Proceedings of the Academy of Natural Sciences of Philadelphia 136: 67-97, 1984

African Crickets (). 5. East and South African Species of Modicogryllus and Several Related Genera (, Modicogryllini)

DANIEL OTTE Academy of Natural Sciences of Philadelphia 19th and the Parkway, Philadelphia, PA, 19103

WILLIAM CADE Biological Sciences, Brock University St. Catharines, Ontario L25 3AI

ABSTRACT.-This paper is part of a series of preliminarypapers on the African cricket fauna. We discuss all nominal species of Modicogryllus from the Afrotropicslisted by Chopard(1967) in his catalogue, as well as fourteen new species discovered in eastern and southernAfrica. We have examined and comparedthe types of all species. Nine nominal species are moved out of this genus (see list below). Five additionalAfrican species have not been studied, either because they are from northAfrica or because the types are females; their generic status remains uncertain.Two species are moved into Comidogryllus,previously known only from Australasia. One species, royi, is moved to a new genus Modicoides. [Africa, crickets, Gryllidae, Modicogryllini, morphology, new taxa, ,songs, systematics]

This paper began when we attemptedto de- two species of Comidogryllus, and one of the termine which Modicogryllus species we had new genus Modicoides. recorded and collected in eastern and southern The paper is merely an interim reporton the Africa. Since the descriptions of previously present status of these genera. It is highly prob- described species rarely included the necessary able that numerousadditional species will even- diagnostic characters it became necessary to tually be discovered. study all of the types from the Afrotropical We have not examined many of the speci- zone. We have made new illustrations of the mens which have accumulated in various male genitalia, figured the forewings of males, museums. We believe these are best studied and counted the file teeth. Some type specimens after the songs from the respective areas are came without male genitalia-evidently they also known. While the male genitalia seem to had dropped off the points to which they were be reasonably species-specific in these genera, glued. it is often the case in Gryllinae that different This paper does not include a key to species. species have identical genitalia, leading one It does, however, include a table of comparison therefore to incorrectly identify species when and the illustrationsof male genitalia necessary no song data are available. for identification. Positive identification of The following taxa are treatedand discussed specimens from outside of those regions where here: songs have been recorded will be difficult or impossible. Modicogryllus Chopard 196 lb Chopard's 1967 catalogue lists 86 species of Modicogryllus, at least fifteen of which have 1. segnis n. sp. n. sp. since been moved to other genera (Otte and 2. serengetensis 3. conspersus Schaum 1853 Alexander 1983). In the present paper we dis- 4. Iefevrei Chopard 1938b cuss and illustrate28 species of Modicogryllus, 5. kivuensis Chopard 1939

67

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6. brincki Chopard 1955 The following African Modicogryllus have 7. parilis n. sp. not been studied and their status remains un- 8. meruensis n. sp. certain: 9. elgonensis Chopard 1938a 10. vicinus Chopard 1938b 11. minutus Chopard 1954 guanchicus Krauss 1852 (north Africa) 12. zolotarewskyi Chopard(in Chopardand Kevan 1954) palmetorumKrauss 1902 (north Africa) 13. amani n. sp. tripunctatusWerner 1908 (Tripoli) 14. vittatifrons Chopard 1961a luteus Karny 1907 (a female, Sudan) 15. zinzilulans n. sp. densivervis Chopard(a female, Ituri, Zaire) 16. maliensis n. sp. 17. kenyensis n. sp. 18. laticeps Chopard 1939 METHODS 19. nitidus Chopard 1925 All tape recordingswere made with a Nagra 20. uncinatus Chopard 1938c 21. beibienkoi Chopard 1938a SN tape recorder(8.9 cm/s) and a Sony ECM- 22. perplexus n. sp. 5OPSelectret condenser microphone attached to 23. dewhursti n. sp. a parabolicreflector. Tapes were copied onto 5 24. geonomes n. sp. inch reels by a Uher 4000 Report L tape re- 25. jagoi n. sp. corder. Sonagraphsof songs were made with a 26. garriens n. sp. 27. mombasae n. sp. Kay Elemetrics 7029A Sonograph. 28. alluaudi Chopard 1932 Temperatures were determined by placing the thermometer as close as possible to the Comidogryllus Otte and Alexander 1983 place where the cricket was singing, especially when crickets were singing in dense grass, in 29. consanguineus Chopard 1961a (moved from Mod- burrows, or on the ground. icogryllus) Body parts of crickets are named according 30. vividus Sjosted 1900 (moved from Modicogryllus) to the terminologyof Otte and Alexander 1983. Specimens examined are deposited at the UNCERTAIN STATUS following institutions: LUND, Zoological 31. Modicogryllus rotundipennis Chopard 1938a Museum; PARIS, Museum National d'Histoire Naturelle; PHILADELPHIA(ANSP), Academy of Modicoides n. gen. Natural Sciences; TERVUREN,Musee Royal de STOCKHOLM,Naturhistoris- 32. royi Chopard 19 (moved from Cophogryllus) I'Afrique Centrale; ka Riksmuseet. The following African Modicogryllus sensu Chopard 1967 are moved or will be moved to other genera (see also Otte and Alexander 1983 TAPE RECORDINGAND COLLECTINGLOCALITIES for treatmentof Australianspecies): 2. Tanzania, Mt. Meru, N slope, 12 x 1980 similis Chopard 1938 (to Velarifictorus, Otte ms) 5. Tanzania, Serengeti Park, Seronera, 14 x 1980 rhombiferChopard 1954 (to Velarifictorus, Otte ms) 6. Tanzania, Serengeti National Park, Kirawira area, lepesmei Chopard 1961b (to Velarifictorus, Otte ms) 20 x 1980 affinis Chopard 1948 (to Velarifictorus, Otte ms) 6A. Tanzania, Serengeti Park, Mbalageti R, nr. Kira- fuliginatus Chopard 1961a (to Scapsipedus, Otte and Cade wira, 20 x 1980 1984c) 9. Tanzania, Ngorongoro Crater, 20 x 1980 consanguineusChopard 1961a (to Comidogryllus,Otte and 11. South Africa, Natal, Eshowe, 30 x 1980 Cade, present work) 13. South Africa, KwaZulu, Hlabisa, 9 ix 1980 schultzei Karny 1910 (to new undescribedgenus, Otte ms) 14. South Africa, Natal, St. Lucia, 10 ix 1980 nigrodorsatus Chopard 1961b (synonym of Scapsipedus 15. South Africa, Natal (KwaZulu) Hluhluwe Game fuliginatus, Otte and Cade 1984c) Reserve, 10 xi 1980

