the african species of heliocheilus grote (lepidoptera: noctuidae)

Systematic Entomology (1987) 12,459-473

The African species of Heliocheilus Grote (Lepidoptera: Noctuidae)

MARCUS MATTHEWS Department of Entomology, British Museum (Natural History), London

ABSTRACT. The heliothine genus Heliocheilus Grote is recalled from synonymy with Heliothis Ochsenheimer on the basis of a forewing modification in the male. Raghuva Moore and Canthylidia Butler are synonymized with Heliocheilus. Six species of Heliocheilus are recognized to occur in Africa, the adults of five are redescribed and the sixth described as new. Figures of the adults and genitalia of all species are provided. Lectotypes of Raghuva multiradiata Harnpson, Raghuva thomalae Gaede, Raghuva albipunctella de Joannis and Heliocheilus mekrana Brandt are designated, and twelve specific synonymies are proposed.

ABSTRACT. Le genre Heliocheilus Grote est retabli, sur la base d’une modification de l’aile anterieure chez le m2le, apres avoir ete mis en synonymie avec Heliothis Ochsenheimer. Raghuva Moore et Canthylidia Butler sont mis en synonymie avec lui. Six especes d’Heliocheilus se rencontrent en Afrique, l’une d’entre elles est dkcrite ici pour la premiere fois, les adultes des cinq autres redecrits. Les adultes et les genitalia de toutes les especes sont figures. Des lectotypes de Raghuva multiradiata Hampson, Raghuva albipunctella de Joannis, Raghuva thomale Gaede, et de Heliocheilus mekrana Brandt sont designes. Douze synonymies au niveau specifique sont proposees.

Introduction

The noctuid subfamily Heliothinae contains several species of great importance to world agriculture. Of these, Helicoverpu armigera, Helicoverpa zeu and Heliothis virescens are probably the most serious pests. These species are comparatively well known, and a considerable body of literature exists documenting their damage to crops, and efforts to control them.

The pest status of another heliothine,

Correspondence: Mr Marcus Matthews, Depart- ment of Entomology, British Museum (Natural History), Cromwell Road, London SW7 5BD.

Heliocheilus albipunctella, which damages millet in the sahel savannah of Africa, appears to have developed very recently. This has been coincident with the onset of drought from 1972 (Deeming, 1978; Vercambre, unpublished).

At present there is no way in which workers concerned with the pest can name it reliably, or distinguish it easily from its five African con- geners, three of which occur with it in the sahel. The aim of this paper is to stabilize the nomenclature of these species, and provide a means for their quick identification. To simplify this, a colour plate figuring adults of all six species is provided. Therefore, notes on colour

459

460 Marcus Matthews

in the text are kept to a minimum, except in the case of H.roseus sp.n. where a full description is given.

Genitalia do not provide characters for species differentiation in all cases. I can find no reliable differences in the males between H. confertissirna, H.multiradiata or HMgrnatia. However, these species all vary in the following: the degree to which the valves are recurved distally; the extent of the corona along their ventral (outer) margins; and the size of the scobinate patch and its constituent denticles at the base of the vesica.

In the females I can find no consistent differences between the genitalia of H.confertissima, H.multiradiata, Hstigmatia or H. discalis. However, these species all vary in the number and length of the setae on abdominal segment 8.

Nevertheless, both sexes of H.confertissima may be distinguished by the more heavily sclerotized and notched posterior margin of tergum seven, visible in genitalic preparations. Despite these similarities, figures of the genitalia of all six species are provided. Wherever species differentiation is possible using the genitalia, this is pointed out in the text.

In this paper I recall Heliocheilus from synonymy with Heliothis on the basis of a forewing modification in the male.

The genus is cosmopolitan, and appears to be most diverse in Australia.

Introduction

La sous-famille des Heliothinae (Noctuidae) comprend plusieurs espbces d’une trbs grande importance pour l’agriculture mondiale. Helicoverpa armigera, Helicoverpa zea et Heliothis virescens sont probablement les especes les plus nuisibles aux cultures. Elles sont comparativement bien connues, et il existe une litterature considerable B propos des degats qu’elles occasionnent et des efforts de contr6le.

Le statut de parasite des cultures d’une autre heliothine, Heliocheilus albipunctella, qui s’atta- que au mil dans les savanes saheliennes afri- caines semble s’&tre dkveloppe trbs recemment. I1 a coincide avec le debut de la secheresse en 1972 (Deeming, 1978; Vercambre, non publie).

Actuellement il n’existe, pour les chercheurs concernes, aucun moyen sur pour nommer cette espbce ou pour la distinguer facilement de ses

cinq congeneres africains, dont trois sont egale- ment presents dans la zone sahelienne. Le but de cet article est de clarifier le nomenclature de ces especes et de fournir les moyens de les identifier rapidement. Pour simplifier, les adultes des six especes sont figures sur une planche en couleur . Cependant, dans le texte, les notes sur la colora- tions sont reduites au minimum, excepte dans le cas d’Heliocheilus roseus sp.n. pour laquelle j’ai donne une description complete.

