' f‘ ‘

I’.'w r mt’ii' ,' ' ■ .’••• •

'» !l 1 . •< ’.

■' -"i

t;,.

■ •'<. „•;

'4'

1

i

'4

i

4

V

1

It

n

Annals of the

Cape Provincial Museums

Natural History

Ann. Cape Prov. Mus. (nat. Hist.)

Volume 19 Part 1 30th J anuary 1994

\

1

'I

1

jUH 2 6

Published jointly by the Cape Provincial Museums
j at the Albany Museum, Grahamstown, South Africa

ANNALS OF THE CAPE PROVINCIAL MUSEUMS

These Annals are published in two series, Natural History and Human Sciences, the latter series
covering cultural history, ethnology, anthropology and archaeology. They are issued in parts at
irregular intervals as material becomes available.

The primary objective of these Annals is to disseminate the results of research work undertaken by
staff members of the Cape Provincial Museums. However, a manuscript by any author is eligible
for publication provided it is based wholly or partly on the collection/s of one or more of these
museums or if the material on which the work is based is wholly or partly deposited in one of
these museums.

Parts are obtainable from the Librarian of the Albany Museum. Correspondence in cormection
with these Annals should be addressed to the Editor, Albany Museum, Grahamstown 6140.

Editor

Dr F. W. GESS

Assistant Editor
Dr S. K. GESS

Die Gattung Sphinginopalpus Pic (Coleoptera: Malachiidae)
im siidlichen Afrika

(36. Beitrag zur Kenntnis der Fauna Afrikas)

von

W. WITTMER

(Naturhistorisches Museum, Basel, Schweiz)

INHALT

Seite

Abstract 3

Material und Verdankungen 3

Einfiihrung 4

Incertae sedis 6

Bestimmungstabelle fiir die Arten der Gattung

Sphinginopalpus Pic 8

Beschreibungen (Taxonomic) und Kommentare 16

Sphinginopalpus (Untergattung Sphinginafricanus)

1. S. (S.) atripennis Champion 18

2. S. (S.) pallidicornis sp. nov 19

3. S. (S.) delicatulus sp. nov 21

4. S. (S.) bilineaticeps Pic 22

5. S. (S.) reavelli sp. nov 24

6. S. (S.) complicatus sp. nov 25

7. S. (S.) nerinaensis sp. nov 27

8. S. (S.) endroedyi sp.nov 29

9. S. (S.) tergalis sp. nov 31

10. S. (S.) bimaculiceps sp. nov 33

11.5. (S.) isidengensis sj^.noy 34

12. 5. (S.) martini Pic 36

13. 5. (S.) consimilis sp. nov 37

14. 5. (S.) formicarius (Gorham) 38

15. 5. (S.) recurvus sp. nov 39

16. 5. (S.) recurvus planicornis ssp. nov 40

17. 5. (S.) repandus sp.nov 41

18. 5. (S.) emarginatus sp. nov 43

19. 5. (S.) reductosetosus sp. nov 45

20. 5. (S.) tubulatus sp. nov 46

21.5. (S.) natalensis sp. nov 48

22. 5. (S.) obtusodentatus sp. nov 50

23 5. (S.) unisetosus sp.nov 51

1

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Sphinginopalpus (Untergattung Sphinginopalpus)

24. S. s. str. pallidipalpis sp. nov 53

25.5. s. str. /Zav'omargmato Champion 55

26. 5. s. str. triangulatus sp. nov 57

27. 5. s. str. orientalis sp. nov 58

28. 5. s. str. lesothoensis sp. nov 60

29. 5. s. str. orangensis sp. nov 62

30. 5. s. str. tetrastigma Champion 64

31.5. s. six. hanglipensis sp.noy 66

32. 5. s. str. rufithorax sp. nov 67

33. 5. s. str. nigrithorax sp. nov 69

34.5. s. str. sp. nov 71

35.5. s. six. dentifrons sip. no\ 73

36. 5. s. str. variceps sp. nov 76

37.5. s. str. (Boheman) 77

38. 5. s. str. complexus sp. nov 80

39. 5. s. str. politipennis sp. nov 82

40. 5. s. str. trispinopsis sp. nov 83

41. 5. s. str. oneili Pic 85

42. 5. s. str. leiicosideae sp. nov 86

43. 5. s. str. amatolensis sp. nov 88

44. 5. s. str. nelshoogtensis sp. nov 90

45. 5. s. str. nelshoogtensis transformatus spp. nov 93

46. 5. s. str. quadratipalpus sp. nov 93

47. 5. s. str. trimaculatus sp. nov 95

48. 5. s. str. tridensmimus sp. nov 97

49. 5. s. str. longidens Champion 99

50. 5. s. str. bidens Champion 100

51.5. s. str. sp. nov. 102

52. 5. s. str. nigriceps sp. nov 105

53. 5. s. str. uitsoekensis sp. nov 106

54. 5. s. str. ciin’ispinus sp. nov 109

55. 5. s. str. acutispiniis sp. nov 110

56. 5. s. str. majeri sp. nov 112

57. 5. s. str. expansits sp. nov 114

58. 5. s. str. oculimacidatus sp. nov 114

59. 5. s. str. Zulu sp. nov 117

60. 5. s. six. politus sp. nov 118

61. 5. s. str. levemaculatus sp. nov 120

62. 5. s. str. trispinus sp. nov 123

63. 5. s. str. productus sp. nov 125

64. 5. s. str. nidehecki Wittmer 127

65. 5. s. str. wezaensis sp. nov 129

66. 5. s. str. dentatus sp. nov 131

2

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

67. S. s. str. bedfordensis sp. nov 132

68. S. s. str. zimbabwensis sp. nov 134

69. S. s. str. basutolandicus sp. nov 137

70. S. s. str. breytenbachae sp. nov 139

71. 5. s. str. elongatidens sp. nov 140

72. S. s. str. karkloofensis sp. nov 142

73. S. s. str. nylsvleyensis sp. nov 144

74. S. s. str. tridens sp. nov 146

Literatur 147

ABSTRACT

The species of the genus Sphinginopalpus Pic occurring in southern Africa are revised. From a
total of 13 species placed by Champion (1922) in the genus 11 remain in Sphinginopalpus. The
subgenus Sphinginafricanus is described to include all species in which the third segment of the
maxillary palps is elongate in the female, namely atripennis Champion, bilineaticeps Pic, martini
Pic, formicarius (Gorham) and the new species pallidicornis, delicatulus, reavelli, complicatiis,
nerinaensis, endroedyi, tergalis, bimaculiceps, indengensis, consimilis, recurviis, recw-viis ssp.
planicornis, repandus, emarginatus, reductosetosus, tiibulaliis, natalensis, obtusodentatiis and
unisetosus. The following species remain in Sphinginopapalpus s.str.: flavomarginatus Champion,
tetrastigma Champion, albilabris (Boheman), oneili Pic, longidens Champion, bidens Champion,
rudebecki Wittmer, and the new species described: pallidipalpis, triangulatus, orientalis,
lesothoensis, orangensis, hanglipensis, mfithorax, nigrithorax, capensis, dentifrons, variceps,
complexus, politipennis, trispinopsis, leucosideae, amatolensis, nelshoogtensis, nelshoogtensis
ssp. transformatus, qiiadratipalpiis, trimaculatus, tridensmimus, dwesaensis, nigriceps, uitsoekensis,
curvispinus, acutispinus, majeri, expansus, oculimacidatiis, zidii, politus, levemaculatiis, trispinus,
productus, wezaensis, bedfordensis, zimbabwensis, basutolandicus, breytenbachae, elongatidens,
karkloofensis, nylsvleyensis and tridens. Some species described as females had to be placed under
incertae sedis: Chalicorus collaris Boheman (Sphinginopalpus subgen. Sphinginafricanus),
Sphinginopalpus barkeri Pic (Sphinginafricanus), Chalicorus myrmecodes Boheman
(Sphinginopalpus s. str.), Chalicorus albifrons Boheman (Sphinginopalpus s. str.).

MATERIAL UNO VERDANKUNGEN

Fur die Bearbeitung lag mir u. a. das ausserordentlich reichhaltige Material des Transvaal
Museums, Pretoria, zur Verfiigung, das fast ausschliesslich durch meinen lieben Freund Dr. Sebastian
Endrody-Younga zusammengetragen wurde. Ihm und den vielen anderen Kollegen die mir Material
und Typen zum Studium zur Verfiigung stellten, sei hier herzlichst gedankt.

Meinen besonderen Dank spreche ich auch Herrn Prof. Dr. R. Guggenheim und seinen
Mitarbeitern R. Colombo und Evi Zuberbiihler vom Labor fiir Rasterelektronenmikroskopie der
Universitat Basel aus, fiir die vielen Aufnahmen die ich dort machen durfte. Ebenso gilt mein Dank
Herrn Karel Majer fiir die zahlreichen mit grosser Geduld und Sachkenntnis ausgefiihrten
Strichzeichnungen.

3

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Verwendete Abkiirzungen;

AMG = Albany Museum, Grahamstown, Dr. F. W. Gess

BM = British Museum (The Natural History Museum), London, Mr. D. Kempster
MP = Museum de Paris, Mr. J. J. Menier

NCI = National Collection of Insects, Pretoria, Mr. R. Oberprieler
NHMB = Naturhistorisches Museum, Basel, Dr. M. Brancucci
NMB = National Museum, Bloemfontein, Prof. Dr. S. Louw
RS = Rijksmuseum, Stockholm, Dr. P. Lindskog
SAM = South African Museum, Cape Town, Mrs. M. Cochrane
TMP = Transvaal Museum, Pretoria, Dr. S. Endrody-Younga
ZML = Zoological Museum, Lund, Dr. R. Danielsson

EINFUHRUNG

Champion (1922) revidierte die Vertreter der Gattung Sphinginopalpus Pic aus dem siidlichen
Afrika. Er zahlte 13 Arten auf und beschrieb die Untergattung Chalicorophasis mit der Art rajfrayi.
Diese Untergattung stellt eine eigene Gattung dar and muss won Sphinginopalpus getrennt werden.
Sie steht den mit Colotes verwandten Gattungen nahe und hat wie diese 4-gliedrige Vordertarsen bei
den S , nebst weiteren Merkmalen, auf die in einer separaten Publikation eingetreten wird.

Die von Pic als Sphinginopalpus formicoides nach 1 ? beschriebene Art, gehdrt nicht in diese
Gattung und wird provisorisch zu Chalicorophasis comb. nov. gestellt.

Bei der Bearbeitung der Arten ergab sich, dass sie in zwei gut differenzierte Gruppen aufgeteilt
werden konnen. Als Typus-Art von Sphinginopalpus hat oneili Pic, 1903, zu gelten. Die Arten von
Sphinginopalpus s. str. zeichnen sich alle durch einen mehr parallelen Korper aus. Insbesondere die
Fliigeldecken (Abb. D, E) sind in beiden Geschlechtern weniger aufgewolbt. Bei 6 und ? sind die
Schulterbeulen normal ausgebildet. Als besonderes Merkmal der ? (Abb. F) ist das breite, beilformige
letzte Glied der Maxillarpalpen anzusehen. Die Maxillarpalpen der S sind stark vergrossert und
vielformig gebildet und geben sehr brauchbare Merkmale fiir die Unterscheidung der Arten ab.

Fiir die zweite Gruppe von Arten errichte ich die Untergattung Sphinginafricanus subgen. nov.
mit der Typus-Art recurvus sp. nov. Diese Untergattung hat einige Merkmale mit Sphinginopalpus
s. str. gemeinsam, so z. B. den Bau des Kopfes, der meistens breiter ist als der Halsschild, die
vergrosserten Maxillarpalpen bei den 6 , den meistens langer als breiten Halsschild, dessen vorderer
Teil stark aufgewolbt, zur Basis verengt und abgeflacht ist, sowie die 5- gliedrigen Vordertarsen bei
6 und ?. Als besonderes Unterscheidungsmerkmal der neuen Untergattung ist die Form der
Fliigeldecken zu betrachten, die breit oval und stark aufgewolbt sind (Abb. A, B); Schulterbeulen bei
den 6 vorhanden, manchmal mit kurzem angedeutetem Langswulst dahinter. Bei den $ , die meistens
fliigellos sind, sind die Schulterbeulen stark zuriickgebildet und fehlen ganz, Oder nur als kleines
spitzes Gebilde vorhanden. Das Endglied der Maxillarpalpen ist schmal und lang (Abb. A, C) und
nicht beilformig wie bei Sphinginopalpus s. str. (Abb. F). Die Fiihler sind meistens langer, oder so
lang wie die Fliigeldecken, bei Sphinginopalpus s. str. meistens um 1/4 bis 1/5 kiirzer als diese. Der
Halsschild ist seitlich, auf dem abgeflachten Teil, bei Sphinginafricanus weniger deutlich eingedriickt
(Abb. G) als bei Sphinginopalpus s. str. (Abb. H). Bei beiden Gruppen bilden die Eindriicke eine
kurze Langsrippe an ihrem Innemande, die bei Sphinginopalpus s. str. deutlicher ist als bei der
Untergattung.

4

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

F

Abb. A-F. A-C. Sphinginopalpus (Sphingmafricanus) recurvus sp. nov. 9 : A, Gesamtansicht (xl7); B, idem fast im
Profil (xl7); C, Maxillarpalpus. D-E. Sphinginopalpus s. str. albilabris (Boheman) 9 (x25): D,
Gesamtansicht; E, idem fast im Profil. F. Sphinginopalpus s. str. orangensis sp. nov. 9, Maxillarpalpus.

5

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. G-H. Basis des Halsschildes 9 (xl70); G, Sphinginopalpus
(Sphinginafricanus) recurvus sp. nov.; H, Sphinginopalpus s. str.
albilabris (Boheman).

Ein paar Aiten die nur nach S
beschrieben werden, stelle ich
provisorisch zu Sphinginopalpus s.
str. bis die dazugehorigen 2
aufgefunden werden.

Damit die letzten

Abdominalsegmente etc. untersucht
und gezeichnet werden konnten,
wurde das Abdomen in einer Lbsung
von 15- 20% KOH mit Wasser
wahrend 1 bis 2 Minuten aufgekocht.
Dabei war es besonders bei Sternit 8
nicht zu vermeiden, dass dieses,
wenn es ganz oder teilweise im
rechten Winkel zu Sternit 7 stand,
flachgelegt werden musste, um ein
Zeichnen zu ermoglichen. Dies ist zu
beriicksichtigen, wenn das Sternit 8
am Abdomen mit den Zeichnungen
verglichen wird, ohne es separat
auszubreiten. Das Tergit 8 ist apikal
mehr oder weniger breit nach itmen
umgeschlagen, auf den Zeichnungen
ist diese Stelle mit einer punktierten
Linie oder einer dunklen Stelle
eingezeichnet.

Die in den Beschreibungen
vorkommenden Zahlen in
Klammern (zum Beispiel: 17x15)
zeigen das Verhaltnis der Lange zur
Breite an oder umgekehrt; sie
wurden bei einer Vergrosserung von
32x gemessen.

INCERTAE SEDIS

Verschiedene Arten sind nach $ beschrieben worden. Es ist momentan unmoglich sie zu
bestimmen, oder den Arten zuzuordnen, die nach 6 beschrieben wurden, oder gar zu synonymisieren,
wie es Champion (1922) versucht hat. Es miissten Merkmale gefunden werden, die es erlauben, die
$ zu unterscheiden. Dies ist bisher nicht mbglich gewesen und wir miissen diese Arten vorlaufig
unter “incertae sedis’’ auffiihren.

6

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA; MALACHIIDAE)

Abb. I-K. Sphinginopalpus (Sphinginafricanus) recurvus sp. nov. 6 : I, Gesamtansicht (xl7); K,
Kopf und Halsschild fast im Profil (x75).

7

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Sphinginopalpus (Sphinginafricanus) collaris (Boheman)

Chalicorus collaris Boheman, 1851: 478.

Sphinginopalpus collaris (Boheman) - Champion, 1922: 331.

Holotypus $ im RS.

Sphinginopalpus (Sphinginafricanus) barkeri Pic

Sphinginopalpus barkeri Pic, 1904: 65. - 1911: 175. - Champion, 1922:330.

Holotypus $ im MP.

Sphinginopalpus s. str. myrmecodes (Boheman)

Chalicorus myrmecodes Boheman, 1851: 475.

Sphinginopalpus myrmecodes (Boheman) - Champion, 1922: 330.

Holotypus 2 im RS.

Die von Champion, 1922, als myrmecodes bestimmten und beschriebenen S 2 Exemplare mit
Fundorten “mouth of Umkomaas River” und “Seymour” gehoren zu amatolensis Wittmer. Die
verbleibenden 2 von anderen Fundorten sind unbestimmbar.

Sphinginopalpus s. str. albifrons (Boheman)

Chalicorus albifrons Boheman, 1851: 476.

Sphinginopalpus albip'ons (Boheman) - Champion, 1922: 330.

Der Holotypus dieser Art war trotz eifrigem Suchen im RS nicht aufzufinden, sodass auch sie
unter “incertae sedis ” aufgefiihrt wird. Der Farbung nach konnte es sich um 1 3 handeln, was auch
Champion, 1922, vermutete.

BESTIMMUNGSTABELLE FUR DIE ARTEN DER GATTUNG SPHINGINOPALPUS PIC

1. Letztes Glied der Maxillarpalpen bei den 2 schmal (Abb.C)

Sphinginafricanus subgen. nov 2

- Letztes Glied der Maxillarpalpen bei den 2 stark verbreitert, beilformig (Abb. F)

Sphinginopalpus s. str 24

2. Fiihlerglied 1 bei den 3 einfach, ohne Zahn 3

- Fiihlerglied 1 bei den 3 gezahnt 5

3. Spitzen von Sternit 8 nach irmen gebogen 4

- Spitzen von Sternit 8 nach aussen gebogen (Abb. 6)

1. 5. (S.) atripennis Champion

4. Apex von Tergit 8 mit je einer kurzen Spitze seitlich und einer langen mittleren Spitze (Abb. 12).
Fliigeldecken an den Seiten hell gesaumt

2. S. (S.) pallidicornis sp. nov.

- Apex von Tergit 8 gerunded, fast halbkreisformig, in der Mitte schwach ausgerandet (Abb. 18).

8

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Fliigeldecken einfarbig schwarz

3. S. (S.) delicatulus sp. nov.

5. Wenigstens 1 Fiihlerglied zwischen 3 und 7 mehi oder weniger verbreitert (*) 6

- Keines der Fiihlerglieder 3 bis 7 verbreitert 8

6. Fiihlerglieder 3 oder 4 am breitesten. Apex von Tergit 8 in der Mitte nicht in 2 kleine, kurze

Fortsatze ausgezogen 7

- Fiihlerglied 5 am breitesten (Abb. 24). Apex von Tergit 8 fast gerade abgestutzt, in der Mitte in 2
kleine, kurze Fortsatze ausgezogen (Abb. 25)

4. 5. (S.) bilineaticeps Pic

7. Fiihlerglied 3 am breitesten (Abb. 30). Tergit 8 schwach, breit ausgerandet, in der Mitte kurz,
schmal nach innen gefaltet (Abb. 31). Spitzen von Sternit 8 lang, nach aussen gebogen (Abb. 32)

5. S. (S.) reave Hi sp. nov.

- Fiihlerglieder 3 und 4 am breitesten (Abb. 103). Tergit 8 am Apex jederseits breit gerundet, Mitte
schwach ausgerandet, mit / ccrvmrx iibereinstimmend (Abb. 97). Spitzen von Sternit 8 kraftig, nach
innen gebogen (Abb. 98)

16. S. (S.) recun’us planicornis n.ssp.

8. Apex von Tergit 8 in der Mitte vorstehend 9

- Apex von Tergit 8 gerade bis mehr Oder weniger tief ausgerandet 11

9. Apex von Sternit 8 aus breiten Teilen bestehend 10

- Apex von Sternit 8 aus schmalen, langen Fortsatzen bestehend, die an der Spitze nach innen oder
nach aussen verlangert sind (Abb. 38)

6. S. (S.) complicatus sp. nov.

10. Tergit 8 gegen die Spitze verschmiilert, diese in einen kleinen abgesetzten Fortsatz ausgezogen
(Abb. 43). Sternit 8 zur Spitze verschmalert, aber nicht in eine scharfe Spitze ausgezogen (Abb.
44). Fiihlerglied 1 mit einem scharfen Zahn nach der Mitte (Abb. 42), davor nicht ausgerandet.
Fliigeldecken einfarbig schwarz

7. S. (S.) nerinaensis sp. nov.

- Tergit 8 zur Spitze breiter, Spitze stumpf, wenig vorstehend (Abb. 50). Sternit 8 in 2 scharfe nach
innen gebogene Spitzen ausgezogen (Abb. 51). Fiihlerglied 1 mit einem Zahn an der Basis, davor
ausgerandet (Abb. 49). Fliigeldecken schmal gelblichweiss gesaumt

8. S. (S.) endroedyi sp. nov.

11. Zwischen der Ausrandung von Tergit 8 befinden sich keine zusatzliche, lange, schmale

Fortsatze 12

- Zwischen der Ausrandung von Tergit 8 befinden sich 2 lange, schmale Fortsatze (Abb. 57)

9. S. (S.) tergalis sp. nov.

12. Spitzen von Sternit 8 nach innen gebogen 13

- Spitzen von Sternit 8 nach aussen gebogen 19

(*) Hierher gehort auch S. (S.) reductosetosus Wittmer, desscn Fiihlerglied 5 auch, jedoch nur sehr wenig
verbreitert ist (in dieser Tabelle unter 19. aufgefiihrt). Von bilineaticeps Pic durch die sehr verschiedenen
Terminalia etc. zu unterscheiden, vergleiche mit Abb. 116-123.

9

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

13. Einschnitt am Apex von Tergit 8 ca. 1/3 der Lange erreichend (in der Mitte gemessen) 14

- Einschnitt am Apex von Tergit 8 deutlich kiiizer als 1/3 der Lange 17

14. Innensack des Aedeagus mit mehreren, oder ohne stark sklerotisierte Hornchen. Spitze von

Fiihlerglied 1 nach itmen gerichtet 15

- Irmensack des Aedeagus mit nur 1 stark sklerotisierten Hornchen (Abb. 67-68). Spitze von
Fiihlerglied 1 nach aussen gerichtet (Abb. 62)

10. S. (S.) bimaciiliceps sp. nov.

15. St emit 8 jederseits mit 2 kurzen Borsten 16

- Sternit 8 jederseits mit nur 1 langen Borste (Abb. 73)

11.5. (S.) isidengensis sp. nov.

16. Innensack des Aedeagus mit 2 oder mehreren stark sklerotisierten Hornchen (Abb. 82)

12. S. (S.) martini Pic

- Innensack des Aedeagus ohne stark sklerotisierte Hornchen (Abb. 85)

13. S. (S.) consimilis sp. nov.

17. Aedeagus mit stark sklerotisierten Hornchen 18

- Aedeagus ohne stark sklerotisierte Hornchen (Abb. 92-93)

14. S. (S.) formicarius (Gorham)

18. Tergit 8 am Apex wenig tief eckig eingeschnitten (Abb. 96)

15. S. (S.) recun’us sp.nov.

- Tergit 8 am Apex zuerst nach irmen abgeschragt, dann kurz eingeschnitten (Abb. 107)

17. S. (S.) repandus sp. nov.

19. Sternit 8 mit 1 oder 2 langen Borsten jederseits 20

- Sternit 8 ohne Borsten (Abb. 114)

18. 5. (S.) emarginatus sp. nov.

20. Sternit 8 mit 2 langen Borsten jederseits 21

- Sternit 8 mit 1 langen Borste jederseits 23

21. Die Spitzen am Apex von Sternit 8 sind lang, fast gerade, oder nur leicht nach aussen

gebogen 22

- Die Spitzen am Apex von Sternit 8 sind sehr kurz, hakenartig nach innen gebogen (Abb. 121)

19. S. (S.) reductosetosiis sp. nov.

22. Ausser den beiden nur leicht nach aussen gebogenen Spitzen am Apex von Sternit 8, sind 2 scharfe
nach vorne gerichtete Spitzen vorhanden (Abb. 128)

20. S. (S.) tubidatus sp. nov.

- Nur 2 leicht nach aussen gebogene Spitzen am Apex von Sternit 8 vorhanden, die beiden nach
vorne gerichteten Spitzen fehlen (Abb. 134)

21. S. (S.) natalensis sp. nov.

10

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA; MALACHIIDAE)

23. Sternit 8 lang, beide Teilstxicke vollstandig getrennt (Abb. 142). Tergit 8 dreieckig, ziemlich tief
eingeschnitten (Abb. 141)

22. S. (S.) obtusodentatus sp. nov.

- Sternit 8 kiirzer, die beiden Teilstiicke in der Mitte durch feines Gewebe verbunden (Abb. 148)
Tergit 8 schwach, breit ausgerandet (Abb. 147)

23. 5. (S.) unisetosus sp. nov.

24. Fiihlerglied 1 bei den S ohne Zahn 25

- Fiihlerglied 1 bei den S nach der Basis ausgerandet und mit Zahn versehen 27

25. Fiihlerglied 1 schmal, nirgends stark verbreitert 26

- Fiihlerglied 1 gegen die Mitte stark verbreitert und gegen die Spitze wieder ein wenig verschmalert
(Abb. 152)

24. S. s. str. pallidipalpis sp. nov.

26.

Apex von Tergit 8 drei Spitzen bildend, die mittlere am grossten und langsten (Abb. 160)

25. S. s. six. flavomarginatus Champion

Apex von Tergit 8 drei Spitzen bildend, die mittlere ist kleiner und kiirzer als die seitlichen
(Abb. 167)

26. 5. s. str. triangulatus sp. nov.

27. Halsschild rotbraun oder orange, selten mit einem dunklen Flecken in der Mitte 28

- Halsschild schwarz, abgesetzte Basis mehr oder weniger breit hell, selten ist der ganze Halsschild
schwarz 33

28. Apex von Tergit 8 etwas eingeschnitten oder ausgerandet, oder in zackiger Linie

verlaufend 29

- Apex von Tergit 8 gerundet, ohne Einschnitt (Abb. 178)

27. 5. s. str. orientalis sp. nov.

29. Apex von Sternit 8 mit nach innen gekriimmten Spitzen 30

- Apex von Sternit 8 mit nach aussen gebogenen Spitzen 31

30. Apex von Tergit 8 gerundet eingeschnitten (Abb. 180)

28. S. s. str. lesothoensis sp. nov.

- Apex von Tergit 8 in zackiger Linie verlaufend (Abb. 186)

29. S. s. str. orangensis sp. nov.

31. Die beiden Seiten von Sternit 8 stellenweise zusammen-gewachsen 32

- Die beiden Seiten von Sternit 8 vollstandig voneinander getrennt (Abb. 192)

30. S. s. str. tetrastigma Champion

32.

Kopulationsapparat in einen langen, schmalen Fortsatz ausgezogen, dessen Spitze ventralwarts
gebogen ist (Abb. 195- 196)

31. S. s. str. hanglipensis sp. nov.

Kopulationsapparat normal gebaut, ohne langen, schmalen Fortsatz (Abb. 207)

32. S. s. str. nifithorax sp. nov.

11

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

33. Fliigeldecken mil einem durchgehenden weisslichen oder gelblichen Querband vor der

Mitte 34

- Fliigeldecken anders gezeichnet 35

34. Halsschild einfarbig schwarz. Apex von Sternit 8 mil 2 grossen spitzen Zahnen (Abb. 212)

33. 5. s. sir. nigrithorax sp. nov.

- Halsschild schwarz mil heller Basis. Apex von Sternit 8 mit 2 kleinen hakenartigen Zahnen
(Abb. 218)

34. 5. s. str. capensis sp. nov.

35. Apex von Sternit 8 jederseits in mehr oder weniger lange, meist schmale Fortsatze ausgezogen,

Oder nach innen gerichtete Spitzen bildend 48

- Apex von Sternit 8 jederseits eine mehr oder weniger grosse, breite Flache, nicht in schmale

Fortsatze ausgezogen, apikal keine Spitzen bildend 36

36. Vorderstirne flach, ohne spitzen Zahn in der Mitte 37

- Vorderstirne mit einem kleinen, spitzen Zahn in der Mitte (Abb. 222)

35. S. s. str. dentifrons sp. nov.

37. Apex von Tergit 8 in der Mitte mehr oder weniger tief eingeschnitten, manchmal ist der Einschnitt

schwer sichtbar, z. B. bei albilabris 38

- Apex von Tergit 8 in der Mitte mehr oder weniger vorstehend, bis fast gerade, nicht
eingeschnitten 40

38. Die beiden Teile von Sternit 8 sind nicht durch loses Gewebe miteinander verbunden 39

- Die beiden Teile von Sternit 8 sind durch loses Gewebe miteinander verbunden (Abb. 231)

36. S. s. str. variceps sp. nov.

39. Apex von Tergit 8 in der Mitte kaum merklich eingeschnitten (Abb. 236). Sternit 8 weniger
komplex gebaut (Abb. 237)

37. S. s. str. albilabris (Boheman)

- Apex von Tergit 8 in der Mitte zuerst breit und dann ziemlich tief eingeschnitten (Abb. 243).
Sternit 8 sehr komplex gebaut (Abb. 244)

38. S. s. str. complexus sp. nov.

40. Apikalrand von Tergit 8 in der Mitte eckig vorstehend 41

- Apikalrand von Tergit 8 in der Mitte, wenn vorstehend, datm nur wenig, nicht eckig, sondern

gerundet oder fast gerade 43

41. Apikalrand von Tergit 8 in der Mitte in spitzem Winkel vorstehend, nicht eingeschnitten ....42
- Apikalrand von Tergit 8 in der Mitte vorstehend und kurz eingeschnitten (Abb. 249)

39. S. s. str. politipennis sp. nov.

42. Fiihlerglied 1 fast parallel, schmal mit kraftigem Basalzahn und langem, schmalem Zahn (Abb.
252). Tergit 8 breit (Abb. 253)

40. 5. s. str. trispinopsis sp. nov.

12

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

- Fiihlergiied 1 fast dreieckig, nicht parallel, der Basalzahn fehJt, der vordere Zahn breit, ausgehbhlt
(Abb. 258). Tergit 8 schmal (Abb. 259)

41. 5. s. str. oneiii Pic

43. Die beiden Seitenteiie von Sternit 8 hangen lose zusammen 44

- Die beiden Seitenteiie von Sternit 8 hangen nicht zusammen, sie sind vollstMdig frei 45

44. Fiihlergiied 1 fast parallel, an der Basis kurz ausgehohlt, Zahn kaum angedeutet (Abb. 268)

42. S. s. str. leucosideae sp. nov.

- Fiihlergiied 1 nicht parallel, an der Basis langlich ausgehohlt, Zahn sehr klein, deutlich (Abb.
274)

43. S. s. str. amatolensis sp. nov.

45. Die Aushohlung an der Basis von Fiihlergiied 1 befindet sich vor der Mitte 46

- Die Aushohlung von Fiihlergiied 1 erreicht die Mitte oder iiberragt sie sogar, an Stelle eines
Zahnes ist dieser Teil breit nach unten verlangert (Abb. 280)

44. S. s. str. nelshoogtensis sp. nov.

46. Zahn von Fiihlergiied 1 mehr dreieckig, an der Basis nicht eingeschniirt 47

- Zahn von Fiihlergiied 1 nicht dreieckig, nach der Basis eingeschniirt (Abb. 287)

45. S. s. str. quadratipalpus sp. nov.

47. Zahn von Fiihlergiied 1 breiter (Abb. 284)

45. S. s. str. nelshoogtensis transformatus ssp. nov.

- Zahn von Fiihlergiied 1 zierlicher (Abb. 293)

47. S. s. str. trimaculatus sp. nov.

48. Die Fortsatze von Sternit 8 sind nach aussen gebogen 49

- Die Fortsatze von Sternit 8 sind nach irmen gebogen und spitz 53

49. Jeder Seitenteil von Sternit 8 ausser den Fortsatzen am Apex mit einem zusatzlichen spitzen

Fortsatz auf der Innenseite 50

- Jeder Seitenteil von Sternit 8 ohne zusatzlichen spitzen Fortsatz auf der Innenseite 51

50. Der zusatzliche Fortsatz von Sternit 8 auf der Innenseite ist sehr lang, an der Basis sehr breit
(Abb. 302)

48. S. s. str. tridensmimus sp. nov.

- Der zusatzliche Fortsatz von Sternit 8 ist auf der Innenseite sehr kurz, an der Basis schmal,
dornformig (Abb. 308)

49. S. s. str. longidens Champion

51. Fiihlergiied 1 an der Basis ohne zusatzlichen Zahn 52

- Fiihlergiied 1 an der Basis mit einem breiten zusatzlichen Zahn (Abb. 313)

50. S. s. str. bidens Champion

52. Zahn auf Fiihlergiied 1 gross, er bildet einen schwachen spitzen Winkel (Abb. 321)

51. S. s. str. dwesaensis sp. nov.

13

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

- Zahn auf Fiihlerglied 1 nicht spitz, sondern breit gerundet, nur an der Basis befindet sich eine
kurze Spitze (Abb. 327)

52. S. s. str. nigriceps sp. nov.

53. Sternit 8 ohne lange, nach innen gerichtete Borsten 54

- Sternit 8 mit 1 oder 2 langen, nach innen gerichteten Borsten 64

54. Sternit 8 ohne langen, nach hinten gerichteten Fortsatz 55

- Sternit 8 mit langem, nach hinten gerichtetem, spitzem Fortsatz, der fast parallel mit dem Seitenteil
verlauft (Abb. 335)

53. S. s. str. uitsoekensis sp. nov.

55. Sternit 8 zur Spitze nicht verbreitert, dahinter nicht eingeschniirt 57

- Sternit 8 zur Spitze verbreitert, dahinter eingeschniirt 56

56. Apikaler Teil von Sternit 8 fast parallelseitig, daim jederseits mit einem dorsalwarts gerichteten,
leicht gekriimmmten, spitzen Fortsatz (Abb. 340)

54. S. s. str. curvispinus sp. nov.

- Apikaler Teil von Sternit 8 nicht fast parallel, sondern zur Spitze verbreitert, ohne dorsal
gerichteten spitzen Fortsatz (Abb. 346)

55. 5. s. str. acutispinus sp. nov.

57. Sternit 8 nicht leierformig geformt 61

- Sternit 8 fast leierformig (Abb. 352, 363, 369) 58

58. died 3 der Maxillarpalpen 2 mal so lang oder noch langer als der Vorderrand dieses

Gliedes 59

- died 3 der Maxillarpalpen weniger als 2 mal so lang wie der Vorderrand dieses Gliedes 60

59. Glied 2 der Maxillarpalpen eine breite Lamelle mit fast geradem Distalrand (Abb. 348)

56. S. s. str. majeri sp. nov.

- Glied 2 der Maxillarpalpen distal nicht breit, das Glied ist schmal, etwas gebogen (Abb. 356)

57. 5. s. str. expansus sp. nov.

60. Glied 3 der Maxillarpalpen apikal kaum verschmalert, breit gerundet (Abb. 359)

58. 5. s. str. oculimaculatus sp. nov.

- Glied 3 der Maxillarpalpen apikal verschmalert, ausgerandet (Abb. 365)

59. S. s. str. Zulu sp. nov.

61 . Die beiden Seitenteile von Sternit 8 hangen in der Mitte nicht lose zusammen, sie sind vollstandig

getrermt. Tergit 8 apikal in der Mitte spitz vorstehend 62

- Die beiden Seitenteile von Sternit 8 hangen in der Mitte zusammen (Abb. 375). Tergit 8 apikal
gerundet

60. 5. s. str. politus sp. nov.

62. Spitzen von Sternit 8 stark nach innen gebogen 63

- Spitzen von Sternit 8 schwach nach innen gebogen (Abb. 382)

61. S. s. str. levemaculalus sp. nov.

14

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

63. Fiihierglied 3 ein wenig verbreitert (Abb. 386). Die nach irnien gerichteten Spitzen von Sternit 8
befinden sich kurz vor der Spitze (Abb. 388)

62. S. s. str. trispinus sp. nov.

- Fiihierglied 3 nicht verbreitert (Abb. 392). Die nach innen gerichteten Spitzen von Sternit 8
beginnen an der Spitze (Abb. 394)

63. S. s. str. productus sp. nov.

64. Sternit 8 jederseits mit 1 langen nach innen gerichteten Borste 65

- Sternit 8 jederseits mit 2 langen nach innen gerichteten Borsten, ofters erscheinen diese beiden

Borsten wie zusammengeklebt 68

65. Die beiden Seitenteile von Sternit 8 sind miteinander durch loses Gewebe verbunden 66

- Die beiden Seitenteile von Sternit 8 sind vollstandig getrennt 67

66. Fiihierglied 1 bis ungefahr zur Mitte tief ausgehdhlt (Abb. 398). Tergit 8 apikal gerundet mit
einem kleinen Fortsatz in der Mitte (Abb. 399)

64. S. s. str. rudebecki sp. nov.

- Fiihierglied 1 nur an der Basis kurz ausgehdhlt (Abb. 405). Tergit 8 apikal in Wellenlinie, in der
Mitte ein wenig mehr vorstehend als an der Seiten (Abb. 406)

65. S. s. str. wezaensis sp. nov.

67.

Fiihierglied 1 fast dreieckig mit einem grossen, langen Zahn in der Mitte und einem viel kleineren
an der Basis (Abb. 411)

66. S. s. str. dentatus sp. nov.

Fiihierglied 1 langlich, nicht dreieckig, mit nur 1 Zahn nach der Aushdhlung (Abb. 418)

67. S. s. str. bedfordensis sp. nov.

68. Fiihierglied 1 vor dem Zahn ein wenig ausgehdhlt, aber an der Basis nicht verdickt 69

- Fiihierglied 1 an der Basis verdickt und hier manchmal mit 1 Zahn 70

69. Fiihierglied 1 fast parallel, Aushdhlung klein, Zahn kaum angedeutet (Abb. 424). Tergit 8 apikal
in der Mitte stumpfwinklig vorstehend (Abb. 425)

68. S. s. str. zimbabwensis sp. nov.

- Fiihierglied 1 langlich dreieckig mit kraftigem Zahn (Abb. 430). Tergit 8 apikal regelmassig
gerundet (Abb. 431)

69. S. s. str. basutolandicus sp. nov.

70. Fiihierglied 1 an der Basis mit 1 Zahn oder mit 1 stumpfen Hdcker 71

- Fiihierglied 1 an der Basis ohne Zahn oder Hdcker, hier nur ausgehdhlt, diese Stelle einen
stumpfen Winkel bildend (Abb. 436)

70. S. s. str. breytenbachae sp. nov.

71. Fiihierglied 1 mit einem Zahn an der Basis 72

- Fiihierglied 1 mit einem Hdcker an der Basis, Zahn besonders lang, etwas verdreht (Abb. 442)

71. S. s. str. elo?igatidens sp. nov.

15

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

72. Fiihlerglied 1 an der breitesten Stelle spitz vorstehend 73

- Fiihlerglied 1 an der breitesten Stelle einen leicht stumpfen Winkel bildend (Abb. 448)

72. S. s. str. karkloofensis sp. nov.

73. Die beiden Seitenteile von Sternit 8 sind getrennt (Abb. 455). Fiihlerglied 1 mit einer kurzen
Querleiste neben der Aushohlung (Abb. 453)

73. S. s. str. nylsvleyensis sp. nov.

- Die beiden Seitenteile von Sternit 8 hangen zusammen (Abb. 460). Fiihlerglied 1 mit einem
kleinen zusatzlichen Zahn neben der Aushohlung (Abb. 458)

74. S. s. str. tridens sp. nov.

BESCHREIBUNGEN (TAXONOMIE) UND KOMMENTARE

1. Sphinginopalpus (Sphitiginafricanus) atripennis Champion
Abb. 1-8

Sphinginopalpus atripennis Champion, 1922: 333.

Fiir die Abb. 1 und 2 der Maxillarpalpen diente der Holotypus im BM and fiir die Abb. 3-7 ein
S von Badplaas. Fiihlerglieder 1 bis 4 Abb. 3. Tergit 8 Abb. 4. Sternite 7 und 8 Abb. 5 und 6.
Kopulationsapparat Abb. 7.

Die Art ist auffallig durch das einfache, ungezahnte Fiihlerglied 1 beim S . Die S kommen
gefliigelt und ungefliigelt vor. Die gefliigelten S besitzen normal entwickelte Schulterbeulen, bei

den ungefliigelten Exemplaren treten
die Schultern spitz vor. Die Art is weit
verbreitet.

Abb. 1-2. Sphinginopalpus (Sphinginafricanus) atripennis Champion d : 1-
2, Maxillarpalpen (x220) (1 von oben, 2 von unten).

UNTERSUCHTES MATERIAL: Natal: St
Lucia, 10 m, 23-25. x. 1981 (J. & S.
Klapperich) (NHMB). Transvaal:
Waterval Boven, 24. xi. 1981 (J. & S.
Klapperich) (NHMB); Nelspruit Nat.
Res., groundtraps banana bait and
grass netting, 23. ix. & 19.xii. 1986, E-
Y:2287, 2402 (S. Endrody- Younga)
(TMP); 16 km N Barberton, 25°42'S,
30°57'E, 24.ix.1986 and 10.ii.l987,
ground traps banana bait, E-Y:2287
& 2438 (S. Endrody- Younga) (TMP
& NHMB); Berlin Forest Station,
1500 m, 7.xii.l986, E-Y:2360 (S.
Endrody- Younga) (TMP & NHMB);
Uitsoek Forest Station, 1100 m.

16

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 3-8. Sphinginopalpus (Sphinginafricanus) atripennis Champion S : 3, Fiihlerglieder 1-4; 4, Tergit 8; 5, Sternit 7;
6, Sternit 8; 7-8, Kopulationsapparat (7, dorsal; 8, Profil). Massstab von 4 auch fiir 5-8.

12.xii.l986, E-Y:2379 (S. Endrody-Younga) (TMP); Nelshoogte Devil’s Knuckles, grass netting,
4.xii.l986, E-Y:2350 (S. Endrody- Younga) (TMP); Nelshoogte Forest Station, grass netting,
l.xii.l986, E-Y:2336 (S. Endrody-Younga) (TMP); Soutpansberg Hanglip, 23°02"S, 29°47'E, grass
netting, 14. hi. 1973, E-Y:46 (S. Endrody-Younga) (TMP). Transkei: Fraser Gorge, grass tussocks,
31°24'S, 29°42'E,

E-Y:2584 (S. Endrody-Younga) (TMP). Zululand; Lake Bhangasi, grass netting, 28°07'S, 32°31'E,
18.xii.l976 (S. Endrody-Younga) (TMP).

17

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

2. Sphinginopalpus (Sphinginafricanus) pallidicornis sp. nov.
MAnnchen (Abb. 9-14).

Lange 1,8 - 2 mm.

Kopf schwarz, Vorderkopf mil einem mehr oder weniger grossen gelben Flecken, der bis zur
Mitte der Augen reichen kaim, ohne sie zu beriihren, manchmal auch ein Teil der Wangen hell;
Maxillarpalpen orange, Oberseite des vorletzten Gliedes dunkelbraun, Spitze des letzten sehr oft
schwach gebraunt; Fiihler orange, Spitze des lezten Gliedes oft leicht angedunkelt; Halsschild
schwarz, abgeflachter Teil an der Basis orange; Schildchen orange; Fliigeldecken schwarz, Seiten
begitmend an der Basis bis und mit Spitzen schmal gelb; Hinterschenkel schwarz. Basis breit
aufgehellt, Hintertibien an der Basis schmal hell, alle Tarsen orange, 4 Vorderbeine orange, jedoch
Schenkel teils angedunkelt.

Kopf mit den Augen breiter
als der Halsschild, Stirne leicht
gewolbt; Oberflache glatt.
Maxillarpalpen Abb. 9-10. Fiihler
(Abb. 11) auffallig durch das
ungezahnte Glied 1, 3 nur wenig
langer als 4. Halsschild langer als
breit (17x14); Oberflache glatt,
auf dem basalen Teil neben dem
Seitenrand schmal chagriniert, der
Seitenrand hier schwach
krenuliert, Fliigeldecken oval,
Schulterbeulen schwach gerundet
Oder spitz vortretend; falsche
Epipleuren nur ganz schwach
krenuliert; Punktreihen deutlich,
Punkte erst nach der Mitte an Tiefe
verlierend, Spitze fast unpunktiert.
Abb. 9-10. Sphinginopalpus (Sphinginafricanus) pallidicornis sp. nov. cj : 9- TergitS Abb. 12. Stemit 7 Abb. 13,
10, Maxillarpalpen (x200) (9, von oben; 10, von unten). Sternit 8 Abb. 14.

Weibchen.

Wie das S gefarbt, nur Kopf und Maxillarpalpen einfarbig schwarz. Schulterbeulen spitz
vortretend.

Holotypus S und 4 Paratypen (NHMB), 2 Paratypen S $ (TMP), 2 Paratypen S $ (AMG): South
Africa, Natal: 7 km NO Ladysmith, 21. xi. 1987 (W. Wittmer).

Neben S. (S.) atripennis Champion zu stellen. Die neue Art unterscheidet sich durch die hellen
Fiihler und die gelben Seiten der Fliigeldecken, sowie durch die verschieden gebauten Terminalia.
Es ist bis jetzt die einzige Art dieser Untergattung deren Tergit 8 in drei Spitzen auslauft, sie hat

18

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

deshalb grosse Ahnlichkeit mil S. s. sU.flavomarginatus Champion, doch ist das Sternit 8 verschieden
gebaut.

Abb. 11-14. Sphinginopalpus (Sphinginafricanus) pallidicornis sp. nov. S: 11, Fuhlerglieder 1-4; 12, Tergit 8;
13, Stemit 7; 14, Sternit 8. Massstab von 11 auch fiir 14 und 12 auch fiir 13.

3. Sphinginopalpus (Sphinginafricanus) delicatulus sp. nov.

MAnnchen (Abb. 15-21).

Lange 2,2 mm.

Kopf schwarz; Maxillarpalpen gelb, Spitze des
letzten Gliedes schmal angedunkelt; Fiihler gelb,
letzte 3 bis 4 Glieder dunkel; aufgewolbter Teil des
Halsschildes schwarz, abgeflachte Basis gelblich;
Schildchen, Fliigeldecken und Hinterbeine schwarz,
Hintertarsen leicht aufgehellt, 4 Vorderbeine und
Tarsen gelb, Schenkel teils angedunkelt.

Kopf mit Augen breiter als der Halsschild,
Stirne leicht gewolbt, Oberflache glatt.
Maxillarpalpen (Abb. 15-16) besonders schlank,
letztes Glied apikal zugespitzt. Fiihler (Abb. 17) mit
Glied 1 einfach (ungezahnt), 3 kaum merklich langer
als 4. Halsschild langer als breit (16x13,5), Seiten

Abb. 15-16. Sphinginopalpus (Sphinginafricanus)
delicatulus sp. nov. S : 15-16, Maxillarpalpen
(x200).

19

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

von der Mitte an schwach gerundet; Oberflache glatt, nur neben dem Seitenrand an der Basis des
abgeflachten Teils auf kurzer Strecke runzlig gewirkt. Fliigeldecken langoval, Schulterbeulen normal;
Punkte in den Reihen ziemlich tief, nicht ganz regelmassig; falsche Epipleuren ca. 45% der Lange
einnehmend, nicht krenuliert; gefliigelt. Tergit 8 Abb. 18. Sternit 8 Abb. 19.

Kopulationsapparat Abb.20 dorsal, Abb. 21 Profil.

02

Abb. 17-21. Sphinginopalpus (Sphinginafricanus) delicatulus sp. nov. 6. 17, Fiihlerglieder 1-4; 18, Tergit 8; 19, Sternit 8;
20-21, Kopulationsapparat (20, dorsal; 21, Profil). Massstab von 17 auch fur 19-21.

20

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA; MALACHIIDAE)

Weibchen unbekaimt.

Holotypus (TMP): South Africa, E. Transvaal; Nelshoogte Forest Station, 1400 m, grassnetting,
2.xii.l986, E-Y:2345, S. Endrody- Younga.

Obwohl von dieser Art keine 9 vorliegen, stelle ich sie in die Untergattung Sphinginafricanus,
weil sie so nahe mit atripennis Champion verwandt ist. Sie unterscheidet sich von ihr durch die sehr
schmalen Maxillarpalpen, die Terminalia und den Kopulationsapparat.

4. Sphinginopalpus (Sphingina frican us) bilineaticeps Pic
Sphinginopalpus bilineaticeps Pic, 1929: 2.

Die Art ist aus Natal
beschrieben: Weenen, xii.1923, H.

Abb. 22-23. Sphinginopalpus (Sphinginafricanus) bilineaticeps Pic <S : 22-23, ThomaSSet. Ein weiteres 3

Maxillarpalpen (xl65). wurde von mir ebenfalls in Natal

erbeutet: 4 km S Wasbank, R621,
1030 m, 21. xi. 1987 (NHMB). Dieses Exemplar unterscheidet sich von der Holotype lediglich durch
das etwas starker verbreiterte Fiihlerglied 5. Daim fing Dr. S. Endrody- Younga (E-Y:2787) 1 c? in
Karkloof grassveld, 29°19'S, 30°15'E (TMP).

Mannchen (Abb. 22-27).

Der Holotypus befindet sich
im BM. Schon durch die Farbung
des Kopfes fallt diese Art auf. Die
Kopfbasis is schwarz, von der Mitte
der Augen nach vorne gelb, iiber
jeder Fuhlerwurzel ein kleiner
schwarzer, langlicher Flecken. Die
beigegebenen Abbildungen

werden ein Erkennen erleichtern:
Maxillarpalpen Abb. 22-23. Tergit
8 (Abb. 25) sehr charakteristisch
durch den schmalen, in der Mitte
vorstehenden Apikalrand, der fast
bis zur Basis eingeschnitten ist.
Sternit 8 Abb. 26.
Kopulationsapparat Abb. 27 dorsal.

21

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 24-27. Spliinginopalpus (Sphinginafricanus) bilinealiceps Pic 6 : 24, Fiihlerglieder 1-6; 25, Tergit 8; 26, Sternit 8;
27, Kopulationsapparat (dorsal). Massstab von 25 auch fiir 26, 27.

5. Sphinginopalpus (Sphinginafricanus) reavelli sp. nov.

MAnnchen (Abb. 28-33).
Lange 2, 5-2, 7 mm.

Abb. 28-29. Sphinginopalpus (Sphinginafricanus) reavelli sp. nov. S : 28-29,
Maxillarpalpen (xl30) (28, von oben; 29, von unlen).

Kopf gelb bis orange,
Stirne mil einem mehr oder
weniger blassen, dunkeln
Flecken, der bis zur Mitte der
Augen reicht, SchlMen hell;
Maxillarpalpen hellbraun, oft
mil 1 bis 2 kleinen dunkeln
Flecken; Fiihler gelb, Glieder 3
bis 5 Oder 6 dunkel, Oberseite
von 2 meistens angedunkelt.
Halsschild schwarzlich bis
mehr oder weniger dunkel
braun. Basis orange bis
hellbraun; Schildchen und
Fliigeldecken schwarz, falsche
Epipleuren mit schwacher

22

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Tendenz zu Aufhellung; Hinterbeine schwarz, Hintertibien teils aufgehellt, oder ganz gelb,
Mittelschenkel dunkel, meistens ein wenig aufgehellt, Mitteltibien schwarz bis schwarzlich,
Mitteltarsen gelb, Vorderschenkel und Vordertibien noch heller als die mittleren; alle Trochanteren
und Spitzen der Koxen gelb.

