African Crickets (Gryllidae). 1. Teleogryllus of Eastern and Southern Africa

Transcription

1 Proceedings of the Academy of Natural Sciences of Philadelphia 135: , African Crickets (Gryllidae). 1. Teleogryllus of Eastern and Southern Africa DANIEL OTTE Academy of Natural Sciences of Philadelphia 19th and the Parkway Philadelphia, PA AND WILLIAM CADE Biological Sciences Brock University St. Catharines, Ont. L2S 3A1 ABSTRACT. - This work is based mainly on field work conducted by the authors during three expeditions in 1980 and 1982 to Kenya, Tanzania, and South Africa. Twenty five species are discussed. Of these seven were discovered by the authors, four from Kenya and Tanzania, and three from South Africa. Three species are transferred to Teleogryllus from other genera (afer from Gryllus, pulchriceps and soror from Platygryllus). Fourteen species (burri, wittei, macrurus, adustus, brachypterus, nigripennis, leucostomoides, posticus, leucostomus, angolensis, latifrons, afer, griaulei, and soror) were not encountered in this study, but illustrations and notes on the types are provided. T. typographicus is synonymized under xanthoneurus. The type of T. longecaudatus Schulthess was not found. The discrimination of Teleogryllus species is based mainly on song differences and secondarily on morphological differences. Because Africa is likely to contain a large number of Teleogryllus species, this paper represents a preliminary survey. [Africa, crickets, Gryllidae, morphology, new species, songs, systematics, Teleogryllus, types] POSTSCRIPT.-As this paper was in press it was discovered that aequipennis was a synonym of pulchriceps. In figures read pulchriceps in place of aequipennis. Teleogryllus ranges throughout sub- Saharan Africa, then eastwards through Asia to China and Japan, all of Southeast Asia, Australia, and the Pacific as far east as Hawaii. Possibly only Gryllus among the Gryllinae has a more widespread distribution. Among other groups only Gryllotalpa, Anaxipha, Pteronemobius, and Oecanthus have ranges of equivalent size. Chopard's (1967b) catalogue suggests a gap in the distribution between Africa and India. Since Teleogryllus species live either in grasslands or in woodland leaf litter, their absence between Kenya and India is probably explained by the aridity of these regions. Like other groups of crickets, Teleogryllus species are most reliably separated by their songs. We have found this to be especially true of the group of species which we here place in the Africanus Group. Members of 102 the Aequipennis Group differ from one another both in song and morphology. The two species which Chopard placed in Platygryllus (aequipennis) and Gryllus (afer) had not had their genitalia examined, thus it is not surprising that each was placed in the wrong genus. We did not tape record wittei, burri, brachypterus, macrurus, angolensis, latifrons, adustus, nigripennis, and posticus. Aside from the type localities, the distributions of these species are not known and essentially nothing is known about their biology. Chopard lists 32 species of Teleogryllus in his 1967 catalogue. Otte and Alexander (1983) added one Australian species. The diversity of species is greatest in Africa and lowest in the Pacific. The known number of species by country or area is as follows: SubSaharan Africa, 25 species; Indian sub-

2 AFRICAN CRICKETS 103 continent, and Burma, 11 species; eastern Asia, 6 species; Indonesia, 3 species; Australia, 3 species; Pacific Islands, 2 species. In the present work we discuss 25 species; we add 10 species to the genus -7 are new, 3 are transferred from other genera. The African species are placed into four species groups as shown below. The status of the Griaulei Group remains uncertain. By the male genitalia they seem most similar to Teleogryllus. If they are not members of this genus then they will probably have to be placed in a new genus. Since we know nothing about their songs we leave them in Teleogryllus. AFRICANUS GROUP 1. africanus n. sp. 2. meru n. sp. 3. xanthoneurus Gerstaecker 1869 typographicus Saussure 1877 new synonym 4. leo n. sp. 5. grumeti n. sp. 6. natalensis n. sp. 7. marabu n. sp. 8. wernerianus Kamy 1907 xanthoneuroides Chopard burri Chopard 1961 a 10. wittei Chopard macrurus Walker adustus Karsch brachypterus Chopard 1967a 14. nigripennis Chopard leucostomoides Chopard 1961 a 16. posticus Walker 1869 PULCHRICEPS GROUP 17. pulchriceps Gerstaecker 1869 (moved from Platvgryllus) aequipennis Saussure 1899 new synonym 18. leucostomus Serville angolensis Chopard 196 la 20. zululandicus n. sp. 21. gnu n. sp. 22. latifrons Karsch 1893 OCEANICUS GROUP 23. afer Saussure 1877 (moved from Gryllus) GRIAULEI GROUP 24. griaulei Chopard soror Chopard 1940 (moved from Platygryllus) NOMEN DUBIUM 26. longecaudatus Schulthess 1894 evidently lost, not present in the Schulthess collection in Zurich. METHODS All tape recordings were made with a Nagra SN tape recorder (8.9 cm/s) and a Sony ECM-50PS electret condenser microphone attached to a parabolic reflector. Tapes were copied onto 5 inch reels by a Uher 4000 Report L tape recorder. Sonagraphs of all songs were made with a Kay Elemetrics 7029A Sonagraph. Temperatures were determined whenever recordings were made. The thermometer was placed as close as possible to the place where the cricket was singing, especially when crickets were singing in leaf litter, in burrows, or on the ground. Body parts of crickets are named according to the terminology of Otte and Alexander Abbreviations of museum repositories are as follows: ANSP, Academy of Natural Sciences of Philadelphia; BM, British Museum of Natural History; GM, Museum d'histoire Naturelle, Geneva; PM, Museum National d'histoire Naturelle, Paris; VM, Naturhistorisches Museum, Vienna; MNHU, Museum ftur Naturkunde, Humboldt-Universitat, Berlin, DDR. TAPE RECORDING LoCALITIES 2. Tanzania, Mt. Meru, N slope, 12 x Tanzania, Lake Manyara Park, 13 x Tanzania, Serengeti Park, Seronera, 14 x A. Tanzania, Serengeti Park, Mbalageti R., nr. Kirawira, 20 x Tanzania, Serengeti Park, Grumeti R., Kirawira, 20 x Tanzania, Ngorogoro Crater, 20 x South Africa, Natal, Eshowe, 30 x South Africa, Natal, Mtunzini, 7 ix South Africa, Natal, St. Lucia, 10 ix South Africa, Natal, Hluhluwe Game Reserve, 10 ix South Africa, Natal, Tugela R. at Middle Drift, 15 ix 1980