L‘Ctude des genitalia ne fournit pas dans tous les cas des caractttres pour la differentiation specifique. Je ne peux mettre en evidence de differences sures entre les mtdes d’HeliocheiZus confertissima, Heliocheilus rnultiradiata et d’Heliocheilus stigmatia. Toutefois, chez toutes ces espbces, il existe des variations pour les caractbres suivants: le degre selon lequel les valves sont recourbees distalement; l’extension de la corona le long de leur bord ventral (externe); la taille de la zone denticulee situee a la base de la vesica.

Chez les femelles je n’ai pas mis en evidence de diffkrences notables entre les genitalia d’Heliocheilus confertissirna, Heliocheilus multiradiata, Heliocheilus stigmatia et d’Heliocheilus discalis. Toutefois toutes ces especes different par le nombre et la longueur des soies du huitibme segment abdominal.

MalgrC cela, les deux sexes d’Heliocheilus confertissima se distinguent par le bord post- Crieur de septieme tergite plus fortement sclerotise et entaille (caractkre visible sur les preparations des genitalia). En depit de ces similarites les genitalia des six espbces sont figures. Les distinctions specifiques possibles grace B I’Ctude des genitalia sont indiquees dans le texte.

Dans cet article, le genre Heliocheilus est retabli aprbs avoir Cte mis en synonymie avec Heliothis, sur la base d’une modification de l’aile anterieure chez le mgle.

The forewing modification in Heliocheilus

This stucture is variably developed between species, and in some cases provides a good means of identification. Comparisons in the description that follows refer to unmodified species, e.g. Heliothis viriplaca, Fig. 42.

The costa is bowed and swollen at or just distad of its mid point, above the discal cell and

The African species of Heliocheilus Grote 461

tive to both sexes: after some minutes individuals congregate to form small groups (I noted up to seven together in 1985). These com- prise mating pairs and unpaired males, which continue to buzz. Males behaving in this way are quite easily disturbed, especially by the white light of a hand-lamp, and fly off soundlessly.

basad of the end of Sc, forming a ‘blister’. When the swelling is well developed, lamellae in its lower half cross it at right angles to the costa. These are covered by a thin membrane and the upper half is empty. The subcosta is angled more sharply towards the costa from its base, reaching the costa just basad of the swelling and running closely behind it for the rest of its length. R1 is angled more sharply towards the costa for its basal third, then angled back parallel or sub- parallel with Rs, to form an R1-Rs space. The basal quarter of R1 may be swollen; Rs may be swollen from its base to r-ml. Veins R2, R3, and R4 are variable, even within a species. Occa- sionally R3 does not anastomose with R4, so that the accessory cell is absent. In weakly modified species R2 arises before the apex of the accessory cell, and R3 and R4 are stalked together. R-ml is present, at least as long as a quarter of the distance between the bases of M1 and M2. The discal cell is broad. Two partially hyaline patches are present in moderately to strongly modified species, the wing membrane on both the upper and lower surfaces being ridged within them. The ridges are at right- angles to the costa, and the scales inserted in lines along their tops. One patch lies over the R1-Rs space, extending slightly basad over R1 in strongly modified species. The other lies over the anterior part of the discal cell, immediately behind Rs (Fig. 41).

It has been known for some time that this modification in heliothines is associated with sound production (Cook, 1930; Forbes, 1912; Hebard, 1922). Research on this subject con- tinues, and will be discussed more fully in a later paper. A similar structure occurs in some male agaristids (e.g. Aegocera spp.; Alypia spp.; Hecatesia spp.). This is known, in some in- stances at least, to be associated with sound production (e.g. Hannemann, 1956; Nicholson, 1955). Mr John Grunshaw (Tropical Develop- ment and Research Institute) first noted this unusual behaviour exhibited by H.albipunctella at Mourdiah in Mali, and preliminary observa- tions are as follows. Very shortly after sunset, males may be easily located clinging to low vegetation or millet stems, by a buzzing sound of approximately 130 Hz. The wings are vibrated, not through a full beat, but about 20” each side of the moth’s vertical axis. Very often, the terminal segments of the abdomen are fully extended at the same time. The buzzing is evidently attrac-

La modification de I‘aile anterieure chez Heliocheilus

Le dCveloppement de cette structure est variable suivant les especes, et dans certains cas fournit un bon moyen d’identification. Les comparai- sons dans la description suivante se referent aux especes qui ne presentent pas de modification, e.g. Heliothis viriplaca, Fig 42.