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17. South Africa, Natal (KwaZulu), Tugela River (Mid- dle Drift area), 15 xi 1980 IDENTIFICATIONOF GENERA 18. South Africa, Natal, Howick, 16 xi 1980 Modicogryllus 19. South Africa, Natal, Karkloff area, near Howick, 17 xi 1980 1. Male forewings usually with two harp veins and a well 20. South Africa, Natal (KwaZulu), Mkuze Game Re- developed mirror,which is usually divided by one vein, serve, 19 xi 1980 but is undivided in a minority of species. 21. South Africa, Mkuze Game Reserve, Nsumu Pan, 2. Both sexes with small inner and large outer tympana. 19 xi 1980 3. Epiphalluswithout median lobes or with a single small 22. South Africa, Transvaal, Rust-de-Winter, 20 ix one. 1980 4. Ectoparameressomewhat variable, usually quite long, 46. South Africa, Cape Province, 19 km S of Mid- and extending beyond the epiphallic lobes, never as in delburg on R 57, 11 iii 1982 Figs. 13E,F. 50. South Africa, Natal, Eshowe, 13-30 iii 1982 51. South Africa, Natal (KwaZulu) Mtunzini, 15-16 iii Comidogryllus 1982 54. South Africa, Natal, nr. Empangeni, 17 iii 1982 1. Male forewings similar to Modicogryllus. 55. South Africa, Natal (KwaZulu) ca. 18 km W of 2. Epiphalluswith two small lobes between the two larger Empangeni, 17 iii 1982 lateral lobes. 56. South Africa, Natal (KwaZulu) ca. 15 km NE of 3. Tympana as in Modicogryllus. Eshowe, old Empangeni road, 18 iii 1982 4. Ectoparameresshort, each bilobate, the median lobe 57. South Africa, Natal (KwaZulu) ca. 30 km NE of fleshy. Eshowe, old Empangeni road, 18 iii 1982 58. South Africa, Natal, 18 km W of Empangeni, 18 iii Modicoides 1982 1. Males without wings. 59. South Africa, Natal, Nkwaleni Valley, N of Esho- 2. Both sexes without tympana. we, 19 iii 1982 3. Epiphallusresembles that of some Modicogryllus spec- 60. South Africa, KwaZulu, Hlabisa, 21 iii 1982 ies (Fig. 14A). 61. South Africa, KwaZulu, Hlabisa area, 21 iii 1982 4. Ectoparamereslong and slender similar to most Mod- 62. South Africa, Natal, Hlabisa area, 22 iii 1982 icogryllus. 63. South Africa, Natal (KwaZulu), Hluhluwe Game Reserve, 23 iii 1982 66. South Africa, Natal (KwaZulu), Tugela River at Genus Modicogryllus Chopard Middle Drift, 26 iii 1982 67. South Africa, Natal, Mkuze Game Reserve, 28 iii Modicogryllus Chopard 1961b: 272. Type species: Mod- 1982 icogryllus conspersus Schaum 1853: 776, by original 71. South Africa, Transvaal, Komatipoort, 30 iii 1982 designation. 76A. Kenya, Nairobi, 11 ix 1982 76B. Kenya, Nairobi, 10 x 1982 This nominal genus into which Chopard 77A. Kenya, near Nairobi, Karen, 13 ix 1982 (1967) placed 86 species has been under study Hunter's Nairobi-Mombasa 78. Kenya, Lodge, Road, for some years. Otte and Alexander 14 ix 1982 (1983) 85. Kenya, near Mombasa, Matuga, AgriculturalExpt. moved 16 of Chopard'snames to other genera. Sta. 18 ix 1982 In the presentpaper we revise the genus further 87. Kenya, Waa, 17 km S of Mombasa, 18 ix 1982 by adding 14 new species and transferring8 93. Tanzania, Usa River, nr. Arusha, 25 ix 1982 nominal species to other genera. 99. Tanzania, 28 ix 1982 Dar-es-Salaam, The distribution of the genus 102. Tanzania, East UsambaraMountains, Amani, 30 ix remains in 1982 doubt. We do know that it is widespread from 103. Tanzania, East Usambara Mountains, Sigi, below Africa to India, and at least one species known Amani, 30 ix 1982 to belong to this genus occurs on Pacific islands 109. Tanzania, ManyaraNational Park, 7 x 1982 (Fiji, Hawaii). All of the species listed from 111. Gibb off Tanzania, Karatu, Farm, Manyara- Australia(Chopard 1967) to Ngorongoro road, 8 x 1982 belong the closely 112. Tanzania, Arushaarea, Usa River, game sanctuary, related genus Comidogryllus, which we have 9 x 1982 just discovered also occurs in Africa.

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DIAGNOSTIC CHARACTERS. Forewings with brown (or black) and pale, lateral lobes often two harp veins. Foretibiaewith small inner and dark along dorsum, pale below. Forewings: large outer tympana (Comidogryllus similar). Harp with two veins (very rarely 3); mirror Male genitalia: Epiphallusbilobate, with broad divided by one vein; forewings always longer median U-shaped gap between lobes, occa- than pronotum in both sexes. Hindwings: sionally with small median lobe (Fig. 11G) or micropterousor macropterous;hindwings vary with two small median lobes (Fig. 11D); some- in length from less than pronotal length to times quite strongly modified (Fig. 13A). Ecto- where they extend well beyond abdomen. Some parameresoften long and slender, often extend- species have both micropterousand macropter- ing beyond epiphallic lobes and usually upcurv- ous forms, others may always be micropterous. ing at distal end. Front and Middle Legs: usually pale with con- OTHER CHARACTERS. Head: Occiput and ver- trasting dark markings and with stout bristles. tex usually with 6 longitudinal stripes (some- Hind femora: always stout and usually with times with additional narrow median streak); dark oblique stripes on outer face. Hind tibiae: stripes 2 and 3 often joined by oblique stripe. with 5 inner and 5 or 6 outer subapical spurs. Forehead usually with pale stripe connecting BIOLOGY. Of the species we have collected lateral ocelli. Pronotum often variegated dark most live either in thickets of grass and in leaf

Aumy. m.

FIG. 1. A, Modicogrvtllus zinzilulans male. Most Modicogrvllus species resemble this species. B, M. meruensis male.

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some of the dominant species in open savanna especially in moister depressions and along stream margins. In cities they are common in watered lawns and gardens.

1. Modicogryllus segnis n. sp.

HOLOTYPE. 6. South Africa, Natal, Eshowe. 30 x 1980 (D.Otte) ANSP.

This species and M. kivuensis have es- sentially indistinguishable songs. The separa- tion into two species is based upon the fact that the male genitalia between Tanzanian-Kenyan and South African specimens are considerably different and consistently so. When intervening populations are studied it may be discovered that a single species should be recognized. RECOGNITION. Figs. 3DE, 4, 6CD, 8AB. Tables 1, 2. Head without a transverse pale band on vertex (present in conspersus). Occiput with or without 6 pale stripes-absent in es- pecially dark individuals. Frons usually with pale stripe between lateral ocelli, indistinct to 1 t 'l~~~~~~~~~~~~~~~~~~absent in especially dark individuals. Cheeks mostly brown, especially behind eyes (pale in conspersus). Pronotum usually dark brown on dorsum, rarelydistinctly patterned;lateral lobes usually pale only in lower anteriorquarter (not forming distinct horizontal band as in (col- spersus). Legs pale brown, but without strong black markings. Hind femora, pale brown to brown, with oblique stripes, but these usually not very strongly defined. Dorsum of abdomen brown to dark brown (not patternedas in some conspersus). FIG;.2. Modcloides rovi male. DISTRIBUTION. Presently known only from South Africa but probably extends northwards litter;often we found them in soil cracks and in into East Africa. shallow excavations. None of the species makes HABITAT. Open grassy areas, often found in extensive burrows. The songs are quite vari- dry soil cracks along roadsides and open acacia able, but all have a rather rapid pulse rate, woodland. Usually found on the surface under usually delivered in short buzzy chirps. Most litter and grass clumps. species producegroups of chirps. Except possi- SONG. Widely spaced buzzy chirps. The chirp bly for kenvensis and dewhursti which were rate varies considerablysuggesting that the spe- collected in forest or forest edges, most species cies specific components are contained within a live in open grassy areas. These are usually single chirp. See Fig. 5.

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D F F

FIG.3. Modicogryllus.A, B, C, conspersusneotype; D, E, segnis paratype locality 19; F, garriensholotype.

SPECIMENS. Holotype d ANSP. Paratypes(all ANSP): Local- SOUTH AFRICA:Loc. 11, I 9 . Loc. 13, I d . Loc. 14, 3 d ity p/s ch/s p/ch KHz ?C 39. Loc. 19, 6d 79. Loc. 20, 1 9. Loc. 21, Id. Loc. 46, 11 88-98 0.2-0.3 26-40 4.7-5.0 20 lcd.Loc.54, 1d I9.Loc.55, 1d I9.Loc.57, l9.Loc. n = 4 58, 4d. Loc. 60, Id. Loc. 63, 4d 39. Loc. 66, 2d. For above. 14 108-117 0.4-0.6 25-34 5.2-5.5 23 tape recordinglocalities see tableof songs = n 4 2. Modicogryllusserengetensis n. sp. 15 108-115 26-40 5.0 24 n = 3 HOLOTYPE. c, Tanzania,Serengeti National Park, Kirawira 17 105 0.6 36 4.4 17 area, 20 x 1980 (D. Otte)ANSP. 18 72-78 0.5 32-38 4.7 15 n = 2 RECOGNITION.Figs. 4, 6E, 8C. Tables 1, 2. 19 73-86 0.3-0.4 23-35 4.5-4.8 15 Almost indistinguishable from kivuensis and n = 4 segnis, but smaller and with distinctive male 21 105-120 0.4-0.8 22-34 5.6-6.0 genitalia. The song is quite different from those n = 8 species. 22 105 0.6 27 5.2 21 DISTRIBUTION.Presently known only from the 46 90-109 0.6-0.7 33-35 5.4-5.6 23 Serengeti Plains. = n 3 HABITAT. Open grassland. At Kirawira the 54 132-143 0.4-0.5 28-29 6.2-6.4 25 species was found on the surface among clumps n = 3 of grasses cropped very short by wildebeest and 56 152 0.7 26 6.2 ca. 20 zebra. At Seroneramales were singing in longer 0.2-0.9 30-36 5.7-6.8 25 58 143-158 (Kopjes). n = 6 grass near rock outcroppings Each trill consists 61 100 0.6 41 5.3 21 SONG. Widely spaced trills. which leads 63 132, 137 0.5, 0.7 33, 36 6.2 25 of two pulse rates, slower part n = 2 suddenly into a rapid part. The two parts are 71 127 0.7 39 6.0 25 often nearly equal in duration. Trills are re- peated every 1.5 to 2 sec.