3 104 DANIEL OTTE AND WILLIAM CADE 20. South Africa, Natal, Mkuze Game Reserve, 19 ix South Africa, Transvaal, Rust-de-Winter, 20 ix South Africa, Transvaal, Magaliesberg Mountains, 3 iii South Africa, Natal, Richards Bay, 17 iii South Africa, Natal, nr. Empangeni, 17 iii South Africa, Natal, 18 km W Empangeni, 18 iii South Africa, Natal, Hlabisa area, 22 iii South Africa, Natal, Mkuze Game Reserve, 28 iii South Africa, Transvaal, Badplaas, 30 iii South Africa, Transvaal, Clanor, nr. Skukusa Gate of Kruger National Park, 31 iii South Africa, Transvaal, Kruger Nat. Park, nr. Pretoriuskop, 2 iv Kenya, Hunter's Lodge, Nairobi-Mombasa Road, 14 ix Kenya, plain and valley S of Sagala Hills, nr. Voi, 17 ix Kenya, Sagala Hills, south slope, 17 ix Kenya, Matuga, 18 ix Kenya, Shimba hills, W slope, 19 ix Kenya, Waa, 17 km S Mombasa, 18 ix Kenya, Shimba Hills, 22 ix Tanzania, Usa River, nr. Arusha, 25 ix Tanzania, Gonja, South Pare Mts., 26 ix Tanzania, Mombo, 27 ix Tanzania, Mombo, W side, 27 ix Tanzania, Dar-es-Salaam, 28 ix Tanzania, Amani, East Usambara Mts., 30 ix Tanzania, ca. 30 km W Kirogwe, 2 x Tanzania, 40 km W Handeni, 3 x Tanzania, 15 km S Babati, nr. Bonga, 6 x Tanzania, Manyara National Park, 7 x 1982 Genus TELEOGRYLLUS Chopard Teleogryllus Chopard 196 ib: 277. Type species: Gryllusposticus Walker 1869: 30, by original designation. (The designation of posticus as the type species is unfortunate because the holotype is a female and no males have been positively assigned to the species.) Holotype of type species examined. DIAGNOSTIC CHARACTERS (MALES). Teleogryllus species can be separated from other Gryllinae by the male genitalia and by the song. Genitalia: Epiphallus elongate, ending as a single median process; virga extremely long and slender, emerging from an especially long spermatophore tube mold; ectoparameres very long compared to the re- lated genera Platygryllus, Acanthogryllus, Gryllus, and Scapsipedus. Tympana: Inner tympanum small and round (brachypterus without inner tympanum); outer one large and oval. Song: Calling song consists of two parts, for convenience termed part I and part II. Most species produce the two parts together, with a chirp or trill of part I followed by one or more chirps or trills of part II; two species only occasionally sing part II (pulchriceps, zululandicus); most species may sing only part I chirps in the early evening. Only T. wernerianus possess a onepart song. OTHER FEATURES. See Otte and Alexander 1983 for terminology. Body color variable-reddish brown, brown, or black. Vertex and occiput with or without longitudinal pale stripes. Body length varies from about 15 to 25 mm in males, and 17 to 30 mm in females. Head as wide as pronotum or narrower depending on group and condition of hindwings. When hindwings are well developed for flying the head is usually narrower than the pronotum. Head with 3 ocelli, median one usually smaller than lateral ones. Forewings with file teeth; several veins connect veins 2A and 3A; vein 1A thin, usually connected to stridulatory vein (Cu2) with more than 7 veinlets; basal area of forewing with a network of small veins; 4 chords; 2-8 (usually 4 or more) harp veins; mirror almost always divided into two cells (rarely 3 or more); apical area welldeveloped; forewings always more than.8 and less than 1.3 times as long as hind femora. Hindwings variable in length, apparently never less than half as long as forewings; extending to ends of cerci in some specimens of some species. Cerci more than half as long but rarely longer than hind femora. Hind femora between 1.05 and 1.33 times as long as hind tibiae; the latter with 4-7 (usually 5 or 6) inner subapical spurs and 4-8 (usually 6-8) outer subapical spurs and with 3 inner and 3 outer apical spurs. BIOLOGY. We have some familiarity with

4 AFRICAN CRICKETS 105 the habits of 11 African Teleogryllus species. These inhabit two or three main habitat types. None of the species are burrowers, but grassland species may exploit existing holes in the ground. Species of the Africanus Group (except the somewhat aberrant wernerianus) are closely associated with dry leaf litter, often along river beds, but also in upland woodlands. T. wernerianus may be the only known member of the group which lives in dry savanna grasslands; it obviously tolerates the driest conditions. Species in the Pulchriceps Group live in moist grasslands. The Australian and Pacific species (oceanicus, commodus, and marini) are all grassland species, suggesting that the morphologically similar T. afer lives in similar habitats. Nothing is known about the biology of posticus, wittei, burri, macrurus, brachypterus, adustus, leucostomus, angolensis, latifrons, and leucostomoides. Teleogryllus are some of the most acoustically conspicuous animals wherever they occur. Anyone who has camped along dry river beds or in open woodlands in east and South Africa is familiar with their two part songs, although they are probably rarely seen by anyone but collectors. Members of the Africanus Group must have a good bed of larger dry leaves to hide. They are especially abundant in Ficus forests along rivers in east Africa. In towns and villages species of this group are abundant in banana groves, hedges, and sugar cane fields. In South Africa they are found mainly in the Natal subtropical coastal forests and in the Natal and Transvaal lowveld thorn bush. Judging from the length of the hindwings and present distributions we believe that most Teleogryllus species are good fliers and readily colonize new areas. T. oceanicus has been used in flying phonotaxis studies in the laboratory and is a powerful flier (Pollack and Hoy 1981), as are other field crickets with long hind wings (Cade 1979). Of the species treated here only brachypterus lacks hindwings; other species have the hindwings not less than half as long as the forewings. NOTES ON SONGS. The sound produced during one closure of the wings is termed a pulse. A closely spaced series of pulses which obviously forms a group is termed a chirp. A continuous train of pulses is termed a trill. SPECIES GROUPS Africanus Group 1. With a pale ring wholly or partly surrounding each eye. 2. Male epiphallus with elongate, non-fleshy parameres. 3. Dorsum of epiphallus with a narrow fleshy area. 4. Body color reddish brown. 5. Dorsum of head black in anterior half, reddish from vertex to occiput; face patterned with dark and pale marks. 6. Part II of song consisting of trills. 7. Cu2 of forewing without a strong bend. 8. Lower half of lateral lobes no lighter than dorsal half. Pulchriceps Group 1. Without a pale ring around each eye. 2. Male epiphallus with fleshy parameres. 3. Dorsum of epiphallus with a wide fleshy area. 4. Body color variable, black and pale, brownish or reddish brown. 5. Dorsum of head black or brown, with or without longitudinal pale bands; face largely pale. 6. Part II of song consisting of short chirps. 7. Cu2 of forewing with a strong bend. 8. Lower half of lateral lobes much lighter than dorsal half. Griaulei Group 1. Without a pale ring around each eye. 2. Male epiphallus variable (Fig. 12). 3. Dorsum of epiphallus with wide fleshy area. 4. Body color pale brown or straw colored. 5. Dorsum of head brown, with distinct pale stripes. 6. Song not known. 7. Cu2 of forewing without a strong bend at mirror. 8. Lower half of lateral lobes lighter than dorsal half. Oceanicus Group 1. Without a pale ring around each eye. 2. Male epiphallus with wide, non-fleshy parameres. 3. Dorsum of epiphallus largely concave. 4. Body color black. 5. Dorsum of head black, with faint longitudinal pale stripes on occiput. Face black. 6. Song not known.

5 106 DANIEL OTTE AND WILLIAM CADE 4JF FIG. 2. Teleogryllus wernerianus. FIG. 1. Teleogryllus xanthoneurus. All members of the Africanus Group are very similar in appearance except brachypterus and wernerianus. 7. Cu2 of wing not strongly bent. 8. Lateral lobes entirely black. KEY TO MALES Note: Females of closely related species are especially difficult to distinguish; because of this and because females of some species are not known, a key is not included here. Because most of Africa remains unstudied, tentative keys are provided only for Kenya, Tanzania, and South Africa. These may prove to be useful for Mozambique as well. KENYA AND TANZANIA 1. Face largely black. Palpi entirely white. Legs black. Mirror longer than wide. Body length of males less than 18 mm... wernerianus Face with pale coloration ventrally. Palpi not entirely white. Legs brown, reddish, or yellowish. Body length of males more than 18 mm Lower half of pronotal lateral lobes pale (Fig. 3) Lower half of pronotal lateral lobes dark (Fig. 3) Dorsum of head brown, striped (Fig. 3, top right). Dorsum of pronotum brown. File with fewer than 250 teeth... pulchriceps Dorsum of head black, unstriped (Fig. 3, middle right). Dorsum of pronotum brown. File with over 300 teeth... :... gnu 4. Hind femur length less than three times greatest width Hind femur length more than three and a half times greatest width File with teeth... grumeti File with teeth... marabu 6. Hind femur length more than 3.9 times greatest width. File with more than 210 teeth... meru Hind femur less than 3.8 times greatest width. File with fewer than 210 teeth Male genitalia with wider ectoparameres (Fig. 10)... africanus Male genitalia with narrower ectoparameres... xanthoneurus