La costa est arquCe et enflee au niveau ou peu apres la milongueur, au dessus de la cellule discale, avant Ie point de rencontre de la costa et de la sous-costa (Sc); elle forme une ‘cloque’. Lors- qu’elle est tres enflee, des lamelles perpen- diculaires traversent la moitiC infkrieure. Celles ci sont recouvertes d’une fine membrane, la partie superieure est vide. A partir de la base, la sous-costa (Sc) converge plus nettement vers la costa, qu’elle atteint B la base du renflement, elle longe ensuite ce dernier sur le reste de sa longueur. Sur son premiers tiens, R1 se dirige plus nettement vers la costa, puis s’incline vers I’arrikre, devient plus ou moins parallele a Rs, formant ainsi un espace R1-Rs. Le quart initial de R1 peut Ctre enflC, de m&me que Rs depuis la base jusqu’g r-ml, R2, R3, et R4 sont variables, mCme a 1’intCrieur d’une espece. Parfois R3 ne s’anastomose pas avec R4, de ce fait la cellule accessoire est absente. Chez les especes peu modifiCes R2 apparait avant I’extremitC de la cellule accessoire, et R3 et R4 ont une base commune. R-ml est present, au moins aussi long que le quart de la distance entre les bases de M1 et M2. La cellule discale est large. Deux zones partiellement hyalines sont prksente chez les espkces moyennement a fortement modifiCes, la membrane de I’aile y est sillonnCe aussi bien a la surface supCrieure qu’i la surface infCrieure. Les sillons forment des angles droits avec la costa et les Ccailles sont insCrCes en ligne sur les som- mets. Une des zones est situCe sur I’espace R1-Rs, et s’Ctend lCg&rement sur la base de R1 chez les espkces fortement modifiCes. L‘autre est situCe sur la partie anterieure de la cellule discale immkdiatement derriere Rs (Fig. 41).

462 Marcus Matthews

L‘on sait depuis quelque temps que cette modification chez les hkliothines est associee a une Cmission sonore (Cook, 1930; Forbes, 1912; Hebard, 1922). Ceci constitute le sujet de recherches en cours et sera discute plus longue- ment dans un prochain article. Une structure similaire se rencontre chez quelques mfiles d‘agaristides (e.g. Aegocera spp.; A l y p i a spp.; Hecatesia spp.). Elle est connue, au moins dans certains cas, pour Ctre associCe une Cmission sonore (e.g. Hannemann, 1956; Nicholson, 1955). M. John Grunshaw (Tropical Develop- ment and Research Institute) a Cte le premier B noter ce comportement inhabituel de H.alb i - punctella A Mourdiah au Mali, et les observa-a tions prkliminaires sont les suivants: peu de temps apres le coucher du soleil, les mfiles peu- vent &re localisds facilement, alors qu’ils grim- pent dans la vkgdtation basse ou sur les tiges de mil, par un bourdonnement d’approximative- ment 130 Hz. Les ailes vibrent, non pas sur toute leur amplitude mais B environ 20” de chaque c6te

de I’axe vertical du corps. Trbs souvent les segments terminaux de l’abdomen sont en pleine extension pendant ce temps. Le bourdonnem- ment est, a l’bvidence, attractif pour les deux sexes: apres quelques minutes les individus forment de petits agrCgats (j’ai note jusqu’a sept individus ensemble en 1985). 11s sont formes de couples s’appariant et de mfiles seuls qui con- tinuent a bourdonner. k s mhles se comportant de cette mannikre sont facilement dkranges, en particulier par la lumiere blanche d’une lampe de poche, et s’envolent sans bruit.

Abbreviations

The following abbreviations are used for depositories: MNHN, Museum national d’Histoire naturelle, Paris, France; ZMB, Zoologisches Museum, Berlin, DDR; RMS, Naturhistoriska Riksmuseet, Stockholm, Sweden. BMNH, British Museum (Natural History), London, England.

The coloured illustrations of Heliocheilus are numbered consecutively from the top, i.e. the first column is Figs. 1-8, the second column is Figs. 9-16, etc. Les illustrations en couleur des Heliocheilus sont numerotkes du haut en bas a partir de la premitre colonne, c’est a dire l t r e colonne Figs. 1-8, 2tme colonne Figs. 9-16, etc. FIG. 1. H.roseus sp.n. 6, PARATYPE. MALI; Mourdiah. FIG. 2. H.roseus sp.n. d. HOLOTYPE. MALI; Mourdiah. FIG. 3. H.roseus sp.n. @. PARATYPE. MALI: Mourdiah. FIG. 4. H.roseus sp.n. 6. PARATYPE. MALI: Mourdiah. FIG. 5. H.roseus spa. 9. PARATYPE. MALI: Mourdiah. FIG. 6. H.roseus sp.n. 9. PARATYPE. MALI: Mourdiah. FIG. 7. H.roseus sp.n. 9 . PARATYPE. MALI: Mourdiah. FIG. 8. H.roseus sp.n. 9. PARATYPE. MALI: Mourdiah. FIG. 9. H.rnu1tiradiata 0 . SOUTH AFRICA: Transvaal. FIG. 10. H.rnultiradiata 0 . KENYA: Suna. FIG. 11. H.multiradiata d, KENYA: Suna. FIG. 12. H.multiradiata 0“. KENYA: Suna. FIG. 13. Hdgrnatia d. NAMIBIA: Windhoek. FIG. 14. H.stigrnatia d. NAMIBIA: Windhoek. J3G. 15. H.stigmafia 9 . NAMIBIA: Hoffnung. FIG. 16. H.stigmatia 9 . NAMIBIA: Hoffnung. FIG. 17. H.aZbipuncteZ1a @. MALI: Mourdiah.