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4-) 116 I1

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5.3. S 1 qlF |zinzilulans 5 1bc

6.5 4 * * conspersus 14 23C 5.4 ~ im segnis 46 23C

4.5 @ 1 llN ~~~~~~~~~~~segnsn16 19 15C 6.8 | 1I segns

2 males 58 26C

5.4 |parilis1 1

78 25C

5.2 parilis 102 20C

5.7 | ;! 1i ama9ni 4 90 24C

5.6 ,,,lll!lllllllgllltl\ ll R 11|11111 1 garriens 99 24C

7.4 mombasae 85 24C

6.9 | 11 l l t 1111111111111 11111%111|111 |perplexus| 58 24C

5'75.7 | | | |||||||||||||\||||||||||lli Ilililililillilililililill | perplexus~~~~~~~~~~~~~~~~~~~~~~~~~21

5.2 serenget.

5.6 geonomes | 77A 21C

im b11Ikmm~~~~~~~~~~~~~~~~~~~93 ll111* 8ijl48|1*|1llllll1lllgeonomes~ 20C

4.8 dewhursti . II"'g g v ' v g s 1 . 775 21C

6.0 * | t | unknown 0 ~~~~~~~~~~~~~~~~~~~6722C

6.0 |1 1 | | | t| || ||unknown | 73 0.5sec

FIG.4. Modicogryllussongs. Numbersbeneath species names refer to tape recordinglocality and temperature.Number on left is frequency in kiloHerz.

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p/s chirp Local- length -~~~~~~~~ ity part 1 part 2 ch/s (sec) KHz ?C

5 79-83 123-139 0.5-0.8 0.44-0.61 5.2-5.8 19 n= 5 160 - 0 A 6 80, 84 135, 134 0.6, - 0.56, 0.52 5.6, 6.0 19 n = 2 00 o o

140 -0 o 0 A SPECIMENS. Holotype d ANSP. Paratypes:same locality as z 0 holotype, 1d 1 Y ANSP. o cooA 0 U) wn A u120 - a 0 * * A * zinzilulans 0 U 3. Modicogryllus conspersus (Schaum) _ o 0 segnis cP * parilis conspersus Schaum 1853: 776. Neotype G, here ZA conspersus 100 - designated, South Africa, Natal, Eshowe, 13-30 iii 1982 100 (Otte and Cade) ANSP. Modicogryllus, Randell 1964. 00* _ cu. .u A The name conspersus could be assigned to 80 A one of a number of similar species known to occur in eastern Africa from where the species I l l l l I 2 4 6 is described. Since the type is lost we have CHIRPS/SECOND chosen the species whose male genitalia most closely match those illustrated by Chopard FIG.5. Scatter diagram showing some similarities and differences in song parametersamong four Modicogryllus 1961b to bear the name conspersus. The name species with short chirps and fast pulse rates. The scatter appearsquite frequentlyin the literaturebut we within species is due to temperatureeffects. The pairs believe that it is probably applied to a number segnis-parilis and conspersus-zinzilulans have similar of species. By designating a neotype whose songs, but they are morphologically distinct and have song is known we can make the determination allopatric distributions. of the species more certain. RECOGNITION. Figs. 3ABC, 4, 5, 6AB, 8D- G. Tables 1, 2. Head with pale band across vertex (absentin segnis). Occiput always with 6 DISTRIBUTION. Presently known only from stripes. With distinct pale line connecting later- South Africa. al ocelli. Face with pale streakdescending from HABITAT. Open grassy areas. In South Africa median ocellus, often widening in lower half of inhabiting grassy areas from the low coastal frons. Cheeks usually pale, including area be- thorn scrub to the cool eastern short grass veld. hind eye. Dark below eye. Pronotumstrongly In Tanzania we have found the species only in variegatedon dorsum; lateral lobes with black the Ngorongoro and Serengeti areas. We sur- horizontal band on upper 1/2 to 3/4. Dorsum of mise that it is usually more abundantin upland abdomen often patterned with pale and dark grassy areas than in low hot regions. brown. Legs usually very pale with black spots SONG. Usually groups of buzzy chirps (3-7 and markings. All specimens listed below were chirps per group) but occasionally producinga micropterous. Female ovipositor distinctly continuous train of chirps. Chirp groups sepa- longer than in segnis (Table 2). rated by 2 to 6 seconds. See also Fig. 5.

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K F G H

L M N 0 P

Q ~~~R S T

FIG.6. Modicogryllusmale forewings. A, conspersus, loc. 18; B, conspersus, loc. 11; C, segnis, loc. 21; D, segnis, loc. 11; E, serengetensis holotype; F, lefevrei holotype; G, kivuensisholotype; H, kenyensisholotype; J, brinckiholotype; K, parilis, loc. 78; L, parilis, loc. 9; M, meruensis paratype;N, elgonensis holotype; 0, vicinus holotype; P, minutus holotype; Q, zolotarewskyiholotype; R, beibienkoi holotype; S, amani holotype; T, vittatifronsholotype; U, zinzilulans holotype.

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F ~~~~~~H JK ~

N

L ~~~M

FIG.7. Modicogryllus and Comidogryllusmale forewings. A, M. laticeps holotype; B, M. nitidus holotype; C, M. uncinatus, Sierra Leone; D, M. perplexus, loc. 58; E, M. dewhursti holotype; F, M. geonomes, loc. 22; G, M. jagoi holotype; H, M. garriens holotype; J, M. mombasae holotype; K, M. alluaudi holotype; L, undescribedspecies from SierraLeone; M, Comidogryllusconsanguineus holotype; N, M. finoti holotype;0, Comidogryllusvividus holotype; P, M. conspersus, loc. 52; Q, M. maliensis holotype.

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TABLE2. Comparisonof females. Key to abbreviationsin Table 1. Italic numbers are means.

No. subapical

BL F3L FWL OL spurs (mm) (mm) PL F3L inner outer

Modicogryllus conspersus 13.0-16.5 6.5-9.0 2.82 1.14-1.60 4,5 5,6* n = 20 1.37 segnis 10.5-16.0 6.5-8.5 2.61 1.01 5,6 6,7* n = 19 brincki 14 6.8 2.39 1.47 5 5 parilis 10.5-12.0 6.0-7.0 1.7-2.3 1.07-1.25 5 5,6* n = 3 meruensis 12.0-14.0 6.0-7.5 1.6-2.0 ca. 1.00 5 5 n = 6 serengetensis 12.0-,12.5 6.5,7.0 2.5-2.5 1.00-1.08 5 6 n= 2 beibienkoi 13 7.5 2.66 ca 1.00 4 5 dewhursti 15 8.7 0.3 1.15 5 5 geonomes 13.0-14.0 6.0-7.0 2.7-3.3 0.93-1.17 4,5* 5,6* n = 3 perplexus 12.0-13.5 6.0,- 3.0-3.3 1.17,- 5 6 n = 2 zinzilulans 9, 11 7.0,7.5 ca. 2.7 0.57,0.53 5 6 n = 2 amani 16.5 9.3 1.7 0.86 5 6 rotundipennis 14.5 8.5 1.0 0.94 5 5 HT alluaudi 12.0 7.0 2.50 0.93 5 5 Comidogryllus vividus (PLT) 16.5 11.5 1.24 0.78 5 5

*Most common condition.

SPECIMENS. Neotype d ANSP. SOUTH AFRICA (all Local- ANSP): Loc. 11, 3d 49. Loc. 14,3d lY. Loc. 18,4d 8Y. ity p/s ch/s p/ch KHz ?C Loc. 50, 3d 5?. Loc. 52, 2d. Loc. 54, 16. Loc. 58, 1Y. Little Berg, 1860 m. 1d. Port St Johns, 2 i 1981 (R. Toms) 11 123 3.1 19 6.0 18 1 Y. For tape recording localities see table of songs. 11 150 4.3 15-18 6.3 21 11 135-157 3.6-4.7 13-20 6.1-7.3 23 4. Modicogryllus lefevrei (Chopard) n = 3 14 132-152 3.4-4.5 16-19 6.5-6.9 23 Grylluluslefevrei 1938: 342. Holotype 6, Congo, Urundi, n = 3 Kitega (Lefevre) TERVUREN. Modicogryllus, Chopard 50 165 3.8 16-21 7.5 25 1961: 273. Type examined.

