ventral tube with 4 + 4 distal setae and 2 + 2 basal setae. Tibiotarsi I, II, and III with 14, 14, and 13 setae respectively without seta M. Claw with out teeth, empodium triangular, empodial appendage absent (Fig. 35). Femora I, II and III with 12, 11 and 11 setae respectively, trochanters with 6 setae each, coxae I, II and III with 3, 8 and 7 setae respectively, subcoxae “1” I, II and III with 0, 4 and 4 setae respectively, subcoxae “2”: as above.

T ype material . Lectotype preadult c?, presently designated, in MNHN. T ype locality . Tanzania, Kilimanjaro, Shira Plateau, 12,450 ft. (approx. 3,800 m), soil from heath for mation, 29.xi.1948, lgt. G. Salt. Other material . Tanzania, Ngorongoro Conservation Area, 2,200 m a.s.l., brink of the crater, dry for est near Sopa Lodge, 28.viii.1996, lgt. B. & K. Kowalski, 6 specimens (juvenile and 9).

D iscussion . Tullbergia kilimanjarica Delamare Deboutteville, 1953 appears to be close to T. africana Martynova, 1979 described from Democratic Republic of Congo, because of the same type of chaetotaxy on abdominal tergum IV (2 + 2 microchaetae between macro chaetae a4 in a-row). T. kilimanjarica differs from T. africana by pseudocellar formula

232 33 1

t M W A " 4 ^ ">• 1 l J i, • - j r ' ?>!,!/ (4lV‘f t ^ ° i \ i l

- ^ . / h f i y y

Figs 33-37. Tullbergia kilimanjarica (Delamare Deboutteville, 1953). 33 - dorsal chaetotaxy (scale 0.1 mm); 34 - postantennal organ and pseudocellus (scale 0.01 mm); 35 - tibiotarsus III with claw (scale 0.01 mm); 36 - antennal segment III and IV dorsally (scale 0.01 mm); 37 - abdominal sterna I-VI (scale 0.1 mm).

233 (11/111/11121 in T. kilimanjarica and 11/011/11121 in T. africana). Further characters were found in chaetotaxy. On thoracic terga II and III, setae m3 are present in T. kiliman jarica (absent in T. africana). In ventral chaetotaxy, unpaired setae aO and mO on abdomi nal sternum II are present in T. kilimanjarica (absent in T. africana), and the arrangement of setae on abdominal sternum IV is different: seta p4 in T. africana is in the position of m’5. Remarks . Delamare Deboutteville (1953) wrongly placed the present species in the ge nus Mesaphorura Bomer, 1901, whereas it belongs to the genusTullbergia Lubbock, 1876 sensu stricto. However, Tullbergia kilimanjarica is not a new combination since Paclt (1959: 38), who used the generic name Tullbergia in a broad sense, listed this species as Tullbergia (Mesaphorura) kilimanjarica.

Tullbergia sp.

M aterial examined . Tanzania, East Usambara Mts, Amani, 1,000 m a.s.l., evergreen tropical forest, v.1995, lgt. L. & P. Grandcolas, 3 and juvenile. Remarks . Probably a new species, but we do not have enough material to describe it. The specimens have 2, 2, 2, 5 and 2 setae on abdominal pleura I-V (3, 3, 3, 5 and 2 in T. kilimanjarica). Moreover, setae s are shorter and more distinct than inT. kilimanjarica.

Family Isotomidae Cryptopygus thermophilus Axelson, 1900

M aterial examined . Tanzania, East Usambara Mts, Amani, 1,000 m a.s.l., evergreen tropical forest, v.1995, lgt. L. & P. Grandcolas; 16 specimens. Geographical distribution . Probably a cosmopolitanspecies.

Folsomides parvulus Stach, 1922

M aterial examined . Tanzania, Zanzibar, tropical humid forest, 3.ix,1996, lgt. B. & K. Kowalski, 2 specimens; Ngorongoro Conservation Area, 2,200 m a.s.l., brink of the crater, dry forest near Sopa Lodge, 28.viii.1996, lgt. B. & K. Kowalski, 2 specimens; Kenya, near Massai Mara National Reserve, savanna, fissure in granite rock with moss and liverworts, 25.viii.1996, lgt. B. & K. Kowalski, 6 specimens. Geographical distribution . Probably a cosmopolitan species.