FIG. 18. H.albipunctella d. MALI: Mourdiah. FIG. 19. H.albipunctella d. MALI: Mourdiah. FIG. 20. H.albipuncte1la d. MALI: Mourdiah. FIG. 21. H.albipunctella 9 . MALI: Mourdiah. FIG. 22. H.albipunctella $2. MALI: Mourdiah. FIG. 23. H.albipunctella 9 . MALI: Mourdiah. FIG. 24. H.albipunctella 9 . MALI: Mourdiah. FIG. 25. Hdiscalis 6. ETHIOPIA: Dire Dawa. FIG. 26. H.discalis 6. MALI: Mourdiah. FIG. 27. H.discalis d. MALI: Mourdiah. FIG. 28. H.discalis 6. MALI: Mourdiah. FIG. 29. H.discalis 9. MALI: Mourdiah. FIG. 30. H.discalis 9. MALI: Mourdiah. FIG. 31. H.discalis 0 . MALI: Mourdiah. FIG. 32. H.disca1is 9. MALI: Mourdiah. FIG. 33. H.confertissirna d. MALI: Mourdiah. FIG. 34. H.confertissirna d. MALI: Mourdiah. FIG. 35. H.confertissirna d. MALI: Mourdiah. FIG. 36. H.confertissirna d. MALI: Mourdiah. FIG. 37. H.confertissirna 9. MALI: Mourdiah. FIG. 38. H.confertissirna 9. MALI: Mourdiah. FIG. 39. H.confertissirna 9 . MALI: Mourdiah. FIG. 40. H.confertissirna 0 . MALI: Mourdiah.


Heliocheilus Grote

Heliocheilus Grote, 1865: 328-329. Type species: Heliocheilus paradoxus Grote, by monotypy.

Raghuva Moore, 1881: 326. Type species: Leucania confertissima Walker, by monotypy. Syn.n.

Canthylidia Butler, 1886: 406. Type species: Canthylidia pallida Butler, by monotypy. Syn.n. Diagnosis. Species of Heliocheilus may be dis-

tinguished from all other Heliothinae, e.g. Heliothis viriplaca (Fig. 42), by the modified forewing in the male (Figs. 44-49). The male of Masalia perstriata Hampson possesses a modified forewing (Fig. 43), but this differs from

that of all species of Heliocheilus as follows. The small costal swelling lies between Sc and R l , beyond the apex of the discal cell, not basad of Sc above the discal cell as in species of Heliocheilus. An additional small thickening of the wing membrane occurs over the bases of R2 and R3. Sc and R1 are not displacd. R-ml, although present, is very short, less than one quarter the distance between the bases of M1 and M2. There are no hyaline patches in M . perstriata.

Diagnose. Les especes dfieliocheilus se distinguent de tous les autres Heliothinae, e.g. Heliothis viriplaca (Fig. 42) par l'aile anterieure modifiee du male (Figs. 44-49). Le m2le de Masalia perstriata Hampson presente Cgalement une modification aux .ailes anterieures, mais

464 Marcus Matthews

FIGS. 41-49. Male right forewings. Fig. 41, Heliocheilus albipunctella; Fig. 42, Heliothis viriplaca; Fig. 43, Masalia perstriata; Fig. 44, Heliocheilus roseus sp.?; Fig. 45, Heliocheilus confertimima; Fig. 46, Heliocheilus rnultiradiata; Fig. 47, Heliocheilus stigmatia; Fig. 48, Heliocheilus albipunctella; Fig. 49, Heliocheilus discalis.


celles-ci different de celles de toutes les especes d’Heliocheilus de la manniere suivante. Le petit renflement de la costa est situC entre Sc et R1, au dela de I’extrCmitC de la cellule discale et non en deGa de Sc et audessus de la cellule discale comme chez les Heliocheilus. Un petit epaississement supplementaire de la membrane de l’aile est situe sur les bases de R2 et R3. Sc et R1 ne sont pas deplackes. Udc bien que present, est tres court, moins du quart de la distance separant les bases de M1 et M2. I1 n’y a pas de zones hyalines chez M.perstriata.

Key to males of African Heliocheilus

1 Forewing with obvious hyaline patches . . . . . . 2 Forewing without hyaline patches (Fig. 1-4) . . .

costal blister (Figs. 17-20, 25-28,48, 49) . . . . . 3

46,47) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

-

6 roseuv span. 2 Forewing strongly modified, broadest opposite

- Forewing strongly or less strongly modified, broadest towards termen (Figs. 11-14,33-36,4.5,

3 Hindwing with well-marked discal spot (Figs. 25-28), .......................... . 5 . discalis

- Hindwing without well-marked discal spot (Figs. 17-20) ...................... . 4 . albipunctella

4 Forewing moderately strongly modified (Figs. 13, 14,4?) . . . . . . . . . . . . . . . . . . . . . . . . . . 3 . stigmatia

Darker species, with whitish streaks on forewing on ends of veins to termen, hindwing not pale (Figs. 11, 12). . . . . . . . . . . . . . . . . 2 . multiradiata

- Paler species, without whitish streaks on forewing

1 confertissima

- Forewing less strongly modified than this . . . . . 5 5

on ends of veins, hindwing pale (Figs. 33-36). . .