52 180 5.4 16-18 8.0 27 NOTES ON HOLOTYPE. Figs. 6F, 9A. Table 1. 18 82-92 1.8-2.2 21-23 4.8-5.6 18 Similar to other species in conspersa complex. n = 4 Dorsum of head with six pale stripes, with 19 75-84 1.6-1.7 14-23 4.7-4.9 15 transversepale band crossing head along vertex

This content downloaded from 142.66.3.42 on Tue, 23 Jul 2013 17:24:35 PM All use subject to JSTOR Terms and Conditions MODICOGRYLLUS AND RELATED GENERA 79 connecting posteriormargins of eyes. Face pale occiput; pubescent on occiput; forehead with aroundmedian ocellus and with triangularpale pale stripe (reddish stripe between the lateral markingabove clypeus. Pronotum:dorsum var- ocelli). Face black but with narrowreddish line iegated; lateral lobe black along upper two- descending from median ocellus. Cheeks black thirds, pale below. Abdomen: dorsum dark but with horizontal reddish streak just above brown to black, venter pale brownto ivory. File jaws. Pronotum pubescent, lateral lobes with with ca. 110 teeth. Hindwings about two thirds pale strip along lower anterior corner. Fore- as long as forewings. wings: file with ca. 98 teeth; dark brown to ALLOTYPE (FEMALE). Similar to male in color blackish, right wing with 3 veins (inner or Cu2 but more pubescent; perhapsbelongs to differ- lacking) right wing chord Cu2 present but con- ent species even though collected at same local- necting to mirror.Hindwings hidden, abouthalf ity. as long as forewings. Abdomen black on dor- sum and venter. Legs all black. Foretibiaewith large outer and small inner tympanum. Hind tibiae with 5 inner and 5 outer subapical spurs. 5. Modicogryllus kivuensis (Chopard) Cerci dark brown. Gryllulus kivuensis Chopard1939: 10. Holotype 6, Albert ALLOTYPE (FEMALE). Black like male but dor- National Park, Kivu, Rutshuru,15-25 ix 1937 (G. F. de sum of pronotum with definite light and dark Witte) TERVUREN. Modicogryllus, Chopard 1961b. Type markings. Lateral lobes with pale area along examined. lower margin, widest at anterior end. Hind- NOTES ON HOLOTYPE. Figs. 6G, 9B. Table 1. wings extending beyond forewings. Similar to other species in conspersus group. Head: dorsum with six stripes on occiput and vertex, and with broadpale band on vertex that 7. Modicogryllus parilis n. sp. crosses head and extends onto cheeks; black on forehead but with pale stripe connecting lateral HOLOTYPE. d, Kenya, Hunter'sLodge, Nairobi-Mombasa ocelli; face with triangular pale area below road, 14 ix 1982 (D. Otte) ANSP. median ocellus; cheeks pale, becoming black at RECOGNITION. Figs. 4, 6KL, 8DEF. Tables 1, pronotal margin. Pronotum:dorsum variegated 2. This species is virtually indistinguishable black and pale brown; lateral lobes black in from segnis. The two species differ only in the upper two thirds, ivory below. Abdomen: dor- male genitalia. sum black, venter medium brown. Forewings DISTRIBUTION. Extremewestern Zaire to east- brownish, translucent, black along upper side ern Kenya and Tanzania, but probably wide- of lateralfield. Hindwingstwo thirdsas long as spread through east Africa. The contact be- forewings. tween parilis and segnis remains to be studied (see introductionto segnis). HABITAT. Grassy areasespecially where grass is short and forms thick ground cover. Found 6. Modicogryllus brincki (Chopard) both in lowland and upland areas. At Hunters Acheta brincki Chopard 1955. Holotype o, South Africa, Lodge, Kenya, found in dense grasses near Cape Province, Cape Flats, 1 mi E of Zeekoevlei, 8 xii water but not in adjacent dry grassland. In 1950 (Brinck, Rudebeck) LUND. Modicogryllus, Randell Nairobi common in lawns and along roadsides. 1964. Type examined. SONG. Virtually identical to that of segnis. NOTES ON HOLOTYPE. Figs. 6J, 9C. Tables 1, The separation into two species is based on 2. Small black micropterouscricket. Head: dor- differences in the male genitalia. See also Fig. sum with stripes 1, 2, and 3 faint on vertex and 5.

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Local- HABITAT. Ground litter in tall cloud forest, ity p/s ch/s p/ch KHz ?C along jeep road, near crater basin. SONG. Not known. 9 83-91 0.4-1.0 15-18 4.3-4.9 19 SPECIMENS. Holotype d ANSP. Paratypes: n = 5 same locality as holotype, 1 d 6 Y ANSP. 76A 81-91 0.7 14-17 4.4-4.7 20 n = 2 76B 91-97 0.8-1.4 13-17 4.1-4.6 20 9. Modicogryllus elgonensis (Chopard) n = 2 Gryllulus elgonensis Chopard 1938: 125. Holotype 6, 78 104-114 0.8-1.3 17-25 5.2-5.7 21 Kenya, Elgon Saw Mill, Mt. Elgon, Ver Est, Camp II, n = 4 2470 m, 1932-1933 (Mission de l'Omo) PARIS. Mod- 78 100-118 0.8-1.4 19-28 5.4-5.5 25 Day icogryllus, Randell 1964. Type examined. _ c n = J NOTES ON TYPE. Figs. 6N, 15D. Table 1. 85 158 0.8 35-37 6.5 24 Genitalia not with specimens; the figure given 93 92-101 0.6-1.0 14-25 4.7-5.2 20-21 n = 3 in Chopard 1961b of this species indicates that 102 102-122 0.2-0.6 33-35 5.0-5.3 17 it is distinct from all other species discussed n = 4 here, for it has bifurcatelateral epiphallic lobes. 109 115, 116 1.3-1.0 19-21 5.7, 5.5 25-29 Small dark brown cricket. Dorsum of head n = 2 blackish, with 6 strong occipital stripes, these 111 88 0.9 21-22 4.3 20 reach anteriorto vertex. With white stripe be- tween lateral ocelli. Cheeks dark brown. Pro-

SPECIMENS. Holotype d ANSP. Paratypes(all ANSP): TAN- notal disk mostly dark brown but with lighter ZANIA: Loc. 9, 4 3 ?. Loc. 102,1 d. Loc. 103, 1J. Loc. spots. Laterallobes darkbrown except for small 109, 1 1?. Loc. 111, Id. Loc. 112, 1d 3Y. KENYA: pale patch in lower anteriorcorner. Hindwings Loc. 76A, 2d 5Y. Loc. 76B, Id I Y. Loc. 77A, Id. Loc. hidden. All femora pale brown with distinct 78, 5 d 7 ?. For tape recordinglocalities see table of songs. dark brown patches and stripes. Dorsum of abdomen black; venter brown, becoming dark 8. Modicogryllus meruensis n. sp. distally.

