). 1. Teleogryllus of Eastern and Southern...

26
Proceedings of the Academy of Natural Sciences of Philadelphia 135: 102-127, 1983. African Crickets(Gryllidae). 1. Teleogryllus of Easternand SouthernAfrica DANIEL OTTE Academy of Natural Sciences of Philadelphia 19th and the Parkway Philadelphia, PA 19103 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 en- countered 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 mor- phological 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 pul- chriceps. 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 Ha- waii. Possibly only Gryllus among the Gryl- linae has a more widespread distribution. Among other groups only Gryllotalpa, An- axipha, Pteronemobius, and Oecanthus have ranges of equivalent size. Chopard's (1 967b) catalogue suggests a gap in the distribution between Africa and India. Since Teleogryl- lus 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 the Aequipennis Group differ from one another both in song and morphology. The two species which Chopard placed in Platy- gryllus (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 posti- cus. Aside from the type localities, the dis- tributions 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- 102 This content downloaded from 142.66.3.42 on Tue, 23 Jul 2013 17:16:28 PM All use subject to JSTOR Terms and Conditions

Transcript of ). 1. Teleogryllus of Eastern and Southern...

Proceedings of the Academy of Natural Sciences of Philadelphia 135: 102-127, 1983.

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

DANIEL OTTE

Academy of Natural Sciences of Philadelphia 19th and the Parkway

Philadelphia, PA 19103

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 en- countered 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 mor- phological 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 pul- chriceps. 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 Ha- waii. Possibly only Gryllus among the Gryl- linae has a more widespread distribution. Among other groups only Gryllotalpa, An- axipha, Pteronemobius, and Oecanthus have ranges of equivalent size. Chopard's (1 967b) catalogue suggests a gap in the distribution between Africa and India. Since Teleogryl- lus 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

the Aequipennis Group differ from one another both in song and morphology. The two species which Chopard placed in Platy- gryllus (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 posti- cus. Aside from the type localities, the dis- tributions 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-

102

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AFRICAN CRICKETS 103

continent, and Burma, 11 species; eastern Asia, 6 species; Indonesia, 3 species; Aus- tralia, 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 noth- ing about their songs we leave them in Te- leogryllus.

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 1932 9. burri Chopard 1961 a

10. wittei Chopard 1939 11. macrurus Walker 1869 12. adustus Karsch 1893 13. brachypterus Chopard 1967a 14. nigripennis Chopard 1948 15. 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 1839 19. 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 1961 25. 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 Na- gra 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 accord- ing to the terminology of Otte and Alex- ander 1983.

Abbreviations of museum repositories are as follows: ANSP, Academy of Natural Sci- ences of Philadelphia; BM, British Museum of Natural History; GM, Museum d'Histoire Naturelle, Geneva; PM, Museum National d'Histoire Naturelle, Paris; VM, Naturhis- torisches Museum, Vienna; MNHU, Mu- seum ftur Naturkunde, Humboldt-Univer- sitat, Berlin, DDR.

TAPE RECORDING LoCALITIES

2. Tanzania, Mt. Meru, N slope, 12 x 1980 4. Tanzania, Lake Manyara Park, 13 x 1980 5. Tanzania, Serengeti Park, Seronera, 14 x 1980 6A. Tanzania, Serengeti Park, Mbalageti R., nr.

Kirawira, 20 x 1980 7. Tanzania, Serengeti Park, Grumeti R., Kirawira,

20 x 1980 9. Tanzania, Ngorogoro Crater, 20 x 1980

11. South Africa, Natal, Eshowe, 30 x 1980 12. South Africa, Natal, Mtunzini, 7 ix 1980 14. South Africa, Natal, St. Lucia, 10 ix 1980 15. South Africa, Natal, Hluhluwe Game Reserve,

10 ix 1980 17. South Africa, Natal, Tugela R. at Middle Drift,

15 ix 1980

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104 DANIEL OTTE AND WILLIAM CADE