Abb. 30-33. Sphingiiwpalpus (Sphinginafricamis) reavelli sp. nov. S : 30, Fuhlerglieder 1-5; 31, Tergil 8;
32, Sternit 8; 33, Kopulationsapparat (Profil). Massstab von 30 auch I'iir 32, 33.

Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberflache chagriniert.
Maxillarpalpen Abb. 28 von oben, Abb. 29 von unten. Fiihler (Abb. 30) urn etwas mehr als 1/10
1 anger als die Fliigeldecken, Zahn auf died 1 deutlich gebogen, 3 und 4 abgeflacht, beide fast gleich
lang, 3 breiter als 4. Halsschild langer als breit (22x15), sonst wie bei cunipalpis; Oberflache
regelmassig chagriniert. Fliigeldecken wie bei curvipalpis, Punkte weniger tief. Tergit 8 Abb. 31.
Sternit 8 Abb. 32.

Kopulationsapparat Abb. 33 Profil.

Weibchen.

Kopf einfarbig schwarz, Fiihler einfarbig gelb, beim Halsschild ist auch der dunkle Teil dunkler
als beim 6 , Beine dunkler als beim 6 . Fuhlerglied 12 schmal, zur Spitze wenig verbreitert, iibrige
Glieder fast parallel. Fliigeldecken breiter als beim S ; ungefliigelt.

Holotypus and 2 Paratypen (NHMB): South Africa, Natal: Grass edge at Glinza Forest, Eshowe,
19.xii.l985, P. E. Reavell; idem 1 Paratypus (NCI); idem 25. iv. 1956, H. K. Munro, 1 Paratypus

23

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

(NCI); idem 23.- 31.iii., 1.- 22.iv., 23.- 30.iv.l926, R. E. Turner, 5 $ Paratypen (BM), 3 $ Paratypen
(NHMB).

Diese neue Art gehort zu den wenigen mit verbreiterten Fiihlergliedern und ist neben bilineaticeps
Pic zu stellen. Sie unterscheidet sich von dieser Art durch die verbreiterten Fiihlerglieder 3 und 4,
wahrendem bei bilineaticeps die Glieder 3 bis 5 verbreitert sind. Weitere Unterschiede zeigen die
Maxillarpalpen, die Terminalia und der Kopulationsapparat.

6. Sphinginopalpus (Sphinginafricanus) complicatiis sp. nov.

MAnnchen (Abb. 34-39).

Lange 2,5 mm.

Abb. 34-35. Sphinginopalpus (Sphinginafricanus) complicatus sp. nov, dunkel. Rest mit den Schienen gelb, 4
6 : 34-35, Maxillarpalpen (xl30) (34, von oben; 35, von unten). Vordertarsen gelb.

Halsschild, Stirne leicht gewolbt, Oberflache fein chagriniert. Maxillarpalpen Abb. 34-35. Fiihler
(Abb. 36) ein wenig kiirzer als die Fliigeldecken, Glied 2 fast so lang wie 3, 4 so lang wie 3, 3 bis 10
fast parallel. Halsschild ungefahr in der Mitte an breitesten; Oberflache fein chagriniert, sonst wie
bei curvipalpis. Fliigeldecken wie bei curvipalpis, Punktierung etwas weniger tief, in Reihen
angeordnet. Tergit 8 Abb. 37. Sternit 8 (Abb. 38) mit sichelfdrmigen Verlangerungen, die
iibereinander greifen.

Kopulationsapparat Abb. 39 dorsal.

Holotypus und 9 Paratypen (NHMB), 2 Paratypen (AMG): South Africa, Transvaal: Blydepoort,
20.xi.l981, J. &. S. Klapperich.

Wegen dem sehr abweichenden Sternit 8 mit keiner anderen Art vergleichbar. Am ehesten noch
neben curvipalpis Wittmer zu stellen, bei dem das Sternit 8 kurze Ansatze einer Verlangerung der
Spitzen nach aussen aufweist.

Kopf schwarz, von der Mitte der
Augen nach vorne gelb; Maxillarpalpen
schwarz; Fiihlerglieder 1 bis 5 gelb, 1
auf der Oberseite und an der Basis kurz
dunkel, 6 und folgende zunehmend
dunkel; Halsschild schwarz. Basis
gelbbraun; Schildchen und
Fliigeldecken schwarz, letztere seitlich
schmal mit Tendenz zu Aufhellung;
Hinterschenkel und grosster Teil der
Hinterschienen schwarz, Hintertarsen
gelb mit den beiden Basalgliedern
schwach dunkel; Mittelschenkel und
Mittelschienen angedunkelt,

Vorderschenkel auf der basalen Halfte

Kopf mit den Augen breiter als der

24

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 36-39. Sphinginupalpus (Sphinginafricanus) complicaius sp. nov. 6 : 36, Fuhlerglieder 1-4; 37, Tergit 8; 38, Sternit 8;
39, Kopulationsapparat (dorsal). Massstab von 39 auch fur 36, 38.

7. Sphinginopalpus (Sphinginafricanus) nerinaensis sp. nov.

MAnnchen (Abb. 40-46).

Lange 2,5 mm.

Kopf Schwarz, von der Mitte der Augen nach vorne, inklusive der Wangen gelb; Maxillarpalpen
gelb; Fiihler angedunkelt, erste' 4 bis 5 Glieder gelb, 1 auf der Oberseite angedunkelt; Halsschild
schwarz. Basis nicht sehr breit hell; Schildchen, Fliigeldecken und Beine schwarz, bei den 4 vorderen
schwache Tendenz zu Aufhellung.

Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewdlbt; Oberflache chagriniert.
Maxillarpalpen Abb. 40-41. Fuhler (Abb. 42) ca. 10% kiirzer als die Fliigeldecken, Glied 1 nicht

25

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 40-41. Sphinginopalpus (Sphinginafricanus) nerinaensis sp. nov. 6 ; 40-41, Maxillarpalpen (xl55) (40, von
oben, Glied 3 von den iibrigen getrennt und verkehrt liegend; 41, von unten),

ausgerandet, fast auf der Unterseite und erst nach der Mitte gelegen ein spitzer Zahn, 3 nur wenig
langer als 4. Halsschild liinger als breit (25x17), Seiten bis zur Halfte fast parallel, dann verengt, sonst
wie curvipalpis. Fliigeldecken wie bei curvipalpis, jedoch Punktierung schwacher, falsche Epipleuren
deutlich krenuliert. Tergit 8 Abb. 43. Sternit 8 Abb. 44. Spiculum Abb. 45.

Kopulationsapparat Abb. 46 Profil.

Weibchen.

Kopf einfarbig schwarz, sonst wie das S gefarbt; Fiihlerglied 1 sehr lang und schmal;
Schulterbeulen durch eine schwache Erhohung kaum angedeutet; ungefliigelt.

Holotypus 3 und 1 Paratypus ? (TMP): South Africa, Transvaal: Nerina Nature Reserve, 23°42'S,
30°16T, 22.iii.1975, E-Y:761, S. Endrody-Younga.

Neben reavelli Wittmer zu stellen, jedoch ohne verbreiterte Fiihlerglieder und sehr verschieden
gebaute Terminalia.

26

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 42-46. Sphinginopalpus (Sphinginafricanus) nerinaensis ,sp. nov. cJ : 42, Fuhlerglicder 1-4; 43, Tergil 8; 44, Stemit 8;
45, Spiculum; 46, Kopulationsapparat (Profil). Massstab von 44 auch fiir 45, 46.

8. Sphinginopalpus (Sphinginafricanus) endroedyi sp. nov.

Mannchen (Abb. 47-53).

Lange 2,2 - 2,3 mm (ohne Abdomen).

Kopf orange bis gelb, ein schwarzer Stirnflecken reicht bis zur Mitte der Augen und bedeckt
meistens auch die Schlafen; Maxillarpalpen orange, bei 1 Exemplar braun; Fiihler gelb, die letzten
4 bis 5 Glieder dunkel; Halsschild schwarz, abgeflachte Basis orange; Schildchen und Fliigeldecken
Schwarz, letztere mil einem gelben Seitenband, das von der Basis bis fast zu den Spitzen reicht;
Hinterbeine schwarz, manchmal sind die Hintertibien leicht aufgehellt, Hintertarsen gelb, 4 vordere
Beine gelb.

Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewdlbt; Oberflache glatt.
Maxillarpalpen Abb. 47-48. Fiihler (Abb. 49), Glied 2 verhaltnismassig lang, 3 so lang wie 4.
Halsschild langer als breit (19x14), Seiten nach vorne leicht gerundet verengt, gegen die Basis verengt,
Seitenrand deutlich, Oberflache auf dem abgeflachten Teil zwischen der kurzen, seitlichen Liingsleiste
und dem Seitenrand chagriniert. Rest glatt. Fliigeldecken oval; Schulterbeulen als kleine Beulen
vortretend, auf der eine lange Borste sitzt; falsche Epipleuren nur ganz schwach krenuliert; Punkte

27

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 47-48. Sphinginopalpus (Sphinginafricanus) endroedyi sp. nov. S : Maxillarpalpen (x200) (47, von oben; 48, von unten).

28

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

bis etwas iiber die Mitte in regelmassigen Reihen angeordnet, dann etwas verworren und an Tiefe
abnehmend; ungefliigelt. Ter git 8 Abb. 50. Sternit 8 Abb. 51.

Kopulationsapparat Abb. 52 dorsal, Abb. 53 Profil.

Weibchen.

Kopf einfarbig schwarz, an den Fiihlern sind die letzten Glieder manchmal leicht angedunkelt,
sonst wie das 3 gefarbt.

Holotypus 3 and 8 Paratypen (TMP), 5 Paratypen (NHMB), 1 Paratypus (AMG): South Africa, S
Natal: Impetyene grassveld, Weza, 25. xi. 1989, 30°37"S, 29°42'E, E-Y:2717, S. Endrody- Y ounga;
E Transvaal: Nelshoogte Forest Station, grassnetting. Knuckles rock forest, l.xii.l986, 25°47"S,
30°50'E, E-Y:2338, S. Endrody- Younga, 4 Paratypen (TMP).

Eine weitere Art mit hellgesaumten Seiten der Fliigeldecken, die sich von den anderen durch die
Form der Maxillarpalpen deren letztes Glied an der Spitze stark verbreitert und dann stark eingeschniirt
ist, die Form der Fiihler, der Terminalia und des Kopulationsapparates unterscheidet.

Mannchen (Abb. 54-59).

Lange 2,5 mm.

Kopf schwarz, von der Mitte
der Augen nach vorne,
einschliesslich der Wangen,
gelb, iiber jeder Fiihlerwurzel ein
kleiner brauner Flecken;
Maxillarpalpen schwarz, nur
Glied 1 leicht aufgehellt;
Fiihlerglieder 1 und 2 gelb, 3 und
folgende gebraunt, 3 bis 5 oder 6
auf der Unterseite aufgehellt,
folgende mehr oder weniger
gebraunt; Halsschild schwarz,
ungefahr die basale Halfte des
abgeflachten Teils gelbbraun;
Schildchen, Fliigeldecken und
Unterseite schwarz; Beine

Kopf mit den Augen ein wenig breiter als der Halsschild, Stirne leicht gewolbt; Oberflache fein
chagriniert. Maxillarpalpen Abb. 54- 55. Fiihler (Abb. 56) fast so lang wie die Fliigeldecken, Glied
3 so lang wie 4. Halsschild langer als breit (18x17), also verhaltnismassig kurz; Seiten auf dem
vorderen Teii stark gerundet, dann zur Basis ziemlich stark verengt; Oberflache auf dem basalen Teil
etwas starker chagriniert als vorne. Fliigeldecken oval; falsche Epipleuren ca. 70% der Lange

9. Sphinginopalpus (Sphinginafricanus) tergalis sp. nov.

Abb. 54-55. Sphinginopalpus (Sphinginafricanus) tergalis sp. nov. S : 54-55,
Maxillarpalpen (xl50) (54, von oben; 55, von unten).

schwarz, nur die Tarsen gelb, 4 vordere Tibien kaum aufgehellt.

29

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

eiimehmend, kaum merklich krenuliert, sons! wie bei curvipalpis; gefliigelt. Tergit 8 (Abb, 57)
auffallig lang und tief ausgeschnitten und ausserdem mil 2 langen, schmalen, spitz zulaufenden
Fortsatzen versehen. Sternit 8 Abb. 58.

Kopulationsapparat Abb. 59 dorsal.

Holotypus (TMP), 1 Paratypus (NHMB): South Africa, Natal; Weza, Bangeni forest, sifted forest
litter, 30°38'S, 29°39'E, 21. xi. 1989, E-Y:2706, S. Endrody-Younga, I. IClimaszewski.

Die neue Art ist auffallig durch ihren verhaltnismassig kurzen Halsschild, der fast so breit wie
lang ist und das Tergit 8 mit den beiden langen, schmalen Fortsatzen in der Mitte.

Abb. 56-59. Sphinginopalpus (Sphinginafricanus) lergalis sp. nov. 6 : 56, Fiihlerglieder 1-4; 57, Tergit 8; 58, Sternit 8;
59, Kopulationsapparat (dorsal). Massstab von 56 auch fiir 58, 59.

30

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

10. Sphinginopalpus ( Sphinginafrican us) bimaculiceps sp. nov.

MANNCHEN (Abb. 60-68).

Lange 2,5 - 2,7 mm.

Kopf schwarz, nebenjedem Auge ein kleiner gelber Flecken, der oft bis zur Fiihlerwurzel reicht;
Maxillarpalpen vorwiegend schwarz, teils kaum aufgehellt; Fiihler vorwiegend gelb, died 1 auf der
Oberseite schwarz, letzte 1 bis 4 Glieder manchmal mehr oder weniger angedunkelt; Halsschild
schwarz. Basis gelb bis braunlich; Schildchen und Fliigeldecken schwarz, Tibien seltener aufgehellt,
alle Tarsen gelb, nur bei 1 Exemplar sind die hinteren Tarsen angedunkelt.

Abb. 60-61. Sphinginopalpus (Sphinginafricanus) bimaculiceps sp. nov. d : 60-61, Maxillarpalpen
(xl55) (60, von oben; 61, von unten).

Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberflache fein chagriniert
bis fast glatt. Maxillarpalpen (Abb. 60-61). Fiihler (Abb. 62) ungefahr so lang wie die Fliigeldecken,
Glied 1 mit einem spitzwinkligen Zahn vor der Basis, iibrige fast parallel. Halsschild langer als breit
(21x18), sonst wie curvipalpis j edoch etwas weniger tief und weniger dicht punktiert. Ter git 8 Abb.
63. Sternit 8 Abb. 64, Spiculum Abb. 65. Tegmen Abb. 66.

Kopulationsapparat Abb. 67 dorsal. Abb. 68 Profil.

Weibchen.

Kopf einfarbig schwarz, Fiihler gelb, nur die Spitze von Glied 11 kurz dunkel, sonst wie das S
gefarbt. Schulterbeulen nur angedeutet, wahrscheinlich ungefliigelt.

Holotypus (TMP); South Africa, E Transvaal: Graskop, xii.1974, P. E. Reavell; Nelshoogte, Devils
KnucUe, grassnetting, 4.xii.l986, 25°47'S, 30°49'E, E-Y:2350, S. Endrody-Younga, 1 Paratypus
(TMP); Bothasnek, 1450 m, 23. xi. 1987, W. Wittmer, 2 Paratypen (NHMB); Mac-Mac 10 km N of

31

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Sabie, 17. i. 1989, 25°02'S, 30°48'E, E. Grobbelaar, 2 Paratypen (NCI), 2 Paratypen (NHMB); Berlin
Gorge- edge, beating bushes, grassnetting, 25°32"S, 30°44'E, 8.xii.l986, 3.-4.ii.l987, E-Y:2365,
2405, 2413, S. Endrody-Younga, 3 Paratypen (IMP), 1 Paratypus (AMG); Nelspruit Nat. Res. rivulet
vail. 25°29"S, 30°55'E, 5.xii.l986, E-Y:2355, S. Endrody-Younga, 1 Paratypus (TMP).

Die neue Art is neben isidengensis Wittmer zu stellen, von der sie sich durch das verschieden
gebaute Fiihlerglied 1, die Terminalia und den Kopulationsapparat unterscheidet.

Abb. 62-68. Sphinginopalpus (Sphinginafricanus) bimaculiceps sp. nov. 6 : 62, Fuhlergliecler 1-4; 63, Tcrgit 8; 64, Sternit 8;
65, Spiculum; 66, Tegmen; 67-68, Kopulationsapparat (67, dorsal; 68, Profil). Massstab von 63 auch fiir 64-68.

32

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

11. Sphinginopalpus (Sphinginafricanus) isidengensis sp. nov.

MAnnchen (Abb. 69-74).

Lange 2,2- 2,3 mm.

Kopf schwarz, von der Mitte der Augen nach vorne, inklusive der Wangen, gelb; Maxillarpalpen
schwarz; Fiihler gelb, letzte 1 bis 2 Glieder manchmal angedunkelt, 1 mit einem langlichen, schwarzen
Flecken gegen die Spitze auf der Oberseite; Halsschild schwarz, abgeflachter basaler Teil orange;
Schildchen, Fliigeldecken und Unterseite schwarz; Hinterbeine schwarz, Hintertarsen gelb, 4

Vorderschenkel schwarz, an
den Knien mehr oder weniger
aufgehellt, 4 Vordertibien
dunkel, bis zur Halfte oder
dariiber hinaus gelb, 4
Vordertarsen gelb.

Kopf mit den Augen
breiter als der Halsschild,
Stirne leicht gewolbt;
Oberflache chagriniert.

Maxillarpalpen Abb. 69-70.
Fiihler (Abb. 71) nur wenig
kiirzer als die Fliigeldecken,
Glied 3 so lang wie 4.
Halsschild langer als breit
(19x14), sonst wie bei
cunnpalpis. Fliigeldecken
oval, falsche Epipleuren ca.
80% der Lange einnehmend,
schwach krenuliert; sonst wie
bei cun’ipalpis. Tergit 8 Abb.
72. Sternit 8 Abb. 73.

Fiihlerglied 1 ohne schwarzen Flecken, sonst wie das 6 gefarbt. Fiihlerglied 1 lang und schmal,
ohne Zahn. Fliigeldecken mit kleiner, spitzer Schulterbeule, welche nach hinten ein wenig als stumpfer
Kiel verlangert ist (starker ausgepragt als beim 6 ), ungefliigelt.

Holotypus S und 2 Paratypen (TMP): South Africa, Cape Prov.: Amatole, Isidenge Forest Station,
grass tussocks and grassnetting Arum fl., 32°4LS, 27°15'E, 18. xi. 1987, E-Y:2524, S. Endrody-
Younga.

Die neue Art ist neben recurvus Wittmer zu stellen, vergleiche Abbildungen der Terminalia und
des Kopulati ons apparat es .

Abb. 69-70. Sphinginopalpus (Sphinginafricanus) isidengensis sp. nov, 6 : 69-70,
Maxillarpalpen (xl50) (69, von oben; 70, von unten).

Kopulationsapparat Abb. 74 dorsal.
WEIBCHEN.

33

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 71-74. Sphinginopalpus (Sphinginafricanus) isidengensis sp. nov. cj : 71, Fiihlerglieder 1-4; 72, Tergil 8; 73, Sternit 8;
74, Kopulationsapparat (dorsal). Massstab von 72 auch fiir 73, 74.

12. Sphinginopalpus (Sphinginafricanus) martini Pic
Sphinginopalpus martini V\c, 1904; 12.

Mannchen (Abb. 75-83).

Im MP werden 2 Syntypen S $ dieser Art aufbewahrt, die ich als Lectotypus undParalectotypus
bezeichne, Fundort; Natal, Dr. Martin. Fiir die Abbildungen diente der Lectotypus mit Ausnahme
von Abb. 75, fiir die 1 Exemplar von De Rust verwendet wurde. Maxillarpalpen Abb. 75-77. Fiihler
Abb. 78. Tergit 8 Abb. 79. Sternite 7 und 8 Abb. 80- 81. Kopulationsapparat Abb. 82 dorsal.

34

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 75-77. Sphinginopalpus (Sphinginafricanus) martini Pic 6 : 75-77, Maxillarpalpen (xl30)
(75, von oben; 76, von oben in anderem Winkel; 77, von unten).

Abb. 78-83. Sphinginopalpus (Sphinginafricanus) inariini Pic 6 : 78, Fuhlerglicder 1-4; 79, I’crgil 8; 8U, Slcrmt 7; 8i, 8lcrnil
8; 82, Kopulationsapparal (dorsal); 83, Fiihlerglieder 1-4. Mas.sstab von 78 auch lur 80-83. Die Abbildungen 75 und
83 von De Rust Valley, die iibrigen vom Leclotypus.

35

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Untersuchtes MATERIAL: Natal: Weza, Impetyene grassveld, flowering grass, 30°3TS, 29°42"E,
21-25.xi.l989, E-Y:2680, 2712, 2717, S. Endrody-Younga, I. Klimaszewski, 2 Ex. (TMP), 2 Ex.
(NHMB); Karkloof Falls Nature Reserve, 22 km N Pietermaritzburg, 6.x. 1986, W. Wittmer (NHMB);
St Lucia, 10 m, 25-27.X.1981, J. & S. Klapperich, 54 Ex. (NHMB), bei weiteren 12 Ex. mit Fundort:
Cape Prov., De Rust (Valley), 29. xi. 1981, J. & S. Klapperich, muss angenommen werden, dass
derselbe umichtig ist und die Tiere ebenfalls aus St Lucia stammen.

Die Form von Fiihlerglied 1 weicht bei dem ganzen Material von dem des Lectotypus (Abb. 83)
ab, da jedoch der Kopulationsapparat und die Terminalia keine Unterschiede zeigen, stelle ich das
Material zu martini.

13. Sphinginopalpus (Sphinginafricanus) consimilis sp. nov.

Mannchen (Abb. 84-85).

Lange 2,3 mm.

Diese Art ist ausserlich und auch aufgrund der Terminalia nicht von martini zu unterscheiden.
Der Kopulationsapparat (Abb. 85) ist jedoch sehr verschieden gebaut undbesteht somit kein Zweifel,
dass consimilis einer anderen Art angehort. Fiihler (Abb. 84) iihnlich wie bei martini.

Holotypus (NHMB): Natal: Nkandla Forest, 6.i.l982, P. E. Reavell.

Diese neue Art ist neben martini Pic zu stellen.

Abb. 84-85. Sphinginopalpus (Sphinginafricanus) consimilis sp. nov. <S: 84, Ftihlerglicder 1-4; 85, Kopulationsapparat
(dorsal).

36

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

14. Sphinginopalpus (Sphinginafricanus) formicarius (Gorham)

Chalicorus formicarius Gorham, 1901: 358. Sphinginopalpus formicarius Gorham - Champion,
1922: 330, T. 5, f. 22.

MAnnchen (Abb. 86-93).

Abb. 86-87. Sphinginopalpus (Sphinginafricanus) formicarius (Gorham) 6 : 86-87, Maxillarpalpen
(xl85) (86, zeigt links died 3 von unten, daneben died 2 von innen; 87, von unten).

Im BM wird der Holotypus, neu als Lectotypus bezeichnet, aufbewahrt, der von Malvern, Natal,
stammt. Vom gleichen Fundort und gleichen Datum (vi.l897) befindet sich auch 1 c? im BM, dessen
Maxillarpalpen von Champion abgebildet werden. Zwei weitere $ von Malvern sind im MP als
Typen bezeichnet und jetzt von mir mit Paralectotypen-Etiketten versehen worden.

Untersuchtes MATERIAL: Natal: Durban. F. Muir, 2 6 (BM); St Lucia, 10 m, 25.-27.X.1981, J. &
S. Klapperich, 22 Ex. (NHMB); Eshowe, Glinza Eorest, 19.xii.l985, P. E. Reavell, 12 Ex. (NHMB);
Transkei: Dwesa coast, coastal grassland, 32°17'S, 28°51'E, 16.xii.l979, 28. ii. 1985, E-Y:1687, 2175,
S. Endrody-Younga, 6 Ex. (TMP), 3 Ex. (NHMB).

MAnnchen.

Lange 2 - 2,1 mm.

Kopf schwarz, von der Mitte der Augen nach vorne, einschliesslich der Wangen, gelb;
Maxillarpalpen schwarz; Fiihler schwarz, Glieder 1 und 2 gelb, 1 mit einem Langsflecken auf der
Oberseite gegen die Spitze, 2 meistens mehr oder weniger angedunkelt auf der Oberseite; Halsschild
schwarz, abgeflachter Teil an der Basis gelblich; Schildchen, Fliigeldecken und Beine schwarz, mit
Ausnahme der Basis der Schenkel und der Spitzen der Tibien bei den Mittel- und Vorderbeinen, die
gelblich sind, alle Tarsen gelb, die hinteren oft stark angedunkelt.

37

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberflache feinstens
chagriniert, fast glatt, Maxillarpalpen Abb. 86-87. Fiihler (Abb. 88) ungefahr so lang wie die
Fliigeldecken, Spitze von died 1 ein wenig iiber die Basis von 2 reichend, 3 so lang wie 4. Halsschild
langer als breit (18x13), vor der Mitte am breitesten, dann nach hinten zuerst leicht gerundet, dann
fast gerade verschmiilert; Seitenrand vor der Basis erloschend, Basalrand nur angedeutet; Scheibe
auf den vorderen 2/3 stark aufgewolbt, basaler Teil abgeflacht; ganze Oberflache chagriniert.
Fliigeldecken oval, ungefahr in der Mitte am breitesten; Schulterbeulen deutlich vorstehend; Punkte
ziemlich tief in Reihen angeordnet, erst kurz vor der Spitze erloschend; falsche Epipleuren ca. 70%
der Lange eiimehmend, auf dem vorderen Teil krenuliert; ziemlich zahlreiche, lange, aufstehende
Haare vorhanden; gefliigelt. Hinterschienen gegen die Spitze allmahlich schwach verbreitert. Tergit
8 Abb. 89. Sternit 8 Abb. 90. Spiculum Abb. 91.

Kopulationsapparat Abb. 92 dorsal, Abb. 93 Profil.

Weibchen.

Wie das 6 gefarbt, mit Ausnahme des schwarzen Kopfes und der Fiihler, die heller sind, dunkle

02

Abb. 88-93. Splii/igiiiopalpus (Sphinginafricunus) formicarius (Gorham) 6 : 88, Fiihlergliedcr 1-4; 89, Tcrgit 8; 90, Sternit
8; 91, Spiculum; 92-93, Kopulationsapparat (92, dorsal; 93, Profil). Massstab von 89 auch fiir 90-93.

38

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA; MALACHIIDAE)

Farbung auf Glied 1 fehlt oft, oder ist hochstens ungedeutet, 2 und 3 oder auch weitere Glieder hell.
Fiihlerglied 1 sehr lang und schmal, ohne Zahn. Fliigeldecken mit kleiner spitzer Schulterbeule;
ungefliigelt.

15. Sphinginopalpus (Sphinginafricanus) recurvus sp. nov.

MAnnchen (Abb. I, K, 94-100).

Lange 2,5 mm.

Kopf schwarz, Vorderkopf ungefahi von der Mitte der Augen an gelb;
Maxillarpalpen hellbraun, wenige Stellen angedunkelt; Fiihler schwarz,
Glieder 1 und 2 geib, auf der Oberseite angedeutet dunkel, oder nur 1 auf
der Oberseite gegen die Spitze mit einem kurzen schwarzen Wisch, 3 bis
5 auf der Unterseite aufgehellt; Halsschild schwarz. Basis gelbbraun;
Schildchen und Fliigeldecken schwarz; Schenkel schwarz, die 4 vorderen
in variablem Umfang gelb, Hinterschienen schwarz, die 4 vorderen z. T.
fast vollstandig hell, alle Tarsen gelb.

Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewolbt;
Oberflache fein chagriniert. Maxillarpalpen Abb. 94. Fiihler (Abb. 95)
ungefahr so lang wie die Fliigeldecken, Zahn auf Glied 1 wenig lang, einen
schwachen spitzen Winkel bildend. Halsschild langer als breit (21x17),
sonst wie curvipalpis, Oberflache fein chagriniert, ohne glatte Stellen,
feinste Rippen nur angedeutet. Fliigeldecken wie curvipalpis, falsche
Epipleuren deutlicher krenuliert. Tergit 8 (Abb. 96) wenig tief, breit
ausgerandet. Sternit 8 (Abb. 98) jederseits mit einem kraftigen, nach innen
gekriimmten Haken, Sternit 7 (Abb. 97) mit leicht gewelltem Apikalrand.
Gesamtansicht Abb. I, K.

Kopulationsapparat Abb. 99 dorsal. Abb. 100 Profil.

Weibchen (Abb. A-C, G).

Kopf einfarbig schwarz, sonst wie das S gefarbt. Ubrige Beschreibung
mit repandus iibereinstimmend. Gesamtansicht Abb. A, B. Maxillarpalpen Abb. C. Basis des
Halsschildes Abb. G.

Holotypus und 42 Paratypen (TMP), 17 Paratypen (NHMB), 4 Paratypen (AMG): South Africa, SW
Natal: Weza, indigenous forest, 30°35"S, 29°45"E, grassnetting, 24.xii.1985, E-Y :2160, S. Endrody-
Y ounga.

Die Maxillarpalpen erinnern etwas an emarginatus Wittmer, neben welchen die neue Art zu
stellen ist, doch ist das Fiihlerglied 1, die Terminalia und der Kopulationsapparat verschieden gebaut.

Abb. 94. Sphinginopalpus
(Sphinginafricanus)
recurvus sp. nov. S :
94,

Maxillarpalpus
(xl30) (von oben).

39

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb, 95-lUO. Sphinginopalpus (Sphinginafricanus) recurvus sp. nov, 6 : 95, Fuhlerglicder 1-4; 96, Tergil 8; 97, Sternit 7; 98,
Sternit 8; 99-100, Kopulationsapparat (99, dorsal; 100, Profil). Massstab von 96 auch fiir 97 und 98 auch fiir 99, 100.

16. Sphinginopalpus (Sphinginafricanus) recurvus planicornis ssp. nov.

Abb, 101-102. Sphinginopalpus (Sphinginafricanus) recurvus
planicornis ssp. nov. 6: 101-102, Maxillarpalpen(xl30)(101,
von oben; 102, von unten).

MANNCHEN (Abb. 101-103).

Lange 2,5 mm.

Diese Rasse ist weit verbreitet und
unterscheidet sich von der Nominatform
hauptsachlich durch den Bau der Fiihler
(Abb. 103), died 1 ist weniger schlank,
nach dem etwas kraftigeren Zahn
schwach bauchig verdickt, 3 bis 5 sind
deutlich abgeflacht und ein wenig breiter
als die folgenden, auch 6 zeigt noch eine
leichte Verbreiterung. Die Farbung der
Fiihler ist durchwegs dunkler, bereits ab
died 3 schwarz, 2 oft mit einem kleinen
verschwommenen Flecken auf der
Oberseite, ebenso 1, oder 1 und 2 sind
einfarbig gelb. Die Maxillarpalpen
(Abb. 101-102) haben grosse

40

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Ahnlichkeit mit der Nominatform, vergleiche Abbildungen, sie sind jedoch etwas langer und breiter.
Die Terminalia weichen im Bau nur geringfiigig ab. Ich mochte nicht entscheiden, ob es sich um eine
Rasse, oder um eine eigene Art handelt, letzteres scheint mit eher wahrscheinlich.

Holotypus und 10 Paratypen (TMP), 6 Paratypen (NHMB), 1 Paratypus (AMG): South Africa, Natal:
Karkloof forest and grassveld, 29°WS, 30°15'E, 2.-7.xii.l989, E-Y:2729, 2737, 2748, 2749, 2758,
S. Endrody-Younga, I. Klimaszewski; Doreen Clark Nature Reserve, sifted forest litter, 29°34"S,
30°17"E, 8.xii.l989, E-Y:2750, S. Endrody- Younga, I. Klimaszewski, 1 3 Paratypus (TMP);
Pietermaritzburg, Town Bush Forest, 29°36'S, 30°23'E, 27.ii.1984, C. G. E. Moolman, 16 Paratypen
(NCI), 4 Paratypen (NHMB), 2 Paratypen (AMG); Karkloof Falls near Howick, 29°25"S, 30°17"E,
26. xi. 1984, R. Oberprieler, C. G. E. Moolman, 6 Paratypen (NCI).

17. Sphinginopalpus (Sphinginafricanus) repandus sp. nov.

Mannchen (Abb. 104-109).

Abb. 104-105. Sphinginopalpus (Sphinginafricanus) repandus sp. nov. 6 : 104-105,
Maxillarpalpen (xl30) (104, von oben; 105, von unten).

Lange 2,5 mm.

Kopf schwarz,

Vorderkopf ungefahr von
der Mitte der Augen an, gelb;
Maxillarpalpen schwarz,
teils gelb, seltener zum
grossten Teil gelb; Fiihler
gelb, manchmal sind die
letzten Glieder schwach
angedunkelt; Halsschild
schwarz. Basis ziemlich
breit gelb bis orange;
Schildchen und

Fliigeldecken schwarz; 4
Vorderbeine gelb,

Hintertibien schwarz,
Hinterschenkel oft teils gelb
(Basis), Hintertarsen gelb.

Kopf mit den Augen
breiter als der Halsschild,
Stirne leicht gewolbt;

41

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Oberflache glatt. Maxillarpalpen Abb. 104-105. Fiihler (Abb. 106) nur wenig langer als die
Fliigeldecken, fast parallel. Halsschild langer als breit (19x15), vorderer Teil stark aufgewolbt, Basis
breit abgeflacht; Seiten fast bis zur Mitte leicht gerundet, dann schwach verengt; Rand an der Basis
breiter als die Seiten, kurz vor der Basis erloschen; Oberflache fein chagriniert, auf der Mitte der
Scheibe fast glatt, gegen die Seiten oft mit 2 feinen Rippen. Fliigeldecken langlichoval, Schultern
normal entwickelt; falsche Epipleuren lang, scharf, feinstens krenuliert; Oberflache glatt, Punkte
ziemlich grob, besonders an der Basis teils in Reihen angeordnet, Punkte gegen die Spitze fast ganz
erloschen; einzelne lange, abstehende Haare vorhanden; gefliigelt. Tergit 8 (Abb. 107) apikal schrag
abgestutzt, an der Spitze kurz ausgerandet. St emit 8 Abb. 108.

Kopulationsapparat (Abb. 109) dorsal, neben dem Innensack sind 3 kleine, stark sklerotisierte
Hornchen sichtbar.

Abb. 106-109. Sphinginopalpus (Sphinginafricanus) repandus sp. nov. 3 : 106, Fiihlerglieder 1-4; 107, Tergit 8; 108, Stemit
8; 109, Kopulationsapparat (dorsal). Massstab von 106 auch fiir 108 und 107 auch fiir 109.

Weibchen.

Kopf einfarbig schwarz, sonst wie das S gefarbt. Letztes died der Maxillarpalpen zur Spitze
schwach verbreitert. Fiihlerglied 1 einfach, schmal und lang, fast 4 mal langer als 2. Fliigeldecken
breiter als beim c?, Schulterbeulen ein wenig reduziert; ungefliigelt.

Holotypus und 1 Paratypus S (NHMB): South Africa, Transvaal: Bothasnek, 1450 m, 23. xi. 1987,
W. Wittmer. Natal : Cathedral Peak, Drakensberg, 6.xi. 1981, J. & S. Klapperich, 4 Paratypen (NHMB);
idem Mike’s Pass, 1700 m, 28°48'S, 29°14'E, 24.ii.1984, C. G. E. Moolman, 5 Paratypen (NCI); 2
Paratypen (NHMB); idem area above Mike’s Pass, 1973 m, 28°59'S 29°14'E, 19. -23. ii. 1973, B.
Grobbelaar, 2 Paratypen (NCI); idem Arensig Mt., 20.xii.l986, P. Reavell, 1 Paratypus (NHMB);
idem Mt. Nhlosane near Dargle, 1600 m, 29°33'S, 29°58'E, C. G. E. Moolman, 10 Paratypen (NCI),
2 Paratypen (AMG); Nelshoogte Forest Station, Knuckles grassveld, l.-4.xii.l986, 25°47'S,

42

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

30°49/50'E, E-Y:2339, 2350, S. Endrody-Younga, 2 Paratypen (TMP), 1 Paratypus (NHMB).

Diese neue Art ist nahe mit ohtusodentatus Wittmer verwandt. Sie unterscheidet sich durch die
verschieden gebauten Maxillarpalpen, Fiihler und Terminalia.

18. Sphinginopalpiis (Sphinginafricanus) emarginatus sp. nov.

Mannchen (Abb. 110-115).

Lange 2, 2-2, 4 mm.

Kopf schwarz,
Vorderkopf von
ungefahr der Mitte
der Augen,

inklusive der
Wangen, gelblich;
Maxillarpalpen
schwarzbraun, teils
aufgehellt; Fiihler
dunkel, Glieder 1
und 2 gelb, 3 und 5
aufgehellt, oben
s c h w a c h
angedunkelt, 1 mit
einer kleinen
dunkeln Stelle an
der Spitze auf der
O b e r s e i t e ;
H a 1 s s c h i 1 d

schwarz. Basis fast auf der ganzen abgeflachten Stelle gelb; Schildchen und Fliigeldecken schwarz;
Beine schwarzlich, Vorderschenkel innen aufgehellt, die iibrigen an der Basis kurz gelb, 4
Vordertarsen gelb.

Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberflache chagriniert.
Maxillarpalpen Abb. 110-111. Fiihler (Abb. 112), das Glied 3 so lang wie 4, auf der Zeichnung ist 4
etwas zu lang. Halsschild langer als breit (22x15), vorderer Teil stark aufgewolbt. Basis breit
abgeflacht; Seiten bis zur Mitte gerundet, dann verengt; Oberflache auf dem abgeflachten Teil starker
chagriniert als vorne. Fliigeldecken langlichoval, Schultern normal entwickelt; falsche Epipleuren
lang, scharf, fein krenuliert; Oberflache glatt, Punkte ziemlich grob, einzelne Punktreihen erkennbar,
Punktierung nach hinten schwacher; gefliigelt. Tergit 8 (Abb. 113) mit breit aber wenig tief
ausgerandeter Spitze. Sternit 8 Abb. 114.

Kopulationsapparat Abb. 115 dorsal.

Weibchen.

Kopf einfarbig schwarz, Fiihlerglieder 1 bis 6 einfarbig gelb, sonst wie das 6 gefarbt.

Abb. 110-111. Sphinginopalpus (Sphinginafricanus) emarginatus sp. nov. 8: 110-111,
Maxillarpalpen (xl50) (110, von oben; 111, von unten).

43

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Fliigeldecken breiter als beim S , Schulterbeulen angedeutet.

Holotypus and 5 Paratypen (TMP), 2 Paratypen (NHMB): South Africa, Transvaal: Uitsoek Forest
Station, beating, 12.ii. und 7.viii.l986, 1100-1200 m, E-Y:2430, 2380, S. Endrody-Younga;
Nelshoogte, riverine forest litter, 1000 m, 2.xii.l986, E-Y:2341, S. Endrody-Younga, 17 Paratypen
(TMP), 7 Paratypen (NHMB), 2 Paratypen (AMG).

Die neue Art ist neben tubulatus Wittmer zu stellen und aufgrund der Unterschiede gemass der
Abbildungen leicht zu unterscheiden.

Abb. 112-115. Sphinginopalpus (Sphinginafricanus) emarg'matus sp. nov. <J ; 112, Fiihlerglieder 1-4; 113, Tergit 8;
114, Stemit 8; 115, Kopulationsapparat (dorsal). Massstab von 112 auch fiir 114-115.

44

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

19. Sphinginopalpus (Sphinginafricanus) reductosetosus sp. nov.

Mannchen (Abb. 116-123).

Lange 2,5 mm.

Abb. 116-118. Sphinginopalpus (Sphinginafricanus) reductosetosus sp. nov. cj : 116-1 18, Maxillarpalpen (1 16,
von oben, xl30; 117, Ausschnitt aus der Oberseite, x550; 118, von unten, xl30).

Kopf schwarz, von der Mitte nach vorne, inklusive der Wangen, gelb; Fiihler schwarz, Glied 1
gelb, an der Spitze auf der Oberseite ein kleiner schwarzer Flecken, 2 bis 4 ganz und Basis von 5
kurz gelb; Halsschild schwarz, Basis ziemlich breit gelb; Schildchen und Fliigeldecken schwarz; alle
Tarsen gelb, Hinterbeine schwarz, Mittel- und Vorderbeine gelb bis braun, teils leicht angedunkelt.

Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberflache feinstens
chagriniert, teils fast glatt. Maxillarpalpen Abb. 116-118. Fiihler (Abb. 119) so lang wie die
Fliigeldecken, Glieder 3 bis 10 am Innenrand schwach gerundet, 5 ein wenig verbreitert. Halsschild
1 anger als breit (18x14,5), sonst wie curvipalpis; Chagrinierung sehi fein, auf der Scheibe fast glatt.
Fliigeldecken langlichoval, Schulterbeulen reduziert, leicht spitz vorstehend, sonst wie bei
curvipalpis, jedoch ungefliigelt und falsche Epipleuren deutlicher. Tergit 8 Abb. 120. Sternit 8 Abb.
121. Spiculum Abb. 122.

Kopulationsapparat Abb. 123 dorsal.

Holotypus und 33 Paratypen (TMP), 18 Paratypen (NHMB), 4 Paratypen (AMG): South Africa, E
Transvaal: Berlin, 24°32'S, 30°44"E, gorge edge, beating bushes, 3.ii.l987, E-Y:2405, S. Endrody-
Younga; Nelshoogte, 1000 m, riverine forest litter, 2.xii.l986, E-Y:2341, S. Endrddy- Y ounga, 1
Paratypus (NHMB).

45

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 119-123. Spliinginopalpus (Sphingmajricanus) reduaosetosiis sp. nov. d: 119, Fuhlergliedcr 1-5; 120, Tergit 8; 121,
Sternit 8; 122, Spiculum; 123, Kopulationsapparat (dorsal). Massstab von 119 auch fiir 121-123.

Der Widerhaken an died 2 der Maxillarpalpen ist fiir diese Art sehr charakteristisch. Aufgrund
des verbreiterten Fuhlergliedes 5 gehdrt sie eigentlich zu bilineaticeps, reavelli und recurvus
planicornis, siehe Fussnote auf S. 9 der Bestimmungstabelle.

20. Sphinginopalpus (Sphinginafricanus) tubulatus sp. nov.

Mannchen (Abb. 124-129).

Lange 2,5 mm.

Kopf schwarz mil einer kleinen gelben, dreieckigen Makel auf dem Vorderkopf, Spitze gegen
die Stirne gerichtet; Maxillarpalpen schwarz; Fiihler schwarzlich, Glieder 2 und 3 gelb, 4 bis 6 leicht

46

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 124-125. Sphinginopalpus
(Sphinginafricanus) tubulatus sp.
nov. 6: 124-125, Maxillarpalpen
(xl30) (124, von oben; 125, von
unten).

Abb. 126-129. Sphinginopalpus
(Sphinginafricanus)
tubulatus sp. nov. 6:

126, Fiihlerglieder 1-4;

127, Tergit 8; 128,

Sternit 8; 129,

Kopulationsapparat
(dorsal). Massstab von
126 auch fiir 128, 129.

47

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

angedunkelt; Halsschild schwarz, Basis ein wenig aufgehellt; Schildchen, Fliigeldecken und Beine
schwarz, alle Tarsen gelb.

Kopf mil den Augen breiter als der Halsschild, Stirne leicht gewdlbt; Oberflache fein chagriniert,
teils fast glatt. Maxillarpalpen Abb. 124-125. Fiihler (Abb. 126) ungefahr so lang wie die
Fliigeldecken, Glied eins 2 1/2 mal so lang wie 2, 3 bis 10 zur Spitze nur ganz schwach verbreitert,
fast parallel. Halsschild ungefahr in der Mitte am breitesten; abgeflachter Teil etwas deutlicher
chagriniert als der vordere, sonst wie bei curvipalpis. Fliigeldecken wie bei curvipalpis, falsche
Epipleuren ohne Krenulierung. Tergit 8 Abb. 127. Sternit 8 Abb. 128.

Kopulationsapparat Abb. 129 dorsal.

Holotypus (TMP): South Africa, Natal Middld.: Karkloof, intersept, trap 10 days, 1300 m, 29°18"S,
30°13'E, 2.xii.l989, E-Y:2729, S. Endrody-Younga, 1. Klimaszewski.

Die stark ausgeschnittenen Seiten von Glied 3 der Maxillarpalpen erinnern an curvipalpis, doch
sind die Terminalia und der Kopulationsapparat mit den parallen Innenteilen sehr verschieden.

21. Sphinginopalpiis (Sphinginafricanus) natalensis sp. nov.

Mannchen (Abb. 130-136).

Lange 2,7 mm.

Kopf schwarz, von der Mitte der Augen nach vorne gelb; Maxillarpalpen schwarz; Fiihler
schwarz, Glied 1 gelb mit einem schwarzen Langswisch auf der Oberseite, seltener ganz gelb, 2 bis
5 weitgehend gelb; Halsschild schwarz, Basis nicht sehr breit gelbbraun; Schildchen, Fliigeldecken
und Beine schwarz, nur die Tarsen gelb.

Kopf mit den Augen breiter
als der Halsschild, Stirne leicht
gewolbt; Oberflache fein
chagriniert. Maxillarpalpen Abb.
130-131. Fiihler (Abb. 132)
ungefahr so lang wie die
Fliigeldecken, Glied 1 sechs mal
so lang wie 2, 3 bis 10 am
Innenrand ganz schwach
gerundet. Halsschild langer als
breit (21x17), sonst wie bei
curvipalpis, Fliigeldecken wie
bei curvipalpis, jedoch falsche
Epipleuren deutlich krenuliert.
Tergit 8 Abb. 133. Sternit 8 Abb.
134.

Kopulationsapparat Abb.
135 dorsal. Abb. 136 Profil.

Abb. 130-131. Sphinginopalpus (Sphinginafricanus) natalensis sp. nov. 6 ; 130-
131, Maxillarpaipen (xl30) (130, von oben; 131, von unten).

48

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 132-136. Sphinginopalpus (Sphinginafricanus) natalensis sp. nov. 6 : 132, Fuhlerglieder 1-4; 133, Tergit 8; 134, Stemit
8; 135-136, Kopulationsapparat (135, dorsal; 136, Profil). Massstab von 132 auch fiir 134 und 133 auch fiir 135, 136.

Weibchen.

Kopf ganz schwarz; Fiihler dunkei, erste 4 bis 5 Glieder gelb, oder 1 auf der Oberseite schmal
angedunkelt, sons! wie das d gefarbt. Fliigddecken wie bei repandus; ungefliigelt.

Holotypus und 7 Paratypen (TMP), 5 Paratypen (NHMB): South Africa, Natal Middld.: Karkloof
grassveld with river, 29°19'S, 30°15'E, 7.- 9.xii.l989, E-Y:2737, 2747, 2749, 2753, 2758, S. Endrody-
Younga, I. Klimaszewski; Doreen Clark Nature Reserve, beating in forest, 29°34"S, 30°17'E, 8. und
ll.xii.l989, E-Y;2750, 2760, S. Endrddy- Younga, I. Klimaszewski, 3 Paratypen (TMP), 2 Paratypen
(NHMB).

Neben recurvus Wittmer zu steilen.

49

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

22. Sphinginopalpus (Sphinginafricanus) obtusodentatus sp. nov.
MAnnchen (Abb. 137-143).

Lange ca. 2,3 mm.

Kopf schwarz, von der Mitte der
Augen nach vorne gelb; Maxillarpalpen
dunkelbraun; Fiihler bis zu died 10(11
fehlt) gelb; Halsschild dunkelbraun,
Basis ziemlich breit gelb; Schildchen,
Fliigeldecken und Beine braun, Tarsen
gelb.

Kopf mil den Augen breiter als der
Halsschild, Stirne leicht gewolbt;
Oberflache fein chagriniert.
Maxillarpalpen Abb. 137-138. Fiihler
(Abb. 139) ungefahr so lang wie die
Fliigeldecken, Zahn auf died 1 breit
und stumpf, 3 bis 10 am Innenrand ganz
schwach gerundet. Halsschild langer
als breit (21x16), sonst wie bei
curvipalpis; Oberseite fein chagriniert.
Fliigeldecken wie bei curvipalpis.
Tergit 8 (Abb. 140), apikaler Rand umgeschlagen (auf der Abbildung durch eine punktierte Linie
bezeichnet). Sternit 7 Abb. 141; Sternit 8 (Abb. 142) nur mit einer langen Borste versehen.
Kopulationsapparat Abb. 143 dorsal.

Abb. 137-138, Sphinginopalpus (Sphinginafricanus) obtusodentatus sp.

nov. 6 : 137-138, Maxillarpalpen (xl30) (137, von oben; 138,
von unten).

Holotypus (BM): South Africa, Natal: Malvern, ix.l897, G. A. K. Marshall, von Champion als
Sphinginopalpus collaris (Boheman) bestimmt.

Eine neue Art die neben tubulatus Wittmer gestellt werden karm, Fiihlerglied 1 ahnlich gebaut,
jedoch mit verschieden gebauten Maxillarpalpen, Terminalia und Kopulationsapparat.

50

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA; MALACHIIDAE)

Abb. 139-143. Sphinginopalpus (Sphinginafricanus) obtusodentatus sp. nov. cj : 139, Fiihlerglieder 1-4; 140, Tergit 8; 141,
Stemit 7; 142, Stemit 8; 143, Kopulationsapparat (dorsal).

23. Sphinginopalpus (Sphinginafricanus) unisetosus sp. nov.

MAnnchen (Abb. 144-149).

Lange 2,5 mm.

Kopf und Maxillarpalpen schwarz; Fiihler gelb, ab Glied 7 verstarkt angedunkelt, 1 mil
schwarzem Langswisch auf der Oberseite; Halsschild schwarz, Basis rotbraun; Schildchen und
Fliigeldecken schwarz; alle Tarsen gelb, Hinterbeine schwarz, Vorder- und Mittelbeine schwarz, Knie
wenig, Tibien an der Spitze breiter gelb.

Kopf mil den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberflache glatt.
Maxillarpalpen Abb. 144-145. Fiihler (Abb. 146) um ca. 1/10 kiirzer als die Fliigeldecken, Glieder
3 bis 10 zur Spitze kaum merklich verbreitert. Halsschild Finger als breit (21x16), ungefahr in der
Mitte am breitesten, sonst wie bei cunipalpis, jedoch die Oberflache des aufgewolbten Teiles ist

51

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 144-145. Sphinginopalpus (Sphinginafricanus) unisetosus sp. nov. 6: 144-145, Maxillarpalpen
(xl30) (144, von oben, Glieder 1 und 2 aufgeklappi; 14^ von unten).

Abb. 146-149. Sphinginopalpus (Sphinginafricanus) unisetosus sp. nov. 6 ; 146, Fiihlerglieder 1-4; 147, Tergit 8; 148, Sternit
8; 149, Kopulationsapparat (dorsal). Massstab von 146 auch fiir 148 und 147 auch fiir 149.

glatt, die abgeflachte Basis feinstens chagriniert. Fliigeldecken wie bei curvipalpis, aber etwas
schwacher punktiert. Tergit 8 Abb. 147. Sternit 8 Abb. 148.

Kopulationsapparat Abb. 149 Profil.

Holotypus (NCI): South Africa, Natal: Cathedral Peak area, above Mike’s Pass, 1973 m, 28°59'S,
29°14'E, 19.-23.i.l986, B. Grobbelaar.

52

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA; MALACHIIDAE)

Eine der wenigen Arten bei denen die beiden Teilstiicke von Sternit 8 durch feines Gewebe lose
verbunden sind, sie kartn deshalb neben natalensis Wittmer gestellt werden. Sons! von dieser Ait
sehi verschieden, vergleiche mit Abb. 130-136.