HOLOTYPE. 6, Tanzania, Mt. Meru, N side cloud forest, 12 x 1980 (D. Otte) ANSP. 10. Modicogryllus vicinus (Chopard)

RECOGNITION. Figs. 6M, 9E. Tables 1, 2. Gryllulus vicinus Chopard 1938: 343. Holotype 6, Congo Dark cricket with head wider than pronotum. [Zaire] Ituri-Nioka, 1934 (Lefevre) TERVUREN. Mod- Dorsum of head with 6 distinct stripes on occi- icogryllus, Chopard 1961: 273. Type examined. put, withouttransverse band across vertex, with NOTES ON HOLOTYPE. Figs. 60, lOA. Table 1. strong white stripe between lateral ocelli. Pro- Size medium for genus. Body, especially pro- notal disk generally dark but distinctly pat- notum, abdomen, and legs, very pubescent. terned with light and dark areas, lateral lobes Head:dorsum with 6 pale stripeson occiput; on pale at lower anteriorportion, sometimes pale vertex a broad pale band crosses head and area is minute. Dorsum of abdomendark brown descends behind eyes onto cheeks; forehead to blackish or patternedlight and dark. Front black but with ivory band connecting lateral and middle legs with large dark patches; hind ocelli; face with vertical dumbell-shapedpale femur with strong dark oblique stripes. Fore- area, lower part triangular.Pronotum: dorsum wings in female meeting at midline, about 1.5- variegated; lateral lobes black in upper two 2.0 times as long as pronotum;hindwings invis- thirds, ivory below. Forewings black only ible. along uppermargin of lateralfield; with ca. 110 DISTRIBUTION. Known only from type local- teeth. Hindwing about one half as long as ity. forewing. Abdomen: dorsum black, venter

This content downloaded from 142.66.3.42 on Tue, 23 Jul 2013 17:24:35 PM All use subject to JSTOR Terms and Conditions MODICOGRYLLUS AND RELATED GENERA 81 pale. Forelegs with irregulardark markingson ing as follows: vertex without transverse pale inner and outer faces, densely pubescent. Hind band; cheeks posterior to eyes dark brown femur with dark brown oblique stripes. Cercus (largely pale in conspersus). 0.82 times as long as hind femur. DISTRIBUTION. Known only from the type locality. 11. Modicogryllus minutus (Chopard) HABITAT. In lawns in forest clearing, on sur- face under clumps of grass. Scapsipedus minutus Chopard 1954: 91. Holotype , SONG. Groups of short chirps; 3-5 chirps per Guinea, Nimba, xii-vii 1951 (Lamotte and Roy) PARIS. Modicogryllus, Chopard 1967: 78. Type examined. group.

NOTES ON TYPE. Figs. 6P, lOB. Table 1. Small dark brown to blackish cricket. Face Locality p/s ch/s p/ch KHz ?C long. Dorsum of head with 6 pale stripes on 102 88 2.2 6 5.3 19 occiput and with distinct white band between 99 11 2.7 9-10 5.7 24 lateral ocelli. Cheeks blackish. Pronotal disk with brownmarks; later- darkbut patterned pale SPECIMENS. Holotype 5 ANSP. Paratypes:same locality as al lobes blackish but with small pale area in holotype, 25 1 Y ANSP. lower anterior quarter. Abdomen blackish. Front and middle legs with large darkbrown to 14. Modicogryllus vittatifrons (Chopard) blackish markings. Hind femur with oblique Gryllus vittatifronsChopard 1961a: 19. Holotype 5, An- Hindwings tiny. dark stripes. gola, Dundo, PARIS. Modicogryllus, Chopard1961: 274. Type examined. 12. Modicogryllus zolotarewskyi (Chopard) NOTES ON TYPE. Figs. 6T, IIB. Table 1. Acheta zolotarewskyi Chopard 1954: 344. Holotype 5, Dorsum of head with strong occipital stripes. Batha, Ati, French Chad, 13?14'N, 18018'E, 14 xi 1933 With white stripe between lateralocelli. Cheeks (Mission d'Etudes de la Biologie de Acridiens) PARIS. dark posterior to eyes (largely pale in zinzilu- Modicogryllus, Chopard 1961. Type examined. lans). Dorsum of pronotumpatterned with light NOTES ON TYPE. Figs. 6Q, IOC. Table 1. and darkbrown, but mostly dark. Laterallobes Body strongly pubescent. Dorsum of head uni- mostly dark brown, but with light patch in formly dark reddish brown; occiput without lower anteriorquarter (entire lower marginpale pale stripes; without pale line between lateral in very similar zinzilulans). Hindwings extend- ocelli. Pronotum dark brown but with slightly ing beyond ends of cerci. lighter anterior margin and somewhat paler areas at posterior lateral corners of disk and 15. Modicogryllus zinzilulans n. sp. lower anteriorcorners of laterallobes. All three 14 x femora dark brown, but hind femur pale along HOLOTYPE. 5, Tanzania, Serengeti Plains, Seronera, 1980 (D.Otte) ANSP. venter. Hindwings extending well beyond end of abdomen. Head wider than pronotum. RECOGNITION. Figs. 1, 4, 6U, lIC. Tables 1, A male of this species was collected at Kara 2. Small crickets with contrastingmarkings on Macina, Mali, on 4 ix 1983 by Mungai and head and pronotum and pale legs. Dorsum of Jago, ANSP. head blackish with 6 conspicuous occipital stripes. Stripe between lateral ocelli variable, 13. Modicogryllus amani n. sp. sometimes prominent, sometimes very thin. Pronotal disk patterned with light and dark HOLOTYPE. 5, Tanzania, East Usambara Mountains, markings; lateral lobe with horizontal black Amani, 30 ix 1982 (D. Otte) ANSP. band along dorsal half, ivory below (in holo- RECOGNITION.Figs. 4, 6S, lODE. Table 1. type pale area confined to lower anteriorquar- Very similar in color to conspersus, but differ- ter). Femora pale, whitish, with few dark spots

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FIG.8. Modicogryllusmale genitalia, showing dorsal, ventraland lateralaspects. A, segnis, lOC.2 1; B, segnis, lOC.I 1; C, serengetensis, loc. 6; D, conspersus, lOC.I 1; E, conspersus, lOC.52; F, conspersus lOC.18; G, conspersus? lOC.102. and markings. Hind femora dark at distal end, posterior to eyes with large pale areas. with dark brown oblique stripes on outer face. DISTRIBUTION. Known only in the Serengeti Abdomen dark brown to black on dorsum, pale Plains, Tanzania. on venter. HABITAT. Short grass plains. Very similar to vittatifrons but differing as SONG. Groups of chirps. Chirps vary in num- follows: genitalia somewhat narrower,ectopar- ber from 2 to 21 per group, but usually there are ameres less upcurved; lateral lobes with con- 3 to 5 per group. See also Fig. 5. tinuous pale band across the ventral third; area

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SPECIMENS. Holotype d ANSP. Paratypes:same locality as Local- holotype, 2d 2 ANsP. ity p/s ch/s p/ch KHz ?C 5 87.9 2.24 15-16 5.25 19 16. Modicogryllusmaliensis n. sp. 6 83.9-95.3 1.6-2.7 15-24 5.3-6.8 19 HOLOTYPE. 6, Mali, Kara Macina, 4 xi 1983 (Mungai and n = 4 Jago) Asp. 6A 79.1 1.1-1.2 27-28 4.7 25 RECOGNITION. Figs. 7Q, I ID. Table 1. Generally pale cricket with dark brown mark-

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FIG. 9. Modicogryllusmale genitalia.A, lefevreiholotype; B, kivuensisholotype; C, brinckiholotype; D, parilis paratype,Joe. 78; E, parilis parataype, Joc. 9; F, parilis paratype, loc. 103;G, meruensis holotype.

ings. Head: dorsum dark brown to blackish but with 5 large dark brown patches; lateral lobes with 6 distinct pale stripes on occiput and ver- with blackish band along dorsal third, ivory tex; on vertex they coalesce laterally to form below. Forewings medium brown, translucent continuoustransverse band which runs between in dorsal field, pale below. Hindwings at least inner posterior margins of eyes but does not twice as long as forewings. Abdomen: dorsum continue behind eyes; forehead with broad pale transverselybanded with dark brown and ivory; band between lateral ocelli. Face largely ivory venter pale brown. Front and middle legs: fore- colored, but dark above median ocellus, along tibiae with large outer and small inner tym- inner margins of antennal socket and medially panum; femora almost entirely ivory colored just above epistomal suture;mouth parts ivory; with dark bristles. Hind femur with dark brown cheeks ivory, but with dark brown dorsal post- oblique stripes on outer face; inner face with ocular band, which does not reach pronotal two dark patches in distal half. margin. Pronotum: dorsum largely pale and Very similar to zinzilulans and vittatifrons;

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differs from the former in having differently 17. Modicogryllus kenyensis n. sp. shaped epiphallic lobes, a transverse band HOLOTYPE. Kakamega Forest Reserve, 26 x across the top of the head; differs from vittati- 6, Kenya, 1982 (N. D. Jago) ANSP. frons in shape of epiphallic lobes, in having pale cheeks, a mostly pale pronotaldorsum; and RECOGNITION. Figs. 6C, 1 A. Table 1. Based a pale transverseband across top of head. on one male. Dorsum of head with 6 stripes on DISTRIBUTION. Presentlyknown only from the occiput, these fuse indistinctly on vertex type locality. suggesting transverse band as in conspersus. White band between lateralocelli. Frons entire- SPECIMENS. Holotype d ANSP. ly black. Cheeks largely black. Pronotal disk