Key to females of African Heliocheilus

1 Forewing almost uniform reddish-brown or dark reddish-brown, with a postmedian line of white dots on the veins; hindwing uniform grey with a small white crescent-shaped discal marking (Figs. 21-24) ..................... . 4 . aibipunctella

- Forewing not almost uniform reddish-brown or dark reddish-brown, may or may not have a postmedian line of white dots on the veins; hindwing not uniform grey, discal spot dark when present 2

2 Pinkish or buff species; without antemedian fascia even in darker, more strongly marked specimens; without paler spots on hindwing in terminal band on either side of Cul (Figs. 5-8) .6 . roseus sp.n.

- Not pinkish, may be buff; generally with antemedian fascia; generally two paler spots on hindwing in terminal band on either side of C u l . . . . . . . . 3

3 Discal spot on hindwing large and strongly marked (Figs. 2Y-32) . . . . . . . . . . . . . . . 5 . disculis

- Discal spot on hindwing not large or strongly marked . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4 White streaks present on forewing on ends of veins to termen (Figs. 9,10) . . . 2 . multiradiatu, or (Figs. 15, 16) 3. stigmatia

- No white streaks on forewing on ends of veins to termen (Figs. 37-40) . . . . . . . . . . 1. cwnfertissima

1. Heliocheilus confertksima c0mb.n. (Figs. 33-40, 45, 51, 57,63, 69)

Raghuva confertissima (Walker), 1865: 625. Holotype cf [INDIA] South Hindostan: Coimbatoor (Walhouse) (BMNH). [Examined.]

Heliothis hyalosticta Hampson, 1894: 507. Holotype [PAKISTAN] Karachi (Swinhoe) (BMNH). [Examined.] Syn.n.

Raghuva perdentata Hampson, 1903: 30. Holotype cf [KENYA] B[ritish] E[ast] Africa: Kikuyu, Nairobi Plains 5400’, 3.v.1899 (Crawshay) (BMNH). [Examined.] Syn.n.

Heliocheilus designata Brandt, 1941: 853. [IRAN]: Fort Sengan [Sangan], 30.iv (Brandt) (Not in RMS). [Not examined.] Syn.n.

Heliocheilus mekrana Brandt, 1941: 852-853. LECTOTYPE cf (here designated) (RMS prep. No. 5723) IRAN: Baloutchistan [Baluchestan], Bender Tchahbahar [Chah Bahar], i-ii.1938 (Brandt) (RMS). [Examined.] Paralectotype 9 same data as lectotype (RMS). [Examined.] Syn.n.

Raghuva ennedica Herbulot & Viette, 1952: 88. Holotype 0 (genitalia prep. C. Herbulot No. 1795) [CHAD]: Ennedi, Oued, [near] prks An Djeress, 2O.viii.1948 ( d e Mir t ) (MNHN). [Examined.] Syn.n.

Male: Figs. 33-36. Female: Figs. 3740. Male forewing moderately strongiy modified (Fig. 45). Very variable in both sexes, but especially the females which are darker and more distinctly marked.

Genitalia. Male. Figs. 51, 57, 63. Not distinguishable from multiradiata or stigmatia. Female. Fig. 69. Ovipositor valves soft and pad- like. Not distinguishable from muttiradiata, s@- matia, or discalis. However, both sexes of con- fertissinia may be separated from these species by the more heavily sclerotized and notched

466 Marcus Matthews

FIGS. 50-55. Male genitalia of Heliocheilw. Valves. Fig. 50, H.roseus sp.n.; Fig. 51, H.confertissima; Fig. 52, H.multiradiata; Fig. 5 3 , H.stigmatia; Fig: 54, H.albipunctella; Fig. 55, H.discalis.


posterior margin to tergum seven of confertissima (visible in genitalic preparations). See male Fig. 63; (female not figured).

Additional material examined. [56d, 1899) SENEGAL: Kaolack. MALI: Aguel-Hoc; Mourdiah; 64 km E Tombouctou. ALGERIA: 70 miles N Bordj Moktar. NIGER: Aderbissinat (N Damergou); 130 km S Agadez; Aouderas (Air); Aouderas River (Air); Aouderas

(Asben); Atara Valley (SW Air); N of Azzal; Dogondontchi; Tanout (S Air); Tanout (Marandet, Air). ETHIOPIA: Dire Daoua [Dire Dawa]. KENYA: Archers Post (Meru); Chandler Falls (Uaso River); Isiolo; Kajiado; Kibwezi; Ngara Mara NFD; Voi. OMAN: Khasab; Sharjah. INDIA: Ajmere; Bhuj- Kutch; Cuddapah; Deesa; Gooty. BURMA: Myingyan; Thayetmayo.

FIGS. 5661. Male genitalia of Heliocheilus. Aedeagus and vesica. Fig. 56, H.roseus sp.n.; Fig. 57, H.confertissima; Fig. 58, H,multiradiata; Fig. 59, Hstigmatia; Fig. 60, H.albipunctel1a; Fig. 61, H . discalis.