24. Sphinginopalpiis s. str. pallidipalpis sp. nov.

Mannchen (Abb. 150-156).

Lange gut 2 mm.

Kopf gelb mit einem dunkeln Flecken an
der Kopfbasis, der seitlich bis ungefahr zur
Mitte der Schlafen reicht und nach vorne bis
zur Mitte der Augen; Maxillarpalpen gelb;
Fiihler gelb, die letzten 4 bis 5 Glieder
angedunkelt; Halsschild dunkelbraun,
abgeflachter Teil gelbbraun; Fliigeldecken
dunkelbraun mit einem gelblichen Querband
vor der Mitte, das an den Seiten nach hinten
verlangert ist; Hinterbeine schwarzlich, alle 6
Tarsen gelb, 4 vordere Schenkel grosstenteils
hell, 4 vordere Tibien gelb.

Kopf mit den Augen kaum merklich
breiter als der Halsschild, Stirne leicht
gewolbt, Oberflache fein chagriniert.
Maxillarpalpen Abb. 150-151. Fiihler (Abb.
152) um ca. 1/10 kiirzer als die Fliigeldecken,
Glied 1 ohne Zahn, gleich nach der Basis stark
verbreitert und dann zur Spitze verschmalert,
3 bis 10 zur Spitze nur ganz wenig verbreitert, 3 so lang wie 4. Halsschild langer als breit (19,5x17,5),
sonst wie capensis. Fliigeldecken langoval, falsche Epipleuren starker herausstehend, ca. 60% der
Lange einnehmend, an der breitesten Stelle ganz schwach krenuliert; Punkte etwas grober als bei
capensis, sonst wie diese gebaut. Tergit 8 Abb. 153. Sternit 7 Abb. 154, Sternit 8 Abb. 155.
Kopulationsapparat Abb. 156 Profit.

Abb. 150-151. Sphinginopalpiis s. str. pallidipalpis sp. nov. 3 :
150-151, Maxillarpalpen (xl70) (150, von oben; 151,
von unten).

Holotypus (BM): South Africa, Cape Prov.: Dunbrody, 33°28'S, 25°33'E, under seeds, 6.vi.l903,
O’Neil.

Diese neue Art wurde von Champion, 1922, als oneili Pic bestimmt. Sie gehort zu den wenigen
Arten mit durchgehendem, weisslichem Querband auf den Fliigeldecken und gleicht ausserlich etwas
capensis Wittmer, doch sind bei letzterer die Fliigel, der Halsschild und die Beine dunkler, nebst sehr
verschieden gebautem Fiihlerglied 1 und Terminalia.

53

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 152-156. Sphinginopalpus s. sli. palliilipalpii .sp. nov. cj : 152, Fiihlerglicdcr 1-4; 153, Tergil 8; 154, Stcrnil 7; 155,
Sternit 8; 156, Kopulationsapparat (Profil). Massstab von 152 auch fiir 155, 156 und 153 auch fiir 154.

54

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

25. Sphinginopalpus s. sir . flavomarginatus Champion
Sphinginopalpus flavomarginatus Champion, 1922: 334.
MAnnchen (Abb. 157-163).

Abb. 157-158. Sphinginopalpus s. str. flavomarginatus Champion 6 ■ 157-158,
Maxillarpalpen (x220) (157, von oben; 158, von unten).

Am Originalfundort: Frere, Natal, fing ich am 8.xi.l992, ein weiteres 6 . Fiir die Abbildungen
wurde der Holotypus im BM verwendet. Maxillarpalpen Abb. 157-158. Fiihlerglieder 1-4 Abb. 159.
Tergit 8 Abb. 160. Sternit 8 Abb. 161. Kopulationsapparat Abb. 162 dorsal,Abb. 163 Profil.

Dem ganzen Aussehen nach konnte diese Art leicht in die Untergattung Sphinginafricanus
gehdren.

55

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

0.2

Abb. 159-163. Sphinginopalpus s. sir. flavomarginatus Champion S: 159, Fiihlerglieder 1-4; 160, Tergit 8; 161, Sternit 8;
162-163, Kopulationsapparat (162, dorsal; 163, Profil). Massstab von 159 auch fur 161 und 160 auch fiir 162, 163.

56

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA; MALACHIIDAE)

26. Sphinginopalpus s. str. triangulatus sp. nov.
MANNCHEN (Abb. 164-169).

Lange 1,8 mm.

Kopf schwarz, Vorderkopf mil
einem dreieckigen gelben Flecken
zwischen den Fiihlerwurzeln;
Maxillarpalpen gelb, vorletztes died
teilweise braunlich; Fiihler gelb, letzte 5
bis 6 Glieder schwarz; Halsschild
schwarz, abgeflachter Teil gelbbraun;
Schildchen und Fliigeldecken schwarz,
letztere mil einem weissen Saum
jederseits, der unter den Schultern
beginnt und fast bis zur Spitze reicht;
Hinterschenkel dunkel, auf der
Unterseite aufgehellt, die 4 vorderen
braun, Tibien und Tarsen gelblich.

Kopf mit den Augen so breit wie der
Halsschild, Stirne leicht gewolbt;
Obertlache glatt. Maxillarpalpen Abb.
164-165. Fiihler (Abb. 166) fast urn 1/3
kiirzer als die Fliigeldecken, Glied 1 einfach, 3 deutlich langer als 2, 4 ein wenig breiter als 3. Halsschild
knapp langer als breit (14x13), Seiten nach vorne fast in gerader Linie schwach verbreitert;
Seitenrander besonders an der Stelle der Einschniirung stark entwickelt, erhoht, fein krenuliert;
Oberflache glatt. Fliigeldecken langoval, wie bei elongatidens; Punktreihen bis weit nach hinten
sichtbar. Tergit 8 Abb. 167. Sternit 8 Abb. 168.

Kopulationsapparat Abb. 169 dorsal.

Abb. 164-165. Sphinginopalpus s. str. triangulatus sp. nov. d
165, Maxillarpaipen (x250).

164-

Holotypus and 1 Paratypus S (TMP), 1 Paratypus S (NHMB): South Africa, Natal; Weza, Impetyene
grassveld, flowering grassveld, 30°37'S, 29°42'E, 21. und 25. xi. 1989, E-Y:2709, 2718, S. Endrody-
Younga, I. Klimaszewski.

Eine der wenigen Arten mit einfachem, nicht gezahntem Fiihlerglied 1, neben flavomarginatus
Champion zu stellen.

57

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb, 166-169. Sphinginopalpus s. str. triangulaius sp. nov. 3 : 166, Fiihlerglieder 1-4; 167, Tergit 8; 168, Stemit 8; 169,
Kopulationsapparal (dorsal). Massstab von 166 auch fur 167-169.

27. Sphinginopalpus s.str. orientalis sp. nov.

MAnnchen (Abb. 170-180).

Lange 2-2,2 mm.

Gelb, orange und braun; Kopf an der Stirne zwischen den Augen oft leicht angedunkelt; Ftihler
und Maxillarpalpen gelb bis orange; Halsschild und Schildchen braun; Fliigeldecken ebenso, auf der
hinteren Halfte ein grosser, dunkler, isolierter Flecken, der an der Naht manchmal ein wenig aufgelost
ist, oft befindet sich eine kleine, kurze Verdunkelung auf der angedeuteten Rippe hinter den
Schulterbeulen; Beine hellbraun, Hinterschienen leicht angedunkelt.

58

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA; MALACHIIDAE)

Abh. 170-171. Sphinginopalpus s. str. orientalis sp. nov. d : 170-171, Maxillarpalpen (x220) (170,
von oben; 171, von unten).

Kopf mil den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberflache fein chagiiniert.
Maxillarpalpen Abb. 170-171. Fiihler (Abb. 172) ca. 10% langer als die Fliigeldecken, died 1 mit
einer kurzen Kerbe gleich nach der Basis und anschliessend mit einem ziemlich langen, schmalen,
spitzen Zahn; folgende Glieder schmal. Halsschild langer als breit (16x12), Seiten gerundet und
anschliessend gegen die Basis gerundet verengt; Chagrinierung des aufgewblbten Teils fein, auf dem
verengten Teil nur wenig starker. Fliigeldecken ca. doppelt so lang wie der Halsschild. nach hinten
verbreitert, hinter den Schulterbeulen eine kurze angedeutete Leiste, Punktierung grob, wenig tief,
zuerst in Reihen, hinten zerstreut. Hinterschienen gekriimmt, schwach verbreitert. Tergit 8 Abb. 173.
Sternit 8 Abb. 174. Spiculum Abb. 175.

Kopulationsapparat Abb. 176 dorsal.

Weibchen.

Wie das 6 gefarbt, aber die kleinen Flecken auf den Rippen der Fliigeldecken fehlen manchmal.
Letztes Glied der Maxillarpalpen langlich, fast oval. Fiihler einfach.

Holotypus und 18 Paratypen (NHMB), 2 Paratypen (NCI), 2 Paratypen (AMG): South Africa, Natal:
St Lucia, 10 m, 25. -27.x. 1981, J. & S. Klapperich. Zululand: Dukuduku Forest Station, 28°22'S,
32°19"E, 5.iv.l974, E-Y:327, S. Endrbdy-Younga, 2 Paratypen (TMP). Cape Prov.: De Rust Valley,
29. xi. 1981, J. & S. Klapperich, Fundort wahrscheinlich unrichtig.

Diese neue Art gehort zur kleinen Gruppe mit braunem bis orangenem Halsschild (lesothoensis,
tetrastigma, hanglipensis, nifithorax). Sie unterscheidet sich von diesen Arten gemass der
Bestimmungstabelle.

59

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19. PT. 1 .10 JANUARY 1994

At>b. n2-il6. Sphinginopalpus s. sir. onentalii sp . nov. 6 :172, Fuhlerglieder 1-4; 173,Tcrgit8; 174, Sterna 8; 175, Spiculum;
176, Kopulationsapparat (dorsal). Massstab von 172 auch fiir 173-176.

28. Sphinginopalpus s. str. lesothoensis sp. nov.

MAnnchen (Abb. 177-182).

Lange 2, 1-2,3 mm.

Kopf und Maxillarpalpen schwarz; Fiihler schwarzlich, erste 5 Glieder gelb; Halsschild braun;
Schildchen schwarz; Fliigeldecken braun mil einem queren schwarzen Flecken an der Basis, die
Seiten nicht erreichend, an der Naht gegen das Schildchen schwach aufgehellt und je einem langlichen,
isolierten schwarzen Flecken hinter der Mitte; Schenkel schwarz, Tibien und Tarsen braun.

Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberflache glatt.
Maxillarpalpen Abb. 177-178. Fiihler (Abb. 179) um fast 1/4 kiirzer als die Fliigeldecken, Glied 3
deutlich langer als 4. Halsschild kaum langer als breit (14,5x14), Seiten vorne gerundet, gegen die
Basis verengt; vorderer Teil nicht sehr stark aufgewolbt; Oberflache glatt. Fliigeldecken langoval;
falsche Epipleuren ca. 70% der Lange einnehmend, Krenulierung fehlt; Punkte weit auseinander
stehend, meistens in Reihen. Tergit 8 Abb. 180. Sternit 8 Abb. 181.

Kopulationsapparat Abb. 182 dorsal.

60

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA; MALACHIIDAE)

Abb. 177-178. Sphinginopalpus s. str. lesoihoensis sp. nov. 3 : 177-178, Maxillarpalpen (x200)
(177, von oben; 178, von iinicn).

Abb. 179-182. Sphinginopalpus s. str. lesothoensis sp. nov. 3 : 179, Fuhlerglieder 1-4; 180, Tergit 8; 181, Sternil 8; 182,
Kopulationsapparat (dorsal), Massstab von 179 auch fiir 181, 182.

61

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Weibchen.

Wie das S gefarbt, Fiihlerglied 1 einfach.

Holotypus 6 und 1 Paratypus ? (TMP): South Africa, Lesotho: Hodson’s Peak, 3000 m, 28°37'S,
29°1TE, ll.iii.l976, E-Y:1068, S. Endrody-Younga.

Neben tetrastictus Champion und rufithorax Wittmer zu stellen. Die neue Art unterscheidet sich
durch die Form der Fiihlerglieder 1 und 2 und die verschieden gebauten Terminalia.

29. Sphinginopalpus s. str. orangensis sp. nov.

Mannchen (Abb. 183-188).

Lange 2, 2-2,5 mm.

Kopfbasis orange, Stirne zwischen dem oberen Teil der Augen mit einer breiteren dunkleren
Makel, die die Augen oft beriihrt, Vorderkopf mit den Maxillarpalpen gelb; Fiihler bis died 6 gelb,
Oberseite von 2 und Basis von 3 leicht angedunkelt, 7 bis 11 dunkelbraun bis schwarz; Halsschild
mit schmaler, fast gelber Basis, daim quer schmal dunkler, seitlich, manchmal bis zum Vorderrand
dunkler, Mitte bis Vorderrand braunlich; Schildchen dunkel, manchmal braunlich aufgehellt;
Fliigeldecken braun bis gelblich, jede Decke mit 2 hintereinander liegenden, ringsum isolierten
braunen bis dunkelbraunen Langsmakeln; Beine gelblich bis braunlich, besonders die Hintertibien
ein wenig dunkler.

Abb. 183-184. Sphinginopalpus s. str. orangensis sp. nov. 3 : 183-184, Maxillarpalpen (xl85) (183,
von oben; 184, von unten).

62

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA; MALACHIIDAE)

1 88

Abb. 185-188. Sphinginopalpus s. str. orangensis sp. nov. 6 : 185, Fuhlerglieder 1-4; 186, Tergit 8; 187, Sternit 8; 188,
Kopulationsapparat (dorsal). Massstab von 186 auch fiir 187-188.

Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewolbt; feinstens chagriniert.
Maxillarpalpen Abb. 183-184. Fiihler (Abb. 185) ungefahr 10% kiirzer als die Fliigeldecken, died
1 mit einer kurzen Kerbe an der Basis und einem kleinen spitzwinkligen Zahn an der Kerbe, 4 und
einige folgende deutlich breiter als 3. Halsschild langer als breit (16x14), Seiten zuerst fast parallel,
daim allmahlich eingeschniirt; aufgewolbter Teil fast glatt bis feinstens chagriniert, dann gegen die
Basis starker chagriniert, matt. Fliigeldecken ca. 2,6 mal langer als der Halsschild, nach hinten etwas
verbreitert, Punktierung grob, in Reihen bis etwa 2/3 der Lange, dann weniger tief und zerstreut.
Hinterschienen ein wenig gekriimmt, gegen die Spitze allmahlich schwach verbreitert. Tergit 8 Abb.
186. Sternit 8 Abb. 187.

Kopulationsapparat Abb. 188 dorsal.

WeibcHEN (Abb. F).

Kopf und Halsschild meistens etwas blasser als beim S . Letztes died der Maxillarpalpen
beilformig (Abb. F). Fuhlerglieder 4 und folgende nur ganz unmerklich breiter als 2 und 3, die dunkle

63

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Farbung an 2 und 3 nur angedeutet oder ganz fehlend. Halsschild langer (18/19x15).

Holotypus und 34 Paratypen (NHMB): South Africa, Richtersveld: Alexander Bay, river, 28°32'S,
16°33'E, 6.-7.ix.l987; idem E-Y: 2478, S. Endrody-Younga, 56 Paratypen (TMP), 4 Paratypen
(AMG).

Von den iibrigen Arten mit braunem bis orangenem Halsschild unterscheidet sich diese Art durch
den mehr oder weniger stark leicht dunkel gefleckten Halsschild, das Fiihlerglied 1 und dieTerminalia.

30. Sphinginopalpus s.str. tetrastigma Champion

Sphinginopalpus tetrastigma Champion, 1922: 333.

MAnnchen (Abb. 189-194).

Der Holotypus befindet sich im BM, er wurde fiir die
Abbildungen verwendet.

Das letzte Glied der Maxillarpalpen (Abb. 189) ist vorne
viel starker verbreitert als bei den folgenden 2 Arten und der
Aussenrand verlauft in gerader Linie nach hinten, bei den
anderen Arten ist er gerundet ausgeschnitten. Fiihlerglieder
1 bis 4 (Abb. 190). Tergit 8 Abb. 191. Sternit 8 Abb. 192.
Kopulationsapparat Abb. 193 dorsal. Abb. 194 Profil.

Abb. 189. Sphinginopalpus s. str. tetrastigma
Champion cJ : 189, Maxillarpalpus
(x220).

64

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 190-194. Sphinginopalpus s. str. leirastigma Champion 6 : 190, Fuhlerglicder 1-4; 191, Tergit 8; 192, Sternit 8; 193-
194, Kopulationsapparat (193, dorsal; 194, Profil). Massstab von 190 auch fiir 192 und 191 auch fiir 193, 194.

65

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

31. Sphinginopalpus s. str. hanglipensis sp. nov.
Mannchen (Abb. 195-201).

Lange 2mm.

Kopf orange, Stirne angedunkelt, die dunkle Farbung zieht sich schmal nach vorne, bis zur Halfte
neben den Augen und Wangen orange; Maxillarpalpen, Halsschild, Schildchen und Beine orange bis
hellbraun; Fliigeldecken hellbraun, jede Decke mit einem basalen dunkelbraunen Flecken, beim
Schildchen praktisch an die Naht anstossend, die Seiten unter den Schultern nicht erreichend und
einem weiteren, langlichen, isolierten Flecken jederseits hinter der Mitte.

Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberflache glatt, fein,
massig dicht punktiert. Maxillarpalpen Abb. 195-196. Fiihler (Abb. 197) um 1/5 kiirzer als die

Fliigeldecken, Glied 1 nach dem
basalen Einschnitt am breitesten,
gegen die Spitze regelmassig
verschmalert, 3 nur wenig langer
als 4, 2 bis 10 zur Spitze nur
schwach verbreitert. Halsschild
kaum langer als breit
(15,5x14,5), vorderer Teil wenig
aufgewolbt, abgeflachter Teil
ebenfalls schwach gewolbt;
Seiten ungefahr in der Mitte am
breitesten, Randung wie bei
capensis; ganze Oberflache glatt,
Punkte wie auf dem Kopfe.
Fliigeldecken langoval; falsche
Epipleuren sehr lang, fast 90%
der Lange eirmehmend, keine
Krenulierung; Punkte sehr gross,
nicht sehr tief, in regelmassigen
Punktreihen angeordnet; die
aufrechtstehenden Haare

Abb.

195-196. Sphinginopalpus
Maxillarpalpen (x230).

s. sir. hanglipensis sp. nov. S : 195-196,

besonders lang. Tergit 8 Abb. 198. Sternite 7 und 8 Abb. 199.
Kopulationsapparat Abb. 200 dorsal. Abb. 201 Profil.

Holotypus(TMP): South Africa, N Transvaal: Soutpansberg, Hanglip, grassnetting, summit, 30°00"S,
29°54'E, 18.iii.l973, E-Y:68, S. Endrody-Younga.

Eine mit tetrastigma Champion nahe verwandte Art, Unterschiede vergleiche Abbildungen.

66

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 197-201. Sphinginopalpus s. sir. hanglipensis sp. nov. cJ ; 197, Fuhlcrglictler 1-4, 198, Tcrgii 8; 199, Slcrnii 7 und 8,
200- 201, Kopulationsapparat (200, dorsal; 201, Profil). Massstab von 197 auch fiir 199-201.

32. Sphinginopalpus s. str. rufithorax sp. nov.

MANNCHEN (Abb. 202-207).

Lange 2,2 mm.

Kopf schwarz, von ca. der Mitte der Augen nach vorne gelb; Maxillarpalpen braun, teilweise
angedunkelt; Fiihler gelb, ab Glied 5 oder 6 zunehmend dunkel; Halsschild orangebraun; Schildchen
schwarz; Fliigeldecken vorwiegend gelb bis gelbbraun. Basis bis iiber die Schultern schwarzlich.

67

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

jedoch Seiten darunter hell, jederseits je ein ovaler isolierter Flecken beginnt kurz vor der Mitte und
lasst die Spitzen breit hell, Oder die dunkle Farbung ist starker ausgebreitet, die ovalen Flecken grosser,
sie verschmelzen an der Naht und sind schmaler werdend mit der dunkeln Basis verbunden, Spitzen
breit hell; Schenkel mehr Oder weniger angedunkelt, ebenso die hinteren Schienen, 4 Vorderschienen

Kopf mit den Augen nur wenig breiter
als der Halsschild, Stirne leicht gewolbt;
Oberflache glatt, glanzend. Maxillarpalpen
Abb. 202- 203. Fiihler (Abb. 204) ca. 1/4
kurzer als die Fliigeldecken, Glieder 3 bis 10
nur ganz wenig zur Spitze verbreitert, 3 ein
wenig langer als 4. Halsschild langer als breit
(17x15), mit capensis iibereinstimmend,
jedoch vorderer Teil weniger stark
aufgewolbt und Oberflache glatt, glanzend,
nur der abgeflachte Teil chagriniert.
Fliigeldecken langoval, ungefahr in der Mitte
am breitesten, Schulterbeulen normal; falsche
Epipleuren ca. 50% der Lange der Decken
einnehmend, nicht krenuliert; Punktierung
ahnlich wie bei capensis. Behaarung
staubartig, auf den Fliigeldecken nur ganz
vereinzelte, langere, helle, aufstehende
Haare. Tergit 8 Abb. 205. Sternit 8 Abb. 206.
Kopulationsapparat Abb. 207 dorsal.

Holotypus und 9 Paratypen (TMP), 5 Paratypen (NHMB), 1 Paratypus (AMG): South Africa,
Transvaal: Waterberg, Haakdoringboom farm, flood debris, Mogo IR, 24°11'S, 27°50'E, 13. ii. 1976,
E-Y:1039, A. Strydom; Onderstepoort (Pretoria), iv.l987, 1. Pajor, 2 Paratypen (NHMB); Nelspruit
Nature Reserve, rivulet vail., grassnetting, 19.xii.l986, 25°29'S, 30°55'E, E-Y:2402, S. Endrody-
Younga, 1 Paratypus (TMP), 1 Paratypus (NHMB).

Diese neue Art ist neben lesothoensis Wittmer zu stellen, mit der sie den schwarzen Kopf
gemeinsam hat, Unterschiede zeigt das Fiihlerglied 1 und die Terminalia, siehe Abbildungen.

gelb, ebenso alle Tarsen.

Abb. 202-203 . Sphinginopalpus s. str. rufithoraxsY>. nov. S : 202-
203, Maxillarpalpen (xl25) (202, von oben; 203, von
unten)

Weibchen.

Wie das S gefarbt; Maxillarpalpen einfach.

68

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 204-207. Sphinginopalpus s. sir. rufithorax sp. nov. 6 \ 204, Fiihlerglieder 1-4; 205, Tergit 8; 206, Sternit 8; 207,
Kopulationsapparat (dorsal). Massstab von 204 auch fiir 206, 207.

33. Sphinginopalpus s. str. nigrithorax sp. nov.

MAnnchen (Abb. 208-213).

Lange 2,3 mm.

Kopf schwarz, von der Mitte der Augen nach vorne gelb, liber dem Clypeus ein kleiner,
verschwommener, dunkler Flecken; Maxillarpalpen dunkel, teils aufgehellt; Fiihler dunkel, erste 5
Glieder gelb, 1 und 2 auf der Oberseite schwach dunkel; Halsschild, Schildchen und Fliigeldecken
schwarz, letztere mil einem schmalen, weissen Querband vor der Mitte, das an der Naht schmal
unterbrochen 1st; Hinterbeine schwarz, Hintertibien kaum merklich aufgehellt; 4 vordere Schenkel
und Tibien dunkel, Knie und Spitzen der Tibien etwas aufgehellt, 4 vordere Tarsen ziemlich hell.

Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewdlbt; Oberflache glatt.
Maxillarpalpen Abb. 208-209. Fiihler (Abb. 210) um ca. 1/5 kiirzer als die Fliigeldecken, Glieder 3
bis 10 fast parallel, 3 deutlich langer als 4. Halsschild langer als breit (17x14), wie bei capensis,
jedoch aufgewolbter Teil glatt, abgeflachter Teil chagriniert. Fliigeldecken lang, zur Spitze etwas
verbreitert, Schulterbeulen normal; falsche Epipleuren kurz, ca. 30% der Lange einnehmend, wenig

69

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 208-209. Sphinginopalpus s. str, nigrithorax sp. nov. 6 :

Maxillarpalpen (x200) (208, von oben; 209, von unten).

vorstehend, ganz fein krenuliert; Punkte ein wenig grosser als bei capensis; Oberflache glatt. Tergit
8 Abb. 211. Sternit 8 Abb. 212.

Kopulationsapparat Abb. 213 dorsal.

Holotypus (NHMB): South Africa, Cape Prov.; 2 km E Suurfontein, 22. ix. 1986, W. Wittmer, M.
Macpherson.

Bisher die einzige aus Siidafrika bekarmtgewordene Art mit einfarbig schwarzem Halsschild.

70

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 210-213. Sphinginopalpus s. sir. nigriihorax sp. nov. 6 : 210, Fuhlerglieder 1-4, 211, Tergit 8; 212, Stcrnil 8; 213,
Kopulationsapparat (dorsal). Massstab von 210 auch fur 212, 213.

34. Sphinginopalpus s. str. capensis sp. nov.

Mannchen (Abb. 214-219).

Lange 1, 9-2,1 mm.

Kopf schwarz mil einem breiten gelben Flecken am Vorderkopf, der selten bis zu den Augen
reicht, Wangen meistens teils bis ganz gelb; Maxillarpalpen meistens gelb, seltener braun bis
dunkelbraun; Fuhlerglieder 1 bis 5 gelb, seltener 5 teils dunkel, restliche schwarz; Halsschild schwarz,
an der Basis schmal aufgehellt; Schildchen und Fliigeldecken schwarz, letztere mit einem schmalen,
gelben, durchgehenden Querband vor der Mitte; Beine schwarz, Spitzen der Vordertibien seltener.

71

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 214-215. Sphinginopalpus s. str. capensis sp. nov. 3: 214-215, Maxillarpalpen (x200) (214,
von oben; 215, von unten).

Vordertarsen fast immer ganz aufgehellt.

Kopf mit den Augen kaum merklich breiter als der Halsschild, Stirne leicht gewolbt; Oberflache
fein chagriniert. Maxillarpalpen Abb. 214-215. Fiihler (Abb. 216) urn ca. 1/5 kiirzer als die
Fliigeldecken, died 1 mit einem kleinen Zahn an der Basis oben und einem grosseren auf der
Unterseite, 3 ein wenig langer als 4. Halsschild Linger als breit (16x14), vorderer Teil stark aufgewdlbt,
Basis abgeflacht; Seiten gerundet, in der Mitte am breitesten, nach vorne kaum verschmalert, gegen
die Basis stark verengt; Basalrand deutlicher als die Seitenrander, die kurz vor der Basis erloschen;
Oberflache fein chagriniert. Fliigeldecken langoval, nach der Mitte am breitesten; Schulterbeulen
normal; falsche Epipleuren ca. 40% der Lange der Decken einnehmend, nur angedeutet krenuliert;
Punktierung bis zur Mitte, oder dariiber hinaus ziemlich grob, in Punktreihen, nach hinten erloschend,
verworren. Behaarung des Korpers fein staubartig, die Fliigeldecken mit einzelnen langen, hellen,
aufstehenden Haaren. Tergit 8 Abb. 217. Sternit 8 Abb. 218.

Kopulationsapparat Abb. 219 dorsal.

Weibchen.

Kopf einfarbig schwarz, sonst wie das S gefarbt.

Holotypus und 6 Paratypen (NHMB): South Africa, SW Cape Prov. ; 3 km S Lamberts Bay, 22.ix. 1984,
W. Wittmer; 50 km S Kleinsee, 30°10'S, 17°14'E, 10.ix.l987, W. Wittmer, 6 Paratypen (NHMB);
20 km N Sout Pan, 12.ix.l985, W. Wittmer, 1 Paratypus (NHMB); near Strandfontein, 31°45'S,
18°14"E, 16. ix. 1986, W. Wittmer, R. Oberprieler, 28 Paratypen (NHMB), 15 Paratypen (NCI), 4
Paratypen (AMG);Papendorp dunes, 31°38'S, 18°12'E, 22. viii. 1981, E-Y: 1827, S. Endrody-Younga,
Paratypen (TMP); Seweputs coast, 31°39'S, 18°17'E, E-Y:1836, S. Endrody-Younga, 8 Paratypen
(TMP); Hondeklipbaai, 30°19"S, 17°17"E, 14.ix.l986, W. Wittmer, R. Oberprieler, 10 Paratypen
(NHMB); 5 km E Port Nolloth, 29°17^S, 16°55^E, 9.ix.l986, W. Wittmer, R. Oberprieler, 20
Paratypen (NHMB), 3 Paratypen (NCI).

Diese neue Art ist neben pallidipalpis Wittmer zu stellen.

72

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA; MALACHIIDAE)

Abb. 216-219. Sphinginopalpus s. str. capensis sp. nov. 6 : 216, Fuhlcrglicder 1-4; 217, Tergit 8; 218, Slernit 8; 219,
Kopulationsapparat (dorsal). Massstab von 216 auch fiir 218, 219.

35. Sphinginopalpus s. str. dentifrons sp. nov.

MANNCHEN (Abb. 220-226).

Lange 2,5 mm.

Kopf schwarz, von der Mitte der Augen nach vorne zuerst braun aufgeheilt, dann inMusive
Wangen gelb; Halsschild schwarz, Maxillarpalpen gelbbraun; Fiihler schwarzlich, Glieder 1 und 2
gelb, 3 bis 5 hellbraun; Halsschild schwarz, ungefahr die Halfte des abgesetzten Teils an der Basis
gelbbraun; Schildchen und Fliigeldecken schwarz, letztere mil einem weisslichen, antimedianen,
dreieckigen Seitenflecken, der sich etwas mehr nach hinten als nach vorne verlangert und die Naht
nicht erreicht; Hinterbeine schwarz, Mittelbeine dunkelbraun, Mitteltarsen ein wenig heller.

73

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 22Q-221. Sphinginopalpus s. str. dentifrons sp. nov. tj : 220-221, Maxillarpalpen (xl70)
(220, von oben; 221, von unten).

Vorderschenkel dunkel, bei den Knien heller, Vordertibien und Tarsen hellbraun.

Kopf (Abb. 222) mil den Augen breiter als der Halsschild; Vorderstirne mil einem kleinen,
spitzen Zahn in der Mitte; Oberflache glatt. Maxillarpalpen Abb. 220-221. Fiihler (Abb. 223) urn ca.
15% kiirzer als die Fliigeldecken, died 3 deutlich langer als 4. Halsschild langer als breit (20x18);
Seiten nach vorne kaum merklich verengt; Oberflache vorne glatt, abgeflachter Teil zum grossten
Teil chagriniert. Fliigeldecken langoval; falsche Epipleuren ca. 70% der Lange einnehmend, nicht
krenuliert; 2 Punktreihen neben der Naht auf 1/3 der Lange regelmassig, nach hinten nehmen die
Punkte an Tiefe und Dichte ab. Tergit 8 Abb. 224. Sternit 8 Abb. 225.

Kopulationsapparat Abb. 226 dorsal.

Holotypus (TMP): South Africa, Natal: Indumeni For., Bergville Distr., humus, x.1960, N. Leleup.

Die einzige bis jetzt bekanntgewordene Art, deren Vorderstirne mit einem kleinen, spitzen Zahn
bewehrt ist.

74

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 222-226. Sphinginopalpus s. str. dentifrons sp. nov. d ; 222, Kopf; 223 Fuhlerglieder 1-4; 224, Tergit 8; 225, Stemil 8;
226, Kopulalionsapparat (dorsal). Massstab von 223 auch fiir 225, 226.

75

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Kopf schwarz, Vorderkopf mil den
Wangen variabel gelb bis orange gefarbt,
manchmal beginnend am Vorderrand der
Augen, Oder etwas weiter vorne ein dunkles
gebogenes Querband, beginnend am
Vorderrand der Fiihlerwurzel, in alien Fallen
sind die Wangen hell; Maxillarpalpen gelb;
Fiihler gelb, die letzten 5 bis 6 Glieder
zunehmend dunkel, bei einem Exemplar ein
dunkler Langswisch auf 1 basal; Halsschild
schwarz, grbsster Teil der abgeflachten Basis
hell; Schildchen und Fliigeldecken schwarz,
letztere jederseits vor der Mitte mil einem
kleinen, weisslichen Flecken; Hinterbeine
schwarz, Hintertibien manchmal leicht
aufgehellt, 4 Vorderbeine gelb, Schenkel bis
kurz vor den Knien schwarz, alle 6 Tarsen
gelb.

Kopf mil den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberflache glatt, einige
zerstreute, feine Haarpunkte sichtbar. Maxillarpalpen Abb. 227-228. Fiihler (Abb. 229) fast um 1/3
kiirzer als die Fliigeldecken, Glieder 3 bis 10 gegen die Spitze kaum erweitert, 3 so lang oder ein
wenig langer als 4. Halsschild ungefahr so lang wie breit, sonst wie bei capensis, jedoch Oberflache
glatt, nur die abgeflachte Basis chagriniert. Fliigeldecken langoval, wie bei capensis, aber starker
punktiert, falsche Epipleuren ca. 50% der Lange einnehmend, praktisch nicht krenuliert. Tergit 8
Abb. 230. Sternit8Abb. 231.

Kopulationsapparat Abb. 232 dorsal.

Weibchen.

Kopf einfarbig schwarz, ebenso die Maxillarpalpen, sonst wie das 6 gefarbt.

Holotypus und 14 Paratypen (TMP), 9 Paratypen (NHMB), 2 Paratypen (AMG): South Africa, E
Transvaal: Nelshoogte Forest Station, 1.- 7.xii.l986, 8.iv.l987, 25°47'S, 30°49"E, grassnetting, E-
Y:2335, 2338, 2350, 2360, 2456, S. Endrody-Younga; Cycas Farm, grassnetting, 5.xi.l985, E-Y:
2262, S. Endrody-Younga, 1 Paratypus (NHMB). Natal: Richards Bay, dense grass, 28°50'S, 32°05"E,
13.x. 1984, P. E. Reavell, 1 Paratypus (TMP).

Diese neue Art ist neben albilabris (Boheman) zu stellen. Sie ist von gleicher Farbung
(ausgenommen der Kopf) und Korperform, das Fiihlerglied 1 ist etwas verschieden gebaut, weitere
grosse Unterschiede zeigen die Terminalia, vergleiche Abbildungen.

36. Sphinginopalpus s. str. variceps sp. nov.

Mannchen (Abb. 227-232).

Lange 2 mm.

227 228

Abb. 227-228. Sphinginopalpus s. str. variceps sp. nov. 6 : 227-
228, Maxillarp^pen (x200) (227, von oben; 228, von
unten).

76

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 229-232. Sphinginopalpus s. str. variceps sp. nov. <5: 229, Fiihlerglieder 1-4; 230, Tergit 8, 231, Sternit 8; 232,
Kopulationsapparat (dorsal). Massstab von 229 auch liir 231, 232.

37. Sphinginopalpus s. str. albilabris (Boheman)

Chalicorus albilabris Boheman, 1851: 477.

Sphinginopalpus albilabis Boheman - Champion 1922: 332, parte.

MANNCHEN (Abb. 233-239).

Der Holotypus ( c? ) stimmt mit Exemplaren iiberein, die mir von mehreren Fundorten vorliegen.
Fiir die Abbildungen wurde 1 Ex. von Weza, Impetyene, E- Y :2680, verwendet: Maxillarpalpen Abb.
233-234. Fiihler Abb. 235. Tergit 8 Abb. 236. Sternit 8 Abb. 237. Kopulationsapparat Abb. 238
dorsal, Abb. 239 Profil.

77

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb, 233-234. Sphing'mopalpus s. str. albilabris (Boheman) <3: 233-234, Maxillarpalpen
(233, von oben, x300; 234, von unten, x260).

Weibchen (Abb. D, E, H).

Ausserdem verweise ich auf die Gesamptaufnahme eines $ von Nelshoogte, E-Y:2350, Abb.
D, E und die Basis des Halsschildes Abb. H.

Eingesehenes MATERIAL: South Africa, Natal: Frere, ix. und x.1892, G. A. K. Marshall, 3 $ (BM),
det. G. C. Champion; Weza, Impetyene grassveld, flowering grass, grass tussocks and intersept, trap
12 days, 30°37'S, 29°42'E, 16.-21.xi.l989, E-Y:2680, 2709, 2712, 2718, S. Endrody-Younga, I.
Klimaszewski, 8 Ex. (TMP), 5 Ex. (NHMB). Transkei: Fraser Gorge, grass tussocks, 31°24'S,
29°42'E, 27. xi. 1988, E-Y:2584, S. Endrody-Younga, 10 Ex. (TMP), 5 Ex. (NHMB). E Transvaal:
Uitsoek Forest Station, 1100 m, grassveld, 12.xii.l986, 6. v. 1987, E-Y:2379, 2426, 25°18'S, 30°55^E,
S. Endrody-Younga, 39 Ex. (TMP), 11 Ex. (NHMB); Gods Window, grassnetting, 24°35"S, 30°5HE,
12.xi.l980, E-Y:1734, S. Endrody-Younga, 14 Ex. (TMP).

Die von Champion (1922) als albilabris bestimmten Exemplare aus Salisbury (jetzt Harare)
(Zimbabwe) gehoren einer neuen Art an, die ich hier als majeri Wittmer beschreibe.

78

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 235-239. Sphinginopalpus s. str. albilabris (Boheman) S : 235, Ftihlerglieder 1-4; 236, Tergit 8; 237, Stemit 8; 238-239,

Kopulationsapparat (238, dorsal; 239, Profil). Massstab von 235 auch fiir 237-239.

79

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

38. Sphinginopalpus s. str. complexus sp. nov.

MAnnchen (Abb. 240-245).

Lange 2 mm.

Eine in Korperform und Farbung mil albilabris (Boheman) iibereinstimmende Art. Sie
unterscheidet sich durch die Form der Maxillarpalpen (Abb. 240-241), die zum grossten Teil gelb
sind, nur died 1 ist teils leicht angedunkelt, Fiihler Abb. 242, Tergit 8 Abb. 243, Sternit 8 Abb. 244,
Kopulationsapparat Abb. 245 dorsal, vergleiche mit den Abbildungen 233-239.

Holotypus und 28 Paratypen (NHMB), 2 Paratypen (TMP), 2 Paratypen (AMG): South Africa,
Transvaal; Bothasnek, 1450 m, 23. xi. 1987, W. Wittmer.

Diese neue Art is neben albilabris (Boheman) zu stellen, von der sie sich durch die verschiedenen
Terminalia unterscheidet. Die Maxillarpalpen (Abb. 240-241) gleichen sehr albilabris.

Abb. 240-241. Sphinginopalpus s. sir. complexus sp. nov. 6 ■ 240-241, Maxillaq>alpen
(x250) (240, von oben; 241, von unten).

80

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 242-245. Sphinginopalpus s. str. complexus sp. nov. 6 : 242, Fiihlerglieder 1-4; 243, Tergit 8; 244, Sternit 8; 245,
Kopulationsapparat (dorsal). Massstab von 242 auch fiir 244, 245.

81

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

39. Sphinginopalpus s. str. politipennis sp. nov.

MAnnchen (Abb. 246-251).

Lange fast 2,5 mm.

Kopf schwarz, von der Mitte der Augen nach vorne, inklusive der Wangen, gelb; Maxillarpalpen
Schwarz; Fiihler gelb, die letzten 4 bis 5 Glieder schwarzlich; Halsschild schwarz, ein Teil der
abgesetzten Basis gelbbraun; Schildchen und Fliigeldecken schwarz, letztere jederseits mit einem
schmalen, kurzen, antemedianen weisslichen Flecken; Hinterschenkel schwarz, Hintertibien
schwarzlich, an der Basis kurz aufgehellt, Hintertarsen braun, 4 Vorderschenkel schwarz, an den
Knien kurz heller, 4 vordere Schienen und Tarsen gelb.

Kopf mit den Augen breiter
als der Halsschild, Stirne leicht
gewolbt; Oberflache glatt.
Maxillarpalpen Abb. 246-247.
Fiihler (Abb. 248) um ca. 1/5
kiirzer als die Fliigeldecken,
Zahn an Glied 1 mit sehr langer
undschmaler Spitze, 3 nur wenig
langer als 4. Halsschild langer als
breit (15,5x14), sonst wie bei
oculimaculatus, jedoch

Chagrinierung an der
abgeflachten Basis feiner.
Fliigeldecken langoval, die
falschen Epipleuren konnten
nicht gemessen werden, weil die
Decken beidseitig eingerollt
sind; Oberflache glatt,
Abb. 246-247. Sphinginopalpus s. str. politipennis sp. nov. 6 : 246-247, Punktierung Sehr spiirlich. Tergit
Maxillarpalpen (xl70) (246, von oben; 247, von unten). g Abb. 249. Stemit 8 Abb. 250.

Kopulationsapparat Abb. 251.

Holotypus (TMP), 1 Paratypus (NHMB): South Africa, E Transvaal: Nelshoogte, Devils Knuckle,
grassnetting, 25°47"S, 30°49'E, 4.xii.l986, E-Y:2350, S. Endrody-Younga.

Die stark glanzenden Fliigeldecken mit den verhaltnismassig wenig weit nach hinten reichenden
Punkten, nebst den Terminalia, unterscheiden diese Art von den vielen ahnlich gefarbten.

82

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 248-251. Sphinginopalpus s. str. poUtipennis sp. nov. S : 248, Fiihlerglieder 1-4; 249, Tergit 8; 250, Stemit 8; 251,
Kopulationsapparat (dorsal). Massstab von 248 auch fiir 250, 251.

40. Sphinginopalpus s. str. trispinopsis sp. nov.

MAnnchen (Abb. 252-255).

Lange 2,2 mm.

Line in der Farbung und Kdrperform mil trispimis Wittmer iibereinstimmende Art, nur bei den
Fiihlern fehlt der kleine schwarze Flecken auf Glied 3, dafiir ist 4 bereits angedunkelt, bei trispimis gelb.

83

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Fiihler Abb. 252. Die Maxillarpalpen sind denen von trispinus sehr ahnlich und werden deshalb
nicht abgebildet. Tergit 8 Abb. 253. Sternit 8 (Abb. 254) und der Kopulationsapparat (Abb. 255)
zeigen grossere Unterschiede.

Weibchen.

Fiihlerglieder 1 bis 5 gelb, nur 1 an der Basis ganz wenig dunkel, sonst wie das 6 gefarbt;
ungefliigelt.

Holotypus 6 und 1 Paratypus $ (TMP), 1 Paratypus $ (NHMB); South Africa, E Transvaal: Gods
Window, grassnetting, 24°35'S, 30°51'E, 12. xi. 1980, E-Y:1734, S. Endrody-Younga.

Diese neue Art gehort zu einer kleinen Gruppe deren Fuhlerglied 1 mit drei Zahnen bewehrt ist
(tridens, tridensmimus).

Abb. 252-255. Sphinginopalpus s. str. trispinopsis sp. nov. S : 252, Fiihlerglieder 1-4; 253, Tergit 8; 254, Stemit 8; 255,
Kopulationsapparat (dorsal). Massstab von 252 auch fiir 254, 255.

84

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

41. Sphinginopalpus s. str. oneili Pic.

Sphinginopalpus oneili Pic, 1903: 164. - 1904: 13. - Champion, 1922: 328, T. 5, Fig. 21, parte.
MANNCHEN (Abb. 256-264).

Im MP befindet sich der Lectotypus
und 7 Paralectotypen, im BM 4
Paralectotypen, im SAM 3 Paralectotypen,
im NHMB 1 Paralectotypus.
Originalfundort : Dunbrody (Cape Prov.),
33°28'S, 25°33'E. Ftir die verschiedenen
Abbildungen diente der Lectotypus.
Maxillarpalpen Abb. 256-257. Fiihler
Abb. 258. Tergit 8 Abb. 259. Sternit 8 Abb.
260. Spiculum Abb. 261. Tegmen Abb.
262.

Kopulationsapparat Abb. 263 dorsal.
Abb. 264 Profil.

Abb. 256-257. Sphinginopalpus s. str. oneili Pic 6 : 256-257,

Maxillarpalpen (xl55) (256, von oben; 257, von unten).

Weibchen (Abb. 265).

Champion (1922) stellte eine Anzahl von Exemplaren von Salisbury (jetzt Harare) (Zimbabwe)
zu oneili, die einer anderen, neuen Art angehoren, die ich hier als majeri Wittmer beschreibe.

85

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 258-265. Sphinginopalpus s. sir. oneili Pic 6 : 258, Fuhlerglicdcr 1-4; 259, Tcrgit 8; 260, Sterna 8; 261, Spiculum; 262,
Tegmen; 263-264, Kopulationsapparat (263, dorsal; 264, Profil); 265, Sphinginopalpus s. str. oneili Pic $ :
Maxillarpalpus. Massstab von 258 auch fiir 265 und 259 auch fur 261-264.

42. Sphinginopalpus s. str. leucosideae sp. nov,

MANNCHEN (Abb. 266-271).

Lange 2 mm.

Kopf schwarz, ein gelber Flecken auf dem Vorderkopt, zwischen den Fiihlerwurzeln, die Augen
und Wangen nicht erreichend; Maxillarpalpen schwarzbraun, teils schwach aufgehellt; Fiihler dunkel,
die Glieder 1 bis 5 gelb, Basis von 1 schwarz und mil einem schwarzen Langswisch auf der Oberseite;
Halsschild schwarz, die ganze abgeflachle Basis gelbbraun; Schildchen und Fliigeldecken schwarz,
letztere mil einem weisslichen, antimedianen, dreieckigen Seitenflecken, der sich etwas mehr nach
hinten als nach vorne hinzieht, die Naht wird nicht erreicht; Hinterbeine schwarz, Hintertarsen leicht
aufgehellt, 4 vordere Schenkel schwarz, 4 vordere Tibien und Tarsen gelb.

Kopf mit den Augen ein wenig breiter als der Halsschild, Stirne leicht gewolbt; Oberflache glatt.

86

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 266-267. Sphinginopalpus s. str. leucosideae sp. nov. cj : 266-267, Maxillarpalpen
(xl70) (266, von oben; 267, von unten).

Maxillarpalpen Abb. 266-267. Fiihler (Abb. 268) um fast 1/4 kiirzer als die Fliigeldecken, died 3
ein wenig langer als 4. Halsschild langer als breit (18x15), Oberflache glatt, abgeflachter Teil
chagriniert, sonst wie elongatidens. Fliigeldecken langoval; falsche Epipleuren ca. 70% der Lange
einnehmend, nicht krenuliert; Punkte grob, in Reihen angeordnet, nach der Mitte wenig tief,
verworren, sparlicher und fast ganz erloschend. Tergit 8 Abb. 269. Sternit 8 Abb. 270.
Kopulationsapparat Abb. 271 dorsal.

Weibchen.

Kopf einfarbig schwarz; Fiihlerglied 1 gelb, nur die Basis kurz schwarz, Schulterbeulen stark
ruduziert, wahrscheinlich ungefliigelt.

Holotypus (NMB): South Africa, Orange Free State: Golden Gate Highlands National Park, SE 28
28 BC, 13.-14.xi.l985, ex Leucosidea sericea, Mus. staff. Natal: Nyala Reserve, 10 km inland of
Empangeni, in Acacia veld, 10.ii.l985, P. E. Reavell, 1 Paratypus (NHMB); Ukulu River, swept in
grass, floodplain, P. E. Reavell, 1 Paratypus (NHMB); 4 km S Wasbank, R 621, 1030 m, 21. xi. 1987,
W. Wittmer, 4 Paratypen (NHMB).

Diese neue Art scheint weit verbreitet zu sein, sie ist leicht erkennbar an dem ziemlich parallelen
Fiihlerglied 1 sowie an den Terminalia.

87

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 268-271, Sphinginopalpus s. str. leucosideae sp. nov. d; 268, Fiihlerglieder 1-4; 269, Tergit 8; 270, Stemit 8; 271,
Kopulationsapparat (dorsal). Massstab von 268 auch fiir 270-271.

43. Sphinginopalpus s. str. amatolensis sp. nov.

Mannchen (Abb. 272-277).

Lange 2,5 mm.

Kopf mil den Wangen schwarz, zwischen den Fiihlerwurzeln gelb und einem kleinen gelblichen
Flecken neben jedem Auge, der auch fehlen kann; Maxillarpalpen schwarziich, Glied 1 aufgehellt;
Fiihler schwarz, Glieder 1 bis 5 gelb, Oberseite von 1 schwarz; Halsschild schwarz, abgeflachte Basis
schmal gelbbraun; Schildchen und Fliigeldecken schwarz, letztere mit einem weisslichen,
antimedianen Querband, das an der Naht unterbrochen und seitlich verlangert ist; Hinterbeine

88

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

schwarz, Tarsen teils aufgehellt, 4 Vorderbeine schwarz bis schwarzlich, 4 Vordertarsen gelb.

Kopf mit den Augen ein wenig breiter als der Halsschild, Stirne leicht gewdlbt; Oberflache
chagriniert. Maxillarpalpen Abb. 272-273. Fiihler (Abb. 274) um nicht ganz 1/5 kiirzer als die
Fliigeldecken, Glied 1 mit sehr kleinem, feinem Zahn, 3 kaum merklich langer als 4. Halsschild langer
als breit (18x16), aufgewolbter Teil weniger stark chagriniert als der Kopf, abgeflachter Teil
chagriniert, sonst wie elongatidens. Fliigeldecken langoval; falsche Epipleuren ca. 70% der Lange
einnehmend, ganz schwach krenuliert; Punktierung ahnlich wie bei elongatidens. Tergit 8 Abb. 275.
Sternit 8 Abb. 276.

Kopulationsapparat Abb. 277 dorsal.

Abb. 272-273. Sphinginopalpus s. str. amatolensis sp. nov. S : 272-273,
Maxillarpalpen (xl70) (111, von oben; 273, von unten).

Weibchen.

Kopf einfarbig schwarz, sonst wie das S gefarbt. Schulterbeulen nur angedeutet; falsche
Epipleuren fein krenuliert; ungefliigelt.

Holotypus und 20 Paratypen (TMP), 10 Paratypen (NHMB), 2 Paratypen (AMG): South Africa, Cape
Prov.: Amatole, Isidenge Forest Station, grass tussocks, 32°4LS, 27°15'E, 18.xi.l987, E-Y:2524, S.
Endrody- Younga. Natal: Mouth of Umkomaas River, ix.l897, G. A. K. Marshall, 1 Paratypus (BM),
von Champion als myrmecodeslalbifrons bestimmt.

Diese neue Art ist neben trimaculatus Wittmer zu stellen. Die von Champion (1922) als
myrmecodes Boheman bestimmten und beschriebenen 3 2 mit Fundorten “mouth of Umkomaas
River” und Seymour” gehdren zu amatolensis.

89

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 274-277. Sphinginopalpus s. sir. amatolensis sp. nov. 21 A, Fuhlcrglieiler 1-4; 275, Tergit 8; 276, Sternit 8; 277,

Kopulationsapparat (dorsal). Massstab von 274 auch fiir 276, 277.

44. Sphinginopalpus s. str. nelshoogtensis sp. nov.

MAnnchen (Abb. 278-283).

Lange 2,8 mm.