Folsomina onychiurina Denis, 1931

M aterial examined . Tanzania, Ngorongoro Conservation Area, 2,200 m a.s.l., brink of the crater, dry forest near Sopa Lodge, 28.viii.1996, lgt. B. & K. Kowalski, 4 specimens. Geographical distribution . With wide holotropical distribution.

Isotomiella nummulifer Deharveng & Oliveira, 1990

M aterial examined . Tanzania, East Usambara Mts, Amani, 1,000 m a.s.l., evergreen tropical forest, v.1995, lgt. L. & P. Grandcolas, 7 specimens; Tanzania, Ngorongoro Conservation Area, 2,200 m a.s.l., brink of the crater, dry forest near Sopa Lodge, 28.viii.1996, lgt. B. & K. Kowalski, 3 specimens. Geographical distribution . Described from Manaus (Amazonian Region) in Brazil and found in the Seychelles (Deharveng & Fjellberg, 1993), in Sumatra (Deharveng & Suhardjono, 1994) and in Thailand, southern part of the Malay Peninsula (Bedos & Deharveng, 1994). This is the first record of this species from the African continent.

234 CONCLUSIONS Following a bibliographical analysis, we give the list of valid species belonging to Hy- pogastruridae, Brachystomellidae, Odontellidae, Onychiuridae and Isotomidae from Kenya and Tanzania.

LIST OF SPECIES

Hypogastruridae Hypogastrura manubrialis (Tullberg, 1869); Kenya (Deharveng & Diaz, 1984) Hypogastrura cf. manubrialis (Tullberg, 1869); Kenya (Deharveng & Diaz, 1984) Hypogastrura cf. sahlbergi Reuter 1895; Tanzania (Paclt, 1967) Hypogastrura cf. tullbergi Schaeffer, 1900; Kenya (Deharveng & Diaz, 1984) Ceratophysella cf. denticulata (Bagnall, 1941); Kenya (Deharveng & Diaz, 1984) Ceratophysella cf. gibbosa (Bagnall, 1940); Kenya (Deharveng & Diaz, 1984) Octocanthella aethiopica Martynova, 1961; Kenya (Deharveng & Diaz, 1984) Xenylla cassagnaui Gama, 1983; Kenya (Gama, 1983; Deharveng & Diaz, 1984) Xenylla deharvengi Gama, 1983; Kenya (Gama, 1983; Deharveng & Diaz, 1984) Xenylla gamae Cardoso, 1967; Tanzania Xenylla cf. gisini Cardoso, 1968; Kenya (Deharveng & Diaz, 1984) Xenylla kenyensis Gama, 1983; Kenya (Gama, 1983; Deharveng & Diaz, 1984) Xenylla rhodesiensis Womersley, 1926; Tanzania (Paclt, 1967) Acherontides huetheri Deharveng & Diaz, 1984; Kenya Acherontiella candida (Delamare Deboutteville; 1952); Kenya (Deharveng & Diaz, 1984) Acherontiella kowalskiorum sp. n.; Tanzania Brachystomellidae Brachystomella sp. [Brachystomella parvula (Shaeffer, 1896) sensu Delamare Deboutteville, 1953]; Tanzania. This is certainly a new species, but it cannot be described on the basis of one specimen only. Odontellidae Afrodontella septemlobata (Salmon, 1954); Tanzania Neanuridae Frieseinae Friesea africana Delamare Deboutteville, 1953; Tanzania Friesea vtorovi Tshelnokov, 1977; Tanzania Friesea nigrissima (Salmon, 1954); Kenya (Salmon, 1954) Friesea said (Salmon, 1954); Kenya (Salmon, 1954) Pseudachorutinae Pseudachorutes alluaudi Delamare Deboutteville, 1945; Kenya Pseudachorutes mabirensis Philiptschenko, 1926; Kenya, Uganda (Philiptschenko, 1926) Pseudachorutes niloticus Wahlgren, 1906; Kenya (Delamare Deboutteville, 1945) Pseudachorutes ouatilouensis Najt & Weiner, 1997; Tanzania Aethiopella africana Delamare Deboutteville, 1945; Tanzania (Delamare Deboutteville, 1945) Aethiopella flavoantennata (Philiptschenko, 1926); Kenya, Uganda (Philiptschenko, 1926) Aethiopella maculata Delamare Deboutteville, 1945; Tanzania (Delamare Deboutteville, 1945) Aethiopella mandibulata Delamare Deboutteville, 1945; Kenya (Delamare Deboutteville, 1945) Aethiopella pedifalx (Salmon, 1956); Tanzania Kenyura muldsensa Salmon, 1954; Kenya (Salmon, 1954) Furculanurida grandcolasorum sp. n.; Tanzania Stachorutes dallai sp. n.; Tanzania Linnaniemia bicolor (Delamare Deboutteville, 1945); Tanzania (Delamare Deboutteville, 1945) Linnaniemia gigas Philiptschenko, 1926; Kenya, Uganda, (Philiptschenko, 1926; Delamare Deboutte ville, 1945 from Kenya)