20. South Africa, Natal, Mkuze Game Reserve, 19 ix 1980

22. South Africa, Transvaal, Rust-de-Winter, 20 ix 1980

26. South Africa, Transvaal, Magaliesberg Moun- tains, 3 iii 1982

53. South Africa, Natal, Richards Bay, 17 iii 1982 54. South Africa, Natal, nr. Empangeni, 17 iii 1982 58. South Africa, Natal, 18 km W Empangeni, 18 iii

1982 62. South Africa, Natal, Hlabisa area, 22 iii 1982 67. South Africa, Natal, Mkuze Game Reserve, 28

iii 1982 68. South Africa, Transvaal, Badplaas, 30 iii 1982 72. South Africa, Transvaal, Clanor, nr. Skukusa Gate

of Kruger National Park, 31 iii 1982 75. South Africa, Transvaal, Kruger Nat. Park, nr.

Pretoriuskop, 2 iv 1982 78. Kenya, Hunter's Lodge, Nairobi-Mombasa Road,

14 ix 1982 82. Kenya, plain and valley S of Sagala Hills, nr.

Voi, 17 ix 1982 83. Kenya, Sagala Hills, south slope, 17 ix 1982 85. Kenya, Matuga, 18 ix 1982 86. Kenya, Shimba hills, W slope, 19 ix 1982 87. Kenya, Waa, 17 km S Mombasa, 18 ix 1982 91. Kenya, Shimba Hills, 22 ix 1982 93. Tanzania, Usa River, nr. Arusha, 25 ix 1982 95. Tanzania, Gonja, South Pare Mts., 26 ix 1982 96. Tanzania, Mombo, 27 ix 1982 97. Tanzania, Mombo, W side, 27 ix 1982 99. Tanzania, Dar-es-Salaam, 28 ix 1982

102. Tanzania, Amani, East Usambara Mts., 30 ix 1982 105. Tanzania, ca. 30 km W Kirogwe, 2 x 1982 106. Tanzania, 40 km W Handeni, 3 x 1982 108. Tanzania, 15 km S Babati, nr. Bonga, 6 x 1982 109. Tanzania, Manyara National Park, 7 x 1982

Genus TELEOGRYLLUS Chopard

Teleogryllus Chopard 196 ib: 277. Type species: Gryl- lusposticus 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). Teleo- gryllus 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 espe- cially long spermatophore tube mold; ec- toparameres 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 fol- lowed 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 eve- ning. Only T. wernerianus possess a one- part song.

OTHER FEATURES. See Otte and Alexander 1983 for terminology. Body color vari- able-reddish brown, brown, or black. Ver- tex 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 nar- rower depending on group and condition of hindwings. When hindwings are well de- veloped for flying the head is usually nar- rower than the pronotum. Head with 3 ocel- li, median one usually smaller than lateral ones. Forewings with 139-388 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 well- developed; forewings always more than .8 and less than 1.3 times as long as hind fem- ora. Hindwings variable in length, appar- ently never less than half as long as fore- wings; extending to ends of cerci in some specimens of some species. Cerci more than half as long but rarely longer than hind fem- ora. 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

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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 wer- nerianus) are closely associated with dry leaf litter, often along river beds, but also in up- land 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 grass- lands. The Australian and Pacific species (oceanicus, commodus, and marini) are all grassland species, suggesting that the mor- phologically similar T. afer lives in similar habitats. Nothing is known about the bi- ology of posticus, wittei, burri, macrurus, brachypterus, adustus, leucostomus, ango- lensis, latifrons, and leucostomoides.

Teleogryllus are some of the most acous- tically 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 for- ests 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 hindw- ings not less than half as long as the fore- wings.

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 para-

meres. 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 dor-

sal 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.