Kopf mil den Wangen schwarz, neben jedem Auge ein ganz kleiner, verschwommener heller
Flecken; Maxillarpalpen schwarzlich; Fiihler dunkel, died 1 auf der Unterseite und 2 bis 5 ganz hell;
Halsschild schwarz, Basis nicht ganz bis zur Halfte der abgeflachten Basis gelbbraun; Schildchen
und Fliigeldecken schwarz, letztere mil einem gelblichweissen, antimedianen Querband, das an der
Naht unterbrochen ist, seitlich beidseitig ein wenig verliingert; die 4 hinteren Beine schwarz, Tarsen
schwach aufgehellt, Vorderbeine dunkel, Schenkel gegen das Knie und Tarsen aufgehellt.

90

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 278-279. Sphinginopalpus s. sir. nelsiwogiensis sp. nov. <S : 278-279, Maxillarpalpen(x200) (278,
von oben; 279, von unten).

Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberflache glatt.
Maxillarpalpen Abb. 278-279, Fiihler (Abb. 280) um ca. 1/5 kiirzer als die Fliigeldecken, Glied 3
deutlich langer als 4. Halsschild breiter als lang (20x15,5), sonst wie bei majeri; auf dem abgeflachten
Teil jederseits neben dem Seitenrand eine feine Liingsrippe (bei majeri nicht mit Sicherheit
erkennbar). Fliigeldecken langoval; falsche Epipleuren ca. 70% der Lange einnehmend, keine
Krenulierung; Punktreihen besonders in der Mitte regelmassig, an der Basis und hinter der Mitte
Punkte verworren, gegen die Spitze immer mehr erloschend. Tergit 8 Abb. 281. Sternit 8 Abb. 282.
Kopulationsapparat Abb. 283 dorsal.

Holotypus (TMP): South Africa, E Transvaal : Nelshoogte, grassnetting, 25°47'S, 30°49'E, 8.iv.l987,
E-Y:2456, S. Endrody- Younga.

Diese neue Art is leicht an der Farbung des Kopfes und der Fliigeldecken zu erkennen, bei
letzteren reicht das antimech ane, helle Querband bis fast zur Naht. Das Fuhlerglied 1 hat einen breiten,
stumpfen Zahn.

91

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 280-284. 280-283. Sphinginopalpus s. str. nelshoogiensis sp. nov. 6 : 280, Fiihlerglieder 1-4; 281, Tergit 8; 282, Sternil
8; 283, Kopulationsapparat (dorsal). Massstab von 280 auch fiir 282, 283. 284. Sphinginopalpus s. str. nelshoogiensis
transformatus ssp. nov. 6 '■ Fiihlerglieder 1-4.

92

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

45. Sphinginopalpus s. str. nelshoogtensis transformatiis ssp. nov.

MAnnchen (Abb. 284).

Lange 2,3 mm.

Wie die Nominatform gefarbt, nur der Flecken neben jedem Auge ist grosser, er zieht sich schmal
gegen die Fiihlerwurzeln und um diese herum, seitiich auch ein wenig auf die Wangen; das Fiihlerglied
1 ist nur an der Basis kurz und schwach dunkler und die Maxillarpalpen sind gelb anstatt schwarzlich.

Die Maxillarpalpen sind ahnlich gebaut wie bei der Nominatform, doch bei den Fiihlern (Abb.
284) ist das Glied 1 sehr verschieden. Aufgrund dieser Merkmale konnte man transformatiis wohl
als eigene Art anerkennen, wenn die Terminalia und der Kopulationsapparat nicht so ausserordentlich
ahnlich waren. Deshalb mocht ich transformatiis nur als Unterart von nelshoogtensis beschreiben.

Holotypus (NHMB): South Africa, Natal: Manguzi Forest, Kosi, 26°58'S, 32°44"E, 10.i.l985, edge
of swamp forest, P. E. Reavell.

46. Sphinginopalpus s. str. quadratipalpus sp. nov.

MAnnchen (Abb. 285-290).

Lange 2,3-2,4 mm.

Kopf schwarz mit 3 gelben Flecken, je einem kleinen neben den Augen und einem grossen am
Vorderkopf, die Wangen nicht einschliessend, oder Flecken neben den Augen fehlen (Fundort E-
Y:2584); Maxillarpalpen letztes Glied schwarzlich, vorletztes teilweise stark gelblich aufgehellt;
Fiihlerglieder 1 bis 4, oder 1 bis 6-7 gelb, Oberseite von 1 schwarz und die letzten dunkel; Halsschild
schwarz. Basis mehr oder weniger breit gelbbraun; Schildchen und Fliigeldecken schwarz, letztere

mit einem breiten, weisslichen,
antimedianen Querband, das
an der Naht unterbrochen ist,
an den Seiten verbreitert;
Hinterbeine schwarz, oder die
Hintertarsen schwach

aufgehellt, 4 vordere Tarsen
gelb, 4 Vorderbeine schwarz,
Tibien an der Basis und Spitze
der Schenkel etwas aufgehellt.

Kopf mit den Augen
breiter als der Halsschild,
Stirne leicht gewolbt;
Oberflache chagriniert.

Maxillarpalpen Abb. 285-286.
Fiihler (Abb. 287) um ca. 1/4
kiirzer als die Fliigeldecken,
Glied 3 deutlich langer als 4.

Abb. 285-286. Sphinginopalpus s. str. quadratipalpus sp. nov. cJ : 285-286,
Maxillarpalpen (x200) (285, von oben; 286, von unten).

93

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 287-290. Sphinginopalpus s. str. quadraiipalpus sp. nov. cj : 287 Fiihlerglieder 1-4; 288, Tergit 8; 289, Stemit 8; 290,
Kopulationsapparat (dorsal). Massstab gilt fiir 287-290.

Halsschild langer als breit (18x15,5), Seiten von der Mitte nach vorne fast gerade, kaum merklich
gerundet,leichtverengt; aufgewdlbterTeil glatt,abgeflachterTeilchagriniert.FIugeldeckenlangoval,
falsche Epipleuren ca. 60% der Lange einnehmend, Kienulierung nur angedeutet, kaum sichtbar;
Punkte grob, bis zur Mitte und zum Teil darilbei hinaus in Reihen angeordnet, Punkte nach hinten
mehr erldschend. Tergit 8 Abb. 288. Sternit 8 Abb. 289.

Kopulationsapparat Abb. 290 dorsal.

94

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Holotypus (TMP): South Africa, Transkei: Fraser Gorge, grass tussocks, 31°24'S, 29°42'E,
27. xi. 1988, E-Y:2584, S. Endrody-Younga. Natal Middl.: Karkloof, grassveld, 29°19"S, 30°15"E,
10.xii.l989, E- Y:2758, S. Endrody-Younga, 1 Paratypus (NHMB).

Eine durch die Form der Maxillarpalpen charakterisierte Art, deren Glied 3 distal fast quadratisch
ist; auch bei dieser Art ist das helle, antimediane Querband erst kurz vor der Naht unterbrochen.
Weitere Unterschiede zeigt der Bau von Fiihlerglied 1 und die Terminalia.

47. Sphinginopalpus s. str. trimaculatus sp. nov.

Mannchen (Abb. 291-297).

Lange 2,3-2,5 mm.

Kopf Schwarz mit 3 gelben Flecken, der grosste am Vorderkopf und je ein weiterer neben jedem
Auge, diese beiden manchmal nur angedeutet, Wangen schwarz, der Flecken am Vorderkopf
manchmal stark reduziert, sodass nur ein schmales gelbes Band ganz vorne verbleibt; Maxillarpalpen
vorwiegend schwarz; Fiihlerglieder 1 bis 5 gelb, oft auch 6 teilweise, 7 bis 11 und Oberseite von 1,
dieses teilweise, schwarz; Halsschild schwarz, abgeflachte Basis grdsstenteils gelbbraun; Schildchen
und Fliigeldecken schwarz, letztere jederseits vor der Mitte mit einem weisslichen, fast dreieckigen
Flecken, der die Naht nicht erreicht; Beine schwarz, die vier vorderen ein wenig aufgehellt, die 4
vorderen Tarsen starker aufgehellt.

Abb. 291-292. Sphinginopalpus s. str. trimaculatus sp. nov. <5: 291-292, Maxillarpalpen
(xl70) (291, von oben; 292, von unten).

Kopf mit den Augen praktisch so breit wie der Halsschild, Stirne leicht gewolbt; Oberflache
feinstens chagriniert. Maxillarpalpen Abb. 291-292. Fiihler (Abb. 293) um ca. 1/5 kiirzer als die
Fliigeldecken, Glied 1 beidseitig schwach, fast regelmassig gerundet, Zahn klein, schmal, 3 und 4

95

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

parallel, 3 deutlich langer als 4. Halsschild langer als breit (20x17), wie capensis gebaut; Oberflache
auf der Scheibe glatt, an den Seiten und abgeflachte Basis chagriniert, einzelne Punkte, zum Teil
ziemlich stark erloschen, vorhanden. Fliigeldecken langoval, sonst wie bei capensis, aber starker
punktiert, falsche Epipleuren ca. 60% der Lange einnehmend, Krenulierung sehr fein, nur angedeutet;
lange aufstehende Haare vereinzelt, Tergit 8 Abb. 294. Sternit 8 Abb. 295.

Kopulationsapparat Abb. 296 dorsal. Abb. 297 Innensack leicht verschoben.

Abb. 293-297. Sphinginopalpus s. str. irimaculatus sp. nov. 6 : 293, Fiihlerglieder 1-4; 294, Tergit 8; 295, Sternit 8; 296-297,
Kopulationsapparat (296, dorsal; 297, mit leicht verschobenem Iimensack). Massstab von 293 auch fiir 295-297.

WEIBCHEN.

Kopf einfarbig schwarz, sonst wie das 6 gefarbt. Schulterbeulen stark zuriickgebildet, gerundet;
Krenulierung der falschen Epipleuren wie beim S .

96

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA; MALACHIIDAE)

Holotypus und 5 Paratypen (TMP), 5 Paratypen (NHMB), 2 Paratypen (AMG): South Africa, Natal;
Weza, Impetyene, grassveld, intersept, trap, 30°37'S, 29°42'E, 16.xi.l989, E-Y:2680, S. Endrody-
Younga, I. Klimaszewski; Bergville Distr., Indumeni for., humus, x.1960, N. Leleup, 1 Paratypus
(TMP); Karkloof grassveld, 29°19'S, 30°15T, 7.xii.l989, E-Y:2747, 2756, 2758, S. Endrody-
Younga, 2 Paratypen (TMP), 1 Paratypus (NHMB); Cathedral Peak, Drakensberg, 6.xi.l981, J. & S.
Klapperich, 3 Paratypen (NHMB). E Transvaal: Uitsoek, high altitude grassveld, 25°15"S, 30°34"E,
6.ii.l987, E-Y:2426, S. Endrody-Younga, 2 Paratypen (TMP), 1 Paratypus (NHMB); Berlin, gorge
edge, 24°32'S, 30°44"E, 4.ii.l987, E-Y:2413, S. Endrody-Younga, 2 Paratypen (TMP); Nelshoogte
Forest Station, 25°47^S, 30°50'E, 4.xii. 1987, E-Y:2336, 2340, 2350, 2456, S. Endrody-Younga, 6
Paratypen (TMP), 3 Paratypen (NHMB). Transkei: Dwesa coast, grassland, 32°17'S, 28°51"E,
28. ii. 1985, E-Y:2175, S. Endrody-Younga, 1 Paratypus (TMP). E Cape Prov.; Katberg, 4000 ft,
X.1932, R. E. Turner, 1 Paratypus (BM).

Eine hiibsche kleine Art mit 3 hellen, kleinen Flecken auf dem Vorderkopf, sonst verweise ich
auf die verschiedenen Abbildungen.

48. Sphinginopalpus s. str. tridensmimus sp. nov.

MAnnchen (Abb. 298-303).

Lange 2,3-2,5 mm.

Kopf schwarz, Vorderkopf
mit einer gelben Makel, die die
Wangen nicht erreicht, neben
jedem Auge eine kleine gelbe
Makel; Maxillarpalpen schwarz;
Fiihler schwarz, erste 4 Glieder
und Basis von 5 gelb, 1 an der
Basis kurz dunkel; Halsschild
schwarz. Basis schmal leicht
aufgehellt; Schildchen und
Fliigeldecken schwarz, letztere
mit einem schwachen, seitlichen,
antimedianen Saum, der nur die
erste Punktreihe erreicht; Beine
schwarz.

Kopf mit den Augen breiter
als der Halsschild, Stirne leicht
gewolbt; Oberflache glatt.
Maxillarpalpen Abb. 298-299. Fiihler (Abb. 300) um ca. 1/5 kiirzer als die Fliigeldecken, Glied 1 mit
3 Zahnen, von denen der eine, obere, gegeniiber dem basalen, bei der Ansicht von vorne nicht, oder
kaum sichtbar ist (siehe Abbildung), von oben gesehen, ist ein kleiner stumpier Zahn sichtbar, 3
deutlich langer als 4. Halsschild langer als breit (19x16); Seiten von der Mitte nach vorne praktisch
parallel; Oberflache glatt, nur an den Seiten des abgeflachten Teils fein chagriniert. Fliigeldecken
langoval; falsche Epipleuren ca. 60% der Lange einnehmend, Krenulierung nur angedeutet;

Abb. 298-299. Sphinginopalpus s. str. tridensmimus sp. nov. 6 : 298-299,
Maxillarpalpen (xl70) (298, von oben; 299, von unten).

97

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 300-303. Sphinginopalpus s. sir. tridensmimus sp. nov. S'. 300, Fuhlerglieder 1-4; 301, Tergit 8; 302, Sternit 8; 303,
Kopulationsapparat (dorsal). Massslab von 300 auch fur 302, 303.

Punktreihen bis zur Mitte deutlich, Punkte dann weiter auseinander stehend, weniger tief. Tergit 8
Abb. 301. Sternit 8 Abb. 302.

Kopulationsapparat Abb. 303 dorsal.

Holotypus (NHMB): South Africa, Natal: Golden Gate Park, Drakensberg, 6.xi.l981, J. & S.
Klapperich.

Der Bau von Fiihlerglied 1 erinnert an tridens Wittmer und die glatten Fliigeldecken an
politipennis Wittmer, doch sind letztere bei der neuen Art ein wenig starker und dichter punktiert.
Weitere Unterschiede liefern die Terminalia und der Kopulationsapparat.

98

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

49. Sphinginopalpus s. str. longidens Champion

Sphinginopalpus longidens Champion, 1922: 329.
MAnnchen (Abb. 304-310).

Abb. 304-305. Sphinginopalpus s. str. longidens Champion d ; 304-305, Maxillarpalpcn
(x220) (304, von oben; 305, von unten).

Im BM befinden sich der Lectotypus und 7 Paralectotypen, im SAM 4 Paralectotypen.

Fiir die Abbildungen wurde der Lectotypus verwendet: Maxillarpalpen Abb. 304-305.
Fiihlerglieder 1 bis 4 (Abb. 306). Champion gibt in seiner Beschreibung an, dass das died 3 langer
ist als 4, erwahnt aber nicht, dass es auch ein wenig breiter ist als 2 und 4. Tergit 8 Abb. 307. Sternit
8 Abb. 308. Kopulationsapparat Abb. 309 dorsal. Abb. 310 Profil.

99

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Kopulationsapparat (309, dorsal; 310, Profil), Massstab von 306 auch fiir 308-310,

50. Sphinginopalpus s. str. bidens Champion

Sphinginopalpus bidens Champion, 1922: 328, T. 5, figs 20, 20a.

MAnnchen (Abb. 311-317).

Der Holotypus befindet sich im BM, er stammt aus Frere (Natal) und diente zur Anfertigung
folgender Abbildungen: Maxillarpalpen Abb. 311-312. Fiihlerglieder 1 bis 4 Abb. 313. Tergit 8 Abb.
314. Sternit 8 Abb. 315. Kopulationsapparat Abb. 316 dorsal. Abb. 317 Profil.

100

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 311-312. Sphinginopalpus s.str. bidens Champion 6 : 311-312, MaxillaqDalpen (x220) (311,
von oben; 312, von unten).

Eingesehenes Material: Transvaal: Pretoria, 20. xi. 1981, J. & S. Klapperich, 1 S (NHMB);
Blydepoort, 20.xi.l981, J. & S. Klapperich, 3 S (NHMB); 21/22 km S Lydenburg, 15. xi. 1983, W.
Witlmer, 1 S (NHMB); Irene, 20.ii.l989, van Viegen (TMP); Middelburg, Cycas Farm, grassnetting,
25°31'S, 29°16'E, 5.xi.l985, E-Y:2262, S. Endrody- Younga, 2 3 (TMP), 1 3 (NHMB). Transkei:
S coast Dwesa Forest Station, 32° ITS, 28°50"E, 16.xii.l979, 26. ii. 1985, E- Y : 1687, 2168, S. Endrody-
Younga, 3 c? $ (TMP), 1 3 ? (NHMB).

Die gelblichweissen seitlichen Langsbander sind beim Holotypus hinter den Schultern nur wenig
erweitert. Bei den Exemplaren von den verschiedenen Fundorten linden sich solche mit dem hellen
Seitensaum, wahrenddem die Flecken stark gegen die Naht erweitert sind.

Weibchen.

Kopf einfarbig schwarz.

101

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 313-317. Sphinginopalpus s.str. bidens Champion 6‘. 313, Fuhlerglieder 1-4; 314, Tergit 8; 315, Stemit 8; 316-317,
Kopulationsapparat (316, dorsal; 317, Profil). Massstab von 314 auch fiir 315 und 313 auch fiir 316, 317.

51. Sphinginopalpus s. str. dwesaensis sp. nov.

MANNCHEN (Abb. 318-324).

Lange 2,5 mm.

Kopf inklusive der Wangen schwarz, von der Mitte der Augen nach vorne gelb, oder die gelbe
Farbe liegt zwischen den Fiihlerwurzeln nach vorne und neben jedem Auge befindet sich ein kleiner,
verschwommener, gelber Flecken; Maxillarpalpen braun bis schwarzlich; Fiihler schwarz, Glieder 1
bis 5 Oder 6 gelb, 1 mehr oder weniger breit schwarz auf der Oberseite; Halsschild schwarz, der ganze
abgeflachte Teil ist gelbbraun; Schildchen und Fliigeldecken schwarz; letztere mit einem weisslichen
Seitenflecken, der wenig nach vorne, jedoch lang und schmal nach hinten verlangert ist; Hinterbeine
schwarz, Hinterschienen an der Basis kurz aufgehellt, Hintertarsen gelb, 4 Vorderschenkel
schwarzlich, oder mehr oder weniger aufgehellt, 4 Vordertibien und 4 Vordertarsen gelb.

102

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA; MALACHIIDAE)

Abb. 318-320. Sphinginopalpus s. str. dwesaensis sp. nov. <J : 318-319, Maxillarpalpen (xl30) (318,
vonoben; 319, von unten); 320, Labialpalpen und Maxille (x550).

Kopf mil den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberflache glatt.
Maxillarpalpen Abb. 318-319, Labialpalpen und Maxille Abb. 320. Fiihler (Abb. 321) urn 1/5 kiirzer
als die Fliigeldecken, Glieder 3 bis 10 fast parallel, 3 nur wenig langer als 4. Halsschild langer als
breit (17,5x15), sonst wie bei capensis, jedoch Oberflache glatt, nur die abgeflachte Basis chagriniert.
Fliigeldecken langoval, wie bei capensis, etwas starker punktiert; falsche Epipleuren ca. 80% der
Lange einnehmend, nicht krenuliert. Tergit 8 Abb. 322. Sternit 8 Abb. 323.

Kopulationsapparat Abb. 324 dorsal.

Holotypus und 2 Paratypen (TMP): Transkei: Dwesa Forest, coast, grassnetting, dusk, 32°17"S,
28°50"E, 26.ii.1985, E-Y:2168, S. Endrody-Younga. Transvaal: Irene, iii.1990, van Viegen, 1
Paratypus (NHMB); Rivonia, 23.-27.X.1968, M. 1. Russell, 2 Paratypen (BM), 1 Paratypus (NHMB).

Diese neue Art ist neben nigriceps Wittmer zu stellen, von der sie sich hauptsachlich durch den
verschiedenen Bau der Terminalia und des Kopulationsapparates unterscheidet.

103

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 321-324. Sphinginopalpus s. str. dwesaensis sp. nov. d: 321, Fiihierglieder 1-4; 322, Tergit 8; 323, Stemit 8; 324,
Kopulationsapparat (dorsal). Massstab von 321 auch fiir 323, 324.

104

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA; MALACHIIDAE)

52. Sphinginopalpus s. str. nigriceps sp. nov.

MAnnchen (Abb. 325-330).

Lange 2, 3-2, 5 mm.

Kopf schwarz; Maxillarpalpen schwarz, oder teilweise leicht aufgehellt; Fiihler schwarzlich,

Glieder 1 bis 5 oder 6 gelb, 1 und

2 auf der Oberseite basal oder
ganz dunkel; Halsschild schwarz;
Fliigeldecken mil einem weiss-
lichen, dreieckigen Flecken
jederseits, die Naht nicht er-
reichend, an den Seiten beidseitig
verlangert; Flinterbeine schwarz
bis dunkelbraun, 4 Vorder-
schenkel mehr oder weniger
angedunkelt, 4 Vorderbeine oft
ganz gelb, alle Tarsen gelb.

Kopf mit den Augen breiter
als der Flalsschild, Stirne leicht
gewolbt; Oberflache glatt.
Maxillarpalpen Abb. 325-326.
Fiihler (Abb. 327) um ca. 1/5
kiirzer als die Fliigeldecken, Glied

3 nur ganz wenig langer als 4, 2
bis 10 zur Spitze sehr wenig verbreitert. Halsschild langer als breit (19x15), sonst mit caperisis
iibereinstimmend, jedoch aufgewblbter Teil glatt, abgeflachter chagriniert. Fliigeldecken wie bei
capensis, jedoch falsche Epipleuren ca. 60% der Lange der Decken einnehmend, kaum merklich
krenuliert, nur wenige lange, helle, aufstehende Haare vorhanden. Tergit 8 Abb. 328. Sternit 8 Abb.

Abb. 325-326. Sphinginopalpus .s. .str. nigriceps sp. nov. 6
Maxillarpalpen (x200) (325, von oben; 326, von unten).

325-326,

329.

Kopulationsapparat Abb. 330 dorsal.

Holotypus (TMP), 1 Paratypus (NHMB): South Africa: Transvaal: Nelshoogte, Knuckles grassveld,
grassnetting, 25°4TS, 30°49"E, 24.x. 1986, 4.xii.l986, E-Y:2312, 2350, S. Endrody-Younga; Gods
Window, grassnetting, 24°35"S, 30°51'E, 12. xi. 1980, E-Y:1734, S. Endrody-Younga, 1 Paratypus
(TMP), 1 Paratypus (NHMB); Uitsoek, high altitude grassveld, grassnetting, 25°15'S, 30°34"E, 1500
m, 12.xii.l986, 6.ii.l987, E-Y:2381, 2426, S. Endrody-Younga, 1 Paratypus (TMP), 1 Paratypus
(NHMB).

Eine der wenigen Arten bei denen auch die $ einen einfarbig schwarzen Kopf besitzen. Das
Fiihlerglied 1 erirmert an nelshoogtensis Wittmer, neben den diese neue Art zu stellen ist.

105

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 327-33U. Sphinginopalpus s. sir. nigriceps sp. nov. S : 327, Fuhlerglieder 1-4; 328, Tergit 8; 329, Sternit 8; 330,
Kopulationsapparat (dorsal). Massstab von 327 auch fiir 329, 330.

53. Sphinginopalpus s. str. uitsoekensis sp. nov.

MANNCHEN (Abb. 331-336).

Lange 2,5 mm.

Kopf schwarz; Maxillarpalpen vorwiegend schwarz, teils etwas aufgehellt; Fiihler gelb, Oberseite
der Glieder 1 und 2 und 7 bis 1 1 schwarz; Halsschild schwarz, abgeflachter Teil gelbbraun; Schildchen
und Fliigeldecken schwarz; letztere mil einem weisslichen, leicht langsdreieckigen, seitlichen
Flecken, dessen Spitze gegen die Naht gerichtet ist, ohne sie zu erreichen, er ist nach hinten etwas
mehr als nach vorne verliingert, Totallange wenig mehr als die Halite der Decken; Schenkel schwarz,
die 4 vorderen gegen die Knie ein wenig aufgehellt, Hinterschienen schwarz, die mittleren auf der

106

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 331-332. Sphinginopalpus s. sir. uitsoekensissp. nov. S : 331-332, Maxillaqjalpen
(xl70).

unteren Hiilfte, die vorderen fast ganz hell, Tarsen gelb, nur die hinteren leicht angedunkelt.

Kopf mit den Augen nur wenig breiter als der Halsschild, Stirne leicht gewblbt; Oberflache glatt.
Maxillarpalpen Abb. 331-332. Fiihler (Abb. 333) um ca. 1/4 kiirzer als die Fliigeldecken, Glieder 3
und 4 gleich lang. Halsschild liinger als breit (17x15), sonst wie bei elongatideiis, aufgewblbter Teil
glatt, abgeflachter sehr schwach chagriniert. Fliigeldecken langoval, falsche Epipleuren ca. 60% der
Lange einnehmend, nicht kienuliert; Punktreihen von unterschiedlicher Regelmassigkeit, Punkte auf
den letzten 1/3 fast verschwindend. Tergit 8 Abb. 334. Sternit 8 Abb. 335.

Kopulationsapparat Abb. 336 dorsal.

Weibchen.

Wie das S gefarbt, bei einem Exemplar sind die letzten Fiihlerglieder aufgehellt und die dunkle
Farbung auf 1 und 2 fehlt.

Holotypus S und 2 Paratypen ? (TMP), 1 Paratypus $ (NHMB); South Africa, E Transvaal; Uitsoek
Forest Station, 1100 m, grassnetting, 25°18'S, 30°55'E, 13.xii.l986, E-Y:2384, S. Endrody-Younga.

Neben nigriceps Wittmer zu stellen, mit dem diese neue Art nahe verwandt ist, beide haben einen
schwarzen Kopf und ein ahnlich gebautes Fiihlerglied 1, jedoch sehr verschiedene Terminalia.

107

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 333-336. Sphinginopalpus s. str. uiisoekensis sp. nov. <3: 333, Fuhlerglicder 1-4; 334, Tergit 8; 335, Sternit 8; 336,
Kopulationsapparat (dorsal), Massstab von 333 auch fiir 335, 336.

108

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

54. Sphinginopalpus s. str. curvispinus sp. nov.

Mannchen (Abb. 337-341).

Lange knapp 3 mm.

Kopf inklusive der Wangen schwarz, von der Mitte der
Augen nach vorne gelb; Maxillarpalpen schwarz; Fiihler
schwarz, Glieder 1 bis 4 gelb, 1 und 2 auf der Oberseite teils
ganz schwach angedunkelt; Halsschild schwarz. Basis
schmal aufgehellt; Schildchen und Fliigeldecken schwarz,
letztere jederseits mit einem dreieckigen, antimedianen,
weisslichen Flecken, der sich gegen die Naht ausdehnt, ohne
sie zu erreichen; Beine schwarz.

Kopf mit den Augen breiter als der Halsschild, Stirne
ziemlich flach; Vorderstirne leicht gewolbt und erhoht,
gegen die Stirne durch einen leicht gebogenen Wall von der
Mitte der Augen an abgegrenzt, der die Grenze bildet
zwischen der schwarzen und der gelben Farbung.
Maxillarpalpen Abb. 337. Fiihler (Abb. 338) um ca. 1/5
kiirzer als die Fliigeldecken, Zahn an Glied 1 sehr lang,
Spitze leicht gebogen, 3 kaum merklich langer als 4.
Halsschild langer als breit (21x17), sonst wie bei
tridensmimus, nur dass der abgeflachte Teil chagriniert ist.
Fliigeldecken langoval; falsche Epipleuren ca. 55% der
Lange eitmehmend, Krenulierung kaum wahrnehmbar;
Punktierung ahnlich wie elongatidens. Tergit 8 Abb. 339.
Sternit 8 Abb. 340.

Kopulationsapparat Abb. 341 dorsal.

Holotypus (NHMB): South Africa, Natal: Elandskop, ca. 40 km W Pietermaritzburg, 2930 CA,
10.ix.l979, rocky ridge, grassland and proteas, J. Londt.

Diese neue Art zeichnet sich durch den besonderen Bau des Kopfes und den langen, leicht
gebogenen Zahn an Fiihlerglied 1 aus. Sie ist neben longidens Wittmer zu stellen.

Abb. 337. Sphinginopalpus s. str. curvispinus
sp. nov. 8 : 337, Maxillarpalpus
(xl70) (von unten).

109

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 338-341. Sphinginopalpus s. str. curvispinus sp. nov. d; 338, Fuhlerglieder 1-4; 339, Tcrgit 8; 340, Sternit 8; 341,
Kopulationsapparat (dorsal). Massstab von 338 auch fiir 339-341.

55. Sphinginopalpus s. str. acutispinus sp. nov.

Mannchen (Abb. 342-347).

Lange 2mm.

Kopf inklusive der Wangen schwarz, von der Mitte der Augen nach vorne gelb; Maxillarpalpen
gelbbraun, gegen die Spitzen der Glieder 1 und 2 augedunkelt; Fiihler dunkel, basale 5 Glieder gelb;

110

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 342-343. Sphinginopalpus s. str. acutispinus sp. nov. S'. 342-343,
Maxillarpalpen (xl70) (342, von oben; 343, von unten).

glatt, Chagrinierung kaum angedeutet. Maxillarpalpen Abb. 342-

Halsschild schwarz, ein Teil der
abgesetzten Basis aufgehellt;
Schildchen und Fliigeldecken
schwarz, letztere jederseits mit
einem antimedianen,

dreieckigen, weisslichen

Flecken; Hinterbeine schwarz
mit leicht helleren Tarsen, 4
vordere Schenkel dunkel, gegen
die Knie kurz aufgehellt, vordere
4 Tibien und Tarsen hellbraun
Oder gelblich.

Kopf mit den Augen ein
wenig breiter als der Halsschild,
Stirne leicht gewolbt; Oberflache
343. Fiihler (Abb. 344) um ca. 1/5

Abb. 344-347. Sphinginopalpus s. sir. acuiispinus sp. nov. S : 344, Fiihlerglicder 1-4, 345, Tergit 8; 346, Sternit 8; 347,
Kopulationsapparat (dorsal). Massstab von 344 auch fiir 346, 347.

Ill

.\NN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

kiirzer als die Fliigeldecken, Zahn auf died 1 besonders lang und schmal, 3 kaum merklich langer
als 4. Halsschild langoval; falsche Epipleuren ca. 60% der Lange einnehmend, Kienulierung
angedeutet; Punktierung ahnlich wie bei oculimaculatus. Tergit 8 Abb. 345, Sternit 8 Abb. 346.
Kopulationsapparat Abb. 347 dorsal.

Holotypus (TMP): South Africa, Cape Prov.: Amatole, Isidenge Forest Station, grass tussocks,
32°41'S, 27°15T, 18.xi.l987, E-Y:2524, S. Endrody-Younga.

Eine weitere neue Art mit langem Zahn auf Fiihlerglied 1, die jedoch hinreichend durch die
verschiedene Form des Zahnes, die Terminalia und den Kopulationsapparat von den anderen
differenziert ist.

56. Sphinginopalpus s. str. majeri sp. nov.

Kopf Schwarz mit einem
fast viereckigen gelben
Flecken auf dem Vorderkopf,
der knapp die Mitte der
Augen erreicht, ohne sie zu
beriihren; Maxillarpalpen
braun; Fiihler schwarz, died
1 auf der Unterseite und 2 bis
5 ganz gelb; Halsschild
schwarz, ganzer abgeflachter
Teil gelbbraun; Schildchen
und Fliigeldecken schwarz,
letztere jederseits mit einem
antimedianen, weisslichen
Flecken, der die Naht nicht
erreicht und an den Seiten
etwas mehr nach hinten als
nach vorne verlangert ist. Hinterbeine schwarz, Hinterschienen gegen die Knie und Tarsen aufgehellt;
die 4 Vorderbeine gelb mit den Schenkeln zum grossten Teil schwarz.

Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberflache glatt.
Maxillarpalpen Abb. 348-349. Fiihler (Abb. 350) um ca. 1/5 kiirzer als die Fliigeldecken, died 1
nach dem Zahn nur wenig gegen die Spitze verjungt, fast parallel, Zahn fast quadratisch, 3 nur wenig
langer als 4. Halsschild langer als breit (18x15), Seiten auf dem vorderen Teil nur wenig nach vorne
gerundet verengt; Oberflache auf dem aufgewolbten Teil glatt, abgeflachte Basis fein chagriniert.
Fliigeldecken langoval, falsche Epipleuren ca. 65% der Lange eiimehmend, nicht krenuliert;
Punktreihen bis zur Mitte ziemlich regelmassig, grobe Punkte ziemlich dicht, dann sparlicher, immer
mehr erloschend, Spitze glatt. Tergit 8 Abb. 352. Spiculum Abb. 353.

Kopulationsapparat Abb. 354 dorsal. Abb. 355 Profil.

Mannchen (Abb. 348-355).

Lange 2-2,2 mm.

Abb. 348-349. Sphinginopalpus s. str. majeri sp. nov. 6 ; 348-349, Maxillarpalpen
(xl55) (348, von oben; 349, von unten).

112

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA; MALACHIIDAE)

Abb. 3S0-355. Sphinginopalpus s. str. majeri sp. nov. 6 : 350, Fuhlerglieder 1-4; 351, Tergii 8; 352, Stcrnit 8; 353, Spiculum;
354-355, Kopulationsapparat (354, dorsal; 355, Profil), Massslab von 351 auch fiir 352-355.

Weibchen.

Kopf einfarbig schwarz; Fiihlerglied 1 nur an der Basis kurz, leicht angedunkelt; Fliigeldecken
ohne Quermakel, Seitenbander etwas verbreitert; Punktierung weniger stark.

Holotypus S und 2 Paratypen (BM), 1 Paratypus $ (NHMB): Zimbabwe: Salisbury (jetzt Harare),
Mashonaland, MCH. (March ?), G. A . K. Marshall.

Es freut mich, diese Art meinem lieben Kollegen, Karel Majer, Brno, widmen zu diirfen, dem
ich die zahJreichen Abbildungen in dieser Publikation verdanke.

Diese Ait wurde von Champion (1927) als oneili Pic bestimmt. Die Maxillarpalpen zeigen wenig
Unterschiede zu oneili, unsomehr sind die Fiihler, die Terminalia und der Kopulationsapparat
verschieden.

113

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

57. Sphinginopalpus s. str. expansus sp nov.

Mannchen (Abb. 356-357).

Lange 2,7 mm.

Line ausserlich mil oculimaciilatiis Wittmer iibereinstimmende Ait, es eriibrigt sich, sie noch
einmal zu beschieiben. Auch die Terminalia stimmen weitgehend mit ociilimaculatus iiberein. Grosse
Unterschiede zeigen die Maxillarpalpen (Abb. 356-357), sodass expansus ohne Zweifel als eigene
Art gelten karm.

Abb. 356-357. Sphinginopalpus s. str. expansus sp. nov. i : Maxillarpalpen (xl70) (356, von
oben; 357, von unten).

Holotypus S (TMP): South Africa, E Transvaal: Nelshoogte Forest Station, S. Endrddy-Younga.
58. Sphinginopalpus s. str. oculiniaciilatus sp. nov.

Mannchen (Abb. 358-364).

Lange 2,7 mm.

Kopf schwarz mit einem kleinen gelblichen Flecken neben jedem Auge iiber der Fiihlerwurzel;
Maxillarpalpen schwarz; Fiihler schwarz, die ersten 5 Glieder gelb, 1, manchmal auch 2 ganz wenig
und 5 auf der Oberseite angedunkelt; Halsschild schwarz. Basis kurz gelbbraun; Schildchen und
Fliigeldecken schwarz, letztere mit einer antimedianen Quermakel, die an der Naht unterbrochen ist
und sich an den Seiten wenig ausdehnt; Hinterschenkel schwarz, die 4 vorderen braun, an der Basis
ein wenig dunkler, Hintertibien und Tarsen dunkel, die 4 vorderen Tarsen mehr oder weniger
aufgehellt.

114

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA; MALACHIIDAE)

Abb. 358-359. Sphinginopalpus s. str. oculiinaculatus sp. nov. cJ : Maxillarpalpen (xl55)
(358, von oben; 359, von unten).

Kopf mil den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberflache auf der Stirne
glatt, nach vorne schwach chagriniert. Maxillarpalpen Abb. 358-359. Fiihler (Abb. 360) um ca. 1/10
kiirzer als die Fliigeldecken, Glied 3 deutlich langer als breit (21x17); Oberflache glatt, abgeflachter
Teil chagriniert, sonst wie elongatidens. Fliigeldecken langoval; falsche Epipleuren ca. 70% der
Lange einnehmend, nicht krenuliert; Punktreihen nicht ganz regelmassig, Punkte nach hinten weniger
tief und auf dent letzten 1/4 fast ganz erloschen. Tergit 8 Abb. 361. Sternite 7 und 8 Abb. 362 und
363.

Kopulationsapparat Abb. 364 dorsal.

Weibchen.

In der grossen Serie von Nelshoogte befand sich kein $ , wahrscheinlich weil bis auf 2 Exemplare
alle am Auto abgelesen wurden, wo sie angeflogen kamen. Ein ? befand sich unter dem Material
aus Berlin, das ich jedoch wegen der abweichenden hellen Farbung der Beine und auch wegen den
etwas helleren Fliigeldecken nur mit Vorbehalt zu dieser Art stelle.

Holotypus und 162 Paratypen (TMP), 31 Paratypen (NHMB), 6 Paratypen (AMG): South Africa, E
Transvaal: Nelshoogte Forest Station, grassnetting, beating, but attracted to the car in big number,
25°47'S, 30°50'E, l.xii.l986, E-Y:2336, 2337, 2339, S. Endrody- Younga; Berlin Eorest Station,
1500 m, grassnetting 25°32'S, 30°44'E, E-Y:2360, 7.xii.l986, S. Endrody- Younga, 5 Paratypen
(TMP), 2 Paratypen (NHMB).

Neben acutispinus Wittmer zu stellen, dessen Fiihlerglied 1 sehr ahnlich gebaut ist, sonst aber
dutch die glatten, etwas weniger dicht punktierten Fliigeldecken und die Terminalia sehr verschieden.

115

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

3 6 2

Abb. 360-364. Sphinginopalpus s. str. oculimaculaius sp. nov. 3 : 360, Fiihlerglieder 1-4; 361, Tcrgit 8; 362, Sternit 7; 363,
Stemit 8; 364, Kopulationsapparat (dorsal). Massstab von 360 auch ftir 363, 364 und 361 auch fur 362.

116

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

59. Sphinginopalpus s. str. zulu sp. nov.

MANNCHEN (Abb. 365-370).

Lange 2,1 mm.

Kopfschwarz, Vorderkopfum dieFiihlerwurzelnundWangen
leicht aufgehellt; Maxillarpalpen gelbbraun; Fiihler gelb, die
letzten 5 bis 6 Glieder, sowie die Oberseite von 1 verschwommen
schwarzlich; Halsschild schwarz, ca. die Halfte des abgeflachten
Teils gelbbraun; Schildchen und Fliigeldecken schwarz, letztere
mit einem ziemlich breiten, weisslichen, antimedianen Querband,
das an der Naht unterbrochen und an den Seiten ein wenig
verlangert ist; Hinterbeine schwarz, Tarsen ein wenig aufgehellt,
Schenkel und Schienen der 4 vorderen Beine dunkel, teils
aufgehellt, die 4 vorderen Tarsen gelb, letztes Glied leicht
angedunkelt.

Kopf mit den Augen breiter als der Halsschild, Stirne leicht
gewdlbt; Oberflache glatt. Maxillarpalpus Abb. 365. Fiihler (Abb.
366) um ca. 1/5 kiirzer als die Fliigeldecken, Glied 3 deutlich langer
als 4, fast doppelt so lang wie 2. Halsschild langer als breit (18x15);
Oberflache glatt, abgeflachter Teil chagriniert, sonst wie
elongatidens. Fliigeldecken langoval; falsche Epipleuren ca. 70%
der Lange eirmehmend, ungefiihr in der Mitte, kurz, ganz fein krenuliert; Punktierung in Punktreihen,
die nicht ganz regelmassig sind, Punkte gegen die Spitze fast erloschend. Tergit 8 Abb. 367. Sternite
7 und 8 Abb. 368 und 369.

Kopulationsapparat Abb. 370 dorsal.

Holotypus (TMP): South Africa, Zululand: Lake Bhangasi, grassnetting, 28°07'S, 32°31'E,
18.xii.l976, E-Y:982, S. Endrody- Younga.

Diese neue Art is mit albilabris (Boheman) und variceps Wittmer verwandt, Fiihlerglied 1
ahnlich, Terminalia sehr verschieden, vergleiche Abbildungen.

Abb. 365. Sphinginopalpus s. str. zulu
sp. nov. i : 365,

Maxillarpalpus (xl70) (von
unten).

117

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT, 1 30 JANUARY 1994

60. Sphinginopalpus s. str. politus sp. nov.

Mannchen (Abb. 371-376).

Lange 2, 2-2, 3 mm.

Kopf schwarz, nur die Wangen schmal bis zu den Augen, seltener nur der Vorderkopf schma!
gelb; Maxillarpalpen schwarzlich; Fiihler schwarz, die ersten 5 Glieder gelb, mit der Basis von 1
schwarz, oft ist auch 6 an der Basis aufgehellt; Halsschild schwarz. Basis gelbbraun; Schildchen und

118

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 371-372. Sphinginopalpuss. stT.politussp. nov. S : 371-372, Maxillarpalpen (xl70) (371,
von oben; 372, von unten, Glieder 1 und 2 zuriickgelegt).

Fliigeldecken schwarz, letztere mil einem durchgehenden, gelben Querband vor der Mitte, das sich
an den Seiten verlangert; bei 1 Exemplar 1st das Querband stark reduziert und an der Naht
unterbrochen; 4 Hinterbeine schwarz, Spitzen der Mitteltibien mehr oder weniger aufgehellt, Tarsen
meistens ganz aufgehellt, Vorderschenkel angedunkelt, Vordertibien fast ganz hell.

Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberflache glatt, glanzend.
Maxillarpalpen Abb. 371-372. Fiihler (Abb. 373) um ca. 1/10 kirrzer als die Fliigeldecken, der kleine
Zahn von Glied 1 nahe der Basis ist stark reduziert, vom grosseren Zahn ab ist das Glied fast parallel,
3 ein wenig langer als 4, 4 nach innen schwach gerundet erweitert. Halsschild langer als breit (17x13);
Oberflache glatt, glanzend, nur der abgeflachte Teil schwach chagriniert, sonst mit capensis
iibereinstimmend. Fliigeldecken langoval, Krenulierung der falschen Epipleuren von variabler Starke,
manchmal so gering wie bie capensis; Punktierung wie bei diesem; Behaarung ebenso. Tergit 8 Abb.
374. Sternit 8 Abb. 375.

Kopulationsapparat Abb. 376 dorsal.

Weibchen.

Kopf einfarbig schwarz; Querband auf den Fliigeldecken meistens an der Naht unterbrochen.
Schulterbeulen nur wenig entwickelt; ungefliigelt.

Holotypus und 9 Paratypen (TMP), 4 Paratypen (NHMB): South Africa, SW Cape Prov.: Heuningnes
River, 34°42'S, 20°02'E, 28.x. 1983, E- Y :2028, 2029, S. Endrody-Younga; Clanwilliam-Elandsbaai,
16. ix. 1985, W. Wittmer, 1 Paratypus (NHMB); Verlorevlei farm, 32°19'S, 18°22"E, 28.viii.1981, E-
Y:1857, S. Endrody-Younga, 1 Paratypus (TMP), 1 Paratypus (NHMB).

Diese neue Art hat ein ahnlich paralleles Fiihlerglied 1 mit kleinem Zahn wie leiicosideae
Wittmer, jedoch sehr verschieden gebaute Maxillarpalpen und Terminalia.

119

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 373-376. SphinginopalpuS s. sir. poltnis sp. nov. cJ ; 373, Fiihlerglieder 1-4; 374, Tcrgil 8; 375, Sternit 8; 376,
Kopulationsapparat (dorsal). Massstab von 373 auch fiir 375, 376.

61. SphinginopalpuS s. str. levemaculatus sp. nov.

MAnnchen (Abb. 377-383).

Lange 2,1 mm.

Kopf Schwarz, iiber jeder Fiihlerwurzel ein kleiner, verschwommener, heller Flecken;
Maxillarpalpen schwarz; Fiihler schwarz, die ersten 4 Glieder und die Basis von 5 gelb, 1 auf der
Oberseite an der Basis angedunkelt; Halsschild schwarz, bei letzterem die Seiten, ungefahr in der

120

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 377-379. Sphinginopalpus s. str. levemaciilatus sp. nov. : 377-319, Maxillarpalpen
(377, von oben, xl70; 378, von unten, xl70; 379, died 2, x390).

Breite und Lange der falschen Epipleuren, weisslich; Hinterbeine schwarz, die 4 vorderen schwarzlich
mil den Spitzen der Tibien und Tarsen angedunkelt.

Kopf mil den Augen breiter als der Halsschild, Stirne leicht gewblbt; Oberflache glatt.
Maxillarpalpen Abb. 377-378. Fiihler (Abb. 380) ca. um 1/5 kiirzer als die Fliigeldecken, Glieder 3
bis 10 fast parallel, 3 deutlich langer als 4. Halsschild langer als breit (17x13,5), sonst wie capensis,
nur dass der aufgewolbte Teil glatt und der abgeflachte chagriniert ist. Fliigeldecken langoval,
ungefahr in der Mitte am breitesten; Schulterbeulen normal; falsche Fpipleuren ungefahr 50% der
Lange einnehmend, Krenulierung fehlt; Punkte wenig dicht und wenig tief, in Punktreihen; einzelne
lange, aufrechtstehende Haare vorhanden. Tergit 8 Abb. 381. Sternit 8 Abb. 382.
Kopulationsapparat Abb. 383 Profil.

Holotypus (NHMB): South Africa, Cape Prov.: 5 km E Humansdorp, 7.xi.l988, W. Wittmer, M. J.
D. Brendell.

Fine weitere Art mit langem Zahn auf Fuhlerglied 1, sie kann neben acutispinus Wittmer and
longispinus Wittmer gestelit werden, mit verschieden gebauten Terminalia und Kopulationsapparat.

121

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 380-383. Sphinginopalpus s. sir. levemaculatus sp. nov. <S : 380, Fuhlergliedcr 1-4; 381, Tergit 8; 382, Sternit 8; 383,
Kopulationsapparat (Profil). Mas.sstab von 380 auch fiir 382-383.

DIE GAITUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

62. Sphinginopalpus s. str. trispinus sp. nov,

MANNCHEN (Abb. 384-389).

Lange 2,2 mm.

Kopf schwarz, Wangen
teilweise gelb, ein kleiner
Flecken am Vorderkopf und je
ein kleiner Flecken neben den
Augen, gelb; Maxillarpalpen
schwarz; Fiihler schwarzlich,
Glieder 1 bis 5 gelb, 1 an der
Basis und 3 mit einem mehr oder
weniger grossen Flecken oben;
Halsschild schwarz. Basis fast
bis zu 1/3 des abgeflachten
Teiles gelbbraun; Schildchen
und Fliigeldecken schwarz,
letztere mit einem seitlichen,
antimedianen, weisslichen
Flecken, der die Naht nicht
erreicht, am Seitenrand am
breitesten; Hinterbeine schwarz,
Tarsen schwach heller, 4 Vorderschenkel schwarz, 4 Vorderschienen dunkel, an den Spitzen heller,
4 Vordertarsen gelb.

Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberfliiche fein chagriniert.
Maxillarpalpen Abb. 384-385. Fiihler (Abb. 386) um ca. 1/5 kiirzer als die Fliigeldecken, Glied 3
langer als 4, der dritte Zahn auf 1 ist nicht immer deutlich sichtbar (je nach Stellung). Halsschild
langer als breit (21x15), Seiten nach vorne kaum merklich verengt; Oberflache des aufgewolbten
Teiles fast glatt, oder fein chagriniert, abgeflachter Teil chagriniert, sonst wie elougatidens.
Fliigeldecken langoval, falsche Epipleuren ca. 65% der Liinge einnehmend, Krenulierung sehr fein;
sonst wie elougatidens. Tergit 8 Abb. 387. Sternit 8 Abb. 388.

Kopulationsapparat Abb. 389 dorsal.

Weibchen.

Kopf einfarbig schwarz; Fiihler ohne schwarzen Flecken auf Glied 3, sonst wie das 6 gefarbt.
Fliigeldecken mit starker gerundeten Seiten, Schulterbeulen nur angedeutet; ungefliigelt.

Holotypus and 5 Paratypen (TMP), 3 Paratypen (NHMB): South Africa, Natal Middld.: Karkloof
grassveld with river, 29°19'S, 30°15'E, 7.xii.l989, E-Y:2747, 2758, S. Endriidy-Younga, 1.
Klimaszewski. Cape Prov.: Amatole, Isidenge Forest Station, grassnetting, indigenous forest and
ground traps 25 days with faeces bait, 32°4 1 "S, 27° 14- 16 'E, E- Y :25 10, 25 1 1 , 25 1 4, 2524, S. Endrody-
Younga, 3 Paratypen (TMP), 2 Paratypen (NHMB); idem. 8.xii. 1987, E- Y :2559, S. Endrody- Younga,
netted and Arum flowers, 1 Paratypus (TMP).

Abb. 384-385. Sphinginopalpus s. str. trispinus sp. nov. cJ : 384-385,
Maxillarpalpen (xl70) (384, von oben; 385, von unten).

123

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Eine der wenigen Arten mil verbreitertem Fiihlerglied 3 und sehr breitem 1, das Tergit 8 lauft
in 3 Spitzen aus. Die neue Art is neben productiis Wittmer zu stellen, Fiihler und Sternit 8 sehr
verschieden gebaut.

Abb. 386-389. Sphinginopalpiis s. sir. trispinus sp. nov. 6 : 386, Fuhlerglieder 1-5; 387, Tergit 8; 388, Sternit 8; 389,
Kopulationsapparat (dorsal). Massstab von 386 auch fiir 388, 389.

124

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

63. Sphinginopalpus s. sir. productus sp. nov.

Mannchen (Abb. 390-395).

Lange knapp 2mm.

Kopf schwarz, von der Mitte der Augen nach vorne mil den Wangen gelb; Maxillarpalpen
vorwiegend gelb; Fiihler gelb, letzte 4 bis 5 Glieder schwarz; Halsschild schwarz, abgeflachter Teil
gelbbraun; Schrldchen und Fliigeldecken schwarz, letztere an den Seiten mil einem weisslichen,
langlichen Flecken, der ungefahr die Halfte der Lange einnimmt; Schenkel schwarz, die 4 vorderen
gegen die Knie kurz gelb, Schienen gelb, die hinteren angedunkelt, Tarsen gelb, die hinteren etwas
angedunkelt.

Kopf mil den Augen etwas
breiter als der Halsschild, Stirne
leicht gewolbt; Oberflache an der
Basis fein punktiert, dazwischen
chagriniert. Maxillarpalpen Abb.
390-391. Fuhler (Abb. 392) ca. 1/5
kiirzer als die Fliigeldecken, Glied 3
ein wenig langer als 4 (langer als auf
der Abbildung gezeichnet).
Halsschild wenig langer als breit
(15x13,5), sonst wie bei
elongatidens, aufgewolbter Teil glatt,
abgeflachter chagriniert.

Fliigeldecken langoval, wie
elongatidens, jedoch falsche
Epipleuren kiirzer, nur ca. 60% der
Lange einnehmend, nicht krenuliert
und Punkte auf dem letzten Drittel fast ganz erloschen. Tergit 8 Abb. 393. Sternit 8 Abb. 394.