This content downloaded from 142.66.3.42 on Tue, 23 Jul 2013 17:24:35 PM All use subject to JSTOR Terms and Conditions 86 DANIEL OTTE AND WILLIAM CADE strongly patternedwith pale brown and black; NOTES ON TYPE. Figs. 7B, I IF. Table 1. lateral lobes black, with small pale area in Body color entirely black. Dorsum of head lower anterior quarter. Dorsum of abdomen without occipital stripes and without pale stripe black. Front and middle legs pale and with between lateral ocelli. Hindwing tiny. pronounced black spots and markings. Hind femur with strong dark brown. Oblique stripes on outer face. 20. Modicogryllus uncinatus (Chopard) DISTRIBUTION. Reported from the type local- Scapsipedusuncinatus Chopard 1938: 57 Lectotype 6, here ity and from Bitshumbiand Rutshuruin western designated, SierraLeone, Njala, iii 1930 (E. Hargreaves) Zaire and from the Algada area of Kenya. We PARIS. Modicogryllus, Chopard 1961: 274. record it here from the Kakamega area east of NOTES ON TYPE. Figs. 7C, 11G. Table 1. A Lake Victoria. brown species. Dorsum of head with 6 occipital HABITAT. Jago collected one male in the stripes; and with strong pale band between Kakamega Forest Reserve, apparentlyin leaf lateralocelli. Cheeks brown but with pale band litter. descending vertically from posterior margin of SONG. Not known. eye. Pronotaldisk patternedwith light and dark brown;lateral lobes mostly darkbrown but with SPECIMENS. Holotype d ANSP. pale area in lower anterior quarter. Legs pale with brown spots; hind femur with oblique 18. Modicogryllus laticeps (Chopard) stripes. Lateralfield of forewings darkbrown in upper third, pale below. Grylluluslaticeps Chopard1939: 11. Holotype 6, Ruanda, A male from Tshela, Congo 12 xi Ninda (2150) 21-22 ix 1934 (G. F. de Witte) TERVUREN. [Zaire] Acheta, Chopardand Kevan 1954. Modicogryllus, Cho- 1920, PARIS, was also seen. It resembles the pard 1961: 273, Type examined. lectotype closely.

NOTES ON HOLOTYPE. Figs. 7B, IIE. Table 1. Size small for genus. Head: dorsum shiny 21. Modicogryllus beibienkoi (Chopard) black, with six pale stripes on occiput and vertex; forehead with white stripe connecting Gryllulusbeibienkoi Chopard 1938: 124. Lectotype c, here lateralocelli; face black but pale aroundmedian designated, Ethiopie Merid, Bourie, bord de la Riv. ocellus and with thin pale streak descending Omo, 600 m, (Mission de l'Omo) PARIS. Modicogryllus, 1961b. examined. middle of face; cheeks mostly black but with Chopard Type pale brown line descending from middle poste- NOTES ON TYPES. Figs. 6R, 15E. Tables 1, 2. rior margin of eye to mandibles. Pronotum: Male: Genitalia not with type. We cannot tell dorsum: anteriorhalf black, posteriorhalf var- whetheror not this species is the same as one of iegated black and ivory; lateral lobe all black, the species described here as new. Brownish but with tiny pale spot at lower anteriorquarter. cricket. Dorsum of head with 6 occipital stripes Forewings: with ca. 106 teeth, dark brown on and with pale band running between lateral dorsum, blackish on lateral field. Hindwing ocelli. Dorsum of pronotumlargely brown but about one third length of forewing. Front and with some lighter brown spots; lateral lobe middle legs dark brown to black and with pale mostly brown but slightly paler in lower anteri- spots. Hind femur: outer face blackish. or corner. All femora unicolorous light orange brown. Harp with two veins. Mirror once di- 19. Modicogryllus nitidus (Chopard) vided. Hindwings extending well beyond abdo- men. Gryllus nitidus Chopard 1925: 291. Holotype d, Algeria, Female: Very similar to male in color and Sidi-Bel-Abbes (la Moult) PARIS. Gryllulus, Chopard 1943: 188. Modicogryllus, Chopard 1961: 273. Type size and wing length, but lateral lobes pale in examined. entire lower half.

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FiG. 10. Modicogryllusmale genitalia. A, vicinus holotype; B, minutusholotype; C, zolotarewskyiholotype; D, amani holotype; E, amani paratype, loc. 99.

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FIG. 11. Modicogryllus male genitalia. A, kenyensis holotype; B, vittatifronsholotype; C, zinzilulans holotype; D, maliensis holotype; E, laticeps holotype; F, nitidus holotype; G, uncinatus lectotype.

22. Modicogryllusperplexus n. sp. DISTRIBUTION. Eastern Lowveld of South Africa. HOLOTYPE. 6, South Africa, Transvaal, Komatipoort,30 iii HABITAT. areas, usuallywhere 1982 (Otte and Cade)ANSP. Open grassy there are numerous soil cracks. Common RECOGNITION.Figs. 4, 7D, 12AB. Tables 1, aroundseasonally wet pans. indistinguishable 2. Very similar and largely SONG. Shorttrills consisting of two parts:the from geonomes and segnis, but differing in the firstthird to two thirdsconsists of a simpletrain male genitalia. Also, lateral lobes often with a of pulses;the second partconsists of pairsof large pale area in the lower anteriorquarter, and pulses. pale line connecting the lateral ocelli is usually very narrow.

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p/s trill length Locality Ist 2nd part pairs/s tr/s (sec) KHz ?C

15 60.3 81.4 35.2 .77 .45 5.7 24 21 58,59 79,78 31.4,30.5 0.9,0.9 0.57,0.65 5.7,5.7 21 n = 2 58 66,60 107,101 38.8,37.0 0.8,1.4 0.46,0.53 7.1,6.9 25 n = 2 63 62.9 86.5 35.5 1.2 .55 6.5 25 67 58.1 88.0 35.2 1.0 .53 6.3 22 71 76.4 107.3 40.6 .73 .46 6.7 25'

'Daytime, somewhat overcast, ground temperaturecould have been much higher.

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SPECIMENS. Holotype d ANSP. Paratypes (all ANSP): RECOGNITION. Figs. 4, 7F, 12D. Tables 1, 2. SOUTH AFRICA:Loc. 59, 15 1 Y. Loc. 63, 25 . Loc. 67, Dorsum of head with 6 occipital stripes, with- 1d. Loc. 71, 15 1 Y. For tape recordinglocalities see table out transverse band on vertex; with distinct of songs. stripe between lateral ocelli. Pronotum pat- terned with light and dark brown on dorsum, 23. Modicogryllus dewhursti n. sp. usually with larger pale areas in posterior half; HOLOTYPE. 5, Kenya, Karen (nr. Nairobi) 13 ix 1982 (D. lateral lobes almost entirely dark, but usually Otte) ANSP. with small pale area in lower anteriorcorner. RECOGNITION. Fig. 4, 7E, 12C. Table 1. Legs pale brown with brown markings. Hind Male: Very dark cricket but with pale venter. femora with oblique brown stripes on outer Head generally black but becoming lighter on face. Dorsum of abdomendark brown to black- mouthparts. Occiput with faint traces of 6 ish. This species differs from very similar seg- stripes. With thin brownish line connecting nis mainly in male genitalia, but also in having lateral ocelli. Pronotumlargely black but with more distinct stripes on occiput, more pro- traces of lighter color on disk; lateral lobes nounced pale patterns on pronotal disk and black. Abdomen blackish. Front and middle darker lateral lobes. legs with large blackish area on dorsum and DISTRIBUTION. Widespreadover eastern Afri- proximal areas but containing pale spots; basal ca from Zululand to Kenya. and ventralareas pale. All tibiae blackish. Hind HABITAT. Open grassland. Usually found on femur darkbrown, but pale at base, with strong the surface under clumps of grass or in litter. dark oblique stripes on outer face. Cerci dark SONG. A very rapid succession of 3 and 4 brown. pulse chirps. Chirps delivered either in groups Female: Presumed to be conspecific with or in a continuoustrain. The first and last chirps male. Absence of inner tympanumsuggests she of a series usually with more pulses. may belong to different species. Considerably lighter in color, with pale stripesbetween poste- Locality p/s ch/s p/ch KHz ?C rior margin of eyes and pronotum. Pronotum like male. Forewings tiny, about one third 5 142.6 29.3 3-6 6.0 20 6 158.3 29.8 3-7 6.5 19 length of pronotum.Legs generally pale brown 14 137.0 25.4 3-6 6.1 23 with some dark spots. Hind femur spotted with 17 141.3 28.4 3-4 6.25 17 dark brown on dorsum and striped with dark 20 188.4 38.3 3-12 6.5 25 brown on outer face. 21 152.2 28.1 3-5 6.0 21 22 143.9 28.3 3-5 6.8 22 DISTRIBUTION. Known only from the type 50 155.3 25.8 3-4 6.5 22 locality. 51 168.5 35.4 3-4 6.8 25 HABITAT. In litter and grass in a garden sur- 62 167.2 34.4 3-5 7.0 21 rounded by large trees and forest. Probably a 77A 113.14 20.3 4-7 5.6 18-21 forest floor or forest edge species. 87 162.5 33.0 3-11 6.8 25 6.7 SONG. Not known. Possibly the song shown 93 148.5 28.8 3-10 20 in Fig. 4, third from bottom belongs to this SPECIMENS. Holotype d ANSP. Paratypes(all ANSP): TAN- species. ZANIA: Loc. 6, 1 d. SOUTH AFRICA:Loc. 21, 1Y. Loc. 22, 1 d. Loc. 50, 2/. Loc. 51, 6d 1 Y. Loc. 52, 1d. Loc. SPECIMENS. Holotype d ANSP. Paratype:same locality as 58, 1. Loc. 62, 1d. KENYA: Loc. 77A, 26. Loc. 87 holotype, Y ANSP. 2d.