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106 DANIEL OTTE AND WILLIAM CADE

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 espe- cially difficult to distinguish; because of this and be- cause females of some species are not known, a key is not included here. Because most of Africa remains un- studied, 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

4JF

FIG. 2. Teleogryllus wernerianus.

entirely white. Legs brown, reddish, or yel- lowish. Body length of males more than 18 mm ............. .............. 2

2. Lower half of pronotal lateral lobes pale (Fig. 3) .......... ................. 3

Lower half of pronotal lateral lobes dark (Fig. 3) .......... ................. 4

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, mid- dle right). Dorsum of pronotum brown. File with over 300 teeth .......... : ......... gnu

4. Hind femur length less than three times great- est width ...............5............

Hind femur length more than three and a half times greatest width ...................... 6

5. File with 140-190 teeth ................ grumeti File with 230-290 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 ............. 7

7. Male genitalia with wider ectoparameres (Fig. 10) ......................... africanus

Male genitalia with narrower ectoparameres .............................. xanthoneurus

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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 .... ........ 2

2. Head, pronotum and legs black (Fig. 3, lower left) ............ afer

Head, pronotum, and legs with some pale col- oration ............ 3

3. Lower half of pronotal lateral lobes pale. Disk of pronotum black ........... zululandicus

Lower half of pronotal lateral lobes dark (brown to blackish reddish). Disk of pronotum red- dish brown ............ 4

4. 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 .... 5 5. 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 col- or 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, incom- plete in others; lateral ocelli connected to the eye rings by pale arms; head narrower than pronotum. Pronotum: lateral lobes sometimes much darker than disk, some- times similar in color. Forewings: Veins often yellowish especially in basal and api- cal areas. Abdomen: dorsum reddish brown, venter lighter in color sometimes orange- brown. 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.

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108 DANIEL OTTE AND WILLIAM CADE

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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 22-25 15.5 14.0 17.0 1.10 13.5 (n = 2) 23.5

meru not known

xanthoneurus 25 15.0 11.0 20.0 1.33 16.5 (n= 1)

leo 17.5-26.0 14.0 12.7 19.8 1.41 12.8 (n = 5) 23.0

grumeti 20-22 12-13 12.0 16-17 1.32 11.0 (n= 3) 20.8 12.5 16.5

natalensis 22 15.0 14.0 21.0 1.32 14.0

marabu not known

wernerianus 13-20 10.3 10.9 9.1 (n = 10) 16.1

pulchriceps not known

zululandicus not known

gnu 25 15.0 29.0 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 dif- fers from xanthoneurus, grumeti, leo, and marabu in having more slender hind femora (Tables 1, 2). Head blackish on frons. Lat- eral lobes darker than disk. Hind femur length/width = ca. 3.5.

DISTRIBUTION. Presently known from Na- tal 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, al- though the number singing is reduced dur- ing daylight hours and the songs of day sing- ers is not as loud.

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CZ

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AFRICAN CRICKETS

Pulses/sec No. pulses

Locality I II I/II I II kHz ?C

11 9.0 15.10 .60 4 11 2.7 18 11 9.4 15.14 .62 4 10 2.8 18 11(n= 3) 12.9-13.6 19.4-21.0 .65-.67 4-6 7-14 2.7-3.0 21 11 12.9 (1.14 chirps/sec) - 5-6 - 2.8 21 11(n= 6) 14.1-15.2 23.1-25.5 .59-.62 4-5 7-13 2.8-3.2 27 12 12.0 18.6 .65 4 10-12 2.8 22 14 13.9 20.3 .68 4 6 3.1 23 20 14.7 (1.63 chirps/sec) - 6 - 3.0 25 20 15.2 22.7 .67 4 12 3.3 25 20 15.2 24.3 .63 4 12-16 3.0 25 54 15.2 26.0 .58 4 12-16 3.3 25

108 10.1 15.2 .67 4 16 2.3 22 108 11.5 16.3 .71 5 11 2.8 22 108 11.3 (1.45 chirps/sec) - - - 2.7 22 108 11.0 (1.55 chirps/sec) - - - 2.7 22

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 xanthoneu- rus 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 east- ern slopes of Mount Meru in Arusha Na- tional 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 afri- canus, 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

2 9.2 20.3 .45 6 23 4.0 18 93 9.43 24.0 .39 3 (4) 24 4.3 21

9.43 - 4 - 4.1 21

SPECIMENS. Holotype 8 ANSP.