468 Marcus Matthews

Remarks. Brandt (1941) did not specify which specimen he intended to be the holotype of mekrarta. I have examined two syntypes. One is a male, probably the specimen in Brandt’s plate 27 at the top of the column labelled ‘46’ [his Fig. 461. The other is a female, and corresponds exactly to the specimen at the bottom of the column labelled ‘46’ (just above the number). They both bear red labels reading ‘LEC- TOTYPE 6‘’ and ‘LECTOTYPE 9’ respec- tively. I designate as LECTOTYPE the male, which bears a yellow card label reading: ‘RMprepl5723’.

Dr B. Gustaffson has informed me that the holotype of designata is not in the RMS. Brandt described designata only tentatively as new (‘Heliocheilus designata Sp. ?n.’), and noted its great similarity to mekrana.

2. Heliocheilus multiradiata corn b.n. (Figs. 9-12,46, 52, 58, 64, 70)

Raghuva multiradiata Hampson, 1902: 257. LECTOTYPE 6 (here designated) [ZIM- BABWE] Mashonaland: [Harare] Salisbury, xii.1900 (Marshall) (BMNH). [Examined.] Paralectotype 6 same data as lectotype (BMNH). [Examined.]

Male: Figs. 11, 12. Female: Figs. 9, 10. Male forewing quite weakly modified (Fig. 46), only the lower patch hyaline in some specimens. Claviform stigma generally long, extending from the base of the orbicular to beneath the reniform as a dark patch bisected along its length by paler scales.

Genitalia. Male. Figs. 52, 58, 64. Not distinguishable from confertissima or stigmatia. Female. Fig. 70. Not distinguishable from confertissima, stigmatia or discalis. However, both sexes may be separated from confertissima by the form of tergum 7 (visible in genitalic preparations), see under confertissima.

Additional material examined. (346 , 25 9) UGANDA: SE Ruwenzori. KENYA: Athi fiver; Eb Urru; Kibwezi; Nairobi; Nakuru; Suna (S Kavirondo). MADAGASCAR: Diego Suarez; Maintirano; Sakaramy. ZIMBABWE: ‘Mashonaland’; Salisbury [Harare]. TANZANIA: Kilindi; Ruo Valley. MALAWI: Mt Mlange. SOUTH AFRICA: Pretoria; Shilouvane (Transvaal); Umtata (Transkei); White River (Transvaal).

Remarks. This species is generally darker and slightly smaller than stigmatia, with a more clearly-defined claviform stigma. The males of both species may be identified easily using the forewing modification, but females unassociated with males may be difficult to place. Although I have insufficient material to be certain of the limits of distribution of either species, it appears multiradiata occurs in East Africa and Madagascar, and stigmatiu occurs in South Africa and south-western Africa, with a zone of overlap in eastern South Africa. Unassociated females taken in this zone cannot be identified with certainty at present.

Hampson (1903) described multiradiata from two male syntypes (BMNH) from the same locality One is here designated LECTOTYPE and has been labelled accordingly.

3. Heliocheihs stigmatia c0rnb.n. (Figs. 13-16,47, 53, 59, 65, 71)

Raghuva stigmatia Hampson, 1903: 29. Holotype 6 [SOUTH AFRICA] Cape Col- ony: Transkei (Barrett) (BMNH). [Examined.]

Raghuva biocularis Gaede, 1915: 39. Holotypc 9 [SOUTH AFRICA]: Johannisburg [Johan- nesburg] (ZMH). [Examined.] Syn.n.

Male: Figs. 13, 14. Female: Figs. 15, 16. Male forewing strongly modified (Fig. 47). Females more variable than males, often darker. The claviform stigma is indistinct: a dark ring, or may be absent.

Genitalia. Male. Figs. 53, 59, 65. Not distinguishable from confertissima or multiradiata. Female. Fig. 71. Not distinguishable from confertissima, multiradiata or discalis. However, both sexes may be separated from confertissima by the form of tergum seven (visible in genitalic preparations), see under confertissimu.

Additional material examined. (706 , 479) SOUTH AFRICA: 5 miles NW of Beaufort West; Durban; Isipingo (Nr Bloemfontein); Kenhardt; Orange Free State; Potchefstrom; Pretoria; Upington. LESOTHO. BOTSWANA: 42 miles W Kalkfontein; Kuke Pan; Lake Ngami (12 miles NE Sehithwa). SOUTH WEST AFRICA: Hoffnung; Okahandja; 33 miles ENE Omaruru; Windhoek.

Remarks. Although the present species is generally larger and paler than ~u l t i rud ia tu , with a less well defined claviform stigma, only males can be identified with certainty (using the difference in forewing modification). Unfor- tunately, the holotype of bioculuris is a female from the zone of overlap. However, I place it in synonymy with stigmatia on the basis of its pale colouration, and small claviform stigma.