Kopulationsapparat Abb. 395 dorsal.

Holotypus und 1 Paratypus (TMP), 2 Paratypen (NHMB): South Africa, Natal: Weza, Impetyene
grassland, grass tussocks, 30°37'S, 29°42'E, 25. xi. 1989, E-Y:2718, S. Endrddy-Younga.

Diese neue Art is neben trispinus Wittmer zu stellen, sie hat ein ahnlich gebautes Tergit 8, der
Kopulationsapparat ist von einer einfacheren Bauart, die iibrigen Merkmale wie Fiihlerglied 1,
Maxillarpalpen, Sternit 8 und der Kopulationsapparat sind sehr vergchieden.

Abb. 390-391. Sphinginopalpus s. str. productus sp. nov. S: 390-391,
Maxillarpalpen (xl70) (390, von oben; 391, von unten).

125

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 392-395. Sphinginopalpus s. str. productus sp. nov. 3\ 392, Fiihlerglieder 1-4; 393, Tergit 8; 394, Stemit 8; 395,
Kopulationsapparat (dorsal). Massstab von 392 auch fiir 394, 395.

126

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

64. Sphinginopalpus s. str. rudebecki Wittmer

Sphinginopalpus rudebecki Wittmer, 1960: 117, figs 4-5.

Mannchen (Abb. 396-401).

Die Art ist seit ihrer Entdeckung nicht wieder gefunden worden. Um eine sichere Bestimmung
zu ermdglichen, wurden noch folgende Aufnahmen und Zeichnungen angefertigt; Maxillarpalpen
Abb. 396-397. Fiihlerglieder 1 bis 4 Abb. 398. Tergit 8 Abb. 399. Sternit 8 Abb. 400.
Kopulationsapparat Abb. 401 dorsal.

Abb. 396-397. Sphinginopalpus s. str. rudebecki Wittmer 6 : 396-397, Maxillarpalpen (x200) (396,
von oben; 397, von unten).

127

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 398-401. Sphinginopalpus s. str. rudebecki Wittmer 6: 398, Fiihlerglieder 1-4; 399, Tergit 8; 400, Stemit 8; 401,
Kopulationsapparat (dorsal). Massstab von 398 auch fiir 400, 401.

128

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

65. Sphinginopalpus s. str. wezaensis sp. nov.

MAnnchen (Abb. 402-408).

Lange 3 mm.

Kopf schwarz, von der Mitte nach vorne mit den Wangen gelb; died 3 der Maxillarpalpen
schwarz, 2 schwarzlich; Fiihler gelb; Halsschild schwarz, Basis schmal gelbbraun; Schildchen

Abb. 402-404. Sphinginopalpus s. sir.wezaensis sp. nov. 6 : 402-403, Maxillarpalpen
(xl30) (402, von oben; 403, von unten); 404, Maxille (x 525).

129

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

schwarzlich; Fliigeldecken schwarz, Basis vor den Schultern kurz braunlich, vor der Mitte ein
gelbbraunes, schmales, leicht onduliertes, durchgehendes Querband, das an der Naht bis zum
Schildchen verlangert ist; Schenkel schwarz, die vorderen an der Basis starker als die hinteren
aufgehellt, Vordertibien am wenigsten angedunkelt, die mittleren starker, die hinteren ganz schwarz,
alle Tarsen gelb.

Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberflache chagriniert.
Maxillarpalpen Abb. 402-403. Maxille Abb. 404. Fiihler (Abb. 405) ungefahi so lang wie die
Fliigeldecken, Zahn an der Basis von died 1 sehr kurz, 3 fast doppelt so lang wie 4, 3 bis 10 fast
parallel. Halsschild wie bei repandiis; Oberflache chagriniert. Fliigeldecken wie bei repandus. Tergit
8 Abb. 406. Sternit 8 Abb. 407.

Abb. 405-408. Sphinginopalpus .s. sir. wezaensis sp. nov. d : 405, Fuhlerglicder 1-4; 406, Tcrgit 8; 407, Sternil 8; 408,
Kopulationsapparat (dorsal). Mas.sstab von 405 auch fur 407 und 406 auch fiir 408.

130

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Kopulationsapparat Abb. 408 dorsal.

Holotypus (TMP): South Africa, SW Natal: Weza indigen. forest, grassnetting, 30°35"S, 29°45'E,
24.ii.1985, E-Y:2160, S. Endiody- Y ounga.

Die neue Art ist neben leucosideae Wittmer zu stellen, doch ist das Fiihlerglied 1 weniger parallel
und die Terminal ia verschieden.

66. Sphinginopalpus s. str. dentatiis sp. nov.

Kopf Schwarz, Vorderkopf gelb
ohne die Augen und die Wangen zu
erreichen; Maxillarpalpen dunkelbraun,
stellenweise leicht aufgehellt; Fiihler
schwarz, die ersten 5 Glieder gelb;
Halsschild schwarz, abgeflachte Basis
gelbbraun; Schildchen und Fliigeldecken
schwarz, letztere mit einem
gelblichweissen Seitensaum, sofort
hinter den Schultern beginnend und die
Spitzen fast erreichend; Hinterbeine
schwarz, Hintertarsen schwach
aufgehellt, 4 vordere Schenkel
dunkelbraun, 4 vordere Tibien und
Tarsen gelb.

Kopf mit den Augen breiter als der
Halsschild, Stirne leicht gewolbt;
Oberflache glatt. Maxillarpalpen Abb. 409-410. Fiihler (Abb. 411) um ca. 1/5 kiirzer als die
Fliigeldecken, Glied 3 deutlich 1 anger als 4 (von der Abbildung leider nicht ersichtlich). Halsschild
1 anger als breit (17x14), Seiten nach vorne nur schwach verengt, fast parallel; sonst wie elougatidens,
Fliigeldecken langoval; falsche Epipleuren ca. 70% der Liinge einnehmend, Krenulierung angedeutet;
Punkte in Reihen bis iiber die Mitte gross, dann verworren und praktisch erldschend. Tergit 8 Abb.
412. Sternite 7 und 8 Abb. 413 und 414.

Kopulationsapparat Abb. 415 dorsal.

Holotypus (TMP): South Africa, N Transvaal : Soutpansberg, Hanglip, grassnetting summit, 30°00"S,
29°54T, 18.iii.l973, E-Y:68, S. Endrody-Younga.

Der lange Zahn auf Fiihlerglied 1 erinnert an curvispinus Wittmer und aciitispinus Wittmer,
neben die die neue Art zu stellen ist, doch sind die Maxillarpalpen und die Terminalia verschieden
gebaut.

Mannchen (Abb. 409-415).

Lange 2 mm.

4Gi# 410

Abb. 409-410. Sphinginopalpus s. str. dentatus sp. nov. 6 : 409-410,
Maxillarpalpen (xl70) (409, von oben; 410, von unten).

131

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

0.2

Abb. 411-415. Sphinginopalpus s. sir. deniatus sp. nov. 6 : 411, Fiihlerglieder 1-4; 412, Tergit 8; 413, Sternit 7; 414, Sternit
8; 415, Kopulationsapparat (dorsal). Massstab von 411 auch fiir 412, 414, 415.

67. Sphinginopalpus s. str. bedfordensis sp. nov.

Mannchen (Abb. 416-421).

Lange 2 mm.

Kopf einfarbig schwarz, oder mil einem kleinen, gelblichen Flecken auf dem Vorderkopf,
zwischen den Fiihlerwurzeln, die Augen und Wangen nicht erreichend; Maxillarpalpen dunkelbraun;
Fiihler schwarz, Glieder 1 bis 4 gelb oder aufgehellt, 1 an der Basis, 2 an der Spitze, 3 an der Basis

132

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

schwarzlich; Halsschild schwarz, abgeflachter Teil gelbbraun; Schildchen und Fliigeldecken schwarz,
letztere mil weisslichgelben Seiten, beginnend unter den Schultern bis zu den Spitzen, vor der Mitte
kurz und undeutlich ein wenig nach innen erweitert; Hinterbeine schwarz, Hintertibien wenig,
Hintertarsen ein wenig starker aufgehellt, 4 vordere Schenkel dunkel mit aufgehellten Knien, 4
Vordertibien und 4 Vordertarsen gelb.

4 1 6 41 7

Abb. 416-417. Sphinginopalpus s. str. bedfordensis sp. nov. 6 : 416-417, Maxillarpalpen
(x220) (416, von oben; 417, von unten).

Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberflache glatt.
Maxillarpalpen Abb. 416-417. Firhler (Abb. 418) urn ca. 1/4 kiirzer als die Fliigeldecken, died 3 ein
wenig langer als 4. Halsschild nur wenig liinger als breit (19x18); Sonst wie bei elongatidens.
Fliigeldecken langoval; falsche Epipleuren ca. 80% der Lange einnehmend, nicht krenuliert;
Punktreihen regelmassig, bis iiber die Mitte deutlich, dann Punkte sparlicher, fast erloschend. Tergit
8 Abb. 419. Sternit 8 Abb. 420.

Kopulationsapparat Abb. 421 dorsal.

Weibchen.

Kopf einfarbig schwarz, sonst wie das S gefarbt.

Holotypus und 9 Paratypen (NHMB): South Africa, Cape Prov.; 5-14 km S Bedford (Karoo),
600/730m, ll.x.1984, W. Wittmer. S Transvaal: Robertsdrift, Vaal R., from flood debris, 8.x. 1973,
27°02'S, 29°02'E, E-Y:189, S. Endrddy-Younga, 13 Paratypen (TMP), 4 Paratypen (NHMB), 2
Paratypen (AMG).

Eine kleine Art die dem oneili Pic nahesteht, mit ahnJich gebauten Maxillarpalpen, Fiihlerglied
1 und Terminalia sehr verschieden.

133

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 418-421. Sphinginopalpus .s. sir. bedfordensis sp. nov. 6: 418, Fuhlerglieder 1-4, 419, Tergi! 8; 420, Stemit 8; 421,
Kopulationsapparat (dorsal). Massstab von 418 auch fur 420, 421.

68. Sphinginopalpus s. str. zintbabwensis sp. nov.

Sphinginopalpus alhilabris (Boheman) - Champion, 1922: 332 parte.

Mannchen (Abb. 422-427).

Lange 1,8 mm.

Diese Art die von Champion, 1922, zu albilabris gestellt wurde, stellte sich als neu heraus.
Kopf Schwarz, von der Mitte der Augen nach vorne mit den Wangen gelb; Maxillarpalpen gelb;
Filhler gelb, letzte 3 bis 4 Glieder ein wenig angedunkelt; Halsschild schwarz, abgeflachter Teil
gelborange; Schildchen und Fliigeldecken schwarz, letztere mit ziemlich kurzen, gelblichweissen
Seiten, die die Lange und Breite der falschen Epipleuren eirmehmen; Schenkel schwarz, die vorderen

134

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA; MALACHIIDAE)

und mittleren gegen die Knie aufgehellt, Tibien und Tarsen gelb, die hinteren angedunkelt.

Gestalt und Farbung wie majeri Wittmer. Maxillarpalpen Abb. 422- 423. Fiihlerglieder 1 bis 4
Abb. 424. Tergit 8 Abb. 425. Sternit 8 Abb. 426.

Kopulationsapparat Abb. 427 dorsal.

Abb. 422-423. Sphinginopalpus s. str. zimbabwensis sp. nov. 6 : 422-423, Maxillarpalpen (x200)
(422, von oben; 423, von unten).

Holotypus (BM): Zimbabwe (Mashonaland): Salisbury (Harare), iii.1895, G. A. K. Marshall.

Diese neue Art ist naher mit majeri Wittmer als mit albilabris (Boheman) verwandt, sie hat einen
ahnlich langen Halsschild, doch ist sie durch die verschieden gebauten Terminalia und das Fiihlerglied
1 leicht zu unterscheiden.

135

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1 994

Abb. 424-427. Sphinginopalpus s. str. zimbabwensis sp. nov. cj ; 424, Fiihlerglieder 1-4; 425, Tergit 8; 426, Stemit 8; 427,
Kopulationsapparat (dorsal). Massstab von 424 auch fiir 426, 427.

136

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

69. Sphinginopalpus s. str. basutolandicus sp. nov,

MAnnchen (Abb. 428-433).

Lange knapp 2 mm.

Kopf mit den Wangen
einfarbig schwarz; Fiihler
gelbbraun, Basis von Glied 1
schwarz, Spitze von 2 und 3
fast ganz dunkel; Halsschild
schwarzbraun, fast ganze
abgeflachte Basis gelblich
aufgehellt; Schildchen und
Fliigeldecken schwarzlich,
Seiten der letzteren von der
Basis bis zu den Spitzen
gelblich, vor der Mitte ist die
helle Farbung schmal und
verschwommen gegen die
Naht ausgedehnt; Schienen
und Tarsen, besonders die 4
vorderen vorwiegend

gelblich, Schenkel fast
schwarz.

Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberflache glatt.
Maxillarpalpen Abb. 428-429. Fiihler (Abb. 430) um 1/5 kiirzer als die Fliigeldecken, Glied 3 deutlich

1 anger und ein wenig breiter als 4. Halsschild ein wenig 1 anger als breit (14,5x13,5), Oberflache glatt.
Fliigeldecken langoval, Schulterbeulen gut entwickelt; falsche Epipleuren ca. 80% der Lange
einnehmend, nicht krenuliert; Oberflache glatt, Punktreihen bis iiber die Mitte deutlich, dann Punkte
an Tiefe verlierend, verworren, sparlich. Tergit 8 Abb. 431. Sternit 8 Abb. 432.

Kopulationsapparat Abb. 433 dorsal.

Holotypus (ZML), 1 Paratypus (NHMB): South Africa, Basutoland: Nazareth M. S. 20 miles ESE
Maseru, 24.iii.1951 (No. 245), Swedish South Africa Expedition 1950-1951, Brinck, Rudebeck.

Diese neue Art ist neben bedfordeusis Wittmer zu stellen, von der sie sich durch Fiihlerglied 1,
das verdickte Glied 3 und das verschieden gebaute Sternum 8 unterscheidet, das am Innenrand mit

2 langen Borsten bewehrt ist, bedfordeusis hat nur eine Borste; der Kopul at i onsappar at zeigt weniger
Unterschiede.

Abb. 428-429. Sphinginopalpus s. str. basutolandicus sp. nov. 3 : 428-429,
Maxillarpalpen (x200) (428, von oben; 429, von unten).

137

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 430-433. Sphing'mopalpus s. sir. basutolandicus sp. nov. S : 430, Fiihlerglieder 1-5; 431, Tergit 8; 432, Sternit 8; 433,
Kopulationsapparat (dorsal). Massstab von 430 auch fiir 432, 433.

138

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

70. Sphinginopalpus s. str. breytenbachae sp. nov.
MAnnchen (Abb. 434-439).

Lange 2,2 mm.

Kopf schwarz, ein kleiner verschwommener Flecken auf dem Vorderkopf zwischen und liber
den Fiihlerwurzeln und Wangen gelb, ein schmaler Fortsatz auf die Stirne bis zur Mitte der Augen
verlangert. Maxillarpalpen schwarzlich, teils braunlich aufgehellt; Fiihler gelb, letzte 5 Glieder, sowie
Basis von 1, schwarzlich; Halsschild schwarz. Basis schmal, kaum aufgehellt; Schildchen und
Flugeldecken schwarz, letztere jederseits mit einem weisslichen, antimedianen, dreieckigen Flecken,
der fast bis zur Naht reicht; Hinterbeine schwarz, Hintertarsen etwas aufgehellt, 4 Vorderschenkel
dunkel, an den Knien kurz hell, 4 Vordertibien gelb, an der Basis dunkel, 4 Vordertarsen gelb, letztes
Glied leicht dunkler.

Kopf mit den Augen
breiter als der Flalsschild,
Stirne leicht gewolbt;
Oberflache glatt.

Maxillarpalpen Abb. 434-
435. Fiihler (Abb. 436) um ca.
1/4 kiirzer als die
Flugeldecken, Glied 3 kaum
merklich langer als 4.
Flalsschild langer als breit
(16x14), Oberflache glatt,
abgeflachter Teil chagriniert,
sonst wie bei elongatidens.
Flugeldecken langoval;
Abb. 434-435. Sphinginopalpus s. str. breytenbachae sp. nov. 3: 434-435, falsche Epipleuren ca. 55%
Maxillarpalpen (xl70) (434, von oben; 435, von unten). , , • u j

^ ^ ^ der Lange eirmehmend,

Krenulierung nur stellenweise kaum angedeutet; Punktreihen anfangs regelmassig, nach hinten
aufgelost, Punkte erldschend. Tergit 8 Abb. 437. Sternit 8 Abb. 438.

Kopulationsapparat Abb. 439 dorsal.

Holotypus (TMP): South Africa, N Karoo; Baviaankloof, beating, 33°38'S, 24°26'E, 13. i. 1979, W-
B:, W. Breytenbach.

Eine neue Art mit tief ausgehohltem Fiihlerglied 1 an der Basis mit schwach stumpfem Zahn,
ausserdem gekennzeichnet durch das in der Mitte kurz, ziemlich tief ausgerandete Tergit 8.

139

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19. PT. 1 30 JANUARY 1994

Abb. 436-439. Sphinginopalpus s. str. breytenbachae sp. nov. <3 : 436, Fiihlerglieder 1-4; 437, Tergit 8; 438, Stemit 8; 439,
Kopulationsapparat (dorsal). Massstab von 436 auch fiir 438, 439.

71. Sphinginopalpus s. str. elongatidens sp. nov.

Mannchen (Abb. 440-445).

Lange 2,5 mm.

Kopf mil den Wangen schwarz, Vorderkopf beginnend von der Mitte der Augen, gelb;
Maxillarpalpen dunkel, letztes Glied, besonders an der Spitze aufgehellt; Fiihler schwarzlich, Glieder
1 bis 4 gelb; Halsschild schwarz, die ganze abgeflachte Basis hellbraun; Schildchen und Fliigeldecken
schwarz, Seiten von der Basis bis fast zu den Spitzen schmal gelb, Spitzen kurz, schwach aufgehellt,
antimediane gelbe Quermakel an der Naht unterbrochen oder durchgehend; Hinterbeine schwarz,
Hintertarsen leicht aufgehellt, Mittel- und Vorderschenkel schwarz, Mitteltibien etwas weniger stark

140

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 440-441. Sphinginopalpus s. str. elongatidetis sp. nov. S : 440-441, Maxillarpalpen (440, von obcn,
xl30; 441, died 1, x 625).

aufgehellt als die vorderen, 4 Vordertarsen weitgehend gelb.

Kopf mil den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberflache glatt.
Maxillarpalpen (Abb. 440-441), Glied 1 auf der Oberseite mil ziemlich tiefen und breiten Gruben
bedeckt, ahnlich wie bei tridens (Abb. 456). Fiihler (Abb. 442) verhaltnism^sig kurz, fast um 1/3
kiirzer als die Fliigeldecken, Glied 1 durch einen sehr langen, allmahlich verschmalerten, leicht
verdrehten Fortsatz ausgezeichnet, der fast in der Mitte des Gliedes steht, auf der Hohe der Ausrandung
steht eine kurze, quere Lamelle, 3 und 4 ungefahr gleich lang. Halsschild langer als breit (17x15,5),
in der Mitte am breitesten, Seiten nach vorne schwach gerundet verengt, gegen die Basis eingeschniirt;
Seitenrand die Basis nicht ganz erreichend, Basalrand deutlich; Oberflache glatt, an den Seiten der
abgeflachten Basis chagriniert. Fliigeldecken langoval, Schulterbeulen normal, falsche Epipleuren
lang, ca. 80% der Lange eirmehmend, nicht krenuliert; Oberflache glatt, auf dem basalen Teil mit ca.
6 Punktreihen, die an den Seiten weniger deutlich sind, auf dem hinteren Teil Punkte verworren.
Tergit 8 Abb. 443. Sternit 8 Abb. 444.

Kopulationsapparat Abb. 445 dorsal.

Holotypus (TMP): South Africa, E Transvaal: Nelshoogte Forest Station, grassnetting, 25°50"S,
30°50'E, l.xii.l986, E-Y:2335, S. Endrody-Younga; Berlin Forest Station, grassnetting, 1500 m,
25°32'S, 30°44'E, 7.xii.l986, E-Y:2360, S. Endrody-Younga, 3 Paratypen (TMP), 2 Paratypen
(NHMB). Transkei (Pondoland): Port St. John, 7.-13.viii.l923, R. E. Turner, 1 Paratypus (BM).

Diese neue Art ist neben curvidens Wittmer zu stellen, sie unterscheidet sich durch das
verschieden gebaute Fiihlerglied 1, die Maxillarpalpen und die Terminalia. Die Gruben auf der
Oberseite von Glied 1 der Maxillarpalpen sind ahnlich wie bei tridens.

141

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

Abb. 442-445. Sphinginopalpus s. str. elongatidens sp. nov. <? : 442, Fuhlerglieder 1-4; 443, Tergit 8; 444, Stemit 8; 445,
Kopulationsapparat (dorsal). Massstab von 442 auch fiir 444, 445.

72. Sphinginopalpus s. str. karkloofensis sp. nov.

Mannchen (Abb. 446-451).

Lange 2, 2-2, 3 mm.

Kopf Schwarz; died 3 der Maxillarpalpen schwarz, leicht aufgehellt; Fiihler schwarz, Glieder
1 bis 3 gelb, wobei 1 beginnend an der Basis mehr oder weniger stark angedunkelt ist; Halsschild
schwarz, abgeflachter Teil braun; Schildchen und Fliigeldecken schwarz, letztere mit schmalen.

142

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

weissen Seiten, beginnend unter den Schulterbeulen bis fast zu den Spitzen, bei 1 Ex. mit einer
antimedianen Quermakel, die etwas verschwommen ist und die Naht nicht erreicht; alle Schenkel
schwarz, Tibien und Tarsen gelb, Oder die hinteren leicht angedunkelt.

Abb. 446-447. Sph'mginopalpus s. str. karkloofensis sp, nov. d : 446-447,
Maxillarpalpen (xl70) (446, von oben, 447, von unten).

Kopf mit den Augen breiter als der Halsschild, Stirne leicht gewolbt; Oberflache glatt mit feinen
Punkten, die auf der vorderen Halfte deutlicher sind. Maxillarpalpen Abb. 446-447. Fiihler (Abb.
448) um ca. 1/4 kiirzer als die Fliigeldecken, died 3 nur wenig langer als 4, 4 ein wenig breiter als
5. Halsschild langer als breit (16x15), sonst wie bei elougatidens, jedoch Oberflache variabel, beim
Exemplar von Karkloof fast ganz glatt, beim anderen von Isidenge auf der abgeflachten Stelle
weitgehend chagriniert. Fliigeldecken langoval, wie bei elougatidens, Punktreihen etwas deutlicher,
weiter nach hinten reichend, falsche Epipleuren lang, kaum merklich krenuliert, fast glatt. Tergit 8
Abb. 449. Sternit 8 Abb. 450.

Kopulationsapparat Abb. 451 dorsal.

Holotypus (TMP): Natal: Karkloof grassveld with river, 29°19'S, 30°15"E, 7.xii.l989, E-Y;2747, S.
Endrody-Younga. Cape Prov.: Isidenge Forest Station, 32°41'S, 27°15"E, 18. xi. 1987, E-Y:2522,
grassnetting and Arum fl., S. Endrody-Younga, 1 Paratypus (NHMB).

Diese neue Art ist auffallig durch ihr fast dreieckiges Fiihlerglied 1, das an der Basis tief
ausgerandet ist, die Terminalia und hier besonders das Sternit 8 sind sehr charakteristisch.

143

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19. PT. 1 30 JANUARY 1994

Abb. 448-451. Sphinginopalpus s. str. karkloofensis sp. nov. i : 448, Fuhlerglicder 1-4; 449, Tergit 8; 450, Sternit 8; 451,
Kopulationsapparat (dorsal). Massstab von 448 auch fur 450, 451.

73. Sphinginopalpus s. str. nylsvleyensis sp. nov.

Mannchen (Abb. 452-456).

Lange 1,8-2 mm.

Kopf schwarz, von der Mitte der Augen nach vorne inklusive der Wangen gelb; Maxillarpalpen
braun bis dunkelbraun; Fiihler schwarzlich, erste 4 bis 5 Glieder gelb; Halsschild schwarz,
abgeflachter Teil gelbbraun; Schildchen und Fliigeldecken schwarz, letztere mil langem gelbem
Seitenband, das antimedian ein wenig verbreitert ist; Flinterschenkel schwarz, mittlere Schenkel
ebenso, gegen die Knie aufgehellt, Mitteltibien und Mitteltarsen gelb (iibrige Tarsen fehlen).

Kopf mit den Augen nur ganz wenig breiter als der Halsschild, Stirne leicht gewolbt; Oberflache
glatt mit feinen zerstreuten Haarpunkten. Maxillarpalpus Abb. 452. Fiihler (Abb. 453) um 1/4 kiirzer
als die Fliigeldecken, Glied 3 nur ein wenig Linger als 4. Halsschild Linger als breit (15,5x14), sonst
wie bei elongatidens. Fliigeldecken langoval; falsche Epipleuren ca. 60% der Lange eirmehmend,
keine Krenulierung; Punktreihen regelmassig, Punkte weit auseinander stehend. Tergit 8 Abb. 454.

144

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 452. Sphinginopalpus s. str. nylsvleyensis
sp. nov. 6 : 452, Maxillarpalpus (x220,
von oben).

Sternit 8 Abb. 455.

Kopulationsapparat Abb. 456 dorsal.

Holotypus (TMP): South Africa, N Transvaal: Nylsvley, at
river, ground trap with banana bait, replication 4, 24°40"S,
28°42"E, 3.ix.l975, E-Y :875, S. Endrody-Younga; Roberts
Drift, Vaal R., from flood debris, 27°02'S, 29°02'E,
8.x. 1973, E-Y:189, S. Endrody- Younga, 1 Paratypus
(TMP), 1 Paratypus (NHMB).

Eine kleine Art, die neben dwesaensis Wittmer zu
stellen ist, mit ahnlich gebautem Fiihlerglied 1, jedoch sehr
verschiedenen Terminalia.

Abb. 453-456. Sphinginopalpus s. str. nylsvleyensis sp. nov. 3 : 453, Fiihlerglieder 1-4; 454, Tergit 8; 455, Sternit 8; 456,
Kopulationsapparat (dorsal). Massstab von 452 auch fur455, 456.

145

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 1 30 JANUARY 1994

74. Sphinginopalpus s. str. tridens sp. nov.

Mannchen (Abb. 457-461).

Lange 2 mm.

Kopf schwarz, Wangen um die Fiihlerwurzeln,
verbunden mit einer schmalen Verlangerung bis zur
Mitte der Augen und ein kleiner Flecken iiber den
Fiihlerwurzeln, gelblich; Maxillarpalpen gelbbraun;
Fuhler schwarzlich, die ersten 5 Glieder gelb;
Halsschild schwarz, abgeflachter Teil an der Basis
gelbbraun; Schildchen und Fliigeldecken schwarz,
letztere mit schmalen gelben Seiten, unter den
Schultern beginnend und fast die Spitzen erreichend,
antimedian gegen die Naht verbreitert, ohne sie zu
erreichen; Schenkel schwarz, die vorderen gegen die
Knie aufgehellt, Schienen und Tarsen gelb, die
hinteren leicht angedunkelt.

Kopf mit den Augen breiter als der Halsschild,
Stirne leicht gewblbt; Oberflache glatt.
Maxillarpalpen (Abb. 457) haben auf der Oberseite
von Glied 1 ziemlich tiefe und breite Gruben, ahnlich
wie bei elongatidens (Abb. 439-440). Fuhler (Abb. 458) um 1/4 kiirzer als die Fliigeldecken, Glied
3 kaum merklich langer als 4. Halsschild so lang wie breit, sonsl wie bei elongatidens. Fliigeldecken
langoval, falsche Epipleuren ca. 80% der Lange einnehmend, Krenulierung fehlt; Punktierung viel
weniger dicht als bei elongatidens und weniger deutlich in Langsreihen angeordnet. Tergit 8 Abb.
459. Sternit 8 Abb. 460.

Kopulationsapparat Abb. 461 dorsal.

Holotypus und 1 Paratypus (TMP), 1 Paratypus (NHMB); South Africa, Natal Middld.: Karkloof
grassveld, 29°19'S, 30°15'E, 7.xii.l989, E-Y:2737, 2747, S. Endrody-Younga, I. Klimaszewski.

Diese neue Art ist neben elongatidens Wittmer zu stellen, Fiihler von ahnlicher Bauart,
Maxillarpalpen, Terminalia und Kopulationsapparat sehr verschieden.

Abb. 457. Sphinginopalpus s, str. tridens sp. nov. i:
457, Maxillarpalpus (xl70, vonoben).

146

DIE GATTUNG SPHINGINOPALPUS PIC (COLEOPTERA: MALACHIIDAE)

Abb. 458-461. Sph in gin opalpus s. str. iridens sp. nov. 6 : 458, Fiihlerglieder 1-4; 459, Tergit 8; 460, Stemit 8; 461,
Kopulationsapparat (dorsal). Massstab von 458 auch fiir 460, 461.

LITERATUR

Boheman, C. H. 1E51. Insecta Caffraria annis 1838-1845 aJ.A. Wahlbergcollecta. Coleoptera. Holmiae: Fritzeund Norstedt.
Champion, G. C. 1922. Revision of the African species of Dinometopus, Tro glops, Chalicorus, Colotes, Helcogaster, and the
allied genera, with an account of their accessory 6 -characters (Coleoptera). Annafa and Magazine ofNatural History
(9)10(58): 309-358.

Gorham, H. S. 1901. Descriptions of genera and species of Coleoptera from South Africa. Annals and Magazine ofNatural
History (7)7(46): 349-365.

Pic, M. 1903. Malachides de I’Afrique australe. Echange 19: 164-165.

Pic, M. 1904. Un nouveau Malacoderme de I’Afrique australe. Bulletin de la Societe Entomologique de France: 12-13.

Pic, M. 1929. Nouveautes diverses. Melanges exotico-entomologiques S3: 1-36.

WiTTMER, W. 1960. Coleoptera: Cantharidae, Malachiidae & Dasytidae. In: Hanstrom, B. et al., eds, South African Animal
Life. 7. Uppsala: Almquist and Wiksels, pp. 113-127.

147

INSTRUCTIONS TO AUTHORS

LAYOUT of recent issues should be examined for details of acceptable style and format.

MANUSCRIPTS should be submitted on floppy disk accompanied by three print-outs
(spacing I/2 or 2). In exceptional cases typed manuscripts will be accepted. These should be
submitted in triplicate, typed on A4 size paper, double-spaced, on one side of the paper only
and with wide margins all round.

Only generic and specific names should be underlined to indicate italics. All other marking
up should be left to the editor.

Footnotes should be avoided as far as possible.

All measurements should be metric.

Systematic papers should follow the rules of the International code of zoological
nomenclature or International code of botanical nomenclature, as applicable.

ILLUSTRATIONS should bear the author’s name, title of paper and number in soft pencil on
the reverse side. Diagrams, line drawings and photographs are all regarded as figures, should
be termed as such and should be numbered in a single series.

In addition to the original illustrations a photocopy of each should be submitted.

Diagrams and line drawings should be reducible to a size not exceeding 14 x 16 cm (17,5 cm
including captions). Photographs for half-tone blocks should be final size, of high contrast
and printed on glossy paper. Colour illustrations are only printed under special circumstances
and their financing must be arranged by the author.

A scale should appear with all illustrations or the magnification should be given in the
caption.

A list of captions should be submitted on a separate sheet.

TABLES should be submitted on separate sheets. Foldouts and tables printed sideways should
be avoided.

REFERENCES should be listed alphabetically at the end of the paper. Abbreviations of
periodical titles should conform with those in the World list of periodicals (4th ed.). Layouts
must be as follows:

Ai.e.xander, J. 1969. The indirect evidence for domestication. In: Ucko, P. J. and G. W.
Dimbleby eds, The domestication and exploitation of plants and animals. London:
Duckworth, pp. 123-129.

De Winter, B. 1969. Plant taxonomy today. S.AfrJ.Sci. 65 (8): 229-242.

JuBB, R. A. 1967. Freshwater fishes of southern Africa. Cape Town: Balkema.

GALLEY PROOFS are sent (when practicable) to authors for checking printer’s errors.
Adjustments of language, style and punctuation and any other amendments of the original
manuscript, if made at this stage, may be charged to authors.

AUTHOR’S COPIES (50) are issued free of charge. When more than one author is involved
the fifty free copies will be sent to the senior author. Additional copies ordered before printing
are supplied at a reduced cost.

CADAR. P E.-K887

SMITHSONIAN INSTITUTION LIBRARIES

ill

II

III

1

III

III

IS

088 0

1429 5638

*611:.

^ -■ ■' -1"'

: . :”;iL i.-.-s-—

!* ■

:.= ;•(
l[’•'i

■■ .,'H"

f-^=' .

• .m;-;

v>. :<!.

“S

3^! : ■■ ■

*.

ISSN 0570-1880

S>H

\6/3

w

Annals of the

Cape Provincial Museums

Natural History

Ann. Cape Prov. Mus. (nat. Hist)

Volume 19 Part 2 30th June 1994

'^^THSOaT^

26 I99§

iiSRARitS

I

1

. Published jointly by the Cape Provincial Museums
at the Albany Museum, Grahamstown, South Africa

ANNALS OF THE CAPE PROVINCIAL MUSEUMS

These Annals are published in two series, Natural History and Human Sciences, the latter series
covering cultural history, ethnology, anthropology and archaeology. They are issued in parts at
irregular intervals as material becomes available.

The primary objective of these Annals is to disseminate the results of research work undertaken by
staff members of the Cape Provincial Museums. However, a manuscript by any author is eligible
for publication provided it is based wholly or partly on the collection/s of one or more of these
museums or if the material on which the work is based is wholly or partly deposited in one of
these museums.

Parts are obtainable from the Librarian of the Albany Museum. Correspondence in coimection
with these Armais should be addressed to the Editor, Albany Museum, Grahamstown 6140.

Editor

Dr F. W. GESS

Assistant Editor
Dr S. K. GESS

Seasonal occurrence and distribution of nuptial tubercles in two African
minnows, Pseudobarbus afer (Peters, 1864) inhabiting clear water and
Pseudobarbus asper (Boulenger, 1911) inhabiting turbid water
(Cypriniformes, Cyprinidae).

by

J. A. CAMBRAY

(Albany Museum, Grahamstown, 6140, South Africa)

ABSTRACT

Pseudobarbus afer (Peters, 1864) and P. asper (Boulenger, 1911) are a sister species pair which
inhabit different sections of the Gamtoos River system. Pseudobarbus afer occurs in the clear mountain
streams of the Cape Fold Belt andP. asper in the turbid Karoo section of the river system. The seasonal
occurrence and disappearance of tubercles for both species followed the reproductive cycle and was
probably under seasonal hormonal control. Pseudobarbus afer males had the more numerous and
larger tubercles when compared to P. asper males. It is suggested that the enlarged, white head
tubercles of P. afer were used for display prior to breeding, whereas during breeding the tubercles
on the scales and fin rays and the larger pectoral fins were used for maintaining contact with females
in the riffle spawning habitats. The relatively smaller number and size of the head tubercles on P.
asper males possibly indicated that in turbid waters visual characters have less of a selective advantage
than in clear mountain streams. The smaller number and size of tubercles on P. asper males would
have required less resources. The remainder of the resources would then be available for other
reproductive activities. Pseudobarbus afer and P. asper males had larger definitive head tubercles
than the rudimentary tubercles observed on the females. Only the large females of both species had
head tubercles which were never definitive and their presence on reproductively active females was
difficult to explain. The lack of tubercles on the smaller females indicated the expected trade-off
constraint with the energy diverted to egg production rather than sexually dimorphic characters. The
smaller pectoral fins of the females, which had few if any tubercles, also indicated that it was more
beneficial for the female to put reproductive effort into progeny. The development of secondary sexual
characteristics of P. afer and P. asper (epidermal tubercles, breeding coloration and larger pectoral
fins) is favoured on the males because the sexual characteristics all require resources which must be
diverted away from growth, survival requirements and the production of gametes.

INTRODUCTION

Epidermal breeding tubercles are temporary, roughened structures which have evolved in
primitive euteleostean fishes and have been recorded in at least 25 families of bony fishes in 7 orders
(Wiley and Collette, 1970). In addition tubercles occur in some species of the Coregonidae,
Thymallidae (Salmoniformes) and Gadidae (Gadiformes) (Kratt and Smith, 1978; Vladykov et al.,
1985; Witkowski, 1982 in Witkowski and Rogowska, 1991). Structural differences indicate that
tubercles have evolved independently in a number of fish groups (Collette, 1977). Tubercle patterns
can be used to infer similarities or differences in behaviour patterns (Wiley and Collette, 1970). The
head and forepart of the body of many species of the family Cyprinidae may be covered with
excrescences (Breder and Rosen, 1966) and the fish may have a bright nuptial coloration during the
spawning season (Banarescu and Coad, 1991). Tubercles can be valuable systematic characters in

149

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 2 30 JUNE 1994

cyprinids (Wiley and Collette, 1970; Collette, 1977). The distribution, number, size and developmental
patterns of the head tubercles represent critical characters in the separation of some species (Lachner
and Jenkins, 1971). Tubercles have also been considered as important evolutionary characters
(Lachner and Jenkins, 1971). They might have evolved to enable spawning individuals to maintain
contact in fluviatile environments. Tubercles may also be used by some fish species to defend nests
and territories or possibly to stimulate females to breed (Wiley and Collette, 1970; Collette, 1977).

In African cyprinids tubercles have usually been recorded in bariliine and labeine species and
less so for barbine species (Howes, 1980; Reid, 1985). Skelton (1980) noted the presence, size and
patterns of tubercles on 25 Barbus species from southern Africa. Conical tubercles occur in all seven
of the flexible-rayed redfin minnows (= Pseudobarbus) including P. afer (Peters, 1864) and P. asper
(Boulenger, 1911). In comparison to the Barbus species, the flexible-rayed redfin tubercle pattern
consists of large tubercles and there are also tubercles on the borders of the scales which occur in
regular rows (Skelton, 1980). Barnard (1943) and Jubb (1967) recorded the presence of tubercles and
smaller ‘pimples’ on redfin mirmow species in southern Africa and used the presence or absence of
tubercles as a major character to separate species in southern Africa. Tubercles have been identified
as important taxonomic characters in the Pseudobarbus lineage (Skelton, 1980, 1988).

The tubercles in the seven redfin minnow species of the genus Pseudobarbus have a common
distribution pattern. The size and pattern of the tubercles in P. burchelli Smith, 1841, P. burgi
(Boulenger, 1911), P. afer, P. asper, P. tenuis (Barnard, 1938) and vestigially in P. phlegethon
(Barnard, 1938) was found to be unique for African fish and represent a very distinct synapomorphy
(Skelton, 1980). The head tubercles of P. quathlambae (Barnard, 1938) are more numerous and
smaller than in the other flexible-rayed redfins but are arranged in the same basic pattern (Skelton,
1974). As in other cyprinids the tubercles of the flexible-rayed redfin minnows are deciduous,
hypertrophied, epidermal structures with a distinct keratin cap.

Data on the seasonal occurrence of breeding tubercles are usually not collected (Collette, pers.
comm., October 10, 1985) but see Collette (1965). Field studies to note seasonality of tubercle
formation and wear have been rare for African cyprinids. The seasonal occurrence of these structures
has not been studied in any detail for P. afer andP. asper. Skelton (1974) gave a brief account of the
seasonality of the tubercles for oreodaimon (P. quathlambae) and Cambray and Stuart (1985) noted
the seasonality of the head tubercles for P. burchelli over a 12 month period. A SEM micrograph of
a male head tubercle scar in the species P. quathlambae was given by Skelton (1974).

The usual seasonal sequence for breeding tubercles is their appearance at the commencement of the
breeding season and just prior to the spawning season they reach their maximum extent. Following
spawning the structures break off, slough, become eroded, or gradually regress. Cambray and Stuart (1985)
recorded that the next season’s tubercles started to erupt through the scar left from the previous season.

In the fish species chosen for the present study, P. afer inhabits clear, perennial, low conductivity
streams and P. asper inhabits the turbid, intermittent, and high conductivity section of the Gamtoos
River system. Therefore, if the head tubercles of the Pseudobarbus species were used for any visual
signal, such as sex or species recognition, the clear water species (P. afer) should have larger and
possibly more tubercles than the species inhabiting the turbid system (P. asper).

METHODS

Collection sites

Pseudobarbus afer were collected from the Wit River and P. asper from the Groot River in the

150

CAMBRAY: NUPTIAL TUBERCLES IN PSEUDOBARBUS: SEASONAL OCCURRENCE AND DISTRIBUTION

Fig. 1. Distribution of Pseudobarbus afer (Peters, 1864) (dosed circles) and P. asper (Boulenger, 1911) (open circles) in the
Gamtoos River system (Albany Museum records) and collection sites for the present study (Wit River for P. afer and
Smithskraal and Beerpoort for P. asper).

151

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 2 30 JUNE 1994

Gamtoos River system (Fig. 1). The Gamtoos River system has two distinct catchment basins. One
occurs in the Karoo where the Groot River flows over easily erodable Bokkeveld beds. The rivers
are turbid, saline and intermittent (Cambray, 1992). This is the habitat ofP. asper. The other catchment
occurs within the Cape Fold Mountain Belt and is characterised by resistant quartzites. The rivers
are clear, have a low conductivity and are perennial (Cambray, 1992). Pseudobarbus afer occur in
these streams. There is no physical barrier separating these two sister species in the Gamtoos River
system.

Collection and analysis

Fish were collected with a miimow seine net, fixed in 10% formalin and taken to the laboratory
for analysis.

Conical tubercle formation was studied on a monthly basis for the males and females of both
species over a 22 month period (July 1987 to April 1989). Fish were examined each month from a size
range covering the largest fish in the sample to the smallest without any tubercles. Totals of 248 male
and 118 female P. afer and 320 male and 157 female P. asper were examined (Tables 1-4). Height
and basal width of the conical tubercles were measured with an eyepiece micrometer on a stereoscopic
microscope (accuracy 0,05 mm). Volume of the tubercles was considered as the comparative
measurement was calculated from the height and basal width data using the formula for a cone:

(V = 3T r^ h )

3

The terminology used by Lachner and Jenkins (1971) to describe tubercle development was
followed, with several modifications. Five stages were identified, spot, bud, nuptial definitive tubercle,
worn tubercle and scar. Spots were defined as distinct light areas which approach the epidermal
surface. Tubercle buds were small ‘pimples’ which have a rounded or pointed tip. Nuptial definitive
tubercles were mature structures which were hard, pointed and cornified. Worn tubercles were
definitive tubercles which had lost their points and were rounded or flat but still raised above the
surface. Scars were left when the tubercles were lost. Tubercles were easily distinguished from dermal
sense organs simply by lifting the tubercles from the skin with a needle whereupon a small round
depression would remain (Branson, 1962).

Scarming electron micrographs were taken with a JEOL Scanning Electron Microscope operating
at 10 mV. Eruption and wear of tubercles were noted. Pre-spawning and post-spawning head tubercles
were removed to study the microwear on the surfaces of the tubercles using standard SEM techniques.
Microwear on the pectoral fin tubercles was also studied. Microwear is a term used in archaeology
to describe the wear patterns on stone, bone or wooden tools caused by different uses of the tools. It
was thought this method could be used to determine whether the tubercles had been used for butting
females, fighting other males or for digging and making nests. It was envisaged that these activities
would cause different wear patterns on the tubercles of the head and those on the pectoral fins.

The head tubercles were divided into the clusters which Skelton (1988) used to describe the
pattern of tubercles in the flexible-rayed redfin minnows (Fig. 2). In each of the paired clusters the
left and right sides were counted separately to see if there were different growth and/or wear patterns
correlated to side. Regression analyses were run for the total number of tubercles versus standard
length (SL mm) for both sexes of each species. Tests of difference of slopes of the regression lines
of the total tubercle number for P. afer andP. asper males were performed by analysis of covariance
(Zar, 1984). F values yielding probabilities greater than 0,95 were considered to be significant.

152

CAMBRAY: NUPTIAL TUBERCLES IN PSEUDOBARBUS: SEASONAL OCCURRENCE AND DISTRIBUTION

Fig. 2. Diagrammatic representation of the pattern of head tubercles on Pseudobarbus afer (Peters,

1864) and P. asper (Boulenger, 1911); (a) - cluster on snout; (b) - row above nares; (c) - row
above orbit; (d) - anterior dorsal cluster; (e) - posterior dorsal cluster, (from Skelton, 1988).

Comparison of the presence or absence of definitive snout tubercles and the presence of buds on
male P. afer and P. asper was graphed with the mean monthly gonadosomatic indices (data from
Cambray, 1992) to show the relationship between the tubercles and increases in gonad mass. The
presence of definitive tubercles was given the value 1, the presence of buds the value 0,5 and worn
tubercles and scars were given the value 0.

All fins were examined for tubercle formation and presence or absence was recorded over the
22 month period of the study. The number of rows of tubercles on the rays of the pectoral fins was
also recorded. This was not necessary for the other fins since there was only a single row of tubercles
on each ray. Scales were examined for tubercle formation and presence or absence was recorded for
the 838 fish sampled.

153

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 2 30 JUNE 1994

Table 1. Summary of the seasonal pattern of tubercle formation and wear in 248 Pseudobarbus afer
(Peters, 1864) males collected over a 22 month period in the Wit River, Gamtoos River system.

Date

n

Size range
SL (mm)

Tubercles*

Scales**

Rays’

Jul. 1987

11

49,2-71,1

B+Sp

-

-

Aug.

3

45,6-51,8

B-i-Sp

-

-

Sept.

12

39,7-62,8

D-i-W

+

+

Oct.

31

40,3-78,2

Dh- W

+

+

Nov.

12

37,3-70,5

D+W

+

+

Dec.

8

45,9-63,4

D-i-W-i-S

+

+

Jan. 1988

7

46,0-59,2

D-i-W-hS

-

+ (few)

Feb.

12

47,2-68,1

w+s

-

+ (few)

Mar.

6

42,8-63,1

w

-

+ (few)

Apr.

12

39,1-66,0

W-i-S

-

+

May

15

43,1-65,8

w+s

-

-

June

5

46,7-58,0

B+S+Sp

-

-

July

9

51,6-72,5

B+W+S+Sp

-

-

Aug.

7

42,2-64,7

B+Sp

+ (few)

-

Sept.

11

40,7-63,2

D

+

+

Oct.

12

40,6-62,5

D+W

+

+

Nov.

14

43,5-66,2

D+W+S

+

+

Dec.

5

44,2-70,1

D+W

+

+

Jan. 1989

14

37,2-69,6

B+W+S

-

+

Feb.

9

38,0-70,5

B+W

+ (few)

+

Mar.

19

41,4-68,2

B+W+S

-

+

Apr.

14

38,9-67,6

B+W+S

-

+

TOTAL

248

1. B = bud; D

= definitive;

S •- scars; Sp = spots; W

= worn

2. - = absent; + = present

3. - = absent; + = present

RESULTS

Seasonality of the occurrence of the tubercles was evident for males and females of both species
but was more clearly defined in the males (Tables 1-2 and Fig. 3). Definitive snout tubercles were
only present on the males of the two species. The keratinized tips of the definitive tubercles on the
heads of both of the redfin minnow species were pointed and curved posteriorly (Fig. 3). In P. afer
males definitive snout tubercles first appeared in September 1987 and 1988 (Table 1) and there was
a close correlation with the seasonal variation in the gonadosomatic indices (Fig. 4). Definitive snout
tubercles inP. asper males first appeared in October 1987 and in September 1988 (Table 2) and as

154

CAMBRAY; NUPTIAL TUBERCLES IN PSEUDOBARBUS: SEASONAL OCCURRENCE AND DISTRIBUTION

Fig. 3. Definitive tubercles on males otPseudobarbus afer (Peters, 1864) (above) and P. asper (Boulenger, 1911) (below).

155

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 2 30 JUNE 1994

Table 2. Summary of the seasonal pattern of tubercle formation and wear in 320 Pseudobarbus asper
(Boulenger, 1911) males collected over a 22 month period in the Groot River, Gamtoos
River system.

Date

n

Size range
SL (mm)

Tubercles^

Scales^

Rays^

July 1987

8

58,6-67,8

B+S

-

-

Aug.

7

58,8-83,4

B+S

-

-

Sept.

14

43,3-64,0

B

+ (few)

+ (few)

Oct.

15

41,7-64,9

B+D-i-W

+

+

Nov.

20

40,4-72,3

D

+

4-

Dec.

14

45,7-69,9

D

+

+

Jan. 1988

18

45,4-63,1

W

+

+

Feb.

11

42,4-62,0

D+W

+

+

Mar.

6

50,3- 53,7

D

+

+

Apr.

18

34,1-57,6

D+W

+

+

May

10

47,0-58,3

W

-

-

June

16

49,9-65,4

B+S+W

-

-

July

4

48,5-54,5

B+W

-

-

Aug.

10

48,1-70,4

B+S

-

+

Sept.

29

42,8-62,4

B+D

+ (few)

+

Oct.

15

42,4-53,9

D+W

+

+

Nov.

20

44,2-63,6

W

+

+

Dec.

16

36,0-64,5

D

+

+

Jan. 1989

19

37,9-61,2

D

+

+

Feb.

13

45,0-66,1

W

+

+

Mar.

12

48,6-71,9

B+D

+

+

Apr.

25

41,7-66,7

B+W

+ (few)

+

TOTAL

320

1. B = bud; D = definitive; S = scars; Sp = spots; W = worn

2. - = absent; + = present

3. - = absent; + = present

with the P. afer males there was a close relationship between the seasonal formation of tubercles and
the seasonal pattern of the gonadosomatic indices (Fig. 5). The seasonal pattern was also evident in
the presence and absence of scale and ray tubercles for both species. The longer potential spawning
season of P. asper compared to P. afer was evident from the presence of definitive tubercles as late
as April for P. asper males (Figs 4 and 5). In P. afer females buds first erupted in September 1987
and 1988 (Table 3). In P. asper females buds first appeared in September 1987 and 1988 (Table 4).
Tubercles were a dominant feature on the scales and fin rays of the males of both species (Tables 1
and 2) whereas very few females of either species had scale or fin ray tubercles (Tables 3 and 4).

156

CAMBRAY: NUPTIAL TUBERCLES IN PSEUDOBARBUS: SEASONAL OCCURRENCE AND DISTRIBUTION

GSI — t— Snout tub.

Fig. 4. Comparison of the seasonal occurrence of definitive snout tubercles on male Pseudobarbus afer (Peters, 1864)
(n=145) with the mean monthly gonadosomatic index (GSI) of male P. afer for the period July 1987 to April 1989.
(Definitive tubercle = 1; bud tubercle = 0,5).

The smallest male P. afer with tubercles was 41,2 mm SL, collected in October 1987. Five buds
occurred in the posterior dorsal section of this specimen and the pectoral fin rays had tubercles. The
smallest male P. asper was 39,3 mm SL, collected in January 1989, with three tubercle buds (1 snout
and 2 nares).

The smallest female P. afer with tubercles (= spots) was 56,5 mm SL collected in December
1988. Two spots were present, one in the nares position and one in posterior dorsal position. The
smallest female P. afer with buds was a 56,8 mm SL specimen collected in January 1988. There were
6 buds and all were in the posterior dorsal position. In comparison the smallest female P. asper with
tubercles (= spots) was 49,4 mm SL collected in March 1988. The 4 spots were all in the posterior
dorsal position. The smallest female P. asper with buds was a 49,5 mm SL specimen collected in
October 1987. All 6 buds were in the posterior dorsal position.