6 AFRICAN CRICKETS 107 xarnton_urus aequipemnis wernerlanus burrn wittei gnu afer zululandi FIG. 3. Head and pronotal patterns in selected species. Most members of Africanus Group are similar to xanthoneurus. Note: Aquipennis = pulchriceps. SOUTH AFRICA 1. Face black, palpi entirely white. Mirror longer than wide wernerianus Face variable, palpi not entirely white. Mirror wider than long Head, pronotum and legs black (Fig. 3, lower left)... afer Head, pronotum, and legs with some pale coloration Lower half of pronotal lateral lobes pale. Disk of pronotum black... zululandicus Lower half of pronotalateral lobes dark (brown to blackish reddish). Disk of pronotum reddish brown Hind femur length more than 4.0 times greatest width. File averaging more than 220 teeth natalensis Hind femur length less than 3.8 times greatest width. File averaging less than 210 teeth File averaging more than 180 teeth. Epiphallus with short dorsal ridges africanus File averaging less than 180 teeth. Epiphallus with long dorsal ridges leo AFRICANUS GROUP The sixteen species of this group usually have the following characteristics: body color reddish brown; legs reddish brown to dark rusty red. Head: dorsum of head often black or at least much darker in anterior half; face patterned with dark and pale markings (Fig. 3); side of head behind eyes blackish or dark brown, becoming light brown below eyes; most species with a pale ring around the eyes-complete in some species, incomplete in others; lateral ocelli connected to the eye rings by pale arms; head narrower than pronotum. Pronotum: lateral lobes sometimes much darker than disk, sometimes similar in color. Forewings: Veins often yellowish especially in basal and apical areas. Abdomen: dorsum reddish brown, venter lighter in color sometimes orangebrown. Ovipositor long, rarely shorter than hind femora. 1. Teleogryllus africanus n. sp. HOLOTYPE. 8, Eshowe, Natal, South Africa, 30 x 1980 (D. Otte and W. Cade) ANSP.

7 108 DANIEL OTTE AND WILLIAM CADE ~~~~ C +- V C C) o o o o ~~00 IN00 00 r - cc -s V 0 0) O rb 00 ae K? oo oo C's 6t = N 0~ 0 N oo N N N e0 NO 0 c Y N 00 N~~~ ~~~r 0 - ~~~t 00 0Q) 0tr~~~~~~ 0'I. 00o 0N0 00 ~ ri CO r I t > I~ I o I 00 ~ ~ C Cl q Cl* -0 C5 r > > ~oo > u Ev 4u> 0i 06 N Cl >c ooc - N~~~Cl 00 ~ 0 lrl0lcc 0 ~ C - -Cl - C CO~ ~ ~~~~~~~~~~~O 0ot - c mco ^ 3 00 N 00 Cl 00 ~~0 M 0 e C * oo. 0 I I I _tm 00L0 O 000 N D o ~ ^ s r {o m o o a = o < 1N 1 Sv 1 S LL <;t= O0X0U=2=t H4~~~ E Cl 0!.= Z A; E to,~~~_...

8 AFRICAN CRICKETS 109 TABLE 2. East and South African Teleogryllus females. OL, ovipositor length; remaining symbols as in Table 1. Italic numbers are means. BL F3L OL OL Species mm mm FWL mm F3L CL africanus (n = 2) 23.5 meru not known xanthoneurus (n= 1) leo (n = 5) 23.0 grumeti (n= 3) natalensis marabu not known wernerianus (n = 10) 16.1 pulchriceps zululandicus not known not known gnu F3 broken (n= 1) lost afer not known RECOGNITION. See Africanus Group above. Very difficult to distinguish from other members in this group, but with a different song and with fewer file teeth than the most similar species (meru and natalensis). It differs from xanthoneurus, grumeti, leo, and marabu in having more slender hind femora (Tables 1, 2). Head blackish on frons. Lateral lobes darker than disk. Hind femur length/width = ca DISTRIBUTION. Presently known from Natal and north central Tanzania, but probably widespread over eastern Africa. HABITAT. In Natal and Zululand common in the low coastal belt in leaf litter along rivers and in lowland forests behind coastal dunes. Not inhabiting deep forests, but abundant along edges. Prefers sandy soils. In towns common in hedges and among shrubbery in gardens and in banana groves. In northcentral Tanzania the species was found on wooded hillsides near Babati. SONG. T. africanus usually sings a two part song (Fig. 4), but after sundown and in the early evening males often sing a succession of part I chirps, leaving out part II trills altogether. Several males we recorded began with pure part I songs and then began to include longer and longer trills of part II as the evening progressed. By about 9 pm the full song with a normal ratio of part I and part II phrases was restored. The full song was also sung during daylight hours. Pollack and Hoy (1981) demonstrated that female T. oceanicus oriented more often to the part I portion of the song of this species than to the complete song when the signals are broadcast in the laboratory. Males of africanus and other African species may produce part I songs when females are flying early in the evening. The function of part II songs has not been studied. Males sing at all hours of the day, although the number singing is reduced during daylight hours and the songs of day singers is not as loud.

9 c's ^ * *^^s1 CZ l l ll~~~~~~~~~~~~~~~~i s 0a ss E*** *^ *0-* E av i a\.5si

10 AFRICAN CRICKETS Pulses/sec No. pulses Locality I II I/II I II khz?c (n= 3) (1.14 chirps/sec) (n= 6) (1.63 chirps/sec) (1.45 chirps/sec) (1.55 chirps/sec) SPECIMENS. Paratypes: TANZANIA: 15 km S Babati, near Bonga, wooded hillside, 6 x 1982 (Otte) 26 ANSP. SOUTH AFRICA: Natal, Eshowe, 30 x 1980 (Otte and Cade) 11 ANSP. Natal, Ballitoville, 19 viii 1981 (Otte) 16 ANSP. Natal, St. Lucia, 10 ix 1980 (Otte and Cade) 168, 1 9 ANSP. 2. Teleogryllus meru n. sp. HOLOTYPE. 6, coffee plantation at Tengeru a few km E of Arusha, Tanzania, 25 x 1982 (D. Otte) ANSP. RECOGNITION. Based on a unique male. Difficult to distinguish from africanus and natalensis, but not sympatric with those species. Differs from sympatric xanthoneurus in song, number of file teeth, and width of hind femora (Table 1). The only male also has 8 harp veins (xanthoneurus has 3-6 harp veins). Male genitalia as in Fig. 10. DISTRIBUTION. Tape recorded along eastern slopes of Mount Meru in Arusha National Park. Tape recorded and collected at Tengeru along lower southern slopes of Mount Meru. HABITAT. Collected holotype in leaf litter under coffee trees at Tengeru. Several males tape recorded in the morning. In Arusha National Park tape recorded in leaf litter along banks of a small stream in a brushy area. SONG. Song most similar to that of africanus, but ratio of the two pulse rates (part I/part II) is slower (. 45 or less vs.5 8 or more in africanus). Around 10 am at Tengeru one male sang only part I songs, another sang the full song. Late at night in Arusha Park all males were singing the full song. Local- Pulses/sec No. pulses ity I II I/II I II khz?c (4) SPECIMENS. Holotype 8 ANSP. 3. Teleogryllus xanthoneurus (Gerstaecker) Gryllus xanthoneurus Gerstaecker 1869: 212. Lectotype 6, here designated. Specimen bears following FIG. 4. Songs of four species. T. africanus: top two traces show typical day or late night songs. Lowest two traces show pure part I songs. Middle traces show stages in change over from evening to late night songs. T. natalensis: bottom trace is a pure part I early evening song. T. marabu: top trace shows a rarely heard part II phrase.