235 Neanurinae Paleonura aberdarensis Cassagnau, 1996; Kenya (Cassagnau, 1996) Paleonura africana Cassagnau, 1996; Kenya (Cassagnau, 1996) Paleonura cassagnaui sp. n.; Tanzania Pronurafluctuans Cassagnau, 1996; Kenya (Cassagnau, 1996) Pronura kilimanjarica Delamare Deboutteville, 1953; Tanzania (Delamare Deboutteville, 1953) lnameria kenyensis Cassagnau, 1984; Kenya (Cassagnau, 1984) Adbiloba sokolovi (Philiptschenko, 1926); Kenya, Uganda (Philiptschenko, 1926; Cassagnau & Dehar- veng, 1980, as Travura reticulata Cassagnau & Deharveng, 1980 from Kenya) Onychiuridae Tullbergiinae Mesaphorura yosii (Rusek, 1967) [Mesaphorura iowensis (Mills, 1932) sensu Delamare Deboutteville, 1953]; Tanzania (our determination). Tullbergia kilimanjarica (Delamare Deboutteville, 1953); Tanzania Mesaphorura said Delamare Deboutteville, 1953; Tanzania (Delamare Deboutteville, 1953). This spe cies belongs to a new genus (Najt & Weiner in litt.) Isotomidae Isotomiella nummulifer Deharveng & Oliveira, 1990; Tanzania Folsomides parvulus Stach, 1922; Tanzania Ballistura sp. (Baliistura excavata Folsom, 1937 sensu Delamare Deboutteville, 1953); Tanzania Proisotoma caerulea Salmon, 1955; Kenya (Salmon, 1955) Proisotoma (Clavisotoma) sjoestedti Wahlgren, 1908; Tanzania Folsomina onychiurina (Denis, 1931); Tanzania Cryptopygus thermophilus (Axelson, 1900); Tanzania Isotomurus palustris (Mueller, 1776); Tanzania (Paclt, 1967); species dubia for this region.

acknowledgements . We are most grateful to all who have made the materials described here available to us: L. Dessutter-Grandcolas and P. Grandcolas from the Laboratoire d’Entomologie, MNHN (Paris), B. Rzebik-Kowalska and K. Kowalski from the ISEA, Polish Academy of Sciences (Kraków). We are also grateful to G. Hodebert (Laboratoire d’Entomologie, MNHN, Paris) for drawing the habitus ofFurcu- lanurida grandcolasorum sp. n., and to V. Kuznetsova (Zoological Institute, Russian Academy of Sci ences, St. Petersburg) for a loan of the types ofFriesea vtorovi and Tullbergia africana.

REFERENCES

B edos A. & D eharveng L. 1994: The Isotomiella of Thailand (Coliembola: Isotomidae), with description of five new species. Entomol. Scand. 25: 451^460. Cassagnau P. 1958: Les espèces européennes du genre Friesea (Collemboles Poduromorphes).Bull. Soc. Hist. Nat. Toulouse 93: 17-29. C assagnau P. 1984: Introduction à l’étude des Phylliomeriens (Coliembola, Neanuridae): Diagnoses pré liminaires des espèces. Trav. Lab. Ecobiol. Arthropodes Edaphiques (Toulouse) 4: 1-30. C assagnau P. 1996: Collemboles Paleonurini primitifs d’Afrique et de Madagascar. Ann. Soc. Entomol. Fr. 32: 121-161. C assagnau P. & D eharveng L. 1980: Sur l’intérêt biogéographique et cytogénétique d’un nouveau genre de Collemboles Neanuridae: Travura n. g. Trav. Lab. Ecobiol. Arthropodes Edaphiques (Toulouse) 2: 1- 12. D eharveng L. 1981: La famille Odontellidae: phylogénèse et taxonomie.Trav. Lab. Ecobiol. Arthropo des Edaphiques (Toulouse) 3: 1-21. D eharveng L. & D iaz A. 1984: Collemboles du Kenya: I. Liste des stations. II. Hypogastruridae. Bull. Mus. Natn. Hist. Nat. (4e Sér.) 6: 335-348. D eharveng L. & Fjellberg A. 1993: Isotomiella from Seychelles islands (Insecta, Coliembola, Isotomi dae). Spixiana 16: 121-125.