A 70,5 mm SL male P. afer had the most tubercles of all the males examined for both species
with 88 tubercles (14 snout; 7 nares; 12 above orbits; 6 anterior dorsal; 49 posterior dorsal)(Table 5).
The biggest tubercles were on the snout and the largest had a volume of 0,89 mm’(b = 1,5 mm; h =
1,5 mm) (Table 6). In comparison the most tubercles found on male P. asper was 53 (6 snout; 6 nares;

157

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 2 30 JUNE 1994

GSI — I — Snout tub.

Fig. 5. Comparison of the seasonal occurrence of definitive snout tubercles on male Pseudobarbus asper (Boulenger, 191 1)

(n=204) with the mean monthly gonadosomatic index (GSI) of male P. asper for the period July 1987 to April 1989.
(Definitive tubercle = 1; bud tubercle = 0,5).

3 above orbits; 4 anterior dorsal; 34 posterior dorsal) on a 69,4 mm SL specimen collected during
March 1989 (Table 7). There was a highly significant difference (t= 5,50985, d.f. = 18, P < 0,001 of
having equal numbers) between number of tubercles on P. afer males compared to P. asper males.
The largest snout tubercle for P. asper males had a volume of 0,26 mm^ (Table 8). There was a highly
significant difference (t = 12,5016, df = 18, P < 0,001 of having an equal volume) between volume
sizes of P. afer male tubercles compared to the largest P. asper tubercles.

There was an increase in the number of tubercles with increasing length for both males (Fig. 6)
and females of the two species. The lower correlation coefficient for P. asper males and the very low
values for the females of both species probably indicated the lack of significance of tubercles for
these groups compared to P. afer males.

Females P. afery = -12,1037 + 0,233274x (n = 60; r’ = 0,323)

P. asper y = -33,3291 -i- 0,694969x (n = 91; r= = 0,583)
where x = SL (mm); y = number of tubercles

For each separate cluster of tubercles regression analyses were run (Tables 9 and 10). The highest
correlation coefficient in P. afer males was in the snout tubercles to length and the lowest in the

158

CAMBRAY: NUPTIAL TUBERCLES PSEUDOBARBUS: SEASONAL OCCURRENCE AND DISTRIBUTION

Table 3. Summary of the seasonal pattern of tubercle formation and wear in 118 Pseudobarbus afer
(Peters, 1864) females collected over a 22 month period in the Wit River, Gamtoos River System.

Date

n

Size range
SL (mm)

Tubercles^

Scales^

Rays^

July 1987

5

52,4-62,5

-

-

-

Aug.

5

57,6-63,0

Sp (few)

-

-

Sept.

5

60,8-70,7

B+Sp

-

-

Oct.

8

67,0-76,1

B+Sp

-

-

Nov.

5

61,0-68,0

W+Sp

-

-

Dec.

5

64,3-68,7

Sp

-

-

Jan. 1988

5

46,8-51,8

-

-

-

Feb.

5

56,8-72,4

B

-

-

Mar.

5

53,4-68,7

B

-

+ (few)

May

5

54,2-61,2

-

-

-

June

5

55,5-60,0

W (few)

-

-

July

5

59,1-72,8

Sp (few)

-

-

Aug.

5

54,9-68,2

-

-

-

Sept.

5

60,6-64,5

B (few)

-

-

Oct.

5

56,4-61,1

B (few)

-

-

Nov.

5

53,2-64,9

-

-

-

Dec.

5

53,4-70,1

Sp

-

-

Jan. 1989

4

52,7-61,1

-

-

-

Feb.

5

54,0-70,5

-

-

-

Mar.

11

58,0 -79,9

B

-

A (few)

Apr.

TOTAL

5

118

60,3-65,4

B

1. B = bud; D = definitive; S = scars; Sp = spots; W = worn; - = absent

2. - = absent; + = present

3. - = absent; + = present

anterior dorsal cluster. Similarly in the tubercle clusters on the head of P. asper the snout tubercles
had the highest and the anterior dorsal cluster had the lowest correlations with length. In the females
of both species there were low correlations for most clusters which had tubercles (Table 10).

The slope of the regression of total tubercle number on standard length of P. afer males was
significantly different from P. asper males (F = 29,5848; d.f. = 345; P<0,001; Fig. 6).

In the female sample a 72,4 mm SL P. afer had the most tubercles (= spots) for that species.
There were 29 spots of which 3 were snout, 4 nares, 4 above orbits, 0 anterior dorsal and 18 posterior
dorsal. In the P. asper female sample a 72,0 mm SL specimen collected in November 1988 had the
most tubercles (including spots). There were 43 spots and buds positioned as follows, 3 (spots) snout,
7 (spots) nares, and 33 (buds) in the posterior dorsal position.

159

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 2 30 JUNE 1994

Fig. 6. Relationship between body length and the number of head tubercles in malePseudobarbus afer {Pe.le.TS, 1864) (n=145)
and P. asper (Boulenger, 1911) (n=204) collected between September and February 1987/1988 and 1988/1989;

P. afer y = -81,2053 + l,99929x (n = 145; F = 0,878)

P. asper y = -43,659 + l,20292x (n = 204; F = 0,722)
where x = SL (mm); y = number of tubercles

The largest snout tubercle measured was found in the P. afer male sample. The volume of the
tubercle was 0,89 mm’ on a 78,2 mm SL specimen (Table 6). In comparison the largest tubercle in
the P. asper sample was on a 72,3 mm SL male collected in November 1987. The volume of that
snout tubercle was 0,26 mm’ (Table 8). The largest P. afer snout tubercle was therefore 3,42 times
the volume of the largest P. asper tubercle.

In contrast to the males, the female P. asper had the most numerous tubercles (= spots or buds)
(Tables 1 1 and 12). The mean number of tubercles for the P. asper sample was 30,7 (SD = 6,3) (Table
12) as compared to 12,1 (SD = 7,7) for P. afer females (Table 11). Buds or spots were present in the
snout, nares and posterior dorsal positions for most specimens. Few spots or buds for both species
were observed in the above orbits and interior dorsal positions (Tables 11 and 12). Volumetric
calculations were not made for the females as the buds are usually less than 0,002 mm’. The largest
bud measured on a female was on a specimen of P. asper ( v = 0,012 mm’, b = 0,3; h = 0,5).

Development of tubercles was not always symmetrical. In P. afer males, snout tubercles ranged
up to 7 on both sides (70,5 mm SL). Of 248 specimens 72 had more tubercles on one side than the
other (X’ = 145,806, 1 d.f., P < 0,0000). In only two specimens was the difference greater than one.

160

CAMBRAY: NUPTIAL TUBERCLES IN PSEUDOBARBUS: SEASONAL OCCURRENCE AND DISTRIBUTION

Table 4. Summary of the seasonal pattern of tubercle formation and wear in 157 Pseudobarbus asper
(Boulenger, 1911) females collected over a 22 month period in the Groot River, Gamtoos
River System.

Date

n

Size range
SL (mm)

Tubercles^

Scales^

Rays^

July 1987

3

54,1-56,0

-

-

-

Aug.

3

69,8-76,0

-

-

-

Sept.

11

50,1-69,6

B

-

-

Oct.

4

42,9-57,4

B

-

-

Nov.

6

49,3-63,2

B+Sp

-

-

Dec.

4

50,4-74,3

B-i-Sp

-

-

Jan. 1988

5

64,7-70,8

Sp

-

-

Feb.

15

50,1-74,1

B+Sp

+ (few)

+ (few)

Mar.

13

45,1-57,8

B+Sp

-

-

Apr.

8

56,7-61,9

B (few)

-

+ (few)

May

4

50,8-60,1

-

-

-

June

5

67,8-72,3

-

-

-

July

4

54,0-63,5

-

-

-

Aug.

4

61,9-74,4

-

-

-

Sept.

5

60,5-79,9

B

-

-

Oct.

5

51,3-70,0

B-i-Sp

-

-

Nov.

8

65,5-72,2

B-i-Sp

+ (few)

+ (few)

Dec.

16

54,1-70,7

B+Sp

-

-

Jan. 1989

5

54,5-70,9

B+Sp

-

-

Feb.

9

60,7-76,5

B+Sp

-

-

Mar.

12

62,2-75,2

B+Sp+S

-

-

Apr.

TOTAL

8

157

59,1-70,9

B+Sp

1. B = bud; D = definitive; S = scars; Sp = spots; W = worn; - = absent

2. - = absent; + = present

3. - = absent; + = present

Forty-two of the 72 specimens had more tubercles on the left side than on the right (X^ = 2,0, 1 d.f.,
P < 0,157299). In P. asper males snout tubercles ranged up to 4 on each side. Of 320 specimens
examined, 80 had more tubercles on one side than the other (X^ = 200,0, 1 d.f., P < 0,0000). In three
specimens the difference was two tubercles. Thirty-seven specimens had more tubercles on the left
side than on the right side (X^ = 0,225, 1 d.f., P < 0,635256).

The first set of tubercles to show any wear were the snout tubercles followed by the nare clusters
for both species. The posterior dorsal tubercles often did not show any sign of wear. No further
deductions could be made in the SEM microwear part of this study as no clear patterns of differences
in wear were revealed.

161

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 2 30 JUNE 1994

Table 5. Summary of ten Pseudobarbus afer (Peters, 1864) males with the most head tubercles and
their position.

Date

SL (mm)

Snout

Nares

Above

orbits

Ant.

dorsal

Post.

dorsal

Total

Nov. 1987

70,5

14

7

12

6

49

88

Dec. 1988

70,1

7

6

10

5

39

67

Oct. 1987

70,8

10

5

8

5

34

62

Feb. 1989

70,5

9

8

7

5

33

62

Oct. 1987

78,2

11

7

7

7

29

61

Feb. 1989

66,2

8

6

8

5

34

61

Sept. 1987

71,5

6

6

6

4

38

60

Sept. 1988

70,5

6

6

7

7

31

57

Jan. 1989

69,6

6

6

10

4

31

57

Dec. 1987

63,4

9

6

5

7

27

54

Means

70,1

8,6

6,3

8

5,5

34,5

62,9

SD

3,8

2,6

0,8

2,1

1,2

6,3

9,5

Table 6. Volume of the ten largest snout tubercles found on Pseudobarbus afer (Peters, 1864) males
collected in the Wit River, Gamtoos River System.

Date

SL (mm)

Base (mm)

Height (mm)

Volume (mm^)

Oct. 1987

78,2

1,5

1,5

0,89

Nov. 1987

58,0

1,6

1,0

0,67

Nov. 1987

70,5

1,3

1,5

0,67

Sept. 1987

63,2

1,25

1,05

0,43

Dec. 1987

63,4

1,1

1,1

0,35

Dec. 1987

58,1

1,2

0,9

0,34

Dec. 1987

49,5

1,2

0,85

0,32

Oct. 1987

60,9

1,2

0,7

0,26

Oct. 1987

57,3

1,2

0,6

0,23

Nov. 1988

59,7

1,1

0,6

0,19

Means

61,9

1,27

0,98

0,44

SD

7,9

0,16

0,33

0,23

162

CAMBRAY: NUPTIAL TUBERCLES IN PSEUDOBARBUS: SEASONAL OCCURRENCE AND DISTRIBUTION

Table 7. Summary of ten Pseudobarbus asper (Boulenger, 1911) males with the most head tubercles
and their position.

Date

SL (mm)

Snout

Nares

Above

orbits

Ant.

dorsal

Post.

dorsal

Total

Mar. 1989

69,4

6

6

3

4

34

53

Dec. 1987

69,9

6

6

3

6

30

51

Nov. 1987

72,3

8

6

2

2

31

49

Dec. 1987

64,6

6

6

4

4

24

44

Jan. 1988

63,1

6

6

1

1

29

43

Aug. 1988

70,4

8

6

3

5

20

42

Sep. 1987

62,4

6

6

4

1

24

41

Apr. 1989

66,6

6

6

5

2

21

40

Dec. 1988

59,7

6

6

1

8

18

39

Feb. 1989

65,9

8

6

3

0

22

39

Means

66,4

6,6

6

2,9

3,3

25,3

44,1

SD

4,0

1,0

0,0

1,3

2,5

5,4

5,1

Table 8. Volume of the ten largest snout tubercles found on asper (Boulenger, 1911)

males collected in the Groot River, Gamtoos River System.

Date

SL (mm)

Base (mm)

Height (mm)

Volume (mm^)

Nov. 1987

72,3

1,2

0,7

0,26

Dec. 1988

64,5

1,2

0,7

0,26

Feb. 1989

66,1

1,0

0,9

0,24

Jan. 1989

55,5

1,0

0,8

0,21

Sep. 1988

58,8

1,0

0,7

0,18

Dec. 1988

60,4

1,0

0,7

0,18

Dec. 1988

56,4

0,9

0,8

0,17

Feb. 1988

56,0

1,0

0,5

0,13

Dec. 1988

62,3

0,8

0,7

0,12

Dec. 1988

59,7

0,8

0,7

0,12

Means

61,2

1,0

0,72

0,19

SD

5,3

0,14

0,1

0,05

163

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 2 30 JUNE 1994

Table 9. Linear regression analyses of the number of tubercles in the different clusters (see text) on
the heads to length ofmalePseudobarbusafer(?et&is, 1864) andP. asper (Boulenger, 1911).

Tubercle cluster

n

a

b

r

P. afer 6 c?

Total

143

-80,332

1,980

0,878

Snout

143

-12,664

0,307

0,883

Nare

143

-7,951

0,233

0,748

Orbit

143

-12,968

0,296

0,867

Ant. dorsal

143

-7,784

0,181

0,739

Post, dorsal

143

-38,965

0,964

0,773

P. asper cj 6

Total

202

-44,017

1,210

0,722

Snout

202

-6,366

0,206

0,685

Nare

202

-6,127

0,195

0,601

Orbit

202

-3,052

0,067

0,504

Ant. dorsal

202

-3,257

0,075

0,407

Post, dorsal

202

-25,216

0,666

0,643

Table 10. Linear regression analyses of the number of tubercles in the different clusters (see text) on the
heads to length of female Pseudobar bus afer (Peters, 1864) and P. asper (Boulenger, 1911).

Tubercle cluster

n

a

b

r

P. afer ? $

Total

60

-12,104

0,233

0,323

Snout

60

-2,195

0,040

0,404

Nare

60

-6,463

0,117

0,525

Orbit

60

-33,329

0,695

0,583

P. asper ? $

Total

91

-33,329

0,695

0,583

Snout

91

-5,558

0,113

0,484

Nare

91

-4,587

0,093

0,431

Post, dorsal

91

-22,671

0,479

0,523

164

CAMBRAY: NUPTIAL TUBERCLES IN PSEUDOBARBUS: SEASONAL OCCURRENCE AND DISTRIBUTION

Table 11. Summary of t&nPseudobarbus afer (Peters, 1864) females from Wit River, Gamtoos River
system, with the most head tubercles and their position.

Date

SL (mm)

Snout

Nares

Above

orbits

Ant.

dorsal

Post.

dorsal

Total

Feb. 1988

72,4

3

4

4

0

18

29

Dec. 1987

68,4

2

8

2

2

6

20

Feb. 1988

57,0

0

0

0

0

16

16

Mar. 1989

70,7

2

5

0

1

6

14

Feb. 1988

58,1

0

0

0

0

10

10

Mar. 1989

63,9

2

4

0

0

1

7

Sep. 1987

61,7

2

3

0

0

2

7

Feb. 1988

56,8

0

0

0

0

6

6

Sep. 1987

70,3

0

4

1

0

1

6

Oct. 1987

72,6

2

4

0

0

0

6

Means

65,2

1,3

3,2

0,7

0,3

6,6

12,1

SD

6,4

1,2

2,6

1,3

0,7

6,3

7,7

Table 12. Summary of ten Pseudobarbus asper (Boulenger, 1911) females from the Groot River,
Gamtoos River system, with the most head tubercles and their position.

Date

SL (mm)

Snout

Nares

Above

orbits

Ant.

dorsal

Post.

dorsal

Total

Nov. 1988

72,0

3

7

0

0

33

43

Mar. 1989

68,5

6

6

0

0

29

41

Feb. 1988

74,1

2

4

0

0

27

33

Nov. 1988

72,2

4

3

4

0

18

29

Feb. 1989

76,5

6

4

0

0

18

28

Feb. 1988

66,4

2

4

0

0

21

27

Feb. 1988

65,2

2

3

0

0

22

27

Nov. 1988

70,1

6

4

0

0

17

27

Mar. 1989

75,2

4

4

0

0

19

27

Sep. 1987

68,0

2

4

0

0

19

25

Means

70,8

3,7

4,3

0,4

0

22,3

30,7

SD

3,8

1,8

1,3

1,3

0

5,5

6,3

165

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 2 30 JUNE 1994
DISCUSSION

Nuptial or breeding tubercles can develop on the head, fins and bodies of fish. They are present
in regions which come into contact with another male or female fish. The seasonal occurrence and
disappearance of tubercles follows the reproductive cycle and is probably under seasonal hormonal
control. There are several possible functions for these structures on males: fighting other males or
invaders, the white colour of the tubercles might denote spawning readiness; used to maintain body
contact during spawning; stimulation of females during spawning; protection of the body and fins
from damage during nest construction; tubercles on the ventral surface of females may protect the
anus and urogenital papillae against mechanical damage during egg laying; to hold the substrate in
swift waters during the spawning act; and to take care of the eggs, nest and young (Hubbs and Cooper,
1936; Raney, 1940; Fabricius and Gustafson, 1955; Branson, 1961, 1962; Koehn, 1965; Braasch and
Smith, 1967; Wiley and Collette, 1970; Collette, 1977; Kratt and Smith, 1978; Cambray and Stuart,
1985). The tubercles of the males may reflect a pearlescent hue which may attract females. Since
there are different patterns for different species, tubercles may aid in species and sex recognition.
Collette (1977) suggested that breeding tubercles originally evolved to enable breeding individuals
to maintain close contact during spawning. In fishes which breed in fast-flowing waters close contact
would ensure fertilization of the eggs. The possible functions of the nuptial tubercles on the rays,
scales and head for male P. afer and P. asper are summarised in Table 13.

Table 13. Possible functions of the breeding tubercles of Pseudobar bus afer (Peters, 1864) and

P. (Boulenger, 1911).

Function

P. afer
(clear water)

P. asper
(turbid)

Maintain contact during spawning (tactile;
scale and ray tubercles)

Possible

Possible

Holding to the substrate during spawning
in swift flowing waters

Possible

Possible

(scale, ray and head tubercles)
Stimulation of females (head tubercles)

Possible

Possible

Excavating a nest (head tubercles)

Unlikely

Unlikely

Defense of nests (head tubercles)

Unlikely

Unlikely

Fighting other males (head tubercles)

Possible

Possible

Visual threat (head tubercles)

Possible

Unlikely

Sex recognition (head tubercles)

Possible

Unlikely

Species recognition (head tubercles)

Possible

Unlikely

Tubercles were better developed on males of both P. afer and P. asper than females. The largest
tubercles were found on P. afer males and were up to 3,42 times the volume of the largest tubercles
on male P. asper (Tables 6 and 8). As in the present study Kratt and Smith (1978) and Witkowski
(1982 in Witkowski and Ragowska, 1991) found that tubercles were better developed on those fish

166

CAMBRAY: NUPTIAL TUBERCLES IN PSEUDOBARBUS: SEASONAL OCCURRENCE AND DISTRIBUTION

which have spawned several times than on first time spawners. Secondary sexual characteristics, such
as epidermal tubercles and breeding coloration, all require resources which must be diverted away
from growth and survival requirements (Miller, 1979; Wootton, 1985). The formation of and the
subsequent behaviour of fish with tubercles would probably be energetically costly which would
reduce growth. If females developed large tubercles with associated behaviour then this allocation
of energy to secondary sexual characteristics could possibly reduce the future production of offspring.
This would favour the development of these characters on males only (Wootton, 1990). Large females
of P. afer and P. asper have head tubercles which are never definitive and their presence on
reproductively active females was difficult to explain. Why should the larger females of P. afer and
P. asper have invested any energy into producing tubercles? The females with tubercles had high
gonadosomatic indexes and therefore were not senescent and there was no evidence of
hermaphroditism. When tubercles occurred on the head they followed the pattern exhibited on the
males (Fig. 2). That very few females were found to have tubercles on their rays indicated that they
did not use them for maintaining position during mating and the occurrence of rudimentary head
tubercles on the large females may merely indicate a hormonal change as the fish grew older.
Pseudobarbus asper females were found to have more head tubercles than P. afer females. There
was no obvious adaptive significance to the bearer and, if anything, the allocation of resources to
produce weakly developed tubercles would decrease female fitness.

The nuptial tubercles on the pectoral fins of some cyprinids maybe used in a ‘pectoral fin lock’
in which the male clasps the female (Branson, 1962). The tubercles on the fin can be recurved which
would facilitate clasping. Boulenger (191 1) noted sexual dimorphism in the pectoral fins of P. asper
with males having longer pectoral fins than females. Sexual dimorphism of the fins and length of
pectorals relative to the base of the pelvics was later found to be evident in most of the flexible-rayed
redfins (Barnard, 1943). Male P. afer and P. asper have broader and more rounded pectoral fins than
females (Skelton, 1988). The males may use the larger, longer and broader pectoral fins with more
tubercles for clasping the female during spawning (pectoral fin lock). Cambray and Meyer (1988),
Cambray (1991) and Cambray (1992) have noted that P. afer, P. asper and P. quathlambae bred in
mid-charmel riffles after an increase in flow. Clasping of the pectoral fins, aided by increased size
and tubercles on the rays of these fins, would be advantageous to ensure fertilization of the eggs
especially in the fast-flowing water of the spawning habitat. The tubercles present on the scales would
also help to maintain a spawning embrace. The large tubercles on the head must have a different
purpose and would not be used for clasping. It is therefore in the head tubercles that one may expect
differences in size and/or density between species inhabiting clear or turbid waters.

The head tubercles may be for interactions between males to acquire mates (intrasexual selection)
or for interactions between the sexes to choose mates (intersexual selection). The dorsal tubercles of
the head and trunk in many species are usually fairly well developed, possibly because they are used
for fighting invaders or other males, and attracting and nudging females during spawning. Tubercle
rows over the orbits might be a protective mechanism during agonistic behaviour of P. afer or P.
asper males, as was suggested for some Nocomis species (Lachner and Jenkins, 1971). Because of
the highly turbid waters in which P. asper lives, it would be unlikely that this species used head
tubercles for visual threat, sex recognition or species recognition (Table 13). But as P. afer inhabits
clear waters they could be used for the above functions (Table 13). If the tubercles were used to nudge
the female during spawning then that would not help to explain the smaller tubercles of P. asper
males. There was no obvious evidence for how the tubercles were used in the flexible-rayed redfins
and the scanning electron microscopy microwear study did not elucidate the probable cause(s) of the
observed wear of the head and ray tubercles during the reproductive season.

167

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 2 30 JUNE 1994

One of the most notable characters of sexually mature male P. burchelli were the large conical
head tubercles (Cambray and Stuart, 1985) which are more highly developed than in most other
Pseudoharbus and small Barbus species (Skelton, 1980). Cambray and Stuart (1985) have noted the
seasonal occurrence, wear, number and size of the breeding tubercles of both male and female P.
burchelli. As in P. afer and P. asper the head tubercles of P. burchelli were more developed on males
as compared to females. Larger males were found to have larger tubercles. The large conical tubercles
on the heads of the males (> 80 mm FL) began to erupt in May. The tubercles were found to erupt
through the worn tubercle bases of the previous season. Some tubercles were well formed by August
and the first signs of wear occurred in November which coincided with the spawning season. In P.
burchelli the wearing and/or sloughing off of the tubercles continued until April when only the bases
of the tubercles were visible. Cambray and Stuart (1985) suggested that this seasonal cycle of tubercle
formation and wear added support to the hypothesis that the tubercles were associated with
reproduction in redfin minnows (Skelton, 1980). It was proposed by Cambray and Stuart (1985) that
the wearing as the season progressed suggested that the tubercles are used when defending a territory.
The keratin caps get worn off as the breeding season progresses.

The lifetime reproductive success of individuals may be influenced by the individual variation
in tubercle number and/or size. In male P. afer the head tubercles were both volumetrically larger
and more numerous than those of P. asper. That would suggest that in the clear water habitat of P.
afer size and number of tubercles were important for reproductive success and may have influenced
mate choice. The presence of tubercles may signal information about the genetic quality of the bearer,
or it may only be that the presence of large tubercles and bright red coloration makes the bearer more
likely to be chosen by a potential mate. The ultimate factors which have lead to the evolution of the
secondary sexual characters, such as tubercles, are still a subject of debate (Wootton, 1990). In New
World cyprinids, sexual differences are usually more pronounced in those species that are highly
territorial, have some form of parental care, or carry out their life cycles in high-gradient and clear
rivers (Mayden, 1991). There is no known parental care in Pseudobarbus species but most of the
species, such as P. afer and P. burchelli, inhabit clear water systems.

Quantitative differences in secondary sexual characters of a species may play a role in mate
discrimination by reinforcing speciation once populations that had diverged in allopatry came back
into contact (Dobzhansky, 1940). The noted differences in secondary sexual characters of P. afer and
P. asper males may reinforce speciation in the Groot River as there is no physical barrier separating
these two species (Cambray, 1992).

Pseudobarbus afer, P. asper and P. quathlambae (Cambray and Meyer, 1988; Cambray, 1991;
Cambray, 1992) all spawned after an increase in river flow and their spawning habitat was mid-
channel amongst boulders. Ozarkian minnows which were very active spawners or which spawned
in rapidly flowing waters were found to have more extensively developed, scattered tubercles than
sedentary species which had tubercles in lines (Branson, 1962). Cyprinids with unspecialized breeding
habits usually have small tubercles distributed over the head in no particular pattern (Lachner and
Jenkins, 1971). This may indicate that in the Pseudobarbus species with large tubercles, such as P.
afer and P. burchelli, there may be more specialized breeding habits compared to species such as P.
asper and especially P. phlegethon which have smaller or vestigial tubercles. This difference requires
further investigation.

It is suggested that in P. afer and P. asper males large head tubercles are used for display prior
to breeding, whereas during breeding the tubercles on the scales and fin rays maintain contact with
females, as do the larger pectoral fins of the males, in the riffle spawning habitat. The occurrence of
fewer and smaller tubercles on P. asper males may be because in turbid waters visual characters have

168

CAMBRAY; NUPTIAL TUBERCLES IN PSEUDOBARBUS: SEASONAL OCCURRENCE AND DISTRIBUTION

less of a selective advantage than in a clear mountain stream (Table 13). That the head tubercles are
still relatively large in P. asper may be a phylogenetic constraint within the sister species pair.

ACKNOWLEDGEMENTS

Eve Cambray helped in the field and also made suggestions on a number of drafts of this paper.
Professors Mike Bruton and Tom Hecht made many useful comments on a draft of this work. Garth
and Conrad Cambray helped in the field. This work was made possible by a research grant from the
Foundation for Research Development. The Director of the Albany Museum permitted me to
undertake this work. A grant from the Anglo American and De Beers Chairman’s Fund enabled me
to present this paper at an international conference.

REFERENCES

Banarescu, P.andCoAD, B. W. 1991. Cyprinids of Eurasia. In: Winfield, I. J. and J. S. Nelson eds, Cyprinid fishes systematics,
biology and exploitation. London: Chapman & Hall, pp. 127-155.

Barnard, K. H. 1943. Revision of the indigenous freshwater fishes of the S.W. Cape region. Ann. S. Afr. Mas. 36(2): 101-262.
Boulenger, G. a. 1911. Catalogue ofthefresh-water fishes of Africa in the British Museum (Natural History), Part 2. London:
British Museum.

Branson, B. A. 1961. Observations on the distribution of nuptial tubercles in some catostomid fishes. Trans. Kans. Acad. Sci.
64: 360-372.

Branson, B. A. 1962. Observations on the breeding tubercles of some Ozarkian minnows with notes on the barbels of Hybopsis.
Copeia 1962(3): 532-539.

Braasch, M. E. and Smith, P. W. 1967. The life history of the slough darter, Etheostoma gracile (Pisces, Percidae). III. Nat.
Hist. Surv., Biol. Notes. 58: 1-12.

Breder, C. M. and Rosen, D. E. 1966. Modes of reproduction in fishes. Neptune City: T.F.H. Publications.

Cambray, J. A. 1991. The effects of fish spawning and management implications of impoundment water releases in an
intermittent South African river. Regul. Rivers 6: 39-52.

Cambray, J. A. 1992. A comparative study of the life histories of the sister species, Pseudobarbus afer and Pseudobarbus
asper, in the Gamtoos River system. South Africa. Unpublished Ph.D. thesis, Rhodes University, Grahamstown.
Cambray, J. A. and Meyer, K. 1988. Early ontogeny of an endangered, relict, cold-water cyprinid from Lesotho, Oreodaimon
quathlambae (Barnard, 1938). Rev. Hydrobiol. trop. 21(4): 309-333.

Cambray,!. A. and Stuart, C. T. 1985. Aspects ofthe biology of a rare redfin minnow, Barbas burchelli (Pisces, Cyprinidae),
from South Africa. S. Afr. J. Zool. 20: 155-165.

Collette, B. B. 1965. Systematic significance of breeding tubercles in fishes of the family Percidae. Proc. U.S. natn. Mas.
117: 567-614,

Collette, B. B. 1977. Epidermal breeding tubercles and bony contact organs in fishes. Symp. zool. Soc. Lond. 39: 225-268.
Dobzhansky, T. 1940. Speciation as a stage in evolutionary divergence. A/n. Nat. 74: 312-321.

Fabricius, E. and Gustafson, K. J. 1955. Observations on the spawning behaviour of the grayling, Thymallus thymallus (L.).

Anna. Rep. Inst. Freshwater Res. (Drottningholm) 36: 75-103.

Howes, G. J. 1980. The anatomy, phylogeny and classification of bariliine cyprinid fishes. Bull. Br. Mus. nat. Hist. (Zool.)
37(3): 129-198

Hubbs, C. L. and Cooper, G. P. 1936. Minnows of Michigan. Cranbrook Inst. Sci. Bull. 8: 1-84.

JuBB, R. A. 1967. Freshwater fishes of southern Africa. Cape Town: Balkema.

Koehn, R. K. 1965. Development and ecological significance of nuptial tubercles of the red shiner, Notropis lutrensis. Copeia
1965(4): 462-467.

169

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 2 30 JUNE 1994

Kratt, L. F. and Smith, R. J. F. 1978. Breeding tubercles occur on male and female Arctic grayling (Thymallus articus).
Copeia 1978(1): 185-188.

Lachner, E. A. and Jenkins, R. E. 1971. Systematics, distribution and ei'oiution of the Chub genus Nocomis Girard (Pisces,
Cyprinidae) of eastern United States, with descriptions of new species. Smithson. Contrib. Zool. 85: 97pp.

Lande, R. 1982. Rapid origin of sexual isolation and character divergence in a dine. Evolution 36: 213-223.

Mayden, R. L. 1991. Cyprinids of the New World. In: Winfield, I. J. and J. S. Nelson eds, Cyprinid fishes systematics, biology
and exploitation. London: Chapman and Hall, pp. 240-263.

Miller, P. J. 1979. A concept of fish phenology. Symp. zool. Soc. Lond. 44: 1-28.

Raney, E. C. 1940. The breeding behaviour of the common shiner, Notropis cornutus (Mitchell). Zoologica (NY.) 25: 1-14.
Reid, G. McG. 1985. A revision of African species of Labeo (Pisces: Cyprinidae) and a redefinition of the genus. Theses
Zoologicae 6: 1-322.

Skelton, P. H. 1974. The life colours and nuptial tubercles of Oreodaimon quathlambae (Barnard, 1938) (Pisces, Cyprinidae).
Ann. Cape Prov. Mas. (nat. Hist.) 9(12): 215-222.

Skelton, P. H. 1980. Systematics and biogeography of the redfin Barbus species (Pisces: Cyprinidae) from southern Africa.
Unpublished Ph.D. thesis, Rhodes University, Grahamstown.

Skelton, P. H. 1988. A taxonomic revision of the redfin minnows (Pisces, Cyprinidae) from southern Africa. A«n. Cape Prov.
Mus. (nat. Hist.) 16(10): 201-307.

Vladykov, V. D., Renaud, C. B. and Laeramboise, S. 1985. Breeding tubercles in three species of Gadus (Cods). Can. J.
Fish. Aquat. Sci. 42: 608-615.

Wiley, M. L. and Collette, B. B. 1970. Breeding tubercles and contact organs in fishes: their occurrence, structure and
significance. Bull. Am. Mus. nat. Hist. 143(3):143-216.

WiTKOWSKi, A. and Rogowska, M. 1991. Breeding tubercles in some European cyprinid fishes (Osteichthyes, Cypriniformes:
Cyprinidae). Zool. Abh. Mus. Tierd. Dresden 46(9): 131-156.

WooTTON, R. J. 1985. Energetics of reproduction. In: Tytler, P. and P. Calow eds, Fish energetics new perspectives. London:
Croom Helm, pp. 231-254.

Wootton, R. j. 1990. Ecology ofTeleost fishes. London: Chapman & Hall.

Zar, j. H. 1984. Biostatistical analysis. 2nd edn. Englewood Cliffs: Prentice-Hall.

170

INSTRUCTIONS TO AUTHORS

LAYOUT of recent issues should be examined for details of acceptable style and format.

MANUSCRIPTS should be submitted on floppy disk accompanied by three print-outs
(spacing 1!4 or 2). In exceptional cases typed manuscripts will be accepted. These should be
submitted in triplicate, typed on A4 size paper, double-spaced, on one side of the paper only
and with wide margins all round.

Only generic and specific names should be underlined to indicate italics. All other marking
up should be left to the editor.

Footnotes should be avoided as far as possible.

All measurements should be metric.

Systematic papers should follow the rules of the International code of zoological
nomenclature or Xht International code of botanical nomenclature, as applicable.

ILLUSTRATIONS should bear the author’s name, title of paper and number in soft pencil on
the reverse side. Diagrams, line drawings and photographs are all regarded as figures, should
be termed as such and should be numbered in a single series.

In addition to the original illustrations a photocopy of each should be submitted.

Diagrams and line drawings should be reducible to a size not exceeding 14 x 16 cm (17,5 cm
including captions). Photographs for half-tone blocks should be final size, of high contrast
and printed on glossy paper. Colour illustrations are only printed under special circumstances
and their financing must be arranged by the author.

A scale should appear with all illustrations or the magnification should be given in the
caption.

A list of captions should be submitted on a separate sheet.

TABLES should be submitted on separate sheets. Foldouts and tables printed sideways should
be avoided.

REFERENCES should be listed alphabetically at the end of the paper. Abbreviations of
periodical titles should conform with those in the World list of periodicals (4th ed.). Layouts
must be as follows:

Alexander, J. 1969. The indirect evidence for domestication. In: Ucko, P. J. and G. W.
Dimbleby eds, The domestication and exploitation of plants and animals. London:
Duckworth, pp. 123-129.

De Winter, B. 1969. Plant taxonomy today. S.Afr.J.Sci. 65 (8): 229-242.

JuBB, R. A. 1967. Freshwater fishes of southern Africa. Cape Town: Balkema.

GALLEY PROOFS are sent (when practicable) to authors for checking printer’s errors.
Adjustments of language, style and punctuation and any other amendments of the original
manuscript, if made at this stage, may be charged to authors.

AUTHOR’S COPIES (50) are issued free of charge. When more than one author is involved
the fifty free copies will be sent to the senior author. Additional copies ordered before printing
are supplied at a reduced cost.

CADAR. P,E.-K887

/

1'

*> ■■

'i* . - ■

'i.

1

>,

.1

j

1^ .

• ■■ <

. M

ol3

ISSN 0570-1880

Annals of the

Cape Provincial Museums

Natural History

Ann. Cape Prov. Mus. (nat. Hist.)

Volume 19 Part 3 28th February 1995

I ■

f JUM 26 1995

Published jointly by the Cape Provincial Museums
at the Albany Museum, Grahamstown, South Africa

ANNALS OF THE CAPE PROVINCIAL MUSEUMS

These Annals are published in two series, Natural History and Human Sciences, the latter series
covering cultural history, ethnology, anthropology and archaeology. They are issued in parts at
irregular intervals as material becomes available.

The primary objective of these Annals is to disseminate the results of research work undertaken by
staff members of the Cape Provincial Museums. However, a manuscript by any author is eligible
for publication provided it is based wholly or partly on the collection/s of one or more of these
museums or if the material on which the work is based is wholly or partly deposited in one of
these museums.

Parts are obtainable from the Librarian of the Albany Museum. Correspondence in connection
with these Annals should be addressed to the Editor, Albany Museum, Grahamstown 6140.

Editor

Dr F. W. GESS

Assistant Editor
Dr S. K. GESS

A new subspecies of the freshwater halacarid mite Lobohalacarus weberi
(Romijn and Viets) (Halacaridae, Acari) from a Southern Atlantic Ocean island

by

ILSE BARTSCH

(Biologische Anstalt Helgoland, Notkestrasse 31, 22607 Hamburg, Germany)

ABSTRACT

Lobohalacarus weberi tristanensis subsp. nov., found on Inaccessible, Tristan da Cunha Islands,
is described and compared with populations of L. weberi weberi (Romijn and Viets) from Europe.
Representatives of the Lobohalacarus weberi complex primarily inhabit subterranean waters. A short
survey of species and subspecies of the L. weberi complex is given and means of dispersal to
Inaccessible Island are discussed.

INTRODUCTION

Although comprising mostly marine species, the superfamily Halacaroidea includes several fresh
water species. At present, about 700 marine and 50 limnic species are described. The freshwater
halacarid species differ from marine species in having external genital acetabula. The freshwater
species can be assigned to several halacarid subfamilies.

AREA OF INVESTIGATION, MATERIAL AND ABBREVIATIONS

The Tristan da Cunha island group lies about midway between South Africa and South America,
in a belt with prevailing westerly winds and easterly water currents (Baker et al., 1964). The islands,
of volcanic origin, are located east of the mid-Atlantic Ridge. Inaccessible, the most westerly island,
has an area of about 10 km and reaches a height of 550 m (Baker, 1973). The island is resting on an
older eroded volcanic cone, about 18 million years old (Baker et al., 1964; Chevallier et al., 1992).

The halacarid specimens were collected on the west side of Inaccessible Island, from a small,
narrow freshwater stream bordered by Spartina arundinacea. The sampling site is less than 1 km
from the underground source of the stream; the water is clear, pH 7.0 (Barber-James, in prep).

Abbreviations used in the descriptions are: AD, anterior dorsal plate; AE, anterior epimeral plate;
br, bristle; GA, genitoanal plate; GO, genital opening; OC, ocular plate(s); P, palp, P-2, 2nd palpal
segment; pas, parambulacral setae; PD, posterior dorsal plate; PE, posterior epimeral plate; sp, spine.
Legs are numbered I to IV, leg segments 1 to 6 (I-l, 1-2, 1-3, 1-4, 1-5, 1-6 = trochanter, basifemur,
telofemur, genu, tibia and tarsus on leg I).

The type material is deposited in the Albany Museum, Grahamstown, South Africa.

TAXONOMY

Lobohalacarus weberi tristanensis subsp. nov.

Material

One holotype female, Albany Museum, Grahamstown, TDC II H; Inaccessible Island, from

171

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 3 28 FEBRUARY 1995

narrow freshwater stream; collected by H.M. Barber-James, 23 October 1989.

Three paratype females, one larva, Albany Museum, Grahamstown, TDC I G; type locality. Two
females, author’s halacarid collection, type locality.

Description

Holotype feifiale. Idiosoma 310 pm long. Integument of dorsal plates foveate and delicately
punctate. Anterior rounded idiosoma formed by dorsal portion of AE which is fused in midline.
Anterior AD contiguous but not broadly joined with that area. AD 92 pm long; with an area like an
inverted Y slightly raised (Fig. 1). AD with 18-20 foveae in width and 17 foveae in length. OC
elongate, 75 pm long; no cornea present. PD 184 pm long, with two slightly raised longitudinal
costae; transversely about 20-22 foveae wide, longitudinally with 27-29 foveae. Dorsal idiosomatic
setae small; one pair of setae inserted anterior to the AD, one pair on the AD on a level with insertion
of leg I, one pair each in anterolateral edge and near medial edge of OC. PD with three pairs of setae,
one of them on a level with insertion of leg III, one on that of leg IV, and one pair of setae in posterior
PD.

Ventral shield (fused anterior epimeral plate, posterior epimeral plate and genitoanal plate)
delicately foveate and punctate. Area corresponding to AE with three pairs of long ventral setae and
one pair of epimeral pores (Fig. 2); area of PE with one dorsal, one long lateral and one ventral seta.
GO 70 pm long; genital sclerites with two pairs of acetabula. These acetabula in size and morphology
similar to epimeral pores. Area of GA with two and four perigenital setae (asymmetrically ). Ovipositor
short, in rest hardly extending beyond the GO.

Gnathosoma 108 pm long, with rostrum and gnathosoma base similar in length. Both pairs of
maxillary setae inserted near the rostral base. Palps 4-segmented; P-2 with one long dorsal seta; P-3
with medial spine; P-4 with three basal setae (Fig. 3).

Leg I stouter than posterior legs. All legs with basifemora and telofemora subequal in length.
Genu 1 slightly longer than telofemur I (Fig. 5), II-4 slightly shorter than II-3 (Fig. 6), and genua on
legs III and IV longer than telofemora, respectively (Figs 7 and 8). All tibiae longer than genua.
Number of setae, from trochanter to tarsus (pas and solenidion included): leg I, 1, 3, 5, 6, 8, 11; leg
II, 1, 4, 4, 6, 7, 7; leg III, 1, 2, 2, 3, 5, 6; leg IV, 0, 1,2, 3, 6, 5. Genu I ventrally with two short spines,
1-5 with basal pair of short spines and distal pair of bristles. Tibiae II and IV with one slender ventral
bristle and two short and pectinate bristles; tibia III with one seta and one pectinate bristle. Tarsus I
with one ventromedial spine, two short ventral setae and three long dorsal setae, tip of tarsus with
seta-like dorsolateral solenidion, conspicuously large lateral membrane (Fig. 4) and doubled lateral
and medial pas. Solenidion on tarsus II hollow, dorsomedial in position (Fig. 9). Number of
dorsal/ventral setae on tarsi II to IV: 3/1 , 3-4/0- 1 (asymmetrically), 3/0; posterior tarsi with a pair of
single pas.

Pair of claws on tarsus I smaller than on posterior tarsi; claws with accessory process but without
long pecten. Claws on tarsi II to IV with pectens bearing long tines.

Male. Unknown.

Juveniles. Deutonymph and protonymph unknown. Larva 185 pm long. Dorsal plates foveate.
Anterior margin of idiosoma truncate. Number of dorsal setae same as in adults but first pair of setae
in anterior margin of idiosoma, and third pair of setae inserted within the striated integument anterior
to the OC. PD much smaller than in the female. Ventral plates separate. Genital plate lacking. Larva
with three pairs of 5-segmented legs. Genu and tibia I each with a pair of spines but no ventral bristles.

172

BARTSCH: A NEW SUBSPECIES OF THE HALACARID MITE LOBOHALACARUS WEBERl

Figs 1-10. Lobohalacarus weberi trisianensis subsp. nov., female; 1. idiosoma, dorsal; 2. idiosoma, ventral; 3. gnathosoma,
lateral; 4. tarsus I, lateral (medial setae and claw omitted); 5. leg 1. medial; 6. basifemur to tarsus II, medial; 7.
basifemur to tarsus III, medial; 8. basifemur to tarsus IV, medial; 9. posterior tibia and tarsus II, medial (lateral
claw omitted); 10. tarsus III, medial (lateral claw omitted). 1-9, holotype female; 10, paratype female. (AD, anterior
dorsal plate; AE, anterior epimeral plate; ds, dorsal setae; ds(PE), dorsal seta on PE; ep, epimeral pore; gac, genital
acetabula; im, lateral membrane; Is, lateral seta; OC, ocular plate; pas, parambulacral setae; pgs, perigenital setae;
PD, posterior dorsal plate; PE, posterior epimeral plate; so, solenidion; vs, ventral seta)

173

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 3 28 FEBRUARY 1995

Variability

Length of females 310-341 |am (with frontal spine included) or 301-314 pm (frontal spine
excluded). The females proved to be highly variable in external characters. Three of the specimens
have an elongate and spine-like anteromedian idiosomatic extension (Fig. 1 1 ), one specimen has a
short and rounded frontal hood (Fig. 1 2), and in two of the females, the anterior idiosoma is rounded
(Fig. 13). The number of perigenital setae varies from two to four setae and the number of genital
acetabula from one to two on either side. In the majority of species, the PE have one dorsal, one lateral
and one ventral seta, but the dorsal seta may be lacking (Fig. 12). Bilateral asymmetry in the leg
chaetotaxy is fairly common, e.g., in four specimens both genua I have a pair of ventral spines; in
one specimen both genua I bear a long and slender ventrolateral seta and a short ventromedial spine;
another specimen has a pair of spines on the left genu but a ventromedial spine and a ventrolateral
seta on the right genu. Tibiae II each bear one seta-like ventral bristle, and either one ( 1 specimen,
unilaterally) or two wide and bipectinate bristles; tibiae III have one ventral seta-like bristle plus one
(5 specimens) or two ( 1 specimen) wide and pectinate bristles; tibiae IV all have ventrally one seta-
like and two pectinate bristles. Tarsi III generally bear four dorsal setae (Fig. 10), though, in the
holotype, one tarsus has no more than three dorsal setae. One specimen has a ventral seta on both
tarsi III, the other five specimens demonstrate asymmetry with either one or no ventral setae.

Figs 11-13. Lobohalacarus weberi tristanensis subsp. nov., female, dorsal variants of idiosoma; 11. with frontal spine; 12.

with frontal hood (idiosoma with left lateral seta on PE); 13. anteriorly rounded (idiosoma with lateral setae on
PE). (Is, lateral seta) Scale bars = 50 pm.

The specimens from Inaccessible Island undoubtedly belong to the Lobohalacarus weberi
complex, a group of closely allied species and subspecies with records from Europe, Asia, Africa,
Australia, South and North America.

The specimens from Inaccessible are given the rank of a subspecies, viz., Lobohalacarus weberi
tristanensis. The population demonstrates a much higher tendency to variability than found in the
nominate form L. weberi weberi (Romijn and Viets) in Europe. Most remarkable is the anterior
idiosoma, arch-like, with small rounded tip or with long spine-like projection. In contrast, specimens
from Europe, as well as species and subspecies from other geographical regions, always have a spine-

Remarks

174

BARTSCH: A NEW SUBSPECIES OF THE HALACARID MITE LOBOHALACARUS WEBERI

like process. Telofeniora III and tarsi IV have two and three dorsal but no ventral setae whereas in
the European L. w. weberi these segments bear ventral setae.

GEOGRAPHICAL AND BIOLOGICAL SURVEY

The genus Lohohalacariis is world-wide in distribution. It includes species and subspecies which
are closely allied and belong to a single natural complex, the Lobohakicanis weberi group. L. weberi
was described on the basis of a specimen collected in The Netherlands (Romijn and Viets, 1924).
The species proved to be widely distributed in Europe, Africa, North America and the Hawaiian
Islands (Viets, 1950; Imamura, 1981; and unpublished records), generally inhabiting hypogean
(hyporheic) and epigean waters fed by groundwater. L. biicliarensis Jankovskaja, known from
juveniles only, was found in groundwater in Middle Asia (Jankovskaja, 1967, 1972). L. himurong
Harvey is a hyporheic Australian species (Harvey, 1988), and L. hiimmeliucki Viets was taken on the
bank of a river in Venezuela (Viets, 1940). L. processifer (Walter) is known from a juvenile from a
lake in the Andes, at 5000 m (Walter, 1919), and L. weberi gofoeusis Imamura is a stygobiontic
species from lava caves, Japan (Imamura, 1970). L. weberi tristanensis was collected from a small
stream, together with the oribatid mite Trimalocouothrus sp. (Barber-James, in prep.).

Lobohakicanis primarily inhabits subterranean waters and areas where hypogean waters meet
the surface. Lobohakicarus vi’. weberi prefers darkness (unpublished experiments). The specimens
generally live in constantly waterlogged substrates, but they are known to survive in moist sand,
within the water-film, for more than a month (Teschner, 1961 ). L. w. weberi can withstand marine
conditions for a couple of hours, and brackish water with a salinity of 20 or 10 ppt for about one or
two to three weeks respectively (unpublished preliminary experiments with a population from
northern Germany, from a pond with Sphagnum fed by groundwater).

Lobohalacanis weberi is thought to be primarily parthenogenetic (Bartsch, 1981 ). The fecundity
is low, probably a single generation per year and no more than 10 eggs per female (Teschner, 1963).
The eggs are carefully deposited in the substrate, e.g., sand (Teschner, 1963). They are not
conspicuously thick-walled, but still they may be more tolerant to both desiccation and higher salinities
than are adults.

MORPHOLOGICAL VARIANTS AND CHARACTERISTICS IN LOBOHALACARUS

Representatives of the genus Lobohakicanis are characterized by : Idiosoma with seven pairs of
dorsal setae (adanal setae included, dorsal seta on PE excluded); genital sclerites with acetabula; palps
four-segmented; third palpal segment with median spine; slender fourth palpal segment with three
setae in the basal whorl; genu I nearly as long as telofemur or tibia I; tarsus 1 with three dorsal seta,
dorsolateral solenidion, one ventromedial spinelet or seta, pair of ventral setae, and pair of apical
parambulacral setae; lateral membrane of claw fossa enlarged.

Lobohalacanis differs from the majority of halacarids in having seven pairs of dorsal idiosomatic
setae but no gland pores. All closely related marine and freshwater genera, and the majority of halacarid
genera in general, have six pairs of dorsal setae and one to five pairs of gland pores. Five pairs are
thought to be the original number of pores. The first pair of pores are almost always present, while
the other pores may be greatly reduced, absent or replaced by setae. Substitution by setae is found in
the majority of species of the genus Rhombogiiathides and Metarhombognathus (Bartsch, 1 992), and
in at least one species of Lohmannella (Bartsch, 1993). In species of the L. weberi complex, the
anterior pair of setae is believed to correspond with the first pair of dorsal setae regularly present in

175

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 3 28 FEBRUARY 1995

halacarid mites, whereas the pair of setae in the lateral margin of AD may originate from gland pores.

The author’s halacarid collection includes more than a hundred specimens of Lobohalacarus
weberi, from Europe (northern Baltic, northern Germany, Spain), northeastern America (Rhode
Island), and the Hawaiian Archipelago; the specimens were collected from various substrates such
as groundwater wells, ponds and small streams with mosses and pH as low as 4. 5-5.0, and fine sand,
more or less rich on organic material, pH-values of 6. 0-7. 8, and also slightly influenced by brackish
water.

Variability was studied on populations from Europe and the United States, viz., 20 specimens
each from two sites in Germany, and a small number of specimens from northern Baltic, Spain, Rhode
Island (Table 1 ) and Hawaiian Islands.

Characters known to vary in Lobohalacarus are;

Frontal spine. The nominate species L. w. weberi always has a frontal spine.

Genital acetabiila. The majority of subterranean living species have two pairs of genital acetabula
whereas specimens from epigean waters, such as lakes, ditches, and habitats influenced by slightly
brackish waters, have three pairs of genital acetabula. But, generally, within a population (sample)
both variants are present, asymmetry is fairly common. In populations from northern Germany, 77-
97 % of the specimens had genital sclerites with two acetabula. Specimens with a small, primordial
acetabulum beside the one or two well developed genital acetabula are present, too.