24. Modicogryllus geonomes n. sp. 25. Modicogryllus jagoi n. sp.

HOLOTYPE. 5, South Africa, Natal (KwaZulu) Mkuze HOLOTYPE. 6, Kenya, Kakamega Forest Reserve, 26 x Game Reserve 19 xi 1980 (Otte and Cade) ANSP. 1982 (N. D. Jago) ANSP.

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FIG. 12. Modicogryllus male genitalia;A, perplexus, lOc. 58; B, perplexus, lOc. 87; C, dewhursti holotype;D, geonomes, lOc. 22.

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FIG. 13. Modicogryllus male genitalia. A, jagoi holotype; B, garriens holotype; C, mombasae holotype; D, alluaudi holotype.

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RECOGNITION. Figs. 7G, 13A. Table 1. Dark SONG. Two part song beginning with trill and cricket with strongly contrastinglight and dark ending in train of 4-10 short chirps. markingson pronotum,head, and legs. Dorsum of head black, with 6 stripes on occiput, with Locality p/s ch/s p/ch KHz ?C thin pale line connecting lateral ocelli. Face black above clypeus. Cheeks black. Pronotal 99 100.6 11.5 5-6 5.6 24 disk with strongly contrasting light and dark markings; lateral lobes entirely black. Abdo- SPECIMENS. Holotype d ANSP. men black on dorsum and venter. Front and middle legs very pale but with numerouslarge 27. Modicogryllus mombasae n. sp. darkmarkings. Hind femora with strongly indi- HOLOTYPE. 6, Kenya, Matuga, Agric. ExperimentalStation cate oblique stripes. Cerci dark brown. (near Mombasa) 18 ix 1982 (D. Otte) ANSP. DISTRIBUTION. Known only from type local- RECOGNITION. Figs. 7J, 13C. Table 1. Black- ity. ish with pale orange brown cerci. Head HABITAT. in the Kakamega Forest Collected and pronotum uniformly blackish. Abdomen Reserve. blackish on dorsum, brown on venter. Legs all SONG. Not known. dark reddish brown to blackish. Cerci pale

SPECIMENS. Holotype d ANSP. orange brown. DISTRIBUTION. Known only from type local- ity. 26. Modicogryllus garriens n. sp. HABITAT. Under grass clumps on mowed lawn. HOLOTYPE. 6, Tanzania, Dar-es-Salaam, University SONG. Trains of trills of various length from grounds, 28 ix 1982 (D. Otte) ANSP. 0.8 to 2.5 sec long. RECOGNITION.Figs. 3F, 4, 7H, 13B. Table 1. Cricket with strong light and dark patterns on head, pronotum,and legs. Dorsumof head with Locality p/s trill chirp KHz ?C two prominentcomplete occipital stripes poste- 85 46.2 0.8-2.5 7.4 24 rior to each eye, and two indistinctmore medial stripes. No pale stripe between lateral ocelli. SPECIMENS. Holotype d' ANSP. Paratype:same locality as Cheeks black but with pale patch dorsal to holotype, d ANSP. mandibles. Pronotal disk black centrally and with lateral pale bands. Lateral lobes black 28. Modicogryllus alluaudi (Chopard) except for pale area in extreme lower anterior Gryllus alluaudi Chopard 1932 (1934): 334. Lectotype 6, quarter. Forewings brownish on dorsal field, here designated, Kenya, Taveta, 750 m, iii 1922 blackish on lateral field. Dorsum of abdomen (Alluaud and Jeannel) PARIS. Gryllulus, Chopard 1936: pale medially, blackish along lateral margins, 539. Gryllus, Chopard1941: 7. Modicogryllus, Chopard 274. examined. pale on venter, subgenitalplate dark. Front and 1961: Type middle femora whitish with numerous small NOTES ON TYPES. Figs. 7K, 13D. Tables 1, 2. black spots; tibiae with broad dark brown Male lectotype: Head and pronotumuniformly bands. Hind femorawhitish, becoming blackish brown; occiput without stripes; no pale stripe distally especially on dorsum, with black between lateralocelli. Pronotumuniformly red- oblique stripes on outer face. Cerci pale. dish brown. All legs uniformly pale orange- DISTRIBUTION. Known only from type local- brown. Hindwings extending well beyond end ity. of abdomen. HABITAT. Under grass clumps and leaf litter Female paralectotype: Similar to male in under large trees. coloration. Measurementsgiven in Table 2.

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FIG. 14. A, Comidogryllusconsanguineus holotype; B, Comidogryllusvividus holotype; C, Modicoides royi holotype; D, Modicogryllusfinoti holotype; E, undescribedModicogryllus from Sierra Leone (PARIS).

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30. Comidogryllus vividus (Sjostedt) n. comb.

Gryllodes vividus Sjostedt 1900: 29. Lectotype 6, here designated, Camerun, Ekunda, 20 xii 1890 STOCKHOLM. Gryllopsis, Randell 1964: 1571. Type examined. NEW B COMBINATION. A B NOTES ON LECTOTYPEMALE. Figs. 70, 14B. Tables 1, 2. Rusty red cricket with somewhat FIG. 15. Chopard 1938a illustrationsof Modicogryllus backward slanting face and black head. Head: B, beibienkoi. male genitalia. A, elgonensis; dorsummostly blackish but with six pale stripes on occiput and vertex, stripe 3 about four times Genus Comidogryllus Otte and Alexander as broad as stripe 2; forehead with ivory band connecting lateral ocelli; face dark reddish ComidogryllusOtte and Alexander 1983: 90. Type species: brown, but pale aroundmedian ocellus. Cheeks Comidogryllus adina Otte and Alexander 1983: 91, by dark brown but with vertical yellowish stripe original designation. descending from lower posterior margin of eyes. Pronotum:dorsum reddish brown, black- This genus was first describedfrom Australia ish at anteriorlateral corners; lateral lobes en- where 10 species are known. In the present tirely blackish. Abdomen dark reddish brown. study we include two species from Africa pre- Forewings pale translucentbrown in harp area, viously placed under Modicogryllus. Its pres- other areas dark brown; lateral field blackish. ence in west Africa indicates that the genus is Hindwings shorter than pronotum. Front and nearly as widespread as Modicogryllus and we middle legs pale reddish brown with darker surmise that many species in southern and patches on inner and outer faces. Foretibiae southeastern Asia, presently under Mod- with large outer and a very tiny inner tym- icogryllus, will be shown to belong to Com- panum. idogryllus. A paralectotypemale is very similar to the DIAGNOSTIC CHARACTERS. Distinguishedfrom lectotype in color and shape. Foretibiaewithout Modicogryllus only by male genitalia. Shapes inner tympanum. of epiphallus and ectoparameres essentially PARALECTOTYPEFEMALE. Similar to males in identical to those of Australian species (see color. Forewing 1.24 times as long as pro- Identificationof Genera). notum. Ovipositor 0.78 times as long as hind femora. Forewings blackish, especially on 29. Comidogryllus consanguineus (Chopard) lateral field. Abdomen: dorsum with dark n. comb. brown tergites with pale spotted margin;venter Gryllus consanguineus Chopard 1961a: 20. Holotype 6, medium brown. Foretibiae without inner tym- Angola, Dundo, x 1953 PARIS. Modicogryllus, Chopard panum. 1961b: 274. Loxoblemmus, Randell 1964: 1578. NEW COMBINATION.