11 112 DANIEL OTTE AND WILLIAM CADE labels: "Type, Wanga [Zanzibar] v.d. Decken Expedition." MNHU. Teleogryllus xanthoneurus, Chopard 196 lb. Type examined. Gryllus typographicus Saussure 1877: 25. Holotype 6, Zanzibar (Hildebrandt) BM. Teleogryllus typographicus, Chopard 196 lb: 278. Type examined. Type has 180 file teeth. Genitalia as in Fig. 11. RECOGNITION. Difficult to distinguish from other members of the Africanus Group. Differs from meru and africanus by having relatively wider hind femora and narrower ectoparameres. Differs from grumeti by having more file teeth and by having a complete eye ring and by genitalia. Differs from marabu by having fewer file teeth, by its complete eye ring, and by the male genitalia (see Table 1). Holotype has 204 teeth. DISTRIBUTION. Widespread in Tanzania and southeastern Kenya. HABITAT. Leaf litter but occasionally inhabiting thick grass near trees. In the Shimba Hills near Mombasa abundant in grass and litter around edges of small forest patches and in litter on lightly wooded western slopes. At Arusha, Tanzania, common in leaf litter in gardens and under mango trees. At Gonja, Tanzania, inhabiting leaf litter in dense forest. At Lake Manyara, Tanzania, abundant in leaf litter of large broad-leafed trees at northern end of lake. SONG. Males evidently always sing the full song. We did not record pure part I songs anywhere. Males sing all day, but most intensively after sundown. (See chart at bottom of page.) SPECIMENS. TANZANIA: Lake Manyara, camping area at N end of Lake, 13 x 1980 (D. Otte) holotype 6, 26, 1I ANSP. Same place, 7 x 1982 (D. Otte) 26, 2Q ANSP. Usa River, nr. Arusha, 25 ix 1982 (D. Otte) 16 ANSP. 40 km W Handeni, road to Kondoa, 3 x 1982 (D. Otte) 16 ANSP. 4. Teleogryllus leo n. sp. HOLOTYPE. 6, Clanor, near Skukusa Gate of Kruger National Park, Transvaal, South Africa, 31 iii 1982 (D. Otte and W. Cade) ANSP. RECOGNITION. Difficult to distinguish from the two allopatric species africanus and natalensis, but differing from these species as follows: length of hind femur less than 3.3 times its width; male epiphallus with more pronounced dorsal ridges; stridulatory file Pulses/sec No. pulses Locality I II I/II I II khz?c A (n = 3) (n = 6) (n = 4)

12 AFRICAN CRICKETS ~~ 11.5s FIG. 5. T. xanthoneurus: top trace is the usual song; middle trace heard in mid-afternoon; bottom trace recorded in the evening. T. grumeti: both traces recorded at noon. T. leo: lower trace is the usual song; upper song heard mainly in the morning and occasionally late at night. T. wernerianus lacks a part II song; the small pulses may be made by the opening wings.

13 114 DANIEL OTTE AND WILLIAM CADE Pulses/sec No. Local- pulses ity I II I/II I II khz?c sec (0.85 ch/sec) (0.96 ch/sec) (1.33 ch/sec) (1.70 ch/sec) (1.33 ch/sec) prob (1.40 ch/sec) prob (1.50 ch/sec) prob (1.10 ch/sec) prob prob sec 3.0 prob sec 3.0 prob (0.99 ch/sec) with fewer teeth than natalensis (Table 1). Holotype with 181 teeth. DISTRIBUTION. We collected this species at one locality in Zululand and two localities in the eastern Transvaal. HABITAT. Leaf litter in lightly wooded bushveld. At Rus-de-Winter abundant in the garden of Jan Smut's former winter home. SONG. Males usually sing only part I songs, but late at night and in the morning they sing the full two-part song. Unlike most other species, part II is a long trill appended to part I chirps (Fig. 5). (See chart at top of page.) SPECIMENS. Paratypes: SOUTH AFRICA: Transvaal, Rust-de-Winter, 20 xi 1980 (D. Otte) 21, 1Y ANSP. Transvaal, Clanor, nr. Skukusa Gate, 31 iii 1982 (D. Otte and W. Cade) holotype 6, 36, 2Y ANSP. Natal, ca. 30 km NE Eshowe on old Empangeni road, 18 iii 1982 (D. Otte and W. Cade) 26 ANSP. Natal, ca. 18 km W Empangeni, 18 iii 1982 (D. Otte and W. Cade) 1I ANSP. 5. Teleogryllus grumeti n. sp. HOLOTYPE. 8, Grumeti River at Kirawira, Serengeti National Park, Tanzania, 20 x 1980 (D. Otte) ANSP. RECOGNITION. Difficult to separate from xanthoneurus and marabu, but averaging fewer file teeth than either species (Table 1). Male epiphallus with long dorsal ridges (like marabu and leo but unlike xanthoneurus) (Fig. 10). Holotype with 148 teeth. DISTRIBUTION. Collected only at Kirawira in the Serengeti Plains. HABITAT. In leaf litter under fig trees along Grumeti River. SONG. Full two-part songs and pure part I songs heard around noon. Part II was appended directly to part I (similar to day song of leo). Local- Pulses/sec No. pulses ity I II I/II I II khz?c * ** * 1.86 ch/sec. ** 2.0 ch/sec. SPECIMENS. Holotype a ANSP. Paratypes: same data as holotype, 3a, 2Y ANSP. 6. Teleogryllus natalensis n. sp. HOLOTYPE. a, Tugela River at Middledrift, near Kranskop, Natal, South Africa, 15 xi 1980 (D. Otte and W. Cade) ANSP. RECOGNITION. Difficult to distinguish from africanus, but with more file teeth, a slightly shorter forewing (relative to hind femur

14 AFRICAN CRICKETS 115 Pulses/sec No. pulses Locality I II I/II I II khz?c (continuous trill) (n = 3) *, *, * Part I. length) and a very different song. Sometimes allopatric with leo which has relatively broader hind femora and fewer file teeth (Table 1). Holotype with 231 teeth. DISTRIBUTION. To date found only in Zululand between Tugela River at St. Lucia Bay. HABITAT. Leaf litter under fig trees along lower Tugela River; leaf litter at edge of coastal forest back of sand dunes at St. Lucia; sugar cane fields near Hlabisa and Empangeni. SONG. Full song consists of two to four slow pulses (part I) and a trill with a faster pulse rate (Fig. 4). A male at St. Lucia produced a part I song in the early evening consisting of a continuous train of pulses (Fig. 4). Song is most similar to that of marabu, but latter species rarely produces part II song. (See chart at top of page.) SPECIMENS. Paratypes: SOUTH AFRICA: Natal, St. Lucia, 10 ix 1980 (Otte and Cade) LI ANSP. Natal, Tugela River at Middle drift, 15 xi 1980 (Otte and Cade) 16 ANSP. Same place, 26 iii 1982 (Otte and Cade) 26 ANSP. Natal, a few km W Empangeni, 17 viii 1982 (Otte and Cade) 16, I ANSP. 7. Teleogryllus marabu n. sp. HOLOTYPE. 6, Hunters lodge, along Nairobi-Mombasa Road, Kenya, 14 ix 1982 (D. Otte) ANSP. RECOGNITION. Differs from the very similar and allopatric xanthoneurus as follows: male epiphallus with long dorsal ridges; file with more file teeth (Table 1); pale eye rings not complete. Differs from africanus and meru by having wider hind femora. Holotype has 264 teeth. DISTRIBUTION. Collected at Hunters Lodge and near southern base of Sagala Hills near Voi. HABITAT. Debris and grass under large acacia trees. At Hunters Lodge abundant along a dammed spring arising near base of the Chyulu Hills. Especially abundant in guano covered debris under a tree where marabu storks nested. SONG. Males sang one part songs consisting either of pairs or triplets of widely spaced pulses or of continuous trains of pulses. Only one very faint part II song was taped near Voi. We heard males only after sundown. (See chart at bottom of page.) Pulses/sec No. pulses Locality I II I/II I II khz?c (1.02 pairs/sec) (1.35 pairs/sec) ca (.93 pairs/sec) (continuous) (continuous)