Number of perigenital setae. The most common variant in European populations are five pgs on
either side. About 15-25 % of the specimens in a given population, from both Europe and America,
have four pairs of setae. The combinations 2/4 (two setae on one side, four setae on the other side),
as well as 6/5 have been found in a population from Germany.

Genu I. The majority of specimens have the combination sp/br, that is ventromedially a spine
and ventrolaterally a bristle. In the two populations from Northern Germany, 2-6 % had spines both
ventromedially and ventrolaterally, asymmetrically or on both legs. Exceptionally, the spine or seta
is lacking.

Tibia II. Most specimens have one ventral seta and two wide, pectinate ventromedial bristles. A
single bipectinate bristle (aside from the ventral seta) is found in 0-20 % of the specimens in a given
population. One specimen from the Northern Baltic had two pectinate bristles and two smooth and
slender setae.

Tibia III. Most abundant are two ventral bristles, one short and bipectinate, the other longer, seta-
like and very faintly plumose. Though, in populations from northern Germany and northeast America,
30-40 % of the specimens had three ventral bristles, with two of them being wide and coarsely
pectinate.

Tibia IV. They generally have three ventral bristles, one of them is seta-like and slender, two are
wide and pectinate. Exceptions, viz., tibiae with two ventral bristles, one wide and pectinate, the other
slender, seta-like and delicately plumose, are very rare.

Telofemur III. The specimens from the northern Baltic and northern Germany all have the
chaetotaxal combination 2/1 (two dorsal and one ventral setae), whereas specimens from Spain and
from eastern America either had 2/1 or 2/0 setae.

Telofemur IV. Both combinations 2/1 and 2/0 are commonly represented.

Tarsus III. Eour dorsal and one ventral seta is the most common variant, other combinations are
found, though only unilaterally.

Tarsus IV. Generally, 3/1 (three dorsal and one ventral) setae are present. The combination 3/0
has been found, unilaterally, in two specimens from northeastern America. Specimens with two
ventral setae, instead of one, have been seen in European and northern American populations.

176

BARTSCH: A NEW SUBSPECIES OF THE HALACARID MITE LOBOHALACARUS WEBERl

Table 1. Lobohalacams weheri, variants of characters and, in brackets, percentage of variants in
populations from: Northern Germany, population A (groundwater well), population B (bog
with Sphagmim)\ Northern Baltic (muddy sand, brackish water); Rhode Island (ri verities
with Sphagmimy, Inaccessible Island (river with Spartma).

(br = bristle; d/v = dorsal/ventral; sp = spine)

Populations

\from

Character

Northern Germany
A B

Anterior AD

spine (100)

spine (100)

Number of

2(97)

3(19)

genital

1 1/2(3)

2 1/2 (3)

acetabula

2 (77)

Number of

5 (68)

6(4)

perigenital

4(26)

5(77)

setae

3 (6)

4(15)

2(4)

1-4, combination

sp/br(96)

sp/br (94)

of sp and br

sp/sp (2)
0/br(2

sp/sp (6)

II-5, number

2(100)

2(87)

of pectinate
bristles

1 (13)

II-5, number

1 (100)

1 (58)

of pectinate
bristles

2(42)

IV-5, number

2(92)

2(100)

of pectinate
bristles

1 (8)

III-3, number
of d/v setae

2/1 (100)

2/1 (100)

IV-3, number

2/1 (34)

2/1 (25)

of d/v setae

2/0 (66)

2/0(75)

III-6, number
of d/v setae

4/1 (100)

4/1 (100)

IV-6, number

3/1 (97)

3/1 (100)

of d/v setae

3/2(3)

Northern

Baltic

Rhode

Island

Inaccessible

Island

spine (100)

spine (100)

spine (50)
hood (17)
rounded (33)

3(71)

3(5)

2(83)

2 (29)

2(95)

1 (17)

5 (94)

5 (74)

4(42)

4(6)

4(26)

3 (42)
2(16

sp/br ( 100)

sp/br (100)

sp/br (25)
sp/sp (75)

2 (100)

2 (80)

2 (92)

1 (20)

1 (8)

2 (100)

1 (67)

1 (83)

2 (33)

2(17)

2 (100)

2 (100)

2(83)
1 (17)

2/1 (100)

2/1 (90)
2/0(10)

2/0(100)

2/1 (61)

2/1 (48)

2/0(100)

2/0 (39)

2/0 (52)

4/1 (82)

4/1 (100)

4/1 (50)

3/2 (9)

4/0 (42)

2/1 (9)

3/1 (8)

3/1 (100)

3/1 (90)

3/2(5)

3/0(5)

3/0(100)

177

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19. PT. 3 28 FEBRUARY 1995

Basifemur IV. Both variants, with one or no ventral setae, are represented in the material studied.

The Lobohalacarus weberi complex includes species and subspecies which are extremely similar
in their general facies. Descriptions and discrimination of Lobohalacarus, and halacarids in general,
are based on external morphological characters. Ultrastructural investigations or data from
karyological or molecular genetic studies are lacking until now. Studies in invertebrate taxonomy,
using such techniques, proved a number of moiphologically almost identical populations to represent
complexes of subspecies or species (Grassle and Grassle, 1 976; Pietsch and Westheide, 1 985 ; Schlegel
et ah, 1991; Wood and Backus, 1992; Schmidt and Westheide, 1994).

The external characters in the population of Lobohalacarus weberi on Inaccessible Island show
some overlap with those of L. weberi from the northern hemisphere. Differences in the leg
chaetotaxy, the unusual high degree of variability in the Inaccessible population, the geographical
distance between northern Europe (with L. weberi weberi) and the South Atlantic island in view of
low dispersal ability of Lobohalacarus, are reasons to distinguish between the populations and to
give the specimens from Inaccessible Island the rank of a separate subspecies.

DISPERSAL AND COLONIZATION OE OCEAN ISLANDS

Because of the subsurface way of life and wide geographical distribution of the Lobohalacarus
weberi complex (records exist from all continents except for Antarctica which has not been studied),
one may speculate that these freshwater halacarids belong to an ancient taxon which had already
invaded the freshwater of the supercontinent Pangea ( Bartsch, 1 982). However, the Tristan da Cunha
island group and the Hawaiian Islands, both of which have records of Lobohalacarus, are of more
recent origin. Multiple invasion of a formerly marine ancestor and subsequent spcciation and adaption
to fresh water is unlikely; thus, Lobohalacarus must have been transported to these islands.

Marine, brackish as well as terrestrial species can be spread over long distances through rafting,
but Lobohalacarus, unable to live but a few hours in seawater, would certainly not survive a transport,
drenched with seawater and salt spray, to a distant oceanic island. As documented by Maguire ( 1963),
small aquatic organisms can be dispersed by large animals. Lobohalacarus weberi, which can survive
in a moist substrate for weeks (Teschner, 1961), may have colonized Inaccessible Island via air,
attached to birds or insects which visited continental wells, banks of rivers or groundwater-fed epigean
ponds and later rested on Inaccessible Island.

Recent geological investigations by Christie et al. (1992) showed that seamounts along
submarine ridges once had been islands raised above sea level. These eroded and drowned islands,
being considerably older than the present-day volcanic islands in the Galapagos and Hawaiian
archipelago, may have been stepping stones for the biota now found on these archipelagos (Christie
et al., 1992; Carson, 1992). It is tempting to suppose that the Walvis Ridge provided similar stepping
stones and thus eased the colonization of Inaccessible Island.

ACKNOWLEDGEMENTS

1 am most grateful to H.M. Barber-James, Albany Museum, Grahamstown, South Africa, and
Dr D.R. Cook, Detroit, USA, for giving me the opportunity to study these freshwater halacarid
specimens, and 1 acknowledge valuable comments from referees and Dr E.W. Gess, Editor of the
Annals of the Cape Provincial Museum. The expedition to Inaccessible Island was organized by the
Percy Eitzpatrick Institute for African Ornithology, University of Cape Town.

178

BARTSCH: A NEW SUBSPECIES OF THE HALACARID MITE LOBOHALACARUS WEBERl

REFERENCES

Baker, P.E. 1973. Islands of the South Atlantic. In: Nairn, A.E.M. and F.G. Stehli, eels. The Ocean Basins ami Margins.
Vol. I. The South Allantic. Chapter 13: 493-353.

Baker, P.E., Gas, I.G., Harris, P.G. and Le Maitre, R.W. 1964. The volcanological report of the Royal Society Expedition
to Tristan da Cunha 1962. Philosophical Transactions of the Royal Society. London A, 256: 439-578.

Barber-James, H, Freshwater fauna of the South Atlantic island. Inaccessible. In prep.

Bartsch, I. 1981. Meeresmilben der Umgebung von Hamburg (Arach.: Acari: Halacaridae). Verhandliingen des
naturwissenschaftlichen Vereins in Hamburg (NF) 24: 5-18.

Bartsch, I. 1982. Halacaridae (Acari) im SiiBwasser von Rhode Island, USA, mit einer Diskussion liber Verbreitung und
Abstammung der Halacaridae. Gewdsser und Ahwas-ter 6SI69: 41-58.

Bartsch, I. 1992. Lohnwnnella dictyota n. sp. und L. africana n. sp. von der Stidhalbkugel (Arachnida: Acari: Halacaridae).
Senckenbergiana Biologica 72: 457-464,

Bartsch, I. 1993. Synopsis of the Antarctic Halacaroidea (Acari). In: Wiigele, J.W. and J. Sieg, eds, Synop.ds of the Antarctic
Benthos. Vol. 4, 176 pp, Koenigstein: Koeltz Scientific Books.

Carson, H.L. 1992, The Galapagos that were. Nature {London) 355: 202-203.

Chevallier, L., Rex, D.C. and Verwoerd, W.J. 1992. Geology and geochronology of Inaccessible Island, South Atlantic.
Geological Magazine 129: 1-16.

Christie, D.M., Duncan, R.A., Mcbirney, A.R., Richards, M.A., White, W.M., Harpp, K.S. and Fox, C.G. 1992. Drowned
islands downstream from the Galapagos hotspot imply extended speciation times. Nature {London) 355: 246-248.

Grassle, J.P. and Grassle, J.F. 1976. Sibling species in the marine pollution indicator Capitella (Polychaeta). Science 192:
567-569.

Harvey, M.S. 1988. A new species of Lobohalacarus from Australia (Chelicerata: Acarina: Halacaridae). Memoirs of the
Museum of Victoria 49: 363-365.

Imamura, T, 1970, The fauna of the insular lava caves in West Japan. Bidletin of the national Science Museum. Tokyo 13:
455-458.

Imamura, T. 1981. Fresh-water halacarid mites from Oahu Island, \\d,sNtm. Annotcitiones Zoologicae Japonensis 54: 287-292.

Jankovskaja, a. I. 1967. On the fauna of ground-waters in Middle Asia. I. Hydracarina of the fam. Porohalacaridae (Acari)
in the ground-waters in the wells in Kizylkum desert. Trudy Zoologicheskogo Instituta. Akademiya Nauk SSSR.
Leningrad 43: 109-1 17. (in Russian)

Jankovskaja, A. I. 1972. Phreatic and spring fauna of Northern Tien Sha and Southern Kisilkum. Trudy Zoologicheskogo
Instituta. Akademiya Nauk SSSR. Leningrad 51: 5-56. (in Russian)

Maguire, B. 1963. The passive dispersal of small aquatic organisms and their colonization of isolated bodies of water.
Ecological Monographs 33: 161-185.

PiETSCH, A. and Westheide, W. 1985. Flltrastructural investigations of presumed photoreceptors as a means of discrimination
and identification of closely related species of the genus Microphthalmus ( Polychaeta, Hesionidae ). Zoomorphology
105: 265-276,

Romijn, G, and Viets, K. 1924. Neue Milben. Archiv fiir Naturgeschichte 90: 215-225.

SCHLEGEL, M., Steinbruck, G., Kramer, M. and Brockmeyer, B. 1991. Restriction fragment pattern as molecular markers
for species identification and phylogenetic analysis in the genus Enchytraeus (Oligochaeta). Zeitschrift fiir
zoologische Systematik und Evolutionsforschung 29: 362-372.

Schmidt, H. and Westheide, W. 1994, Isozymes and general protein patterns as taxonomic markers in the taxon Nephtyidae
(Annelida Polychaeta). Marine Biology 119: 31-38.

Teschner, D. 1961. Biologische Beobachtungen an Grundwassermilben. Verhandlungen der Deutschen Zoologischen
Gesellschaft, Bonn 1960: 402-409.

179

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 3 28 FEBRUARY 1995

Teschner, D. 1963. Die Biologic, Verbreitung und Okologie der Grundwassermilbe Lobohalacarus weberi quadriporus
(Walter 1947), Limnohalacaridae Acari. Archiv fur Hydwbiologie 59: 71-102.

ViETS, K. 1940. Zwei neue Porohalacaridae (Acari) aus Siidamerika. (Reise des Herrn P. Wagenaar Hummelinck 1936-37
nach den “Benedenwindsche Eilanden”). Zoologischer Anzeiger 130: 191-201.

Walter, C. 1919. Hydracarinen aus den peruanischen Anden und aus Brasilien. Revue Suisse de Zoologie 27: 19-59.
Wood, T.S. and Backus, B.T. 1992. Differentiation of North American and European forms of Fredericella sultana
(Blumenbach) (Ectoprocta: Phylactolaemata). Hydrobiologia 237: 185-193.

180

INSTRUCTIONS TO AUTHORS

LAYOUT of recent issues should be examined for details of acceptable style and format.

MANUSCRIPTS should be submitted on floppy disk accompanied by three print-outs
(spacing I'A or 2). In exceptional cases typed manuscripts will be accepted. These should be
submitted in triplicate, typed on A4 size paper, double-spaced, on one side of the paper only
and with wide margins all round.

Only generic and specific names should be underlined to indicate italics. All other marking
up should be left to the editor.

Footnotes should be avoided as far as possible.

All measurements should be metric.

Systematic papers should follow the rules of the International code of zoological
nomenclature or the International code of botanical nomenclature, as applicable.

ILLUSTRATIONS should bear the author’s name, title of paper and number in soft pencil on
the reverse side. Diagrams, line drawings and photographs are all regarded as figures, should
be termed as such and should be numbered in a single series.

In addition to the original illustrations a photocopy of each should be submitted.

Diagrams and line drawings should be reducible to a size not exceeding 14 x 16 cm (17,5 cm
including captions). Photographs for half-tone blocks should be final size, of high contrast
and printed on glossy paper. Colour illustrations are only printed under special circumstances
and their financing must be arranged by the author.

A scale should appear with all illustrations or the magnification should be given in the
caption.

A list of captions should be submitted on a separate sheet.

TABLES should be submitted on separate sheets. Foldouts and tables printed sideways should
be avoided.

REFERENCES should be listed alphabetically at the end of the paper. Abbreviations of
periodical titles should conform with those in the World list of periodicals (4th ed.). Layouts
must be as follows:

Alexander, J. 1969. The indirect evidence for domestication. In: Ucko, P. J. and G. W.
Dimbleby eds. The domestication and exploitation of plants and animals. London:
Duckworth, pp. 123-129.

De Winter, B. 1969. Plant taxonomy today. S.Afr.J.Sci. 65 (8): 229-242.

JuBB, R. A. 1967. Freshwater fishes of southern Africa. Cape Town: Balkema.

GALLEY PROOFS are sent (when practicable) to authors for checking printer’s errors.
Adjustments of language, style and punctuation and any other amendments of the original
manuscript, if made at this stage, may be charged to authors.

AUTHOR’S COPIES (50) are issued free of charge. When more than one author is involved
the fifty free copies will be sent to the senior author. Additional copies ordered before printing
are supplied at a reduced cost.

CADAR, P E -K887

K.

/■

/

i -f

■t ^

[ it-

I

id

I y? V

■i

•• •

I. •

J H

ISSN 0570-1880

Annals of the

Cape Provincial Museums

Natural History

Ann. Cape Prov. Mus. (nat. Hist.)

Volume 19 Part 4 28th April 1995

FEB 2 7 1996

I

Published jointly by the Cape Provincial Museums
at the Albany Museum, Grahamstown, South Africa

ANNALS OF THE CAPE PROVINCIAL MUSEUMS

These Annals are published in two series, Natural History and Human Sciences, the latter series
covering cultural history, ethnology, anthropology and archaeology. They are issued in parts at
irregular intervals as material becomes available.

The primary objective of these Annals is to disseminate the results of research work undertaken by
staff members of the Cape Provincial Museums. However, a manuscript by any author is eligible
for publication provided it is based wholly or partly on the collection/s of one or more of these
museums or if the material on which the work is based is wholly or partly deposited in one of
these museums.

Parts are obtainable from the Librarian of the Albany Museum. Correspondence in connection
with these Annals should be addressed to the Editor, Albany Museum, Grahamstown 6140.

Editor

Dr F. W. GESS

Assistant Editor
Dr S. K. GESS

A comparison of nesting success and nesting habits in some Afrotropical
aculeate wasps, with particular reference to nest parasites (Hymenoptera:

Sphecidae, Eumenidae)

by

A. J. S. WEAVING

(Albany Museum, Grahamstown)

ABSTRACT

Nest structure, nesting behaviour, nesting success and nest mortality factors of eleven species
of aculeate wasps were compared. Seven species nest in pre-existing cavities: Auplopus femoralis
(Arnold) and A. vitripennis Smith (Pompilidae), Chalybion laevigatum (Kohl), Isodontia
pelopoeiformis (Dahlbom) and I. stanleyi (Kohl) (Sphecidae), Tricarinodynerus guerinii (Saussure)
and Proepipona meadewaldoi Bequaert (Eumenidae). These were studied using trap-nests. Four
species construct aerial mud-nests: Afreumenes aethiopicus (Saussure), Delta hottentottum
concinnum (Saussure) and Synagris analis Saussure (Eumenidae) and Sceliphron spirifex (L.)
(Sphecidae). Study sites were in Natal and the Eastern Cape in South Africa.

Overall nesting success, as measured by the proportions of successful cells, differed in only one
of the species. In some cases interspecific success rates differed significantly but inconsistently
between localities, due mostly to varying levels of nest parasitism and other mortality factors.

The most important identifiable causes of nest/cell failure were attacks by Miltogramminae
(Diptera), Chrysididae and Eulophidae {Melittobia sp.) (Hymenoptera) with the addition of
Osprynchotus sp. (Hymenoptera: Ichneumonidae) in aerial mud-nests. Ceropales punctiilatus Cameron
(Pompilidae) and Leucospidae were very host specific and therefore not of general importance. A
number of other taxa including Stenarella sp. (Hymenoptera: Ichneumonidae), Mutillidae
(Hymenoptera) and Bombyliidae (Diptera) caused mostly low mortality. Endogenous mortality, caused
by a combination of unidentifiable factors, was also important. Chalybion laevigatum and T. guerinii
were particularly susceptible to parasitism by Miltogramminae and Chrysididae respectively.

Possible effects of nesting characteristics on nest parasitism are considered in the light of the
evidence obtained. Certain characteristics of aerial mud-nests and their influence on parasitism are
discussed with reference to nests of S. spirifex.

It was concluded that nesting success depended largely on the activities of parasites. The unique
nesting habits of each aculeate species provided protection only against specific types of parasites
and success was therefore affected by the interaction between nesting habits and the size and species
composition of parasite populations.

INTRODUCTION

The enormous diversity in nest structure and nesting behaviour found amongst the aculeate
Hymenoptera is extensively documented. However, data on nesting success remain sparse. Success rates,
here understood to be the proportions of successful cells, are reported in 11 papers (Rau & Rau, 1916;
Taffe & Ittyeipe, 1976; Freeman, 1973, 1977 and 1982; Smith, 1979; Coville & Coville, 1980; Brooke,
1981; Coville & Griswold, 1983 and 1984; Rosenheim, 1987a), and are deducible with varying degrees
of certainty from a further 27 out of a selection (not necessarily exhaustive) of 38 papers containing
quantitative data; success rates vary between 25 and 100%, averaging 69.1% (Appendix A). However,
each of these papers deals with a single species or group of closely related, often congeneric, species.

181

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

Thus, direct interspecific comparisons of nesting success are either not available or are limited to closely
related species, severely restricting any evaluation of whether nesting habits affect nesting success.

Much of the reported mortality (mean 68.4%, range 30.2 to 98.8%) in nests of aculeate wasps is
due to nest parasites and predators (Freeman & Parnell, 1973; Freeman, 1973 and 1982; Taffe &
Ittyeipe, 1976; Peckham, 1977; Krombein, 1978; Smith, 1979; Coville & Coville, 1980; Kurczewski
&Peckham, 1982; Coville & Griswold, 1983 and 1984; Parker, 1984;Spoffordef«/., 1986; Rosenheim
& Grace, 1987; Rosenheim, 1987a; Weaving, 1989). Many aspects of nesting behaviour and nest
structure are therefore interpreted as being antiparasite in function, although this is based on scant
supporting data (Rosenheim, 1987a). Relevant literature is listed (not exhaustively) in Appendix A.

If these interpretations are correct, nesting success must be governed largely by interactions
between nesting characteristics of aculeates and their predominant parasites. Such interactions have
been described for a number of species, mostly ground-nesters from the genera Aitunophila
(Brockmann, 1985; Hager & Kurczewski, 1985; Rosenheim, 1987a and \9S9), Anoptiiis (Aim &
Kurczewski, 1984),5e/H6ed/;ns(Gess&Gess, 1975), 5e/«6u(Evans, 1957, 1966a and \911),Centris
(Vinson et al., 1987), Cerceris (Linsley & MacSwain, 1956; Evans & Hook, 1982a and 1982b),
Crahro (Matthews et ah, 1979; Evans et al., 1980), Episyron (Endo, 1980), Odynerus (Hungcrford
& Williams, 1912), Oxybelus (Peckham et al., 1973; Peckham, 1977; Hook & Matthews, 1980),
Paralastor (Smith, 1978), Philantluis (Evans & Lin, 1959; Simonthomas & Simonthomas, 1972;
Alcock, 1975; Gwynne, 1981; Evans & O’Neill, 1988), Podalonia (O’Brien & Kurczewski, 1982),
Spliex (Tsuneki, 1963; Brockmann, 1985), Stictia (Sheehan, 1 984) and Tachysphex (Spofford et al.,
1986), with a few aerial mud-nesters, Auplopus (Wcislo et al., 1988), Eumenes (Taffe & Ittyeipe,
1976) and Sceliphron (Freeman & Ittyeipe, 1976; Freeman, 1982) and species that use pre-existing
cavities, Pacliodynerus (Jayasingh, in Taffe & Ittyeippe, 1976), Trypargiliim (Krombein, 1979),
Trypoxylon (Coville & Coville, 1980; Freeman & Ittyeipe, 1993), and bees (Freeman & Ittyeipe,
1993). However, there have been no published attempts to compare nesting success and levels of
associated parasitism in a variety of species differing in their nesting habits.

The results of such a comparison are reported here. It was my intention at the outset to include
aerial mud-nesting species, those that nest in pre-existing cavities and those that dig burrows in the
soil. Unfortunately no ground-nesters were found in sufficient numbers at any locality, so that
comparison was of necessity confined to aerial mud-nesters and species that used trap-nests.

METHODS, STUDY PERIOD AND LOCALITIES

Determining causes of cell failure

Nest parasites often emerged in the field before nests were collected. Their identification was
achieved by comparing residues remaining in the cells with those in cells from which parasites had
been reared. Empty puparia and a characteristic “fouling” of cell walls (readily visible in trap-nests)
were due to Miltogramminae (Fig. lA). The cocoons of Chrysididae (Fig. IB), Mutillidae (Fig. 1C),
Osprynchotus (Ichneumon idae) (Fig. 1 C-D) and Ceropales (Pompilidae) (Fig. 1 E) were disti nguished
by means of their appearance, method of emergence by the adult and stage of the cell’s development
at which they were spun. Large numbers of pupal exuviae more-or-less filling cells or cocoons were
characteristic of parasitism by Melittohia (2A-B). Adult Leucospidae were the only parasites that
drilled their way out directly through the side walls of the trap-nests (2C). Such retrospective
techniques have been used elsewhere both for aerial mud-nests (Freeman, 1973; Freeman & Parnell,
1973; Taffe «& Ittyeipe, 1976) and nests in stem cavities (Danks 1971).

182

WEAVING: A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

Fig. 1. A - Chalybion laevigatum cell parasitised by Miltogramminae, with empty puparia and fouling of cell wails; B -
Trkarinodynerus guerinii cell with chrysidid cocoon; C - Sceliphron spirifex cocoons opened to show i nner cocoons
of mutiliid (upper) and Osprynchotus (lower); D -S. spirifex cell with Osprynchotus cocoon (parasitism during larval
stage); E - Ceropales cocoon with characteristic opening made by adult \n Auplopus vitripcnnis cell.

183

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

Fig. 2. A - Tricarinodynerus giicrinii cell and B - Sccliphron spirifcx cocoon filled with pupal exuviae of Mclittobia\ C -
leucospid emergence holes drilled through wall of trap-nest; D - observation trap-nest with cover of one section
drawn back to expose window.

Aerial mud-nests

Disused nests, collected mostly from buildings and road culverts, were dissected to establish the
fate of the various cells. Active nests were marked for collection at a future date in order to avoid
interfering with parasites and other mortality factors. Observations on behaviour were made whenever
possible at active nests.

CAVITY-NESTING SPECIES

Trap-nests, consisting of bundles of three to eight lengths of cane, were suspended from bushes
and trees up to 1.8 m above ground and from roof beams of outbuildings with at least one side
permanently open. Cavities offered varied from 1 1 to 245 mm in length and 6 to 21 mm in diameter.
More complex trap-nests (Fig. 2D) were also used to allow examination of their contents without

184

WEAVING: A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

disturbance. Used trap-nests were replaced with new ones at six month intervals at which times
observations were made on behaviour at currently active nests. Nests from which the occupants had
emerged were opened to record their fate. Nests still occupied were stored and opened at intervals.
The times elapsed since completion were not known but nests were categorised according to the stage
at which they were opened and whether or not this was followed by further incubation, as follows.

Category 1 - contents emerged or about to emerge; no storage period.

Category 2 - contents consisting of adults and/or pupae; subsequent storage.

Category 3 - contents consisting of various immature stages; subsequent storage.

Category 4 - stored without being opened until emergence.

In the many instances of parasitism by Miltogramminae, Chrysididae, Mutillidae and
Leucospidae that were recorded on the basis of remains left behind after their emergence, it was not
possible to identify actual species except where dead adult parasites were present. However, as
discussed above, the evidence for the different parasite types (i.e. families) was unmistakeable.
Consequently no differentiation between species in the above families was made in the analyses of
levels of parasitism on the assumption that modes of attack were similar within families. In both the
Miltogramminae and Chrysididae there was one dominant species, so that any errors due to differences
in behaviour must be negligible.

The study started in March 1990 and the final samples were collected in May 1992.
Localities and habitats

In the Eastern Cape, aerial mud-nests were collected from a road culvert (Table Farm
33°15’S,26°26’E), the inner walls of a disused fort (Double Drift 33°06’S,26°47’E), below wooden
floors of houses on stilts (Riet River 33°33’S,27°01’E) and within a suburban house (Grahamstown
33°18’S,26°31’E). Trap-nests were set up in trees and bushes along watercourses (usually dry) at
Thursford (33° 1 1 ’S,26°24’E) and Hilton (33°16’S,26°21’E) and beside a disused sandpit at Hilton.

In Natal, aerial mud-nests were collected from roof beams of buildings at Umlalazi Nature
Reserve (28°57’S,31°47’E), Hluhluwe Game Reserve (28°07’S,32°03E), Mkuzi Game Reserve
(27°37’S,32°14’E), Fanies Island (28°06’S,32°26’E) and False Bay Park(27°58’S,32°23’E). At Cape
Vidal (28°07’S,32°33’E) they were collected from underneath houses on stilts as well as from roof
beams. Trap-nests were attached to external roof beams of the same buildings at the above localities
except for Umlalazi Nature Reserve and Cape Vidal. They were also set up in bushes and trees in
woodland at Fanies Island, woodland and savannah habitats at Mkuzi Game Reserve and False Bay
Park, and forest at False Bay Park and Lake Sibaya (27°23’S,32°41 ’E).

SPECIES AND NEST TYPES

Aerial mud-nesters

Afreumenes aethiopicus (Saussure). Nests were single-celled solitary mud pots, 12-14 mm in
diameter with walls approximately 1 mm thick, attached to roof beams, walls and roofing thatch,
usually on vertical surfaces (Fig. 3-2). Oviposition was carried out in the empty nest, followed by
mass-provisioning with caterpillars. The nest remained open during provisioning and was finally

185

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

Fig. 3. Sections through nests of the four aerial mud-nesting species. 1 - Delta liotlenlottum, 1 - Afreumenes aethiopicus, 3b
& c - Synagris analis, 4a & b - Sceliphron spirifex, indicating cell positions (B-basal, 0-outer, l-inner). 3a - frontal
view of Synagris analis nest.

186

WEAVING: A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

plugged with mud. The lip around the entrance was not destroyed and there was no additional daubing
of the exterior with mud. Females were never observed guarding or otherwise caring for their nests.

Delta hottentoltum concitiniun (Saussure). Nests were two-celled mud pots, 1 5- 1 8 mm in diameter
with walls approximately 2 mm thick, with broad areas of basal attachment to the substrate (Fig. 3-
1). Nests were similarly sited to those of A. aethiopiciis, also being found on sheltered parts of tree
trunks. Oviposition took place immediately on completion of the cell and was followed by mass-
provisioning with 2-7 caterpillars. The entrance remained open during provisioning and was finally
plugged with mud, some of which was generated by breaking down the surrounding lip using
regurgitated water. On completion of the second cell, further mud was collected and applied to the
whole nest surface, especially around the base. One timed wasp took 40 min. to build the second cell,
45 min. to provision it with two caterpillars and 75 min. to daub the nest surface. On completion the
female abandoned the nest.

All nests of this species found in Natal were of the above structure. In the Eastern Cape, most
nests (14/18) were single-celled, often in pairs with the extra daubing covering both cells and making
them contiguous. They were usually sited in grooves between bricks on buildings.

Synagris analis Saussure. The substantial mud-nests of this species were multi-celled, sometimes
exceeding 0.5 kg in weight, and were attached to roof beams, walls and branches of trees. The structure
of a typical nest is shown in Fig. 3-3a-c. A maximum of 1 7 cells per nest was recorded. Initially thick
walls followed by extensive daubing resulted in exceptionally thick cell walls and seals, 12.2 mm
(n=30, range 5-22 mm) and 10.3 mm (n=15, range 7-16 mm) respectively. Cells were added
progressively upwards, each new cell being fully incorporated into the overall matrix of the nest by
daubing before provisioning started. Provisioning with caterpillars was progressive and the cell was
sealed once the larva was fully grown, whereupon the next cell was started. Cell entrances were never
temporarily sealed and females were always found inside their cells facing outwards, except when
away hunting or feeding.

Scelipliron spirifex (L.). The large multi-celled mud-nests of this species (Fig. 3-4a & b) were
similarly sited to those of S. analis. A maximum of 41 cells in a nest was recorded. Cells were built
one by one, each being mass provisioned with spiders before the next was started. Extensive daubing
was usually carried out but, in contrast to S. analis, only on completed nests. Wall thickness was
variable due to an absence of daubing on some nests, which were probably not completed. The
thickness of the outer walls of a typical daubed nest ranged from 2.4-4. 9 mm (n=7) and the seals 2.9
to 4.9 mm (n=8). No time was spent inside cells by females and entrances remained open during the
day until provisioning was complete. Cells were temporarily sealed with mud if provisioning was
incomplete at nightfall and were re-opened the following morning.

Trap-nesters

Details of nest structure and aspects of nesting behaviour of Chalybion laevigatum (Kohl),
Proepipona meadewaldoi Bequaert, Tricarinodynenis guerinii (Saussure), Aiiplopus vitripennis
Smith and A. femoralis (Arnold) have already been described (Weaving, 1994a and 1994b). Main
nest and behavioural characteristics of these species are summarised alongside the additional species
described in Table 1.

Isodontia pelopoeiformis (Dahlbom) and /. stanleyi (Kohl). Nest structure and nesting behaviour
in these species has been reported from the Eastern Cape by Gess & Gess (1982) and no notable
differences were observed in this study. These wasps utilised trap-nests sited in vegetation, and the
former species was also found in metal pipes and hollow canes situated horizontally and vertically.

187

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

and also in cavities in branches of trees. In nests of /. pc/opoe/for/Hw partitions were 17.5 mm (n=114,
range 5-35 mm) and end seals 65.6 mm (n=71, range 5-125 mm) in thickness. In /. stanleyi these
measurements were 10.8 mm (n=23, range 5-25 mm) and 54.9 mm (n=13, range 33-102 mm)
respectively.

Table 1 . Summary of main nest and behavioural characteristics of species of aerial mud-nesters and
trap-nesters studied.

Aerial mud-
nesters

Trap-nesters

Af

Dh

Sy

Ss

Cl

Pm

Tg

Av

IP

Is

Auf

Nest type

SP

DP

MC

MC

L

L

L

L

L

L

DM

No. of cells

1

2

n

n

n

n

n

n

n

n

n

Daubing

-

-t-

++

+

-

Partition

m*

m

m

M

V

V

End seal

M*

M

M

VC

VL

VL

Cell lined

-

-

+

-

-

-

Vestibular cell

-

+

+

-1-

-

-

Superstructure

-

-

-

-

-

-

+

-

-

-

-

Cocoon

F

F

F

D

D

F

F

D

D

D

D

Provisioning

M

M

P

M

M

M/P

M/P

S

M

M

S

Temporary plug

-

-

-

-

-

-

-

-

-1-

-1-

-

Guarding

-

-

+

-

-

+

+

+

-

+

+

Nest sharing

-

-

-

-

-

-

-

-

-

-

Legend:-

Species

Nest type

No. of cells

Daubing

Partition

End seal

Coccoon

Provisioning

Other

characteristics

Af = Afreiimenes aethiopicus, Auf = Auplopiis femoralis, Av = Auplopus
vitripennis, Cl = Chalybion laevigatum, Dh = Delta hottentottum, Ip= Isodontia
pelopoeiformis, Is = Isodontia stanleyi, Pm = Proepipona meadewaldoi, Ss =
Sceliphron spirifex, Sy = Synagris analis, Tg = Tricarinodynerus guerinii.

DP = 2-celled mud pots, DM = discrete mud cells in cavity, L = linearly arranged
cells in cavity, MC = multicellular composite mud nests, SP = 1 -celled mud pots,
n = several.

- = nil, + = moderate, -i-H = thick.

m = thin mud, M = thick mud, V = thick vegetable loose.

M = mud, VC = cemented vegetable, VL = loose vegetable.

D = discrete, F = fills cell as lining.

M = mass, P = progressive, S = single.

* coated with uric acid ex bird faeces.

- = negative, + = affirmative.

188

WEAVING: A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

Table 2. Effect of sampling procedure (stage nest opened) on per cent successful emergence of six

trap-nesting species, Tricarin

odynerus

guerinii (Tg), Chalybion laevigatum (Cl), Auplopus

vitripennis (Av), Proepipona

meadewaldoi (Pm) and Isodontia pelopoeiformis

(Ip).

Species

Tg

Cl

Av Pm

Ip

n=5363

n=918

n:

=574 n=417

n=364

Stage %

Stage %

Stage

% Stage %

Stage %

2 67.4’

2 51.8"

3

83.4" 2 72.0"

4

77.2"

4 57.]”

3 45.0"”

2

82.1" 3 51.2"

2

75.0"

3 47 .0‘

4 44.0”

1

58.9” 1 34.6”

3

61.1"

1 39.2^

1 36.6”

4

54.8” 4 18.7”

1

26.6”

Possession of common superscripts indicates no significant difference.

Table 3. Effect of sampling procedure (stage)

on endogenous mortality and inability of adults to

emerge in

trap-nests of Tricarinodynerus guerinii, Clialybion laevigatuin and Auplopus

vitripennis.

Endogenous mortality

Blocked emergence

T. guerinii

C. laevigatum

A. vitripennis

C. laevigatum

Stage %

Stage

%

Stage %

Stage

%

3 28.6"

3

15.8"

4 12.2"

2

8.9"

4 15.7"

2

14.3"

1 6.0”

3

2.3”

1 7.2”

1

8.5”

3 0.0”

4

0.0”

2 6.1”

4

6.9”

2 0.0”

1

0.0”

Possession of common superscripts indicates no significant difference.

EXAMINATION OF THE DATA

Sampling procedure

Chi-square tests were used for each species to assess whether success rates and levels of the
various mortality factors were affected by the sampling procedures categorized above (Table 2).
Lowest success rates were shown in nests of T. guerinii and /. pelopoeiformis in category 1 and in
nests of C. laevigatuin, A. vitripennis and P. meadewaldoi in both categories 1 and 4. Success rates
differed between all categories of nests of T. guerinii and C. kievigatum, although in the latter the
differences were less clear cut.

Mortality factors likely to be affected by the sampling procedure were endogenous mortality
(due to possible changes in the microclimate in cells), inability to emerge (due to fracturing of cell
partitions during opening), and parasitism by Melittobia sp. It was found that this parasite can attack
at any stage of development. Mortality from Mutillidae, Ichneumonidae, psocids and mites was too
low to influence results, whether affected by the sampling procedure or not. Parasitism by

189

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

Tablet. Effect of sampling procedure (stage) on mortality caused by Melittobia in trap-nests of
Tricar inodynerus guerinii, Chalybion laevigatum and Proepipona nieadewaldoi.

T. guerinii

C. laevigatum

P. meadewaldoi

Stage %

Stage

%

Stage

%

1 29.8^

1

9.5“

4

15.6“

3 9.9”

3

3.6”

1

6.5“

4 6.3”

4

3.4”

3

6.2“

2 6.2”

2

2.7”

2

0.0“

Possession of common superscripts indicates

no significant difference.

Table 5. Utilisation of data for analysis of success rates and mortality factors.

Species

Data used

Per cent of

Per cent of

(stage)

whole sample

sample due to

stage 4

T. guerinii

1

46.6

0.0

C. laevigatum

1+4

63.3

3.2

A. vitripennis

1+4

75.0

20.0

P. nieadewaldoi

1+4

63.1

7.7

/. pelopoeiforniis

1

43.3

0.0

Miltogramminae and Chrysididae is achieved prior to the completion of nests or their sampling and
would not therefore have been affected by the sampling.

Endogenous mortality, overall, was not affected by the sampling although there were significant
differences between certain categories in nests of 7. guerinii andC. laevigatum (Table 3). The inability
of adults to emerge from their cells was significantly affected only in nests of A. vitripennis and C.
laevigatum (Table 3).

Mortality caused by Melittobia sp. was highest in nests from category 1 of T. guerinii and C.
laevigatum (Table 4).

Analysis of success rates was therefore confined to data from category 1 samples which could
not be affected by the sampling procedure. Where no significant differences occurred between
categories 1 and 4, as with C. laevigatum, A. vitripennis and P. meadewaldoi, the data from these
two samples were combined (Table 5). Although .some of the individual mortal ity factors were affected
differently in these two categories, category 4 represented only a small proportion of the samples,
except in A. vitripennis (20%), and would have a negligible effect on success rates. Even with this
species, inclusion of data from category 4, with its associated higher mortality due to inability to
emerge, merely reduces the success rate from 58.9% for category 1 alone to 57.7% for categories
1 +4.

Distribution of sample size

Success rates, levels of nest parasitism and dominance of the different parasites inevitably differed

190

WEAVING; A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

between localities. It was important therefore to investigate the consistency of interspecific differences
between localities by making species comparisons within each locality. Wide variation in sample
sizes for species, localities and sample periods limited the range of species that could be compared
at each locality. Sample sizes of less than 20 cells were considered unreliable and were discarded.

Species using trap-nes^s situated in or on buildings were mostly distinct from those in bush sites.
Proepipona meadewaldoi and the four aerial mud-nesters were largely confined to buildings while
both species of Isodontia and both auplopines were limited to bush sites, except for a few nests of I.
pelopoeifonnis. Although C. laevigatiun and T. giierinii were primarily found in building sites,
numbers using bush trap-nests were sufficiently high for making compari.sons between bush sites
and building sites.

RESULTS

Nesting success - species comparisons within localities

Results of Chi-square tests on success rates within localities (Table 6) showed significant
differences between species at seven of 12 sites in/on buildings. Significant differences between trap-

Table 6. Per cent success rates of adequately sampled species at sites in buildings compared within
localities.

FBA

FBG

FBR

FI

FIC

Cl

60.3“

Sy

45.9“

Cl

41.5“

Dh

49.5“

Af

63.5“

Tg

53.6’

Tg

28.6“

Tg

34.4“

Ss

29.9“

Tg

62.8“

Pm

51.4“

Cl

20.1“

Pm

30.7“

Af

25.0“

Pm

20.0“

Af

30.0“

Ss

5.0’

Ss

16.4“

FIG

MK

MKB

HL

UL

Tg

65.9“

Af

37.9“

Tg

38.0“

Af

52.8“

Af

81.2“

Pm

46.1“

Ss

26.7“

Cl

31.0“

Ss

48.9“

Sy

74.2“

Cl

0.0“

Dh

45.2“

Dh

26.7“

Ss

14.7“

CV

GT (Cape)

Ss

89.6“

Ss

61.0“

Af

57.1“

Dh

26.3“

Species abbreviations as in Table 1 .

Locality abbreviations as follows:-

FBA/FBG/FBR - ablution block/garage/rustic huts at False Bay Park; FI/FIC/FIG - rest huts/ablution
block/garage at Fanies Island; MK/MKB - various outbuildings/field offices at Muzi Game Reserve;
HL/UL/CV - outbuildings at Hluhluwe Game Reserve/Umlalazi Nature Reserve/Cape Vidal; GT -
Grahamstown suburban house.

Possession of common superscripts signifies no significant difference.

Species in bold are aerial mud-nesters.

191

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

nesting species occurred only at FIG where C. laevigatiim achieved zero success. Where trap-nesters
and aerial mud-nesters occurred together, S. spirifex was the least successful (at FBR and FBG) and
S. aiialis the most successful (at FBG). Synagris aiialis was again more successful than S. spirifex at
UL. Delta hottentottum was either more or similarly successful compared with S. spirifex at three
Natal sites, but was significantly less successful at GT in the Cape. At four of six localities where S.
spirifex and A. aethiopiciis could be compared, the latter was more successful, significantly so at two.
Flowever, at Cape Vidal the reverse was true.

Table 7. Per cent success rates of adequately sampled species from bush sites compared within
localities.

FBT

FIP

MKH

MKM

MKN

Auf

75.3=

Auf

69.7=

Av

82.9“

Av

67.1"

Cl 69.2=

Av

Cl

55.5^

42.8^

Av

61.1=

IP

Auf

76.9=

44.8'’

Auf

41.7*’

Tg 63.4“
Av 58.8=

MKO

SIB

H

(Cape)

Auf

67.6=

Av

71.4“

Is

57.9=

Ip

56.0=*'

Ip

9.8*’

Ip

0.0*’

Tg

36.4*”

Av

31.6^

Species abbreviations as in Table 1.

Locality abbreviations as follows:-

FBT/FIP - False Bay Park/Fanies Island woodland;

MKH/MKM/MKN/MKO - savannah/woodland/forest/woodland at Mkuzi Game Reserve;

SIB - Lake Sibaya forest;

H (Cape) - trees/bushes bordering sandpit and watercourse.

Possession of common superscripts signifies no significant difference.

Similar tests on data from bush sites (Table 7) showed significant differences between species
at six of eight sites. However, no consistent trends were discernible for any species. For instance, A.
vitripennis was most successful at MKM and SIB but among the least so at MKO and FBT; A.
feinoralis was more successful than A. vitripennis at FBT and MKO but less so at MKH and MKM.

Thus, intra-locality success rates differed significantly at thirteen of the 20 localities with
adequately sized samples (Tables 6 and 7). Whilst there were certain trends in relative success rates,
actual levels for individual species showed much heterogeneity between localities (Fig. 4).

Collectively, trap-nesting species were more successful in bush sites than in buildings, 54.8 and
38.3% respectively (t=3.83 df=7 p<0.01 with arcsin transformation). Trap-nesters in buildings did
not differ significantly from aerial mud-nesters in their collective success rates, 38.3 and 43.3%
respectively (t=1.53 df=8 p>0.10 with arcsin transformation).

Nesting success and main mortality factors

INTER-LOCAIJTY COMPARISONS OF TRAP-NESTING SPECIES

A simultaneous assessment of host species and localities in relation to overall nesting success

192

WEAVING: A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

Bush sites Sites in/on buildings

Fig. 4. Interlocality variation in nesting success in six trap-nesting and four aerial mud-nesting wasps. Mean success is shown
with solid squares.

or mortality due to particular parasites was not possible since locality sets differed for each host as
a result of indequate sample sizes. Instead, data from sites with samples exceeding 20 cells were
compared separately for each species using Chi-square tests. Localities marked with * in Tables 8
and 9 had relatively small samples (<50 cells) which are therefore less reliable.

Chalybion laevigatum. Nesting success varied from zero to 69.2% depending largely on levels
of parasitism by Miltogramminae and Melittobia (Table 8, Fig. 5). Exceptionally high endogenous
mortality contributed to the zero success at FIG but in this case the sample size was minimal. The
high levels of attack by Miltogramminae and Melittobia, and resultant low success rate at FBG, are
probably attributable to the long-standing presence at this locality of a large population of T. guerinii,
with its associated parasites, nesting in a cane partition .

Tricar inody nevus guerinii. A wide range in nesting success was again demonstrated, attributable
in this case mainly to varying levels of parasitism by Chrysididae, Melittobia and unknown factors
(Table 8, Fig. 5). It is noteworthy that, again, success was lowest and parasitism by Melittobia highest
at FBG.

Proepipona meadewaldoi. Varying success rates appeared to be linked to levels of endogenous
mortality and parasitism by Chrysididae (Table 8, Fig. 5). Parasitism by Miltogramminae was high

193

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

Table 8. Success rates and mortality factors in trap-nesters mostly in/on buildings.

Per cent success and parasitism

Site

n

SE

EM

DP

CH

MU

OS

IC

MT

?F

Chalybion laevigatwn
MKN 52 69.2“

1.9

9.6

13.5

0.0

3.8

0.0

0.0

1.9

FBA

73

60.3“

5.5

24.6

5.5

0.0

0.0

0.0

0.0

2.7

FBT

21

42.8“*'

4.8

33.3

0.0

0.0

0.0

19.0

0.0

0.0

FBR

147

41.5^

8.2

28.6

6.8

0.7

2.0

0.7

2.7

3.4

MKB*

29

31.0'"

6.9

41.4

0.0

3.4

0.0

0.0

13.8

3.4

MKO*

36

30.5^-

8.3

16.7

27.8

2.8

2.8

0.0

0.0

2.8

FBG

184

20.T

8.1

47.3

0.0

0.4

0.0

0.5

21.2

0.0

FIG*

21

0.0“

33.3

19.0

0.0

0.0

0.0

4.8

33.3

4.8

Tricarinodynerus
FIG 129

giierinii

65.9“

4.6

0.0

10.8

0.0

0.0

0.0

9.3

4.6

MKN

82

63.4“"

3.6

2.4

20.7

0.0

0.0

0.0

8.5

1.2

FIC

113

62.8“"

6.2

0.9

19.5

0.0

0.0

0.0

8.0

0.9

FBA

420

53.6"

5.0

3.6

24.5

0.9

0.0

0.0

9.8

1.7

MKB

450

38.0‘

6.0

0.2

22.2

0.4

0.0

0.4

24.7

3.8

MKO*

22

36.4““

4.5

0.0

36.4

4.5

0.0

0.0

0.0

13.6

FBR

61

34.4““

3.3

11.5

32.8

0.0

0.0

0.0

0.0

18.0

FBG

1197

28.6“

7.8

6.3

6.3

0.1

0.0

0.0

47.4

0.7

Proepipona meadewaldoi
FBA* 35 51.4“

5.7

22.9

0.0

2.9

0.0

0.0

0.0

2.9

FIG*

36

41.6“"

25.0

2.8

0.0

0.0

0.0

0.0

5.5

8.3

FBR

133

28.6"

25.6

18.8

7.5

0.0

0.0

0.0

10.5

3.8

FBG*

38

18.4"

28.9

2.6

10.5

0.0

0.0

5.3

10.5

2.6

Site abbreviations as in Tables 6 and 7, bush sites in bold.

SE - successful cells; EM - endogenous mortality; DP - miltogramminae; CH - Chrysididae; MU -
Mutillidae; OS - Osprynchotus sp.; IC - Stenarella sp.; MT - Melittobia\ ?F - fate unknown.
Localities in bold are in bush sites.

Localities marked * are inadequately sampled (<50 cells).

Possession of common superscripts indicates no significant difference.

at FBA and FBR but the impact was reduced by low or zero levels of the above two factors at FBA.
Lowest success rates again occurred at FBG.

Aiiplopiis vitripennis. Endogenous mortality was the most consistent factor affecting nesting
success (Table 9, Fig 5). Levels of attack by the various nest parasites varied erratically between
localities but higher mortality from unknown causes, Leucospidae and/or Ceropales was responsible
for the significantly reduced success rates at FBT and MKO.

194

Per cent Per cent

WEAVING: A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

60

40

20

0

Sy At Dh SsN SsC Tg Pm Cl
Aerial mud-nesters Trap-nesters

B ^

Av Auf Tg Cl Ip

Fig. 5. Relative impact of five main mortality factors on nesting success in A - trap-nesters and aerial mud-nesters in/on
buildings and B - trap-nesters in bush sites.

195

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

Table 9. Success rates and mortality factors in trap-nesters confined to bush sites.

Per cent success and parasitism

Site

n

SE

EM

DP

CH

MU

LE

CE

1C

MT

?F

Aiiplopus

vitripennis

FBM*

21

90.5“

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

MKH*

35

82.9“

11.4

0.0

0.0

0.0

0.0

0.0

0.0

2.9

2.9

SIB*

21

71.4“”

14.3

14.3

0.0

0.0

0.0

0.0

0.0

0.0

0.0

MKM

79

67.1“”

19.0

0.0

0.0

6.3

0.0

0.0

0.0

1.3

0.0

FIP

140

6I.4“”

10.0

0.0

1.4

2.1

2.2

6.5

0.0

0.7

7.1

MKN*

34

58.8”

14.7

8.8

0.0

0.0

0.0

0.0

0.0

5.9

0.0

FBT*

45

55.5”

8.9

0.0

0.0

2.2

0.0

0.0

2.2

0.0

15.5

MKO

79

31.6“

15.2

1.3

0.0

2.5

13.9

11.4

0.0

0.0

1.3

Isodontia

pelopoeifonnis

MKH*

26

76.9“

7.7

15.4

0.0

0.0

0.0

0.0

0.0

0.0

0.0

MKO*

25

56.0“

4.0

24.0

8.0

0.0

8.0

0.0

0.0

0.0

0.0

SIB*

41

9.8”

2.4

2.4

9.8

9.9

68.3

0.0

0.0

2.4

4.9

TCape*

23

4.3”

26.1

17.4

8.7

0.0

0.0

0.0

0.0

0.0

34.8

HCape*

43

0.0”

18.6

37.2

20.9

0.0

0.0

0.0

0.0

0.0

9.3

Isodontia

stanleyi

FIP +

MKH*

18

83.3“

5.5

5.5

0.0

0.0

0.0

0.0

0.0

5.5

0.0

HCape*

19

57.9“

15.8

5.3

10.5

0.0

0.0

0.0

0.0

0.0

5.3

Auplopus femora Us

FBT

77

75.3“

10.4

0.0

1.3

0.0

0.0

0.0

0.0

0.0

9.1

MKN*

17

70.6“”

23.5

0.0

0.0

0.0

0.0

0.0

0.0

0.0

5.9

FIP*

33

69.7“”

12.1

0.0

0.0

9.1

0.0

0.0

0.0

0.0

9.1

MKO*

34

67.6“”

29.4

0.0

0.0

0.0

0.0

2.9

0.0

0.0

0.0

MKH*

29

44.8”

24.1

0.0

0.0

17.2

0.0

0.0

0.0

10.3

0.0

MKM*

36

41.8”

22.2

0.0

5.5

8.3

0.0

0.0

0.0

16.7

2.8

Site abbreviations as in Tables 6 and 7.