3. Teleogryllus xanthoneurus (Gerstaeck- er)

Gryllus xanthoneurus Gerstaecker 1869: 212. Lecto- type 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.

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112 DANIEL OTTE AND WILLIAM CADE

labels: "Type, Wanga [Zanzibar] v.d. Decken Ex- pedition." MNHU. Teleogryllus xanthoneurus, Cho- pard 196 lb. Type examined.

Gryllus typographicus Saussure 1877: 25. Holotype 6, Zanzibar (Hildebrandt) BM. Teleogryllus typogra- phicus, Chopard 196 lb: 278. Type examined. Type has 180 file teeth. Genitalia as in Fig. 1 1.

RECOGNITION. Difficult to distinguish from other members of the Africanus Group. Dif- fers from meru and africanus by having rel- atively wider hind femora and narrower ec- toparameres. Differs from grumeti by having more file teeth and by having a complete eye ring and by genitalia. Differs from mar- abu by having fewer file teeth, by its com- plete 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 in- habiting thick grass near trees. In the Shim- ba Hills near Mombasa abundant in grass and litter around edges of small forest patches and in litter on lightly wooded west- ern 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 in- tensively after sundown. (See chart at bot- tom 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 na- talensis, 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

4 16.5 22.6 .73 4 11-14 3.2 25 4 18.0 23.3 .77 4 9-11 3.5 25

109 20.3 30.7 .66 4 11-12 3.7 29 109 15.5-18.0 24.9-28.0 .61-65 4 7-13 3.4-3.7 25

6A 20.3 32.3 62 4 21-22 2.8 25 83 (n = 3) 15.5 23.0-25.6 .61-67 3-4 16-17 3.2-3.4 24 85 16.5 27.8 .59 4 18-21 3.3 25 85 18.8 27.7 .68 4 15-18 3.3 24 86 18.9 29.3 .65 4 16-19 3.5 25-27 86 18.0 28.7 .63 4 17 3.5 27 91 (n = 6) 18.9-20.8 29.8-31.7 .61-68 3-4 18-25 3.3-3.8 26 93 (n = 4) 18.0-20.3 25.7-26.4 .70-76 3-4 17-23 3.6-3.7 26 93 13.9 21.12 .66 4 3.4 20 93 15.8 21.5 .73 5 18 3.5 20 95 18.9 26.4 .72 5 24 3.6 27 96 17.6 23.1 .76 3-4 15 3.3 23

102 15.2 22.0 .69 4 14-16 3.3 17 105 19.8 29.7 .67 5 13-16 3.6 26 106 15.5 25.4 .61 4 11-13 3.3 21

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AFRICAN CRICKETS 113

0 1 ~~ 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.

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114 DANIEL OTTE AND WILLIAM CADE

Local- Pulses/sec No. pulses

ity I II I/II I II kHz ?C

22 19.8 31.7 .62 3 1-5 sec 3.1 21 22 16.5 (0.85 ch/sec) 4-5 2.8 21 22 17.2 (0.96 ch/sec) 4 2.6 21 58 20.3 (1.33 ch/sec) 4 3.0 25 67 19.6 (1.70 ch/sec) 4 3.0 22 72 22.0 (1.33 ch/sec) 4 3.2 prob. 25 72 20.8 (1.40 ch/sec) 4 3.2 prob. 25 72 22.0 (1.50 ch/sec) 4 2.8 prob. 25 72 22.0 (1.10 ch/sec) 4 2.7 prob. 25 72 21.0 3 3.2 prob. 25 72 20.3 35.4 .57 3 1-3 sec 3.0 prob. 25 72 22.0 36.4 .60 3 1-3 sec 3.0 prob. 25 75 18.9 (0.99 ch/sec) 4 3.1 20 75 18.0 4 2.8 20

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 oth- er 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 Na- tional 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 ap- pended directly to part I (similar to day song of leo).