15 116 DANIEL OTTE AND WILLIAM CADE SPECIMENS. Paratypes: KENYA: Hunters Lodge, along Nairobi-Mombasa Rd, 14 ix 1982 (D. Otte) holotype 6, 36 ANSP. Valley on south side of Sagala Hills, near Voi, 17 ix 1982 (D. Otte) 26 ANSP. 8. Teleogryllus wernerianus (Kamy) Acheta wernerianus Karny 1907: 287. Lectogype 6, here designated, Mongalla, VM. Teleogryllus wernerianus, Chopard 196 lb: 277. Type examined. GryllusxanthoneuroidesChopard 1934: 333. Holotype 6, Taveta, Kenya, iii 1912 (Alluaud and Jeannel) PM. Teleogryllus wernerianus, Chopard 196 lb: 277. Type examined. RECOGNITION. Both sexes with white palpi; much smaller than other Teleogryllus, harp with 2 to 4 veins; mirror relatively narrow. Most individuals have very long hindwings and we suspect the species migrates readily about the drier parts of the continent. DISTRIBUTION. Chopard (1967) indicates that the species is from "Tropical Africa." I have collected it in the Tanzania highlands, Kenya coast, Magaliesberg Mountains of the Transvaal, and the coastal grasslands of Zululand. HABITAT. Savanna grasslands. In Serengeti National Park abundant in lawns surrounding SRI bungalows. Near Mombasa, Kenya, found in grassy patches among dry coastal scrub a few hundred meters from beach. In Zululand inhabiting grassland around mouth of St. Lucia Bay. In Transvaal collected on dry stony hillsides of Magaliesberg Mountains. SONG. Song consists of widely spaced pulses. Pulse rate varies quite widely. Occasionally pulses are paired, with the first pulse of a pair considerably more faint; perhaps the first pulse is produced by the opening wing (Fig. 5). Heard only at night. (See top of page.) SPECIMENS. TANZANIA: Serengeti Nat. Park, Seronera, 14 x 1980 (D. Otte) 66, 49 ANSP. Ngorongoro Crater, 20 x 1980 (D. Otte) 16, 49 ANSP. KENYA: Waa, 17 km S Mombasa, 18 ix 1982 (D. Otte) 16 ANSP. SOUTH AFRICA: Natal, Hlabisa, 9 ix 1980 (D. Otte Locality Pulses/sec khz?c (n = 4) (n= 3) (n = 3) (pairs)* (pairs)* pairs/sec 85** ** (n = 4) * Pairs consist of a faint pulse preceding a normal pulse; the first pulse is perhaps produced as the wing is opened. ** No males collected. One female was collected at 87. and W. Cade) 16 ANSP. St. Lucia, 10 ix 1980 (D. Otte and W. Cade) 16 ANSP. Ca. 30 km NE Eshowe 18 iii 1982 (D. Otte and W. Cade) 16 ANSP. Mkuze Game Reserve, 28 iii 1982 (D. Otte and W. Cade) 16 ANSP. Transvaal, Magaliesberg Mountains, 3 iii 1982 (D. Otte and W. Cade) 26 ANSP. 9. Teleogryllus burri (Chopard) Gryllus burri Chopard 196 la: 23. Type 6, Upper Mu- Simoj River, District of Mexico, Angola, 26 ix 1927 (Dr. M. Burr) PM. Teleogryllus burri, Chopard 196 lb: 277. Type examined. NOTES ON TYPE. Similar in appearance to xanthoneurus and africanus. Body color reddish brown, black on head. Head: dorsum of head like xanthoneurus; face as in Fig. 3. Pronotum: disk reddish brown, lateral lobes much darker; narrowing anteriorly; slightly wider than head. File with 217 teeth. Forewing 3.04 times as long as pronotum. Hind femur orange-brown; length about 3 times the width. Hind tibiae with 6 inner and 8 outer subapical spurs. Body length 22 mm; hind femur length 14.5 mm. 10. Teleogryllus wittei (Chopard) Gryllus wittei Chopard 1939: Lectotype 6, here designated, Ninda, Ruanda, 22 ix 1934 (G. F. de Witte) PM. Teleogryllus wittei, Chopard 1961b: 277. Type examined.

16 AFRICAN CRICKETS 117 NOTES ON TYPES. Male: Reddish brown and black, legs dark, reddish. Head: vertex and occiput dark reddish brown, with 6 longitudinal pale stripes; vertex to front of head black. Face marked as in Fig. 3. Pronotum: very slightly wider than head, disk reddish, slightly patterned, lateral lobes blackish, with nearly parallel sides. File with 116 teeth. Forewings with yellow veins mainly in basal area and on lateral field. Forewing 2.4 times as long as pronotum. Hindwing less than half as long as forewing. Hind legs missing. Body length ca. 19 mm. Female: Coloration similar to male, but head blacker on dorsum and with less distinct stripes. Forewing 1.41 times as long as pronotum. Hind femur length ca. 3 times greatest width. Hind tibiae with 6 inner and 7 outer subapical spurs. Ovipositor about 1.1 times as long as hind femur. Hind femur length ca. 13 mm. 11. Teleogryllus macrurus (Walker) Gryllus macrurus Walker 1869, 1: 29. Holotype Y. Specimen bears following original labels. "Fantee [Coast of Ghana, near Cape Coast] 17, Gryllus macrurus, one of Walker's series so named. Type." BM. Teleogryllus macrurus, Chopard 1961: 278. Type examined. NOTES ON TYPE. This large female is in poor condition. By facial markings and color she belongs to the Africanus Group. Hind femur length 17.5 mm. Forewing length 18 mm. Hindwings extending well beyond forewings. Hind femur length 3.23 times greatest width. Hind tibiae with 6 inner and 7 outer subapical spurs. 12. Teleogryllus adustus (Karsch) Gryllus adustus Karsch 1893: 151. Holotype c, Bismarckburg, Togoland (R. Biittner S.) MNHU. Teleogryllus adustus, Chopard 196 1b: 277. Type examined. Gryllus ambiguus Karsch 1893: 151. Synonymized by Chopard 1961: 277. Type not examined. NOTES ON TYPE. Figs. 9 and 10. Superficially similar to other members of the Af- ricanus Group, but file with ca. 356 teeth, veins of wings not yellow. Body length 21 mm; hind femur length 12.7 mm; forewing length 10.8 mm. Hind tibiae with 6 inner and 7 outer subapical spurs. Length of hind femur 3.09 times its greatest width. 13. Teleogryllus brachypterus Chopard Teleogryllus brachypterus Chopard 1967a: 761. Holotype Y, Sibiti, Congo [Zaire], xi 1963, PM. Type examined. NOTES ON ALLOTYPE MALE. Fore tibiae with a large oval outer tympanum but without an inner tympanum. Body color reddish brown and blackish. Legs brownish. Dorsum of head dark reddish brown to blackish but with broad yellowish bands running from median ocellus backwards through lateral ocelli and along the eyes. Face blackish above epistomal suture then with a yellow band across the median ocellus. Side of head dark brown. Pronotum slightly narrower anteriorly, greatest width slightly wider than head. Pronotum: disk dark reddish brown, slightly patterned; lateral lobes blackish. Forewings with two harp veins, with an undivided mirror, a short apical area, and with 123 file teeth. Forewing 3 times as long as pronotum (holotype female forewings about as long as pronotum). Hindwings abseni. Hind femur length 3.1 times maximal width. Hind tibia with 5 inner and 5 outer subapical spurs. Body length 16 mm, hind femur length 12 mm. 14. Teleogryllus nigripennis (Chopard) Gryllus nigripennis Chopard 1948: 112. Holotype Y, Mongbwalu, Kilo, 1939 (Mme Scheitz) PM. Teleogryllus nigripennis, Chopard 1961b: 277. Type examined. NOTES ON TYPE. This female bears a paratype label, but other labels indicate it is the specimen upon which the name is based. Coloration: dark reddish brown, head and pronotum largely black. Forewings with a pale stripe along each wing angle. Tympana:

17 118 DANIEL OTTE AND WILLIAM CADE small innner, large outer. Head largely dark brown to black on dorsum, faint longitudinal bands visible, pale around large ocelli. Face yellowish on labrum, reddish brown on mandibles and clypeus, turning black on upper frons. Cheeks black behind eyes. Without a pale band along posterior margin of eyes. Pronotum uniform dark brown to blackish. Abdomen dark reddish brown. Forewings dark brown, about 1.5 times as long as pronotum. Hindwings present, hidden, about as long as pronotum. Legs all dark reddish brown. Hind femur length about 3.05 times greatest width. Hind tibiae with 6 inner and 7 outer subapical spurs. Ovipositor about 0.95 times as long as hind femur. Body length ca. 17 mm; hind femur length 12 mm; forewing length 5 mm. 15. Teleogryllus leucostomoides (Chopard) Gryllus leucostomoides Chopard 1961 a: 22. Holotype 6, Vila Luso, Angola, 1 xi 1927 (Dr. M. Burr) BM. Teleogryllus leucostomoides, Chopard 196 lb: 277. NOTES ON TYPE. Most similar to leo in number of file teeth and male genitalia. File with about 165 teeth. The main morphological characters which separate this species from leo are as follows: epiphallus smaller and narrower, and ectoparameres not as long in this species; forewing with more numerous small cross veins between the stridulatory and harp veins; lateral lobes light brown (blackish in leo); frons not black; hind tibia with 7 inner and 9 outer subapical spurs. Forewings 3.55 times as long as pronotum and 1.04 times as long as hind femur. Hindwings extending about 21/2 pronotal lengths beyond forewings. Body length 24 mm; hind femur length 13.5 mm. 16. Teleogryllus posticus (Walker) Gryllus posticus Walker Lectotype Y, here designated, Sierra Leone, BM. Teleogryllus posticus, Chopard 1916b. Type examined. NOTES ON TYPE. The type female was described along with three females from Natal and a female from East Africa. These latter females, belonging to a different species (here named africanus), were designated as varieties by Walker and so could not be selected as the lectotype. This is unfortunate because the type specimen of the type species of Teleogryllus is now a female but the definition of the genus is based entirely on males. T. posticus appears to belong to the Africanus Group, but this can only be firmly established when males are collected. PULCHRICEPS GROUP 17. Teleogryllus pulchriceps (Gerstaecker) Gi'vllus pulchriceps Gerstaecker 1869: 212. Holotype a, Mombasa (v.d. Decken) MNHU. Platygryllus, Randell Type examined. NEW COMBINATION. Gryllus aequipennis Saussure 1899: 602. Type a, Zanzibar, GM. Platygryllus, Chopard 1967b: 94. Type examined. RECOGNITION. A large light brown species previously placed under Platygryllus. Face yellow. Dorsum of head with four broader pale longitudinal stripes between eyes (medial pair especially distinct), with a very thin median line, and with a narrow pale line connecting lateral ocelli. Eyes very pale brown or gray. In some specimens upper half of lateral lobes dark, lower half pale. Hind femur very pale with reddish oblique stripes in upper half, dorsal surface with numerous small pale spots. Holotype with 238 file teeth. DISTRIBUTION. Presently known from eastern Kenya, eastern Tanzania, and Zanzibar. HABITAT. Open grassy areas. At Matuga, near Mombasa, males were in burrows in lawns of the Agricultural Experiment Station. In the Shimba Hills males sang in grassy openings in the forest and in grassy roadsides. West of Handeni, Tanzania, the

18 AFRICAN CRICKETS 119 Pulses/sec No. pulses Locality I II I/II I II khz?c (.44 chirps/sec) (.6 chirps/sec) (.9 chirps/sec) (.44 chirps/sec) * * (.56 chirps/sec) (.34 chirps/sec) (.33 chirps/sec) ( chirps/sec) ( chirps/sec) * 2-4 part II chirps per series at ca. 3.3 chirps/sec. ** 3 part II chirps per series at ca. 3.7 chirps/sec. species occupied grass in savanna woodland. SONG. This species usually produces only part I songs. The only part II songs recorded were by several males in the late afternoon in the Shimba Hills. Unlike members of the Africanus Group, part II of the song is a series of short chirps. Chirp rate and chirp lengths of part I songs is variable, suggesting that pulse rate is the most important species distinguishing character. (See chart at top of page.) SPECIMENS. KENYA: Sagala Hills, 17 ix 1982 (D. Otte) 26 ANSP. Matuga, ca. 17 km S Mombasa, 18 ix 1982 (D. Otte) 38 ANSP. TANZANIA: ca. 40 km W Handeni, road to Kondoa, 3 x 1982 (D. Otte) 16 ANSP. 18. Teleogryllus leucostomus (Serville) Gryllus leucostomus Serville 1839: 342. Type 6, Cassamanca, West Africa, GM. Teleogryllus leucostomus, Chopard 1961b: 277. Type examined. NOTES ON TYPE. Body color brown, more reddish or orange on venter. Dorsum of head dark reddish brown with six longitudinal stripes extending forwards to lateral ocelli. Face black above level of median ocellus, pale below. Pronotum dark reddish brown on disk, lighter on lateral lobes. Pronotum with nearly parallel sides. Forewings reddish brown, with dark veins. Hindwings broken, but extending beyond end of abdomen. 19. Teleogryllus angolensis (Chopard) Gryllus angolensis Chopard 1961a: 22. Holotype e, Dundo (Lunda), Angola (A. de B. Machado) PM. Teleogryllus angolensis, Chopard 1961 b: 277. Type examined. NOTES ON TYPE. Large, more or less uniformly reddish brown, forewings with yellow veins. Head reddish on dorsum, without longitudinal pale stripes, becoming blackish on rostrum. Face marked similar to xanthoneurus. Side of head reddish. Head as wide as pronotum. Shape of face more like xanthoneurus than like pulchriceps. Pronotum same color as head, darker on lateral lobes; disk with parallel sides. Forewing brownish, with yellow veins in basal, chordal, apical and lateral fields. Forewing is 3.2 times as long as pronotum. File with 180 teeth. Hindwings extend about 2.5 pro-