The African species of Heliocheilus Grote

4. Heliocheilus albipunctella corn b.n. (Figs. 17-24,41, 48, 54, 60, 66,72)

469

Raghuva albipunctella de Joannis, 1925: 258. LECTOTYPE 0" (here designated) [SENE- GAL] Haute SCnegal (MNHN). [Examined.] Paralectotype 0 [GHANA] Gold Coast: Navaro 11N 1.30W, vi.1923 (Cardinall) (BMNH) . [Examined.]

FIGS. 62-67. Male abdominal segment 8 of Heliocheilus. Fig. 62, H.rosrus sp.n.; Fig. 63, H.confertissirna; Fig. 64, H.multiradiata; Fig. 65, H.stigmatca; Fig. 66, H.albipunctella; Fig. 67, H.discalis.

470 Marcus Matthews

Raghuva vercambrei Laporte, 1977: 429. Holotype (genitalia prep. B. Laporte No. 1084) SENEGAL: Bambey, vi.1975 (Ver- cambre) (MNHN). [Examined.] Syn.n.

Raghuva bordati Laporte, 1977: 430. Holotype 0“ (genitalia prep. B. Laporte No. 1085) SENEGAL: Bambey, 5. ix.1975 (Vercambre) (MNHN). [Examined.] Syn.n.

Raghuva graminivora Laporte, 1977: 430-431. Holotype Q SENEGAL: Bambey, vi.1976 (B. Vercambre) (MNHN). [Examined.] Syn.n.

Raguva brenieri Laporte, 1977: 431. Holotype d (genitalia prep. B. Laporte No. 1089) SENEGAL: Bambey, vi.1976 (Vercambre) (MNHN). [Examined.] Syn.n. Male. Figs. 17-20. Female Figs. 21-24. Male

forewing very strongly modified (Fig. 48). Females more variable than males, often darker.

Genitalia. Male. Figs. 54,60,66. Most similar to discalis; distinguished by: more spines in the corona; no large basal diverticulum just beneath the aedeagus; a heatier scobinate patch at the base of the vesica; a more strongly sclerotized, and notched posterior margin to tergum 7. Female. Fig. 72. Ovipositor valves large, moderately sclerotized, and laterally corn pressed; none of the other five species is like this.

Additional material examined. (42d , 959) MALI: Mourdiah. NIGERIA: Azare; Borno; Kano; Gusau; Sokoto; Zaria. SUDAN: Tembura.

Remarks. De Joannis described albipunctella from two syntypes: a male (MNHN), which I designate as LECTOTYPE, and a female (BMNH).

Laporte (1977) states that the holotype of graminivora is a male, and that the slide on which the genitalia are mounted is No. 1087. The specimen bearing a red ‘HOLOTYPE’ label, and also one reading ‘RAGHUVA graminivora. / n sp Laporte / HOLOTYPE’ is a female albipunctella without abdomen. A white card label with blue ink reading ‘n’est pas / l’holotype’ has been added. The specimen bears a label reading ‘GENIT. / B. / LAPORTE / No.1087’. The genitalia mounted on slide No. 1087 which I have examined are those of a male albipunctella.

In his description of graminivora, Laporte (1977) states the costa of the male’s forewing is straight (‘c6te droite’). This is in contrast to his descriptions of vercambrei, bordati and brenieri,

where he states in the description for each species that the costa of the male forewing is arched (‘arquCe(s)’).

Since the description given fits that of a female albipunctella, I regard the pinned specimen as the holotype. The male genitalia associated with it (possibly by a mistake during the labelling of genitalic preparations and corresponding pinned specimens) probably led Laporte to suppose the holotype was a male.

5. Heiiocheiius discaiis corn b . n . (Figs. 25-32,49, 55, 61,67, 73)

Raghuva discalis Harnpson, 1903: 31. Holotype 9 [KENYA] B[ritish] E[ast] Africa: Athi R[iver], 8.v.1899 (Betton) (BMNH). [Examined.]

Raghuva thomalae Gaede, 1915: 38. LEC- TOTYPE cf (here designated) [TANZANIA] Deutsch Ost-Africa: Bezirk Amani [Amani district], Gomba, 1913 (ZMH). [Examined.] Syn.n.

Heliothis albivenata Hampson, 1916: 103. Holotype 0 [SOMALIA] Somaliland: Man- dera, 47 mi[les] SW of Berbera, 3000 ft open & bush, 20.x.1909 (BMNH). [Examined.] Syn.n.

Male: Figs. 25-28. Female: Figs. 29-32. Male forewing very strongly modified (Fig. 49). Note the well-marked discal spot on the hindwing.

Genitalia. Male. Figs. 55,61,67. Most similar to albipunctella; distinguished by: fewer spines in the corona; a large diverticulum at the base of the vesica, just beneath the aedeagus; a less pronounced scobinate patch at the base of the vesica, and a much less sclerotized, entire posterior margin to tergum 7. Female. Fig. 73. Indistinguishable from confertissima, multiradiata or stigmatia. However, may be separated from confertissima by tergum 7 (visible in genitalic preparations), see under confertissima.