NOTESON TYPE. Fig. 7M, 14A. Table 1. Dark UNCERTAIN GENERIC STATUS reddish brown cricket. Head entirely blackish; no occipital stripes; no stripe between lateral 31. Modicogryllus rotundipennis (Chopard) ocelli. Face short. Pronotum uniformly dark rotundipennisChopard 1938: 126. Holotype Y, Abdomen dark reddish brown Gryllulus reddish brown. Kenya, Marakwet, Elgeyo Escarpment, 2500 m, 1932- on dorsum, pale brown on venter. Legs reddish 1933 (Mission de l'Omo) PARIS. Modicogryllus, Chopard brown, without contrasting spots or stripes. 1961b. Type examined.

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NOTES ON HOLOTYPE. Dorsum of head with 6 C. Amedegnato and M. Donskoff (Museum occipital stripes. With pale stripebetween later- National d'Histoire Naturelle); R. Danielsson al ocelli. Dorsum of pronotum slightly pat- (Zoological Museum, Lund);J. Decelle (Musee terned with lighter and darker brown; lateral Royal de l'Afrique Centrale); R. Leakey lobes pale in lower third, dark brown in upper (National Museum of Kenya); J. Marshall and two thirds. Forewings slightly longer than pro- B. Townsend (British Museum (Natural His- notum, overlapping dorsally, dark brown, but tory)); S., C., and R. Otte; R. Toms (Transvaal pale along forewing angle. Legs unicolorous Museum);M. and W. Astrup, 0. and C. Bour- orange brown. quin; Natal ParksBoard; Kruger National Park; and Serengeti National Park. We especially wish to thank Genus Modicoides n. gen. Nick Jago and the Desert Locust Control Organizationfor East Africa for their help and companionship. TYPE SPECIES. Cophogryllus royi Chopard 1954: 89.

DIAGNOSTIC CHARACTERS. Fig. 2. Male geni- talia similar to Modicogryllus. Entirely wing- LITERATURECITED less. Without auditory tympana. OTHER CHARACTERS. See M. royi. CHOPARD,L. 1925. Descriptions de Gryllides nouveaux (Orthopteres).Annales de la Societe entomologiquede 32. Modicoides royi (Chopard)n. comb. France 94: 291-332. . 1932 (1934). Voyage de Ch. Alluaudet R. Jeannel Cophogryllusroyi Chopard 1954: 89. Holotype c, Nimba, en Afrique Orientale(1911-1912). Gryllidae. EOS 8: Guinee, vii-xii 1951 (Lamotte and Roy) PARIS. Type 325-362. examined. NEW COMBINATION. . 1934. Cataloguesraisonnes de la Faune entomolo- gique du Congo Belge. OrthopteresGryllides. Annales RECOGNITION. Figs. 2, 14C. Head: Dorsum du Musee du Congo Belge, TervurenZoologie. Serie dark brown to blackish. Occiput with six pale 3, Section II, Tome IV, fasc. 1: 1-88. stripes; stripe 3 much wider than other two. . 1938a. Orthoptera. 1. Dictyoptera, Phasmodea, Foreheadwithout pale band connecting lateral . In: Mission Scientifique de'l Omo, Tome ocelli. Face dark brown down to and including IV, Fasc. 33: 89-134. In: M6moires du Museum National d'Histoire Naturelle, Nouvelle s6rie, Tome upper lobe of clypeus. Median ocellus large, VIII. pale. Cheeks dark brown. Pronotum:dorsum . 1938b. Additions et Rectifications au Catalogue darkbrown; lateral lobes blackish in dorsalhalf; des Gryllides du Congo Belge. Revue de Zoologie et ivory-colored in lower half; wider at anterior de Botanique Africaines 31(3-4): 341-350. end than posterior end. Wings entirely absent. . 1938c. On a collection of Gryllidaeand Tridactyli- dae from Sierra Leone. The Annals and Magazine of Abdomen:dorsum darkbrown; venter pale yel- Natural History (1lth series) 1: 49-68. lowish brown. Frontand middle legs: foretibiae . 1939. Explorationdu Parc National Albert. Mis- without tympana, femora pale yellowish brown sion G. F. de Witte (1933-1935) Fascicule 27(2) with dark markings on dorsum. Hind femora Gryllidae, pp. 5-15. with dark brown oblique stripes on dorsum. 1948. Contribution a l'etude des Gryllides du Congo Belge. Revue de Zoologie et de Botanique Hind tibia 0.82 times as long as femur, with 5 Africaines 51(2-3): 109-125. inner and 7 outer subapical spurs. Cerci 1.25 . 1954a. Notes on some southernAfrican crickets. times as long as hind femora. Body length ca. Annals and Magazine of NaturalHistory 7: 913-929. 14 mm; hind femur length ca. 8.3 mm. 1954b. La reserve naturelle integrale du Mont Nimba. III OrthopteresEnsiferes. Memoires de l'Insti- tut francais d'Afrique noire no. 40, fasc. 2: 25-97. ACKNOWLEDGMENTS . 1955. Orthopteraensifera. South African life: Results of the Lund University Expedition in We are grateful to the following institutions 1950-1951. Vol. 2: 266-300. Almquist and Wiksell, and individuals for helping us in this project: Stockholm.

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. 1961a. Orthopteres. Gryllidae et Gryllacrididae OTrE, D. 1983. African Crickets (Gryllidae) 2. Afrogryl- l'Angola. Companhiade Diamantes de Angola, Publi- lopsis Randell andNeogryllopsis n. gen. of easternand cacoes Culturais 56: 13-70. southern Africa (Gryllinae, Brachytrupini). Pro- . 1961b. Les divisions du genre Gryllus basees sur ceedings of the Academy of NaturalSciences of Phil- l'etude de l'appareil copulateur (Orthoptera, Grylli- adelphia 135: 218-235. dae). EOS 37: 267-287. AND W. CADE.1983a. African Crickets (Gryllidae) AND D. K. McE. KEVAN. 1954. Orthoptera- 1. Teleogryllus of eastern and southern Africa. Pro- Ensifera from northernKenya and Jubaland.Transac- ceedings of the Academy of NaturalSciences of Phil- tions of the Royal Entomological Society of London adelphia 135: 102-127. 105: 342-349. AND . 1983b. African Crickets (Gryllidae) GERSTAECKER,A. 1869. Beitrag zur Insekten-Faunavon 3. On the African species of Velarifictorus Randell Zanaibar.No. II. Orthopteraet Neuroptera.Archiv fur (Gryllinae, Modicogryllini). Proceedings of the Naturgeschichte35(1): 201-223. Academy of Natural Sciences of Philadelphia 135: KARNY,H. 1907. Die Orthopteranfauna des agyptischen 241-253. Sudans und von Nord-Uganda (Saltatoria, Gressoria, RANDELL, R. L. 1964. The male genitalia in Gryllinae Dermaptera) mit besonderer Beruchsichtigung der (Orthoptera:Gryllidae) and a tribalrevision. Canadian acridoideengattung Catantops. Sitzungsberichte der Entomologist 96: 1565-1607. Kaiserlichen Akademie der Wissenschaften (Wien). SAUSSURE, H. DE. 1877. Melanges Orthopterologiques. Matematisch-NaturwissenschaftlicheKlasse 116(1): Sme fascicule. III Gryllides. Memoires de la Societe 267-378. de Physique et d'Histoire Naturelle de Geneve 25: KRAUSS,H. 1892. Systematisches Verzeichnis der Canar- 1-352. ischen Dermapterenund Orthopterenmit Diagnosen SCHAUM, H. R. 1853. Bericht uber die zur Bekannt- der neuen Gattungenund arten. Zoologischer Anzeiger machung geeigneten Verhandlungen. Academie der 15: 163-171. Wissenschaften, Berlin, Volume x, 776 pp. . 1902. Beitrag zur Kenntnis der Orthopterenfauna SJ6STEDT,Y. 1900. Mantodeen, Phasmodeen, und Gryllo- der Sahara. Verhandlungen der Kaiserlich- deen aus Kamerunund anderenGegenden westafricas. Koniglichen Zoologisch-Gotanischen Gesellschaft in Bihang Svenska Vet.-Akad. Handlinger25(6): 2-36. Wien 52: 230-254.

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