Local- Pulses/sec No. pulses

ity I II I/II I II kHz ?C

7 33.0 44 .75 5 30-32 3.1 25-30 7 29.3* - 6 3.0 25-30 7 30.4** - 5 3.1 25-30

* 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 Krans- kop, 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

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AFRICAN CRICKETS 115

Pulses/sec No. pulses

Locality I II I/II I II kHz ?C

17 5.3 13.2 .40 3 18 3.5 17 17 5.4 12.9 .42 2 22 3.4 17 14 6.0 (continuous trill) 4.0 23 54 (n = 3) 6.6-6.8 20.1-20.7 .32-34 3 7-13 3.8*, 4.0 25 62 5.7 16.3 .35 3 9 3.8*, 4.0 21

* 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 Zu- luland 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. Lu- cia; sugar cane fields near Hlabisa and Em- pangeni.

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 pro- duced a part I song in the early evening consisting of a continuous train of pulses (Fig. 4). Song is most similar to that of mar- abu, 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 sim- ilar 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. Holo- type 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 consist- ing 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

78 2.93 - - 3 3.9 22 78 2.81 (1.02 pairs/sec) 2 3.8 22 78 3.22 (1.35 pairs/sec) 2 4.0 22 82 3.47 35.2 .11 2 ca. 14 3.8 25 82 3.4-3.8 (.93 pairs/sec) 2 4.0 25 82 3.67 (continuous) 4.0 25 82 3.77 (continuous) 3.7 25

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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, 26.3.05. 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 pal- pi; 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 mi- grates 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 high- lands, Kenya coast, Magaliesberg Moun- tains of the Transvaal, and the coastal grass- lands of Zululand.

HABITAT. Savanna grasslands. In Seren- geti National Park abundant in lawns sur- rounding 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 Trans- vaal collected on dry stony hillsides of Ma- galiesberg Mountains.

SONG. Song consists of widely spaced pulses. Pulse rate varies quite widely. Oc- casionally pulses are paired, with the first pulse of a pair considerably more faint; per- haps the first pulse is produced by the open- ing wing (Fig. 5). Heard only at night. (See top of page.)

SPECIMENS. TANZANIA: Serengeti Nat. Park, Ser- onera, 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

4 10.1 5.3 25 5 (n = 4) 2.5-4.5 5.3-5.5 20 9 (n= 3) 1.9-4.0 5.2-5.5 19-20

26 (n = 3) 2.64-4.55 5.4-6.0 25 26 8.25 (pairs)* 6.0 25 53 1.31 6.2 25 68 9.43 (pairs)* 6.0 25

4.13 pairs/sec 85** 2.36 5.6 23 87** (n = 4) 2.16-2.81 5.8-6.7 25-26

* 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: dor- sum of head like xanthoneurus; face as in Fig. 3. Pronotum: disk reddish brown, lat- eral lobes much darker; narrowing ante- riorly; 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 desig- nated, Ninda, Ruanda, 22 ix 1934 (G. F. de Witte) PM. Teleogryllus wittei, Chopard 1961b: 277. Type examined.

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AFRICAN CRICKETS 117

NOTES ON TYPES. Male: Reddish brown and black, legs dark, reddish. Head: vertex and occiput dark reddish brown, with 6 lon- gitudinal 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 dis- tinct 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 ma- crurus, one of Walker's series so named. Type." BM.

Teleogryllus macrurus, Chopard 1961: 278. Type ex- amined.

NOTES ON TYPE. This large female is in poor condition. By facial markings and col- or 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, Bis- marckburg, Togoland (R. Biittner S.) MNHU. Teleo- gryllus adustus, Chopard 196 1b: 277. Type exam- ined.

Gryllus ambiguus Karsch 1893: 151. Synonymized by Chopard 1961: 277. Type not examined.

NOTES ON TYPE. Figs. 9 and 10. Superfi- cially 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. Ho- lotype 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. Dor- sum of head dark reddish brown to blackish but with broad yellowish bands running from median ocellus backwards through lat- eral 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 un- divided 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 sub- apical spurs. Body length 16 mm, hind fe- mur length 12 mm.