Additional material examined. (22d , 72 0 ) MALI: Mourdiah. NIGER: 64 km E Dogon- dontchi. NIGERIA: Sokoto. ETHIOPIA: Dire Daoua [Dire Dawa]. KENYA: Athi River; Kajiado. TANZANIA: Old Shinyanga; Olduvai Gorge.

Remarks. Gaede (1915) described thomale from two syntypes, a male and a female from the same locality. I have examined only the male, which I here designate as LECTOTYPE.


FIGS. 68-73. Female'genitalia of Heliocheilus. Fig. 68. H.roseus sp.n.; Fig. 69, H.confertissima; Fig. 70, H.multiradiata; Fig. 71, Hstigmatia; Fig. 12, H.albipunctella; Fig. 7 3 , H.disca1i.s.

6. Heliocheilus roseus spn. (Figs. 1-8, 44, 50,56, 62, 68)

Diagnosis. This is the only African Heliocheilus without one or two hyaline patches on the male forewing.

Holotype male Fig. 2; male paratypes Figs. 1, 3, 4; female paratypes Figs. 5-8. Male forewing only weakly modified (Fig. 44), without hyaline

patches. Females more variable than males. Head and thorax uniform pale buff to mixed buff and grey. Thorax laterally with long whitish hairs overlying whitish scales. Legs mixed pink and buff, sometimes with brown scales, expecially on tarsi. Abdomen buff, same as head and thorax. Upperside of forewing ground colour buff; sometimes heavily suffused with pinkish-brown, or with grey, or both, especially

472 Marcus Matthews

between Sc and 1A+2A leaving basal two-thirds of costa and hind margin buff. Orbicular stigma generally absent; when present, grey and indistinct. Reniform stigma grey and moderately defined, to absent. Postmedian fascia a line of very small black dots basally with white streaks or dots distally; either series or both may be absent. A row of small black dots between the veins along the termen, may be absent. Fringe buff, pinkish-brown or grey; a darker line parallel to termen. Upperside of hindwing pale, with a broad darker terminal band of pinkish-brown and sometimes grey scales. Discal spot generally absent, may be an indistinct pinkish-brown or greyish crescent. Fringe buff, with a darker line parallel to termen. Underside of forewing uniform buff, to uniform buff basally with darker pinkish-brown terminal shade beyond postmedian fascia. Sometimes a grey patch between R5-CuA2, not reaching termen. Orbicular stigma present in very heavily marked specimens as a small black dot. Reniform stigma from distinct and dark grey, to indistinct and pinkish-brown; may be absent. Underside of hindwings uniform buff, sometimes a broad slightly darker pinkish-brown terminal band.

Genitalia. Male. Figs. 50,56,62. Vesica with a proportionally larger scobinate patch at its base than the other species treated here. Although the patch is well developed, the constituent denticles are small. Female. Fig. 68. Ovipositor valves well sclerotized and dorsoventrally flat- tened; none of the other five species is like this.

Type material. Holotype Cr" MALI: Mourdiah 14.28N 7.28W (at light) 17.ix.1985 (Matthews) (BMNH). Paratypes (BMNH): all same locality as holotype, 26,49,27.ix.1984 (Jago); lcf,l?, 16.ix.1985 (Matthews); 2 6 , 6 9 , same data as holotype; 5 9 , 18.ix.1985 (Matthews); 16 , 3 9 , 1.x.1985 (Matthews); 2 6 , 2 9 , 7.x.1985 (Jago); 2 9 , 28.x.1985 (Jago).

Additional material examined. lcf [GHANA] Gold Coast: Kete-Krachi (Cardinall).

Acknowledgments

This work was carried out during the tenure of a British Museum (Natural History) studentship, and the author is indebted to the Trustees of the Museum for this assistance. Field-work in Mali in 1985, and part of the cost of the colour plate in this paper, were directly funded by the Overseas

Development Administration in connection with a British Aid programme to control pests on millet in the sahel. The author wishes to thank especially Mr M. R. Honey (BMNH) for his painstaking assistance at all times during the preparation of the manuscript, and Dr M. J. Scoble (BMNH) for his helpful remarks. In addition, Dr I. J. Kitching (BMNH), Dr K. M. Harris (Commonwealth Institute of Entomol- ogy), and Dr J. D. Holloway (Commonwealth Institute of Entomology) contributed in various ways. Mr Francois Gibon kindly translated the French sections while on a short visit to the BMNH. The colour plate was photographed by Mr P. Crabb (BMNH Photographic Unit), and the other photographs taken by Mr P. York (BMNH Photographic Unit). Type material was. kindly made available for examination by Dr J. Minet (MNHN), Dr B. Gustaffson (RMS), and Professor H. J. Hannemann (ZMB). Assistance in the field was given by Dr J. Leishmann, Dr N. D. Jago and Mr J. P. Grunshaw who first observed albipunctella buzzing in 1985.

References

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de Joannis, J. (1925) Une nouvelle espkce de Raghuva Moore [Lep. Noctuidae, Agrotiinae] d’Afrique occidentale. Bulletin de la Sociitt entomologique de France, (1925), 257-260.

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Accepted 2 January 1987

the african species of heliocheilus grote (lepidoptera: noctuidae)

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