19 120 DANIEL OTTE AND WILLIAM CADE CD CL) N IZ1 11 II Ww ** 7 5 FIG. 6. T. pulchriceps (=aequipennis): top two traces represent the usual pure part I songs; mixed part I and part II songs as shown in lower trace were heard only in early afternoon. T. gnu: both songs taped late at night. T. zululandicus: both traces taped about 8 pm. notal lengths beyond forewings. Hind femora uniformly reddish brown, length 3.28 times width, as long as hindwing, and 1.14 times hind tibia length. Body length 28 mm; hind femur length 17 mm. 20. Teleogryllus zululandicus n. sp. HOLOTYPE. 6, Richards Bay, KwaZulu, South Africa, ix 1980 (D. Otte and W. Cade) ANSP. RECOGNITION. Body color dark brown to black on dorsum, pale brown to ivory on venter. Dorsum of head black, with 6 longitudinal stripes visible on occiput; occiput covered with fine pubescence, head smooth shiny black from vertex to frons. Face largely pale below level of antennae, black above this point, but median ocellus connected to pale area below by a pale band. Head dorsoventrally very short (height =.88 times head width). Side of head black above lower margin of eye, ivory below this line. Pronotum black on disk and upper half of lateral lobes; lower half of lateral lobes ivory. Front of pronotum narrower than rear. Forewings dark brown to black between Cu1 and Sc veins; membrane of lateral field pearly white between dark veins. Hindwings extending to ends of cerci. Other characters given in Table 1. DISTRIBUTION. Known only from the type locality. HABITAT. The only male found in waist high grasses near northern margins of Richards Bay among small scattered thorn trees. SONG. Two males heard. One was taped around 8 pm. Sang mostly part I songs, but produced a few phrases of part II. Local- Pulses/sec No. pulses ity I II I/II I II khz?c * * 2 part II chirps in series at 2.24 chirps/sec. SPECIMENS. Holotype 6 ANSP. 21. Teleogryllus gnu n. sp HOLOTYPE. 6, near Kirawira, Serengeti Plains, Tanzania, 20 x 1980 (D. Otte) ANSP. RECOGNITION. Body black on dorsum, ivory on sides, venter, legs, and cerci. Head black above median ocellus ivory below (Fig. 3). Pronotum black on disk, largely pale on lateral lobes. Hindwings extending about two pronotal lengths beyond end of abdomen. DISTRIBUTION. Taped and collected on the Serengeti Plains. Probably heard species on open grassy plains at noon between Kirogwe and Handeni, Tanzania. HABITAT. On Serengeti Plains found in shallow crevices and holes on an open grassland closely cropped by large herds of wildebeest and zebra. The species is probably not a burrower; females have very long ovipositors and both sexes have long hind legs, features characteristic of non-burrowing species. SONG. Part I of song peculiar-consisting of a short chirp in which pulses become progressively longer. Part II consists of a

20 AFRICAN CRICKETS 121 short succession of short chirps (Fig. 6). Most songs consisted only of part I chirps. Taped and collected at night under real or imagined threat of lions. 50 / aequipennis cj ( zululandicus Local- Pulses/sec No. pulses ity I II I/II I II khz?c 6A * * 3-4 part II chirps in series at 2-13 chirps/sec. Cs 40 0 marabu / leo grumeti SPECIMENS. Holotype d and 1Y from same locality, ANSP. 22. Teleogryllus latifrons (Karsch) 0 00 a) 0 (n xanthoneurus gnu a) 20 meru ' 0 Gryllus latifrons Karsch 1893: 150. Holotype 9, Bismarckburg, Togo, , 9.0 (R. Ruittner S.) MNHU. Teleogryllus latifrons, Chopard 196 lb: 278. NOTES ON TYPE. This female belongs to the Aequipennis Group; face and dorsum of head and coloration similar to pulchriceps. Body much smaller than pulchriceps. Hind femur length 3.41 times its greatest width. Hind tibia with 6 inner and 6 outer subapical spurs. Ovipositor as long as hind femur. Body length ca. 19 mm; hind femur 13.5 mm; cerci ca. 15 mm. OCEANIcus GROUP This group contains oceanicus, commodus, and marini from the Australian region. Possibly afer has been introduced to East Africa by man. 23. Teleogryllus afer (Saussure) Gryllus afer Saussure 1877: 159. Type 1, Baie d'algoa, Mozambique, GM. Type examined. NEW COMBI- NATION. RECOGNITION. Body color black, wings brown. Head black. Median ocellus yellow, laterally elongate. Occiput with longitudinal four pale stripes, two behind each eye. Pronotum entirely black and with nearly parallel sides. Forewings brown. Hindwings extending at least two pronotal lengths beyond end of abdomen. Legs all black. Other characters in Table 1. natalensis pulses/second part I FIG. 7. Scatter diagrams comparing songs of all taperecorded species. Note: aequipennis = pulchriceps. DISTRIBUTION. This species is remarkably similar to oceanicus and commodus from Australia, raising the possibility that it was introduced into the old port of Delagoa Bay (Lorenzo Marques) (now Maputo) by man. HABITAT. Not known. The similar oceanicus and commodus inhabit grasslands in Australia. SONG. Not known. SPECIMENS. Holotype 6 GM. GRIAULEI GROUP The two species in this group are much smaller than other species. Both are pale in color, and have the clypeus strongly arched above (Fig. 12). 24. Teleogryllus griaulei Chopard Teleogryllus griaulei Chopard 1961: 281. Holotype 8, Senegal, Rumba-Counda, 1932 (Mission M. Griaule) PM. Type examined.

21 122 DANIEL OTTE AND WILLIAM CADE 40 grumeti CZ V 0Q o CZ A gnu zululandicus CD leo w e a) 0 ** teme xr a nthoneurus CO FIG.O 8sN * 0 05 Bafricanus 0. aequip~cl)ennis meru ~~ V ~~natalensis wernerianus marabu temperature temperature FIG. 8. Pulse rates of part I phrases plotted against temperature. Note: aequipennis= pulchriceps. NOTES ON TYPE. Fig. 12. A small pale cricket (BL ca mm, hind femur length ca. 7 mm). Stridulatory file damaged, not counted. Tympana: large outer, small inner. Head with 6 pale and 5 brown longitudinal stripes. Face yellowish-brown to ivory. Clypeus strongly arched above. Cheeks brown behind eyes, otherwise pale yellowish. Pronotum: disk pale brown, with dark brown spots; lateral lobes with a dark band across upper half; ivory below. Forewings: pale, straw colored, chord 2A brown, diagonal vein and mirror veins brown, with a brown patch in outer half of mirror; mirror without dividing vein, lateral field dark brown. Hindwings extending about one hind femur's length beyond forewings. Abdomen: tergum pale brown medially, dark brown to blackish on lateral faces; sternum straw-yellow. Legs pale, straw-colored, darker on dorsal faces. Hind tibia times as long as hind femur, with 5 inner and 5 outer subapical spurs. Cerci broken. Body length ca mm (16 mm to end of hindwings); hind femur length 7 mm. 25. Teleogryllus soror (Chopard) Gryllulus soror Chopard 1940: 26. Holotype 6, N. Cameroun anglais, Mafinderi, 29 v 1935 (Mission d'6tudes de la Biologie des Acridiens) PM. Platy-

22 AFRICAN CRICKETS 123 brachypterus wittei I africanus adustus leucostomus aequipennis typographicus marabu leucostomoides zululandicus natalensis xanthoneurus gnu wernerianus angolensis burri FIG. 9. chriceps. Wing venation of dorsal field. All but brachypterus drawn to same scale. Note: aequipennis = pul- gryllus, Chopard 1961b. Type examined. NEW COMBINATION. NOTES ON TYPE. Fig. 12. This species is most similar to griaulei. It differs from that species in head and pronotal coloration, in possessing a mirror dividing vein, and in the male genitalia. Holotype with ca. 95 teeth. Tympana: large outer, small inner. Dorsum of head with a broad pale band across the vertex, without distinct longitudinal stripes on vertex or occiput. Lateral ocelli connected by broad pale band. Frons largely pale below clypeus. Upper lobe of clypeus strongly arched dorsally. Side of head pale yellowish brown. Pronotum: disk medium brown; lateral lobes dark in upper two thirds ivory in lower third. Forewings 3.00 times as long as pronotum, straw yellow to pale brown in dorsum, lateral field dark brown along upper margin; pale brown below. Hindwings more than twice as long as forewings. Abdomen: tergum blackish, but each segment with a pale posterior margin; sternum pale. Legs pale brown. Hind tibiae.66 times as long as hind femora, with 5 and 6 inner and 6 outer subapical spurs. Cerci 1.24 times as long as hind femora. Body length ca mm; hind femur length 7.0 mm. NOMEN DUBIUM 26. Teleogryllus longecaudatus Schulthess Type from Somalia is evidently