14. Teleogryllus nigripennis (Chopard)

Gryllus nigripennis Chopard 1948: 112. Holotype Y,

Mongbwalu, Kilo, 1939 (Mme Scheitz) PM. Teleo- gryllus nigripennis, Chopard 1961b: 277. Type ex- amined.

NOTES ON TYPE. This female bears a para- type 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:

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118 DANIEL OTTE AND WILLIAM CADE

small innner, large outer. Head largely dark brown to black on dorsum, faint longitu- dinal 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, hid- den, 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 morpho- logical 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 numer- ous small cross veins between the stridu- latory 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 prono- tum 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 1869. Lectotype Y, here des- ignated, Sierra Leone, BM. Teleogryllus posticus, Chopard 1916b. Type examined.

NOTES ON TYPE. The type female was de- scribed 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 va- rieties by Walker and so could not be se- lected as the lectotype. This is unfortunate because the type specimen of the type species of Teleogryllus is now a female but the def- inition 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, Ran- dell 1964. Type examined. NEW COMBINATION.

Gryllus aequipennis Saussure 1899: 602. Type a, Zan- zibar, 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 (me- dial 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 nu- merous small pale spots. Holotype with 238 file teeth.

DISTRIBUTION. Presently known from eastern Kenya, eastern Tanzania, and Zan- zibar.

HABITAT. Open grassy areas. At Matuga, near Mombasa, males were in burrows in lawns of the Agricultural Experiment Sta- tion. In the Shimba Hills males sang in grassy openings in the forest and in grassy road- sides. West of Handeni, Tanzania, the

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AFRICAN CRICKETS 119

Pulses/sec No. pulses

Locality I II I/II I II kHz ?C

83 13.7 (.44 chirps/sec) 12 - 3.9 18-19 83 12.4 14 - 3.7 18-19 85 18.5 (.6 chirps/sec) 8-10 - 4.4 23 85 17.4 (.9 chirps/sec) 9-10 - 4.3 26 85 19.2 (.44 chirps/sec) 8-9 - 4.4 26 86 20.3 50.8* .40 7 6-7 4.2 26 86 20.0 48.0* .42 6-8 4-6 4.1 27 87 18.9 13 4.2 25 87 18.4 (.56 chirps/sec) 8 4.4 25 87 18.9 8 4.3 25 96 17.0 (.34 chirps/sec) 9 4.2 24 96 16.9 (.33 chirps/sec) 9 4.2 23 96 16.5 12 4.2 23 97 15.5 (.24-.50 chirps/sec) 10-11 4.3 24 97 17.9 10 4.2 24 99 14.9 (.42-.56 chirps/sec) 10 4.3 24

106 14.7 14 4.2 25 * 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 wood- land.

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, Cas- samanca, West Africa, GM. Teleogryllus leucosto- mus, 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 red- dish brown, with dark veins. Hindwings broken, but extending beyond end of ab- domen.

19. Teleogryllus angolensis (Chopard)

Gryllus angolensis Chopard 1961a: 22. Holotype e, Dundo (Lunda), Angola (A. de B. Machado) PM. Te- leogryllus angolensis, Chopard 1961 b: 277. Type ex- amined.

NOTES ON TYPE. Large, more or less uni- formly reddish brown, forewings with yel- low veins. Head reddish on dorsum, with- out 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. Fore- wing 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-

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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 fem- ora 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 lon- gitudinal stripes visible on occiput; occiput covered with fine pubescence, head smooth shiny black from vertex to frons. Face large- ly pale below level of antennae, black above this point, but median ocellus connected to pale area below by a pale band. Head dorso- ventrally very short (height = .88 times head width). Side of head black above lower mar- gin 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 Rich- ards 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

53 23.5 48.0* .49 8-9 4-5 4.3 25 * 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, Tanza- nia, 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 abdo- men.

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 grass- land closely cropped by large herds of wil- debeest and zebra. The species is probably not a burrower; females have very long ovi- positors 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

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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 imag- ined threat of lions.

Local- Pulses/sec No. pulses

ity I II I/II I II kHz ?C

6A 34.4 30.7* 1.12 4-6 5-7 4.0 19 * 3-4 part II chirps in series at 2-13 chirps/sec.