236 D eharveng L. & L ienhard C. 1983: Deux nouvelles espèces du genre Stachorutes Dallai, 1973 (Collem- bola). Rev. Suisse Zool. 90: 929-934. D eharveng L. & S uhardjono Y.R. 1994: Isotomiella Bagnall 1939 (Collembola, Isotomidae) of Sumatra (Indonesia). Trop. Zool. 7: 309-323. D elamare D eboutteville C. 1945: Collemboles récoltés par Ch. Allaud et R. Jeannel en Afrique Orien tale 1911-1912. Première note.Mém. Mus. Natn. Hist. Nat. 21: 153-174. D elamare D eboutteville C. 1952: I. - Collemboles. In: Delamare Deboutteville C. & Paulian R.: Faune de Nids et des Terriers en Basse Côte-d’Ivoire. Encyclopédie Biogéographique Lechevalier,VIII. Paris, pp. 62-77. D elamare D eboutteville C. 1953: Collemboles du Kilimanjaro récoltés par le Dr. George Salt.Ann. Mag. Nat. Hist. (SI2) 6: 817-831. Gama M.M. da 1983: Systématique évolutive des Xenylla. XIII. Espèces provenant du Kenya (Insecta: Collembola). Rev. Univ. Coimbra 29: 249-257. M artynova E.F. 1979: [Two new species of springtails (Collembola) from Central Africa.]Zool. Zh. 58: 440-444 (in Russian, English abstr.). N a jt J. & W einer W.M. 1997: Collemboles Poduromorpha de la Nouvelle-Calédonie. In Najt J. & Matille L. (eds): Zoologia NeocaledonicaMém. Mus. Natn. Hist. Nat. 171: 9^14. Paclt J. 1959: Collembola. In Hanstrom B., Brinck P. & Rudebeck G. (eds): South African Animal Life 6. Almqvist & Wiksells, Stockholm, pp. 24—78. Paclt J. 1967: On South and Central African Collembola. J. Entomol. Soc. Sth. Afr. 29: 135-147. P hiliptschenko J.A. 1926: On the Collembola collected by the expedition of V.A. Dogiel and 1.1. Sok- olow in British East-Africa.Rev. Russe Entomol. 20: 180-196. Salmon J.T. 1954: New genera and new species of Neanurinea (Collembola) from East Africa. Proc. R. Entomol. Soc. Lond. (B)23: 1-9. Salmon J.T. 1955: A new Proisotoma (Collembola) from Kenya.Proc. R. Entomol. Soc. Lond. (B) 24: 33-35. Slawska M. 1996: Stachorutes sphagnophilus n. sp. from Nothern Poland (Collembola: Neanuridae).Ge nus 7: 325-329. T hibaud J.-M. & M assoud Z. 1980: Etude des Collemboles de certains milieux du Maroc et considérations biogéographiques sur la faune du Maghreb. Rev. Suisse Zool. 87: 513-548. T shelnokov V.G. 1977: [Some new and little known species of springtails (Collembola) in the fauna of equatorial Africa.] In Kryzhanovskii O.L. & Galkin A.K. (eds): (Systematics and Faunistics of In sects). Akad. Nauk SSSR, Leningrad, pp. 103-110 (in Russian). W ahlgren E. 1908: Apterygogenea. 1. Collembola.Wiss. Ergebn. Zool. Exped. Kilimandjaro 18: 1-10. W ray D.L. 1952: Some new North American Collembola.Bull. Brooklyn Entomol. Soc. 48: 95-106.

Received May 14, 1997; accepted August 25, 1997

237