23 124 DANIEL OTTE AND WILLIAM CADE (~~~~~~~~~~~~~~~~~~~~~~~~~~~I.2~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~. to1 CC S (I ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ FIG Male genitalia: lateral dorsal, and ventral aspects. lost. It is not in the Schulthess collection at the Entomologisches Institut der ETH, Zurich. ACKNOWLEDGMENTS We wish to thank the following institutions for direct and indirect assistance in

24 AFRICAN CRICKETS \,1e1 - E ' S \ ( } \F xanthoneurus FIG. 11. Male genitalia: lateral, dorsal, and ventral aspects. Note: aequipennis= pulchriceps. this project: National Museums of Kenya (Nairobi), Kenya Agricultural Research Institute (Nairobi), Kenya Agricultural Research Station at Matuga, Desert Locust Control Organization for East Africa (Nairobi and Tengeru), Department of Zoology University of Dar-es-Salaam, Tanzania National Institute for Medical Research (Amani), Centre for Overseas Pest Research (London), Natal Parks Board (Pietermaritzburg), South African Parks Board (Kruger National Park), Hlabisa Hospital, Serengeti

25 126 DANIEL OTTE AND WILLIAM CADE C VAI~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ FIG. 12. Teleogryllus griaulei (a-f) and T. soror (g-i). a, face, showing arched epistomal suture; b, g, forewing; c, banding of head and pronotum; d, e, f, h, i, j, dorsal, ventral, and lateral aspects of male genitalia. Research Institute (Seronera), University of Witwatersrand (Johannesburg), British Museum of Natural History (London), Museum National d'histoire Naturelle (Paris), Museum d'histoire Naturelle (Geneva), Naturhistorisches Museum (Vienna). We also thank the following individuals (in order of appearance) for their assistance or support: Our wives Laurel Otte and Elsa Cade, Richard and Runi Estes, John and Jean Neen, David Ragge, Bruce Townsend, Hugh Paterson, Carl and Ruth Otte, Solveig Otte, Mark and Wendy Astrup, Orty, Camilla, Sven, and Dirk Bouquin, Karl and Margarit Rohwer, Hennie Rohwer, Bobby and Enid R6ttcher, Anna and Jock Rohwer, Engel and Hans Kruse, The Hansel Kruse family, Rob Toms, Lou Taylor, Nick Jago, Charles Dewhurst, Derek Rose, Richard Leakey, and Mark Ritchie. We thank Vickie Backus and Wade French for making numerous measurements on the specimens. We are especially indebted to Nick Jago for making the east African collecting so successful; to Carl Otte and wife Ruth for their hospitality and the use of their car; to Solveig Otte for his companionship and car; to Mark and Wendy Astrup for hospitality and for giving us free access to the Mkuze Game Reserve; to Rob Toms for assistance and companionship in the Kruger National Park. Financial support for W.C. was provided by the Natural Sciences and Engineering Research Council of Canada (Operating grant #A6174).

26 AFRICAN CRICKETS 127 LITERATURE CITED CADE, W Field cricket dispersal flights measured by crickets landing at lights. Texas Journal of Science 31: 1-6. CHOPARD, L Voyage de Ch. Alluaud et R. Jeannel en Afrique Orientale ( ). Gryllidae. Eos, Madrid 8: Gryllidae. Parc National Albert. I. Mission G. F. De Witte Fascicule 27(2): Dictyopt&res et Orthopteres recoltes en Mauritanie et dans la region du Tohad par la Mission d'etudes de la Biologie des Acridiens. Revue FranSaise d'entomologie 7: (1951). Contribution a l'etude des Gryllides du Congo Belge. Revue de Zoologie et de Botanique Africaines 41(2/3): a. Orthopteres. Gryllidae et Gryllacrididae l'angola. Companhia de Diamantes de Angola, Publicacoes Culturais 56: b. Lesdivisionsdugenre Gryllusbasees sur l'etude de l'appareil copulateur. Eos, Madrid 37: a. Contribution a la faune du Congo (Brazzaville). Mission A. Villiers et A. Descarpentries. L. Orthopteres, Grylloidea. Bulletin de Institut Franqais d'afrique noire, XXIX serie A, No. 2: b. Orthopterorum Catalogus. Pars 10. Gryllides. M. Beier, editor. W. Junk, Gravenhage. AND D. K. McE. KFVAN Orthoptera- Ensifera from northern Kenya and Jubaland. Transactions Royal Entomological Society, London 105: GERSTAECKER, A Beitrag zur Insekten-Fauna von Zanzibar. No. II. Orthoptera et Neuroptera. Archiv fur Naturgeschichte 35(1): Zweite Abteilung: Gliederthiere (Insekten, Arachniden, Myriopoden und Isopoden). Baron Carl Claus von der Decken's Reisen in Ost- Afrika. Dritter Band: Wissenschaftliche Ergebnisse. C. F. Wintersche Verlagshandlung. Leipzig und Heidelberg. Pages (section on Gryllodea). KARSCH, F Die Insecten der Berglandschaft Adeli im Hinterlande von Togo (Westafrika) nach dem von den Herren Hauptmann Eugen Kling (1888 und 1889) und Dr. Richard Buttner (1890 und 1891) gesammelten Materiale bearbeitet, mit einem Vorworte von Dr. Richard Buttner und einer Uebersichtskarte des Togogebietes. 1. Abteilung: Apterygota, Odonata, Orthoptera Saltatoria, Lepidoptera Rhopalocera. Berliner entomologischer Zeitschrift 38: KARNY, H Die Orthopteran fauna des agyptischen Sudans und von Nord-Uganda (Saltatoria, Gressoria, Dermaptera) mit besonderer Beruchsichtigung der acridoideengattung Catantops. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften (Wien). Matematisch-Naturwissenschaftliche Klasse 116(1): OTTE, D. AND R. D. ALEXANDER The Australian Crickets (Orthoptera: Gryllidae). Academy of Natural Sciences Monograph 22: POLLACK, G. S. AND R. Hoy Phonotaxis to individual rhythmic components of a complex cricket calling song. Journal of Comparative Physiology, A, 144: SAUSSURE, H. DE Melanges Orthopterologiques. 5me fascicule. III Gryllides. Memoires de la Societe de Physique et d'histoire Naturelle de Geneve 25: Orthoptera. Wissenschaftliche Ergebnisse der Reisen in Madagascar und Ostafrika in den Jahren von Dr. A. Voeltskow. Band 1: Abhandlungen herausgegeben von der Senckenbergischer Naturforschenden Gesellschaft 21, Frankfurt am Main. SCHULTHESS RECHBERG, ANTON VON Die vom Fuirst Ruspoli und Prof. Dr. C. Keller im Somalilande erbeuteten Orthopteren. Zoologische Jahrbuicher. Abtheilung fuir Systematik, Geographie und Biologie der Thiere 8: SERVILLE, JEAN GUILLAUME Histoire naturelle des insects. Orthopteres, Paris: Librairie encyclopedique de Roret. WALKER, F Catalogue of Dermaptera and Saltatoria. Vol. 1. British Museum Publication.