SPECIMENS. Holotype d and 1Y from same locality, ANSP.

22. Teleogryllus latifrons (Karsch)

Gryllus latifrons Karsch 1893: 150. Holotype 9, Bis- marckburg, Togo, 20.9-15.10, 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 pulchri- ceps. 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, commo- dus, 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 be- yond end of abdomen. Legs all black. Other characters in Table 1.

50 / aequipennis

cJ ( zululandicus

grumeti 40

0 marabu / leo Cs

0 00

a) 0 (n xanthoneurus gnu

a) 0

20 meru '

natalensis

10 20 30

pulses/second part I

FIG. 7. Scatter diagrams comparing songs of all tape- recorded 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 oceani- cus 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.

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122 DANIEL OTTE AND WILLIAM CADE

40

A gnu

grumeti

-30 - 30 -

CZ

V 0Q

o CZ zululandicus

CD leo w e

2000 3-0 20 20 a) 0 **

teme xr a nthoneurus CO

FIG.O 8sN 0 aequip~cl)ennis *

05 Bafricanus 0.

10 0 10 0

meru

~~ V ~~natalensis wernerianus

marabu 8 0

0

20 30 20 30

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. 11.5 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 yellow- ish. 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, di- agonal 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. Abdo- men: tergum pale brown medially, dark brown to blackish on lateral faces; sternum straw-yellow. Legs pale, straw-colored, darker on dorsal faces. Hind tibia 0.7 5 times as long as hind femur, with 5 inner and 5 outer subapical spurs. Cerci broken. Body length ca. 11.5 mm (16 mm to end of hind- wings); 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-

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AFRICAN CRICKETS 123

brachypterus wittei I africanus

adustus leucostomus

aequipennis

typographicus marabu

zululandicus

natalensis xanthoneurus leucostomoides

gnu wernerianus

angolensis burri

FIG. 9. Wing venation of dorsal field. All but brachypterus drawn to same scale. Note: aequipennis = pul- chriceps.

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 longitu- dinal 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 yel- low 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 mar- gin; 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. 13.5 mm; hind femur length 7.0 mm.

NOMEN DUBIUM

26. Teleogryllus longecaudatus Schul- thess 1894. Type from Somalia is evidently

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124 DANIEL OTTE AND WILLIAM CADE

(~~~~~~~~~~~~~~~~~~~~~~~~~~~I

.2~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.

to1

CC S (I ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~co

FIG. 1 0. Male genitalia: lateral dorsal, and ventral aspects.

lost. It is not in the Schulthess collection at the Entomologisches Institut der ETH, Zu- rich.

ACKNOWLEDGMENTS

We wish to thank the following institu- tions for direct and indirect assistance in

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AFRICAN CRICKETS 125

E

0

\ ,1e1 - ' 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 In- stitute (Nairobi), Kenya Agricultural Re- search Station at Matuga, Desert Locust Control Organization for East Africa (Nai- robi and Tengeru), Department of Zoology

University of Dar-es-Salaam, Tanzania Na- tional Institute for Medical Research (Amani), Centre for Overseas Pest Research (London), Natal Parks Board (Pietermaritz- burg), South African Parks Board (Kruger National Park), Hlabisa Hospital, Serengeti

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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 Mu- seum of Natural History (London), Mu- seum 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, Ca- milla, 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 nu- merous 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).

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AFRICAN CRICKETS 127

LITERATURE CITED

CADE, W. 1979. Field cricket dispersal flights mea- sured by crickets landing at lights. Texas Journal of Science 31: 1-6.

CHOPARD, L. 1932. Voyage de Ch. Alluaud et R. Jeannel en Afrique Orientale (1911-1912). Gryl- lidae. Eos, Madrid 8: 325-362.

1939. Gryllidae. Parc National Albert. I. Mis- sion G. F. De Witte 1933-1935. Fascicule 27(2): 5-15.

. 1940. Dictyopt&res et Orthopteres recoltes en Mauritanie et dans la region du Tohad par la Mis- sion d'Etudes de la Biologie des Acridiens. Revue FranSaise d'Entomologie 7: 8-30.

1948 (1951). Contribution a l'etude des Gryl- lides du Congo Belge. Revue de Zoologie et de Botanique Africaines 41(2/3): 109-125.

. 1961 a. Orthopteres. Gryllidae et Gryllacri- didae l'Angola. Companhia de Diamantes de An- gola, Publicacoes Culturais 56: 15-69.

. 1961b. Lesdivisionsdugenre Gryllusbasees sur l'etude de l'appareil copulateur. Eos, Madrid 37: 267-287.

. 1967a. Contribution a la faune du Congo (Brazzaville). Mission A. Villiers et A. Descarpen- tries. L. Orthopteres, Grylloidea. Bulletin de In- stitut Franqais d'Afrique noire, XXIX serie A, No. 2: 758-776.

. 1967b. Orthopterorum Catalogus. Pars 10. Gryllides. M. Beier, editor. W. Junk, Gravenhage.

AND D. K. McE. KFVAN. 1954. Orthoptera-

Ensifera from northern Kenya and Jubaland. Transactions Royal Entomological Society, Lon- don 105: 342-349.

GERSTAECKER, A. 1869. Beitrag zur Insekten-Fauna

von Zanzibar. No. II. Orthoptera et Neuroptera. Archiv fur Naturgeschichte 35(1): 201-223.

. 1873. Zweite Abteilung: Gliederthiere (In- sekten, Arachniden, Myriopoden und Isopoden). Baron Carl Claus von der Decken's Reisen in Ost- Afrika. Dritter Band: Wissenschaftliche Ergeb- nisse. C. F. Wintersche Verlagshandlung. Leipzig und Heidelberg. Pages 19-26 (section on Gryllo- dea).

KARSCH, F. 1893. 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 ein- er Uebersichtskarte des Togogebietes. 1. Abtei- lung: Apterygota, Odonata, Orthoptera Saltatoria, Lepidoptera Rhopalocera. Berliner entomolo- gischer Zeitschrift 38: 1-266.

KARNY, H. 1907. Die Orthopteran fauna des agyp- tischen Sudans und von Nord-Uganda (Saltatoria, Gressoria, Dermaptera) mit besonderer Beruch- sichtigung der acridoideengattung Catantops. Sit- zungsberichte der Kaiserlichen Akademie der Wissenschaften (Wien). Matematisch-Naturwis- senschaftliche Klasse 116(1): 267-378.

OTTE, D. AND R. D. ALEXANDER. 1983. The Austra- lian Crickets (Orthoptera: Gryllidae). Academy of Natural Sciences Monograph 22: 1-477.

POLLACK, G. S. AND R. Hoy. 1981. Phonotaxis to individual rhythmic components of a complex cricket calling song. Journal of Comparative Phys- iology, A, 144: 367-373.

SAUSSURE, H. DE. 1877. Melanges Orthopterolo- giques. 5me fascicule. III Gryllides. Memoires de la Societe de Physique et d'Histoire Naturelle de Geneve 25: 159.

1899. Orthoptera. Wissenschaftliche Ergeb- nisse der Reisen in Madagascar und Ostafrika in den Jahren 1889-95 von Dr. A. Voeltskow. Band 1: 569-664. Abhandlungen herausgegeben von der Senckenbergischer Naturforschenden Gesellschaft 21, Frankfurt am Main.

SCHULTHESS RECHBERG, ANTON VON. 1895. Die vom Fuirst Ruspoli und Prof. Dr. C. Keller im Soma- lilande erbeuteten Orthopteren. Zoologische Jahr- buicher. Abtheilung fuir Systematik, Geographie und Biologie der Thiere 8: 67-83.

SERVILLE, JEAN GUILLAUME. 1839. Histoire naturelle des insects. Orthopteres, Paris: Librairie ency- clopedique de Roret.

WALKER, F. 1869. Catalogue of Dermaptera and Sal- tatoria. Vol. 1. British Museum Publication.

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