SE - successful cells; EM - endogenous mortality; DP - miltogramminae; CH - Chrysididae;
MU - Mutillidae; LE - Leucospidae; CE - Ceropales; IC - Stenarella sp.; MT - Melittobia;
?F - fate unknown.

Localities marked * are inadequately sampled (<50 cells).

Possession of common superscripts indicates no significant difference.

Isodontia pelopoeifonnis. Leucospidae were responsible for a substantial and significant
reduction in nesting success at one locality in Natal (Table 9). Otherwise the main mortality factor
was Miltogramminae (Fig. 5). Very low success rates were also recorded at both Cape sites due to
high levels of endogenous mortality, Miltogramminae, Chrysididae and/or unknown factors.

196

WEAVING: A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

Isodontia stanleyi. This species was sampled at only two of the sites in Natal and sample sizes
were well below the minimum of 20 cells. However, at site H in the Cape it was 19. A comparison
was therefore made of the combined Natal sample (18 cells) with the Cape sample. There was no
significant difference in nesting success between the two areas (Table 9). Endogenous mortality and
parasitism by Chrysididae appeared to be greater at the Cape site but much larger samples are required
for this species.

Auplopus femoralis. Success rates were relatively high at four of six localities but only one of
these (FBT) differed significantly from the remaining two less successful ones, MKH and MKM
(Table 9). There were no records of parasitism by Miltogramminae, Leucospidae or Ichneumonidae.
Endogenous mortality was the main cause of failure at all localities (Fig. 5). Mutillidae, Melittobia
and unknown factors reduced success only at some localities.

INTER-LOCALITY COMPARISONS OF AERIAI. MUD-NESTING SPECIES

Afreumenes aethiopicus. Success rates in Natal varied from 25.0 to 81.2% largely due to
fluctuating levels of attack by the three main nest parasites, Miltogramminae, Chrysididae and the
ichneumonid Osprynchotus sp. and, to a lesser extent, endogenous mortality (Table 10, Fig. 5).
Mutillidae were important at only one locality. Data from a well separated locality, Malindi in East
Africa, showed similar trends to those from South Africa.

Delta hottentottum concinnum. Nesting success at four Natal localities was significantly lower
than at FlC due mainly to increased levels of attack by the two principal nest parasites, Chrysididae
and Melittobia (Table 10, Fig. 5). Endogenous mortality was again a consistent cause of failure.
Mortality factors and success rates at GT in the Cape did not differ from those in Natal except for FlC.

Sceliplnon spirifex. The multi-celled aerial mud-nests of this species may contain up to 40 or
more cells. Consequently interlocality comparisons were limited to those localities with samples from
an arbitrary minimum of 10 nests to increase the reliability of the results. Nesting success varied from
11.8 to 48.9% (Table 10). Very high levels of parasitism by Melittobia (Fig 5) were responsible for
the success rates being significantly lowest at RR and MKB. Varying combinations of endogenous
mortality, Miltogramminae, Chrysididae, Osprynchotus, Melittobia and mortality from unknown
causes also significantly reduced success at a further three localities compared with that at HE. All
factors with the exception of Mutillidae and Stenarella were responsible for significant (>10%)
mortality at one or more of the localities. Data from Malindi (East Africa) showed no notable deviations
from the pattern observed in Natal and Eastern Cape. Sample size was, however, inadequate for
inclusion in the analysis.

Synagris analis. Sample sizes were small for this species and it was not possible to apply the
minimum of 10 nests for interlocality comparisons. Chi-square tests were therefore carried out using
data from localities with more than one nest, namely FBG, FIG and UL with 9, 4 and 4 nests
respectively, in order to provide some comparison with S. spirifex. Nesting success was significantly
reduced at FBG due to Miltogramminae and Melittobia which were not recorded from nests of this
species at the other sites, and to a relatively high level of attack by Mutillidae (Table 10). Thus, apart
from FBG where success rates may have been affected by an abnormally high parasite population
emanating from nests of T. guerinii, S. analis appeared to enjoy a high level of nesting success,
Mutillidae being the main parasite.

OVERAIJ. NESTING SUCCESS

Differing pressures from the various mortality factors between localities were primarily

197

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

Table 10. Success rates and mortality factors in aerial mud- nesting species.

Per cent success and parasitism

Site

n

SE

EM

DP

CH

MU

OS

1C

MT

?F

Afreiimenes aethiopiciis
UL* 16 81.2“

6.2

0.0

0.0

0.0

6.2

0.0

6.2

0.0

CV

175

57.1“*’

6.8

1.1

18.3

0.6

6.3

0.0

2.8

5.1

HL*

36

52.8“*"

11.1

5.5

2.8

13.9

11.1

0.0

0.0

2.8

FIG*

17

35. 3*"'

0.0

5.9

41.2

5.9

5.9

0.0

0.0

5.9

FBR

70

30.0^

7.1

21.4

0.0

4.3

27.1

0.0

1.4

8.6

FI*

16

25. 0‘

12.5

12.5

31.2

0.0

12.5

0.0

0.0

6.2

Ml#

30

66.7

0.0

0.0

10.0

3.3

10.0

0.0

3.3

0.0

Delta holtentottiim
FlC 74

63.5“

14.9

0.0

8.1

1.3

5.4

0.0

1.3

4.0

MKE*

22

54.5“*“

13.6

4.5

0.0

18.2

4.5

0.0

0.0

4.5

FI

103

49.5“*’

14.6

1.0

10.7

2.9

7.8

0.0

1.0

7.8

HL*

42

45.2“^

7.1

7.1

2.4

4.8

7.1

0.0

23.8

2.4

FBG*

48

41.7"

6.2

4.2

10.4

4.2

2.1

0.0

16.7

10.4

MK

58

37.9"

13.8

6.9

6.9

1.7

6.9

0.0

13.8

10.3

FIG*

18

27.8"

27.8

5.5

22.2

0.0

0.0

0.0

0.0

5.5

UL*

30

26.7"

13.3

0.0

20.0

0.0

0.0

0.0

36.7

3.3

GT*

19

26.3"

21.0

0.0

36.8

0.0

0.0

0.0

10.5

5.3

Sceliphron spirifex
HL 309 48.9“

8.1

4.5

10.0

1.3

14.6

0.0

4.8

6.1

FI

163

29.9"

15.6

16.9

3.9

0.6

10.4

0.0

13.0

2.6

DD

153

29.4"

17.6

0.0

9.1

0.0

20.9

0.0

0.6

17.0

TF

220

27.7"

9.1

2.3

19.1

0.4

5.9

0.0

18.2

10.9

MKB

178

16.3"

10.1

1.1

1.7

5.1

6.2

0.0

41.6

12.9

RR

279

11.8"

3.6

0.4

1.8

0.0

13.3

0.0

63.1

4.7

Ml#

41

12.2

12.2

9.7

9.7

0.0

19.5

0.0

7.3

24.4

Synagris analis
FIG* 18

77.8“

5.5

0.0

0.0

11.1

0.0

0.0

0.0

5.5

UL*

31

74.2“

25.8

0.0

0.0

0.0

0.0

0.0

0.0

0.0

FBG*

61

45.9"

16.4

8.2

0.0

14.7

0.0

0.0

4.9

8.2

Locality abbreviations as in Tables 6 and 7.

Success and parasite abbreviations as in Tables 8 and 9.

Ml# Data from Malindi not included in analysis.

Localities marked * are inadequately sampled (<50 cells or <10 multi-celled nests).
Possession of common superscripts indicates no significant difference.

198

WEAVING: A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

Table 11. Results of t-tests on overall per cent success rates using arcsin transformation.

Species Nest type Mean % success

(arcsin)

Sites in/on buildings:

Synagris analis

AM

54.6“

Afreumenes aethiopicus

AM

43.5“

Tricar inodynenis guerinii

TN

43.3“

Delta hottentotturn

AM

41.2“

Proepipona meadewaldoi

TN

35.9“

Sceliphron spirifex (N)

AM

33.8“

Chalybion laevigatum

TN

31.5“

Bush sites:

Auplopus vitripennis

TN

54.3“

Auplopus femoralis

TN

51.9“

Tricarinodynerus guerinii

TN

44.9“

Chalybion laevigatum

TN

43.5“

Isodontia pelopoeiformis (N)

TN

42.7“

Cape sites:

Sceliphron spirifex (C)

AM/B

28.2“

Isodontia pelopoeiformis (C)

TN/0

7.6”

AM = aerial mud nest; TN = trap-nest; B = in/on buildings; O = bush sites; (C) = Cape sites; (N) =
Natal sites.

Possession of common superscripts within columns signifies no significant difference. Results of
certain species analyses are indicated in superscript columns c, d and e.

responsible for the intra-specific variations in nesting success. In order to assess whether there were
overall differences in interspecific success rates, data from the relevant localities were combined and
subjected to t-tests (Table 11). No species in either bush or building sites in Natal differed significantly
from any other. At Cape sites, /. pelopoeiformis was significantly less successful than S. spirifex.
Species common to both Cape and Natal sites (5. spirifex and I. pelopoeiformis) and those common
to sites in buildings and bush sites in Natal (T. guerinii and C. laevigatum) also showed no significant
differences in success rate. Thus, with the one exception at Cape sites, species did not differ overall
in nesting success when considered over the range covered by the study.

While recognising the need for more data from localities from as many parts of each species’
distribution as possible, the present data indicate that differences in nest structure and nesting
behaviour do not result in consistent differences in nesting success. They may, however, affect levels
of attack by specific nest parasites and mortality due to other factors as indicated in Table 12. The
relative importance of the various mortality factors varies markedly between species (Fig. 5).

Effects of nest and behavioural characteristics on mortality factors

The main nest structural and behavioural characteristics of the species studied are summarised

199

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

TABLE 12. Interspecific comparisons of mean success rates and mortality factors.

Per cent success and parasitism

Species

SE

EM

DP

CH

MU

LE

CE

OS

IC

MT

?F

Trap-nests in bush sites

Is

70.6

10.6

5.4

5.2

0.0

0.0

0.0

0.0

0.0

2.2

2.6

Av

64.9

11.7

3.0

0.2

1.6

2.0

2.2

0.0

0.3

1.3

3.3

Auf

61.6

20.3

0.0

1.1

5.8

0.0

0.5

0.0

0.0

4.5

4.5

Tg

49.9

4.0

1.2

28.5

2 2

0.0

0.0

0.0

0.0

4.2

7.4

Cl

47.5

3.3

21.4

6.7

0.0

0.0

0.0

1.9

9.5

0.0

0.9

JP

29.4

11.8

19.3

9.5

2.0

15.3

0.0

0.0

0.0

0.5

9.8

Trap-nests in/on buildings

Tg

47.2

5.5

3.7

19.3

0.2

0.0

0.0

0.0

0.1

16.5

4.9

Pm

35.0

21.3

11.8

4.5

0.7

0.0

0.0

0.0

1.3

6.6

4.4

Cl

30.6

12.4

32.2

2.9

0.9

0.0

0.0

0.4

1.2

14.2

2.9

Aerial mud-nests in/on buildings

Sy

66.0

15.9

2.7

0.0

8.6

0.0

0.0

0.0

0.0

1.6

4.6

Af

49.7

7.3

7.7

15.6

4.1

0.0

0.0

11.5

0.0

1.7

1.7

Dh

41.4

14.7

3.2

13.0

3.7

0.0

0.0

3.7

0.0

11.5

5.9

Ss(N)

31.7

11.3

7.5

5.2

2.3

0.0

0.0

10.4

0.0

19.8

7.2

Ss(C)

23.0

10.1

0.9

10.0

0.1

0.0

0.0

13.4

0.0

27.3

10.9

Species abbreviations as in Table 1.

Success and parasite abbreviations as in Tables 8 and 9.

in Table 1 for comparison with the impact of nest parasites on their hosts. The more important parasites
and their recorded hosts are listed in Table 13.

Mil togramminae

Amobia africa Curran was the most commonly recorded species. Senotainia albifrons occurred
in much lower numbers, and any error resulting from grouping the two species together in the analyses
will be negligible. The discussion below therefore refers mainly to A. africa. The apparently more
restricted range of hosts of S. albifrons (Table 13) may be at least partly due to the small sample.
However, these records are of particular interest since this species has apparently never been reared
from an Afrotropical host, and Palaearctic records are from soil-nesting sphecids (Pape, pers. comm.)
compared with trap-nesting species parasitised in this study.

Bush sites. Levels of parasitism were compared using t-tests. Data from all samples were used on the
assumption that attack by Miltogramminae was not affected by the stage at which trap-nests were sampled.
At Natal sites C. laevigatum suffered significantly higher mortality than A. vitripennis, T. giierinii and A.
fenioralis (Table 14). Parasitism of C. laevigatum and /. pelopoeiformis (at both Natal and Cape sites) did
not differ significantly. These species differ in several nesting characteristics (Table 1) but composition
of cell partitions and end seals, cell linings, the presence or absence of vestibular cells and cocoon types
are probably irrelevant, since larviposition by Miltogramminae occurs during cell provisioning. The
composition of cell partitions could, however, affect migration of maggots between cells.

200

WEAVING: A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

Table 13. Host selection by species of principal parasites

Parasite

Av

Bush sites
Auf Ip

Is

Cl*

Tg*

Sites in buildings
Pm Af Dh

Ss

Sy

MILTOGRAMMINAE

Amobia africa

+

-

+

+!

+ +

-

-

Senotainia albifrons

-

+

+

+!

-

-

-

-

CHRYSIDIDAE

Chrysis sp.

wahlbergi grp

+

+

+!

■¥

+ - +

+

+

oculata grp

-

-

-

-

-

-i-

-

-

lincea grp

-

-

-

-

-

.

+

-

smaragdula grp

-

-

-

-

-

-

-

-

succincta grp

-

+

+

-

-

-

-

-

Stilbum cyanururn

-

-

-

-

-

+ +

-

-

Primeuchroeus sp.
ghilianii grp

-

+

-

-

-

-

-

-

MUTILLIDAE

Stenomutilla syrinx

-

-

-

+!

-1-

+ - +

+

+

Dolichomutilla sycorax

-

-

-

-

+ !

.

+

+

D. heterodonta

-

+

-

-

+l

-1-

-

-

Antennotilla sp.

+

+ +

-

-

-

-

-

-

ICHNEUMONIDAE

Osprynchotus sp.

-

-

-

+

-

+ + +

+

-

Stenarella sp.

+

-

-

+

+!

-

-

-

LEUCOSPIDAE

Leucospis varicollis

+

-

-

-

+

-

-

-

L. ornata

-

-

-

-

+

-

-

-

L. fuelleborniana

-

-i-

-

-

-

-

-

-

POMPILIDAE

Ceropales punctidatus

-

-

-

-

-

-

-

EULOPHIDAE

Melittobia sp.

-1- -1-

+

+!

+

+ + -r

+

+

Species abbreviations as in Table 1.

*C. laevigatum and T. guerinii used trap-nests in both bush and building sites.
! indicates parasitism recorded only from building sites.

201

Per cent parastism by Chrysididae Per cent parastism by Miltogramminae

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

Fig. 6. Relation.ship between number of prey per cell and levels of parasitism by Miltogramminae and Chrysididae.

Characteristics that could influence attack by Miltogramminae are the use of temporary nest
plugs, nest guarding and methods of provisioning. The time required to provision a cell completely
will in many cases be longer in speciesusinggreater numbers of prey. This, combined with the greater
number of visits made to respective nests, provides more opportunities for larviposition by
Miltogramminae. However, the correlation between levels of parasitism and mean numbers of prey

.202

WEAVING: A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

Table 14. Results of t-tests on levels of parasitism due to Miltogramminae.

Species

%

parasitism

(arcsin)

Species

pair

t-tests

Prey per cell

t

df

P

mean

range

n

Trap-nesters in bush sites:

lp(C)

31. T

Cl/Av

3.31

9

<0.01

5.1

2 - 14

13

Cl

25.8=

Ip(C)/Ip(N)

1.17

3

>0.1

11.8

4-27

34

Ip(N)

20.4=”

Ip(N)A'g

1.66

3

>0.1

Av

7.3”

lp(N)/Av

2.22

9

>0.05

1.0

Tg

7.3”

16.8

5-35

55

Auf

0.0'

1.0

Trap-nesters and aerial mud-nesters in/on buildings:

Cl

34.3=

Tg/Pm

1.67

8

>0.05

11.8

4-27

34

Pm

18.0”

Cl/Pm

2.74

7

<0.05

no data

Af

14.8”

Ss(C)/(N)

1.74

4

>0.05

no data

Ss(N)

14.1”

9.5

4-30

30

Db

9.8”

no data

Sy

9.3”

no data

Tg

8.5”

16.8

5-35

55

Ss(C)

3.7

Species abbreviations as in Table 1.

Possession of common superscripts indicates no significant difference.

Species in bold are aerial mud-nesters

per cell was not significant (Fig. 6). Any existing correlation may therefore have been masked by
other factors.

The data suggest the following alternative explanations for the different levels of parasitism
observed.

Chalybion laevigatum - many prey per cell with correspondingly longer times to complete
provisioning, the absence of a temporary plug during the day and no guarding of the nest may all
have contributed to higher levels of parasitism. It was observed that the relatively thin cell partitions
also resulted in the invasion of neighbouring cells by miltogrammine maggots, which may account
for this species suffering the highest mortality due to this parasite.

Auplopus vitripennis and A. femoralis - significantly lower parasitism in both species may partly
be due to the use of only one prey per cell. Also, extremely hard and relatively thick partitions in the
former species probably reduce invasion of neighbouring cells while extensive guarding in the latter
species seems to have eliminated Miltogramminae.

Tricarinodynerus giierinii - parasitism was low in spite of prey per cell being significantly higher
than in C. laevigatum (t=3.47 df=89 p<0.001). Extensive nest guarding occurs in this species,
provisioning in some instances being progressive. Also, it is the only one with a nest turret at the
entrance, though the effect of this on Miltogramminae is not known.

Sites in buildings. Similar tests were carried out on data from trap-nesters and aerial mud-nesters

203

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

from buildings (Table 14). Chalybion laevigatum again suffered the highest level of parasitism by
Miltogramminae compared with all the other species including aerial mud-nesters.

Parasitism of the four aerial mud-nesters in Natal did not differ significantly (Table 14), a
surprising result since S. analis provisions progressively and guards its nest extensively, the remainder
mass-provisioning and leaving their nests open and unattended for considerable periods. Other
unidentified factors must have been playing a role.

In general, only three species, C. laevigatum, /. pelopoeiformis and to a lesser degree P.
meadewaldoi, were relatively heavily parasitised by Miltogramminae (Tables 12 and 14). Species
common both to sites in buildings and bush sites showed the same trends, high levels of parasitism
in C. laevigatum and low levels in T. guerinii, indicating a genuine difference in these species’
susceptibility to Miltogramminae rather than being due to habitat effects.

ClIRYSIDIDAE

Again, there was one predominant species, Chrysis sp. wahlbergi group, which showed a wide
host range but excluded the trap-nesting fsodontia spp. and aerial mud-nester A. aethiopicus. This
species of Chrysis was apparently replaced by another of the succincta group in Isodontia nests,
possibly a reflection of the vegetable cell partitions and plugs compared with the mud partitions used
by other species. Stilbum cyanurum (Forster) was apparently confined to the single- or two-celled
aerial mud-nests of A. aethiopicus and D. hottentottum respectively. The remaining species were
uncommon and it is doubtful if their host selection indicated in Table 13 is comprehensive. For the
analysis, it was assumed that modes of attack were similar for all the chrysidids, i.e. during
provisioning.

Bush sites. Data from all samples were subjected to t-tests on the assumption that parasitism was
not influenced by the sampling. Tricarinodynerus guerinii and C. laevigatum were significantly more
heavily parasitised than A. femoralis and A. vitripennis (Table 15). Isodontia pelopoeiformis was
significantly more heavily parasitised than A. vitripennis but did not differ from any of the other species.

As with Miltogramminae, the composition of cell partitions and end seals, cell linings, presence
or absence of vestibular cells or cocoon type are unlikely to affect parasitism by Chrysididae. Nest
guarding by T. guerinii which apparently reduced parasitism by Miltogramminae seemed not to be
effective against Chrysididae. However, in this case there was a significant correlation between
numbers of prey per cell and levels of attack (Table 1 5)(Fig. 6). Minimal opportunities for oviposition
occurred in nests of A. vitripennis and A. femoralis (one prey per cell), coinciding with the lowest
levels of parasitism. With the latter species, nest guarding is likely to be an important factor as well.
Parasitism of I. pelopoeiformis in Natal did not differ significantly from that in the Cape (Table 15).

Sites in buildings. Trap-nesting species in building sites all used several prey per cell and
differences in this characteristic cannot explain the significantly lower parasitism in nests of P.
meadewaldoi and C. laevigatum compared with T. guerinii. Except for C. laevigatum they all guarded
their nests and showed evidence of progressive-provisioning. Therefore, neither of these
characteristics were effective against Chrysididae. The lower level of parasitism in C. laevigatum
compared with T. guerinii is interestingly reflected in bush sites, albeit not significantly. However,
the reason for the difference is not known.

Synagris analis was the only aerial mud-nester not to be affected by Chrysididae (Table 15),
presumably because of progressive-provisioning and extensive nest guarding. The remaining three
species showed similar levels of parasitism, but only S. spirifex and D. hottentottum were significantly
less parasitised than T. guerinii which seemed to be particularly susceptible to chrysidid attack.

204

WEAVING; A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

Although the trend was not significant, parasitism rates appeared to decrease as cells per nest increased
in spite of the learning capabilities known for chrysidids (Rosenheim, 1987b). This could be another
instance of the “selfish herd” mechanism (Wcislo, 1984) where cell is substituted for nest, but more
data are required. As with I. pelopoeiformis in bush sites, parasitism of nests of S. spirifex did not
differ significantly between the Cape and Natal sites.

Table 15. Results of t-tests on levels of parasitism due to Chrysididae.

Species

%

parasitism

(arcsin)

Species

pair

t-tests

Prey per cell

t

df

P

mean

range

n

Trap-nesters in bush sites:

Tg

32.1“

Tg/Auf

7.31

6

<0.001

16.8

5-35

55

Cl

19.9“

Tg/Av

10.18

9

<0.001

11.8

4-27

34

Ip(N)

13.4“'’

Av/lp

3.62

9

<0.01

5.1

2-14

13

Auf

6.8'"

Cl/Auf

2.48

7

<0.05

1.0

Av

3.T

1.0

Ip(C)

22.1“

Ip(N)

13.4“

Trap-nesters and aerial mud-nesters in/on buildings:

Tg

25.6“

Tg/Dh

2.50

12

<0.05

Af

19.3“”

Tg/Ss

2.71

7

<0.05

Dh

16.0'’

Tg/Pm

2.50

12

<0.05

Ss(N)

11.9'’

Tg/CI

4.18

9

<0.01

Pm

10.7'’

Af/CI

1.55

9

>0.10

Cl

8.0'’

Sy

0.0

Ss(C)

16.5“

Ss(N)

11.9“

Species abbreviations as in Table 1.

Possession of common superscripts signifies no significant difference.

Species in bold are aerial mud-nesters.

Chrysididae were therefore more important in aerial mud-nests, parasitising 3 of 4 species at
Natal sites at relatively high levels compared with 1 or 2 of 7 trap-nesting species (Table 12).
Tricarinodynerus giierinii was particularly susceptible both in building sites and bush sites, again
indicating a true species susceptibility (Table 12).

Mutillidae

Four species were recorded, one of which, Antennolilla sp., occurred only in trap-nests in bush
sites (Table 13). The remainder were mostly confined to sites in/on buildings where populations of

205

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

hosts tend to be greater and the location of nests by the wingless females easier. Dolichomutilla
sycorax (Smith), one of the most commonly recorded species, appeared to specialise mostly on large
multi-celled aerial mud-nests, occasionally parasitising trap-nesters such as 7. guerinii that occurred
in close proximity to their main hosts. Stenomiitilla syrinx (Peringuey) appeared to have a wider range
of hosts, but again mostly in building sites.

Table 16. Results of t-tests on levels of parasitism due to Mutillidae and Osprynchotiis sp.

Species

%

parasitism

(arcsin)

Species

pair

t

t-tests

df

P

Mutillidae

- trap-nests in bush sites:

Auf

11.6“

Auf/Cl

1.42

7

>0.10

lp(N)

9.9“

Av

7.6“

Tg

6.T

Cl

3.4“

ls(N)

0.0

Ip(C)

0.0

Mutillidae

- trap-nests and aerial mud-nests in/on buildings:

Sy

15.5“

Sy/Ss(N)

1.27

4

>0.10

At

10.9“

Sy/Cl

2.51

6

<0.05

Dh

10.6“

Sy/Tg

3.72

7

<0.01

Ss(N)

7.6“

Ss(N)/Tg

2.84

7

<0.05

Pm

4.3“'“

Sy/Pm

0.99

9

>0.10

Cl

3.4^

Tg

1.3*’

Ss(N)

7.6“

Ss(N)/Ss(C)

2.31

4

>0.05

Ss(C)

0.8“

Osprynchotus sp. - trap-nests and aerial

mud-nests in/on buildings:

Af

19.1“

Ss(N)/Dh

2.43

9

<0.05

Ss(N)

18.6“

Dh/Cl

3.21

11

<0.01

Dh

11.5'“

Cl

3.9^

Tg/Pm/Sy

0.0

Ss(C)

20.9“

Ss(N)

18.6“

Species abbreviations as in Table 1.

Possession of common superscripts indicates no significant difference.
Species in bold are aerial mud-nesters.

206

WEAVING; A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

Bush sites. Bartlett’s correction (Snedecor & Cochran, 1967) was applied to small samples
(n<50) showing zero parasitism before using the arcsin transformation. The highest level of parasitism
by Mutillidae was 6.7% in A. femoralis, though this did not differ significantly from that in any of
the other species (Table 16). There is therefore no indication that any of the nesting characteristics
listed in Table 1 for trap-nesters affected parasitism by Mutillidae. It is noteworthy, however, that A.
femoralis was the only trap-nesting species that constructed discrete mud cells all of which were
accessible without having to pass through other cells first.

Sites in buildings. Mortality due to Mutillidae was notably grouped according to nest type, aerial
mud-nests suffering significantly higher levels (=>2.3%) than trap-nests (=<0.9%) with the exception
of P. meadewaldoi (Tables 12 and 16). Aerial mud-nests were therefore apparently more susceptible
than trap-nests to attack by Mutillidae. Thickness of daubing seemed to have no influence on levels
of parasitism, suggesting that Mutillidae either are not deterred by thick nest walls or invade nests
while they are still open. If the latter is the case, progressive-provisioning and nest guarding by S.
analis was not effective against Mutillidae.

ICHNEUMONIDAE: Osprynchotus sp.

Bush sites. Attacks in bush sites were confined to nests of C. laevigatiim, though at low levels
(1 .9%) (Table 1 2). This species did not incorporate vestibular cells into its nests and it is significant
that only outer terminal cells were parasitised, a result of the observed fact that Osprynchotus drills
through end seals with its ovipositor. Nests of/, pelopoeiformis and I. stanleyi also had no vestibular
cells, but their very thick end plugs consisting of vegetable materials would deter such methods of
oviposition.

Sites in buildings. All the aerial mud-nesters except S. analis were parasitised by Osprynchotus
sp., A. aethiopicus and S. spirifex at levels significantly higher than D. hottentottum (Table 16). The
exceptionally thick walls and seals of S. analis nests appeared to prevent oviposition. The high level
of parasitism of the single-celled nests of A. aethiopicus was probably partly due to their thin mud
walls facilitating oviposition. The thicker walls of nests of S. spirifex might be expected to have
reduced levels of Osprynchotus. However, many smaller nests were without additional daubing; their
multi-celled construction may also have counteracted any protection by increasing the rate of
parasitisation of cells compared to that in the more widely dispersed single-cell nests of A. aethiopicus.
The 2-celled nests of D. hottentottum were similarly dispersed to those of A. aethiopicus-, however,
their thicker walls may have provided some protection. It is noteworthy that only 1 1.5% (n=26) of
nests attacked by Osprynchotus had both cells parasitised, suggesting that the cryptic design of these
nests reduced such parasitism compared with that of A. aethiopicus through the failure of
Osprynchotus to detect the presence of two cells.

Chalybion laevigatum was again the only trap-nester to be parasitised, albeit at a level
significantly below those of the aerial mud-nesters (Table 16). Levels of parasitism of 5. spirifex did
not differ significantly between Cape and Natal sites.

ICHNEUMONIDAE: Stenarella sp.

Bush sites. Attacks by this ichneumonid were confined to nests of C. laevigatum and A. vitripennis
(Table 12). A meaningful analysis of the results was not possible due to the high proportion of localities
with zero parasitism (Table 8).

Sites in buildings. In contrast to Osprynchotus, this ichneumonid parasitised trap-nesters only

207

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 1 9, PT. 4 28 APRIL 1 995

(Table 12). Again, a meaningful analysis was not possible, but none of the hosts appeared to be
particularly susceptible.

Thus, the hosts of Osprynchotiis sp. were all aerial mud-nesters with the single exception of the
trap-nesting C. laevigahim. On the other hand, parasitism by Stenarella sp. was confined to trap-
nesters. The range of hosts, C. laevigatiim, P. meadewaldoi, T. guerinii and A. vitripennis, includes
species that incorporate vestibular cells in their nests (Table 1), indicating that this nest characteristic
does not prevent parasitism by Stenarella. Also, with C. laevigatiim, a species that does not use
vestibular cells, parasitism by Stenarella was not confined to the terminal cells as it was with
Osprynchotiis. Of the 15 nests affected, inner cells were parasitised in nine, outer cells in four and
all cells in two of them.

Stenarella must therefore either enter nests prior to final sealing to oviposit through the most
recently completed partition or directly on the cell contents of an unsealed cell, or drill through the
side walls of the nest with its ovipositor. The available evidence supports the former possibility. The
immunity to Stenarella shown by /. pelopoeiforniis and I. stanleyi might appear to contradict this.
However, in these species the thick cell partitions and temporary plugs of vegetable material are likely
to impede oviposition and entry respectively. At the same time, this absence of parasitism strongly
suggests that Stenarella does not oviposit through the side walls.

Contrary to expectation, guarding, as observed in P. meadewaldoi and T. guerinii, apparently
did not prevent parasitisation by Stenarella. However, this does not necessarily support the possibility
of oviposition through the side walls. The occurrence of mass-provisioning and its attendant lack of
guarding in some nests of these species is the most likely explanation.

EulophidaE: Melittobui sp.

Due to the influence of storage on infestation levels (Table 4), analysis was confined to data from
sample category I. In buildings, A. aethiopiciis nests suffered significantly lower parasitism than
those of S. spirifex (Table 17). The single-celled mud pots of A. aethiopiciis each have to be located
anew by searching parasites whereas all the cells in a nest of S. spirifex are readily found once the
nest itself has been located. The similarly low levels of parasitism in S. analis nests compared with
S. spirifex, although statistically insignificant (t=l .96 df=4), suggest that particularly thick walls may
reduce entry by Melittobia. Parasitism of trap-nesters appeared to be higher in buildings (Table 17),
but none of the species that were common to bush sites and buildings differed significantly (t for T.
guerinii =0.85 df=6; t for C. laevigatiim =1.81 df=6).

Table 17. Per cent (arcsin) parasitism of cells by Melittobia in trap-nesting and aerial mud-nesting
species.

Trap-nests

Trap-nests

Aerial mud-nests

(bush)

(buildings)

(buildings)

(Tg

1 1 .40

Tg

20.9“

Ss

24.6“

Auf

9.8“

Cl

18.8“

Dh

17.T‘>

(Ip

6.80

Pm

13.0“

sy

8.0“"

Av

6.2“

Af

8.1"

(Cl

3.40

Species abbreviations as in Table 1.

Species in parentheses inadequately sampled.

Possession of common superscripts within each column signifies no significant difference determined
via t-tests.

208

WEAVING; A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

All species were therefore susceptible to Melittobia regardless of nest type and behaviour. The
data fail to demonstrate any significant differences in susceptibility except between single-celled and
multi-celled aerial mud-nests i.e. nests of A. aethiopicus and S. spirifex respectively (Table 17). The
possibility that S. analis nests are less susceptible than those of S. spirifex suggests that single-celled
nests and thick-walled multi-celled nests may offer two alternative ways of reducing mortality from
Melittobia, though more data are required to confirm this.

Table 18. Results of t-tests on parasitism of trap-nests by Leucospidae and Pompilidae: Ceropales.

Parasite

Host

% parasitism
(arcsin)

t-tests

species t df p

Leucospidae

Ip

26.0-

Ip/Av 2.07 9 >0.05

Av

6.7-

Ceropales

Auf

6.7

no t-test

Av

7.8

insufficient samples

Species abbreviations as in Table 1.

Possession of common superscripts indicates no significant difference.

Leucospidae and Pompilidae: Ceropales pimctulatus

Attack by either of these parasites would not be affected by the different sample categories and
thus all data were used in analysing the results (Table 18).

Leucospidae. No conclusions can be drawn with respect to the influence of nest structure or
nesting behaviour on levels of parasitism. Only two species were host to Leucospidae, /.
pelopoeiformis and A. vitripennis. Apart from both being trap-nesters in bush sites, they showed little
in common. Type of prey, amount of provision per cell and materials used for partitions and seals
differed markedly (Table 1). Since oviposition by Leucospidae is probably achieved by drilling
through the side wall of the nest, the nesting characteristics considered here are unlikely to protect
hosts from parasitism. Leucospidae are therefore potentially important agents in reducing nesting
success in the few hosts that they select.

Ceropales pimctulatus Cameron. Nests only of the pompilids A. femoralis and A. vitripennis
were affected and, since oviposition occurs on the prey before it is taken into the nest, none of the
nesting characteristics considered could influence levels of parasitism. Even prey cleaning would be
ineffective against this type of parasite since Ceropales inserts its egg into one of the spider’s book-
lungs where it is i naccessible to the wasp. No observations were available on other aspects of behaviour
during prey transportation. Specialisation on pompilid hosts means that Ceropales is not an important
factor in nesting success of aculeate wasps in general. Insufficient data were obtained to show how
important Ceropales may be to pompilids but levels of parasitism recorded were mostly low (Tables
9 and 12).

Endogenous mortality

Bush sites. Due to the effects of opening and storage of nests, analysis was confined to data from
sample categories 1 and 4 as discussed in ‘Examination of the Data’. Endogenous mortality is due

209

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

to a variety of mostly unknown factors, and the reasons for any differences cannot be established
from the data obtained. Auplopiis femoralis and A. vitripennis suffered the highest mortality of this
type amongst trap-nesters (Table 19). It is not known whether there is any significance in the fact
that they are both Pompilidae.

Building sites. Endogenous mortality amongst trap-nesting species was significantly higher in
P. meadewaldoi compared with T. giierinii (Table 19). Amongst aerial mud-nesters, A. aethiopicus
suffered significantly lower endogenous mortality than 5. analisovD. hottentottiun. Tricarinodynenis
guerinii was the only trap-nesting species that differed significantly from aerial mud-nesters, showing
lower mortality than S. analis or D. Iiottentottum.

Endogenous mortality accounted for significant levels of mortality in all species. Individual
component factors would first need to be identified before being considered in relation to nest structure
and nesting behaviour.

Table 19. Results of t-testson endogenous mortality in trap-nesting and aerial mud-nesting species.

Species

% mortality
(arcsin)

t-tests

species pair

t

df

P

Trap-nesters in bush sites:

Auf

26.4»

Auf/Av

2.43

11

<0.05

A''

19.4'’

Av/Cl

2.24

9

c.0.05

Cl

12.T

IP

12.P

Tg

11.5^

Trap-nesters and aerial mud-nesters in/on buildings:

Pm

26.7“

Pm/Tg

4.26

8

<0.01

Sy

22.7“

SyATg

2.69

7

<0.05

Dh

21.4“

Dh/Tg

3.48

12

<0.01

Ss

19.4“'’

Cl

19.4'"’

Af

14.4"

Tg

13.4"

Species abbreviations as in Table 1.

Possession of common superscripts indicates no significant difference.

Species in bold are aerial mud-nesters

Other mortality factors

The several other factors recorded were mostly unimportant, causing only low mortality except
for sporadic higher levels at certain localities. Levels were also too low for interspecific comparison.
Mean levels of attack are given in Table 20.

Bombyliidae were recorded from nests of T. guerinii, P. meadewaldoi and A. vitripennis and it
is of interest that the presence of a turret {T. guerinii) did not prevent parasitism by this type of parasite.
Bombyliids are known to oviposit in open holes of various kinds (Evans, 1966a; Evans & O’Neill,
1988) and to flip their eggs into nest entrances of trap-nesters while hovering in front of them
(Krombein, 1 967). The turret of T. guerinii might be expected to prevent the latter method; presumably

210

WEAVING: A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

Table 20. Mortality due to various parasites and predators of minor importance averaged for all
species.

Parasite/predator

Mean % mortality

Range

Bombyliidae

1.6

0.1 - 4.6

Unknown parasites

2.6

0.1 - 9.6

Mites*

2.0

0.1 - 11.5

Psocids*

4.9

0.1 - 20.5

Ants

7.7

1.3- 45.0

(2.4)

(1.3- 4.5)

* - uncertain whether actually caused mortality.

a different technique for oviposition was used by the species concerned {Anthrax pitliecius Fabricius).

Unknown parasites collectively caused similar levels of mortality.

Mortality (presumed) from mites was relatively high at only one locality (11.5%), but being
associated with a small sample of only 26 cells of C. laevigatiim from a bush site is of doubtful
significance.

Psocids invaded significant proportions of cells of P. meadewaldoi at two building sites (11.4
and 20.5%), but these figures were based on small samples (39 and 44 cells). Only 1.1% mortality
was recorded at a third site where the sample was substantial (267 cells); otherwise they occurred at
similar levels to those of the other factors considered in this section. It was not definitely established
whether mites and psocids had actually caused the mortality or had invaded cells after the death of
their occupants.

Ants were recorded invading nests only in bush sites with one exception from a building. Mortality
was low except for one site where they caused 45.0% failure of T. guerinii cells, again associated
with a very small sample of 20 cells. However, there was evidence that ants were particularly active
at this site.

It was not possible to demonstrate any species’ particular susceptibility to any of these factors
and therefore no links with particular nesting characteristics can be suggested.

Effects of certain characteristics of aerial mud-nests on levels of parasitism

Sceliphron spirifex was the only species sufficiently well sampled to provide data on the effects
on parasites of certain characteristics of multi-celled aerial mud-nests. Numbers of cells in nests (nest
size), the relative positions of cells within nests and the degree of daubing have been found, or have
been expected, to influence parasitism especially by Melittobia in other species of Sceliphron
(Freeman & Ittyeipe, 1976; Freeman, 1982). Daubing could also be expected to reduce successful
oviposition by parasites that need to drill through cell walls with their ovipositors, such as
Osprynchotus.

Using the data from all localities, a stepwise discriminant analysis of success, with nest size,
presence or absence of daubing and cell position as independent variables, showed all three variables
to influence success significantly. Nest size was the most important (F=22.258 df=l/1301 p<0.0001)
followed by daubing (F=11.103 df=l/1300 p=0.0009) and cell position (F=9.165 df=4/1299
p<0.000 1 ). A 60.6% correct prediction of successes and failures shows that there are other unidentified
factors also regulating success. The significant effects of size, daubing and cell position were

211

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

confirmed by logistic regression and loglinear analyses. The latter further indicated significant
interactions between daubing and nest size (Chi-square=214.05 df=54 p<0.0001) and nest size and
cell position (Chi-square=126.54 df=99 p=0.0324), but none between daubing and cell position (Chi-
square=3.55 df=4 p=0.4701).

1-4 5-10 11-22 >22

Number of cells per nest

Fig. 7. Effect of nest size (cells per nest) on success rate and mortality factors in A - undaubed and B daubed nests of
Sccliphron spirifex. + - Melittobia, □ - Mutillidae, ▲ - Osprynchotiis, * - Chrysididae, ■ - endogenous mortality.
Shaded bars - success rate.

212

WEAVING: A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

Table 21. Effect of daubing on success rates and parasitism of nests of Sceliphron spirifex.

Size (cells/nest)

1-4

Daubed
5-10 11-22

>22

1-4

Undaubed
5-10 11-22

>22

n (nests)

36

41

39

10

39

19

5

1

n (cells)

101

298

577

288

107

170

65

24

Per cent:

Success

10.9

13.1

33.3

19.8

14.0

25.9

41.5

29.2

Endogenous mortality

10.9

7.4

4.2

8.0

11.2

2.3

4.6

0.0

Melittobia

26.7

25.5

28.8

23.3

4.7

22.9

40.0

50.0

Chrysididae

8.9

12.7

3.8

6.9

10.2

5.3

0.0

0.0

Mutillidae

3.0

3.0

0.7

8.3

0.9

3.5

3.1

16.7

Osprynchotus

15.8

12.4

9.3

13.2

14.0

7.0

0.0

4.2

Miltogramminae

4.9

5.7

1.0

6.2

1.9

4.1

1.5

0.0

Chi-squared tests showed that success rates in both daubed and undaubed nests increased
significantly with nest size except in the largest nests (>22 ceils) (Table 21, Fig. 7). While undaubed
nests appeared to be more successful than daubed nests, this was significant only in the 5-10 cell size
category (Chi-squared = 13.83 df=l). Thus, the presence of daubing appeared to have no positive
effect on overall success and, if anything, tended to result in reduced success. However, sample sizes
for large undaubed nests were much smaller than those for daubed nests and additional data are
required to confirm these conclusions.

Endogenous mortality decreased significantly with increasing nest size but did not differ in
daubed and undaubed nests, except those in the 5-10 cell size category in which it was significantly
higher in daubed nests (Chi-squared = 5.31 df=l). Endogenous mortality was therefore more
dependent on nest size than on the presence or absence of daubing.

Parasitism by Melittobia, the most important mortality factor, depended on both nest size and
daubing. Levels of parasitism were not significantly affected by size in daubed nests (Chi-squared =
2.36 df=3) but increased significantly in larger undaubed nests (Table 21, Fig. 7) (Chi-squared =
32.03 df=3). Large undaubed nests were therefore particularly susceptible to Melittobia, presumably
due to easier penetration of the thinner walls combined with the enhanced opportunity for population
increase in large nests. However, this conclusion requires confirmation with larger samples of
undaubed nests, especially in the >22 cell size category. The reason for the low levels of parasitism
in small undaubed nests compared with their daubed counterparts is unknown.

Parasitism by Chrysididae was significantly higher in smaller nests in both daubed and undaubed
nests. Daubing appeared to result in higher chrysidid attack as well, as indicated by the significant
effect in nests of 5-10 cells (Chi-squared = 6.55 df=l) though the reason for this is not known.

Daubing had no significant effect on levels of attack by Mutillidae. However, the largest nests
suffered significantly higher rates of parasitism (Table 21, Fig. 7), but again this is subject to
confirmation from larger samples.

Size had no significant effect on parasitism by Osprynchotiis in daubed nests. However, there
was a significant decline in parasitism with increasing size of larger undaubed nests. Therefore,
contrary to expectation, daubing does not seem to deter oviposition by such parasites.

Generally, then, mortality due to the more important factors declines with increasing nest size
until a certain size is reached after which this trend appears to be reversed, indicating that there is an

213

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

optimum size, probably within the 1 1 -22 cell size category . The most important exception isMelittobia
which increases dramatically with nest size in undaubed nests; although daubing appears to prevent
this effect, it does not prevent a relatively high level of parasitism in all size classes.

Success of cells also varied according to their positions in nests (Table 22). Cells were classified
according to the diagram in Fig. 3-4a. Outer cells, including those in contact with the substrate (basal
outer) were significantly less successful than inner cel Is. Parasitism by Ospry/zcZ/o/tcv was significantly
higher in these cells. Melittohia appeared to be less prevalent in outer cells, but significance was
marginal. However, there was no evidence that cells further from the sub.strate (outer cells) were less
affected by Melittobia than those adjacent to it (basal inner + basal outer cells) as reported by Freeman
& Ittyeipe, 1976. None of the other main mortality factors were affected by cell position. Smaller
nests will tend to have an increasingly large proportion of outer cells until below a certain size the
entire nest consists only of such cells. This could be one factor that helps to explain reduced success
in smaller nests.

Table 22. Effect of cell position on success rates and parasitism of nests of Sceliphron spirifex.

Basal

inner

Inner

Basal

outer

Outer

Chi-

square

P

n (cells)

380

226

348

465

Per cent

successful cells

30.5

29.2

20.1

20.6

13.38

<0.01

Per cent failure

due to:-

Endogenous mortality

6.8

4.9

6.9

6.4

0.99

>0.80

Melittobia

30.5

31.9

22.7

23.4

8.06

<0.05

Osprynchotiis

6.8

4.4

14.6

15.9

28.72

<0.001

Chrysididae

5.3

5.3

8.6

8.6

5.30

>0.10

Mutillidae

1.6

4.9

4.9

3.4

6.98

>0.05

DISCUSSION

Extensive and apparently random variation in nesting success in individual species is one of the
most obvious findings of this study (Fig. 4). Apart from a trend at certain localities for Synagris aiialis
to achieve greater success and Sceliphron spirifex the least success (Tables 6, 1 1 and 12), there is
little support forthe hypothesis that overall nesting success varies according to nesting habit. However,
interlocality variation may have obscured any more subtle interspecific differences. Such variation
has been reported for several species, including Delta maxillosiun (Chapman, 1959), Sceliphron
laetinn (Smith, 1979), Trypoxylon tenoctitlan (Coville & Coville, 1980), Cra6rort/'gu5/m/5'( Matthews
etal., 1979), Tachysphex ter niinatus {Spof ford etal., 1986) and OAy6c/»5 5'cnct?u5( Hook & Matthews,
1980), attributable in each instance to varying levels of attack by certain nest parasites or differences
in types of parasites present.

This study, as well as reports in the literature (Freeman & Parnell, 1973; Taffe & Ittyeipe, 1976;
Peckham, 1977; Freeman, 1982; Spofford et al., 1986; Rosenheim & Grace, 1987), show that nest
parasites and predators are indeed the major cause of nest/cell failure. Mean mortality due to natural

214

WEAVING: A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

enemies for a mixture of host species, expressed as a percentage of total mortality, calculated from
12 papers with suitable data (Appendix A), is 69.2% (range 37.8-98.8%). Figures from the current
study are very similar, 62.5% (range 31.0-94.1%). It seems likely that similar sets of figures would
be obtained wherever such studies are conducted. Nesting success must therefore depend largely on
factors that affect levels and types of parasitism whether or not they are connected with nesting habits.
They fall under three headings, “external” factors not related to nesting habits, nest structure and
nesting behaviour.

External factors. These are particularly prone to variation between localities. Environmental/
climatic conditions (Rau, 1916; Matthews et al., 1979) prior to, during and after nesting are likely to
affect nesting success, but such parameters have never been measured (Rosenheim, 1989).
Characteristics of the nesting site/substrate and vegetational cover (Taffe & Ittyeipe, 1976; Freeman,
1982; Rosenheim, 1988) may also affect success rates and parasitism. However, contributions from
such factors towards observed variations in parasitism and/or nesting success in this study cannot be
assessed because of the lack of data. It would in any event be very difficult to separate the effects of
each factor in the absence of experimental manipulation.

Parasite populations may also depend on the density and extent of the population of the host
species in question (Freeman & Parnell, 1973; Matthews et al., 1979; Wcislo, 1984), which is at least
partly a function of the length of time that the site has been used. Populations of other species at the
same site prior to and during nesting of the host species, especially if they are poorly protected (Evans,

1987) , may also cause a build up of parasites and determine which species are dominant (Spofford
et al., 1986). Such was the case at FBG where there was a long-established, substantial population
of T. guerinii. This species together with others that utilised trap-nests at this site, C. laevigatum and
P. meadewaldoi, as well as the aerial mud-nester S. analis, showed consistently low success and
relatively high levels of attack by Melittobia, Miltogramminae, Mutillidae and/or Chrysididae
compared with other localities (Tables 8 and 10).

Nesting behaviour. Various features connected either with nest structure or nesting behaviour
during nesting have been suggested as factors that affect nest parasitism and, therefore, nesting
success. Most studies deal with ground-nesting species (40/52), only six being concerned with trap-
nesters and six with aerial mud-nesters. Very few, however, are supported by quantitative data.

Behavioural features include diversionary flights by provisioning adults in response to the
presence of parasites (Linsley & MacSwain, 1956; Evans & Lin, 1959; Alcock, 1974 and 1975;
Matthews et al., 1979; Evans et al., 1980; Gwynne, 1981; Hager & Kurczewski, 1985; Evans &
O’Neill, 1988; Kurczewski & Spofford, 1986; McCorquodale 1986, Spofford et al. 1986, Spofford
& Kurczewski, 1 992), nest cleaning in response to parasite presence (Simonthomas & Simomthomas,
1972; Gwynne, 1981; Sheehan, 1984; Hager & Kurczewski, 1985; Rosenheim, 1987a; Evans &
O’Neill 1988, Spofford & Kurczewski 1992), prey cleaning (Linsley & MacSwain, 1956;
McCorquodale, 1986; Spofford etal., 1986; Spofford & Kurczewski, 1992), nest guarding (Peckham
et al., 1973; Peckham, 1977; Coville & Coville, 1980; Hook & Matthews, 1980; Krombein, 1979;
Evans & Hook, 1982a and 1982b; Coville & Griswold, 1984; Wcislo et al., 1988; Spofford &
Kurczewski, 1992), progressive- provisioning (Evans, 1966b and 1977; Evans & Eberhard, 1970;
Krombein, 1978; Hager & Kurczewski, 1985; Gess & Gess, 1989; Weaving, 1989), delayed
provisioning (Rosenheim, 1987b) and method of transporting prey (Evans, 1962; Evans & O’Neill,

1988) .

Nest guarding and progressive-provisioning were the only two of the above behavioural features
for which observations were obtained in this study. In S. analis, the only species that consistently
provisioned progressively, guarding was the outcome of the females’ almost continual presence in

215

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

unsealed cells containing larvae. Nest guarding in A. fenioralis was not associated with progressive-
provisioning but with nest sharing. It is likely that these species’ relatively high success rates are at
least partly attributable to these behaviours.

The possible existence of alternate provisioning strategies in T. giierinii and P. rneadewaldoi is
of particular interest but requires further investigation, as no data were obtained to compare success
rates and levels of parasitism in progressively-provisioned and mass-provisioned nests of these
species. The adaptive significance of alternate provisioning strategies as displayed by T. guerinii and
P. rneadewaldoi has been discussed elsewhere (Weaving, 1994a). The occurrence of (slow)
progressive-provisioning individuals nesting side by side with those that were (fast) mass-provisioners
suggests that neither strategy is necessarily adopted by a nesting population as a whole but by
individuals according to parasite pressures experienced at particular nest sites. This view is supported
by the fact that individuals seemed to be able to switch from one strategy to the other (Weaving,
1994a).

Nest structure. Structural features of nests reported to affect parasitism include accessory burrows
(Tsuneki, 1963; Evans, 1966a, 1966b, 1966c; Sheehan, 1984; Vinson et aL, 1987; Evans & O’Neill,
1988; Evans & Eberhard, 1970), burrow profile (Evans & Lin, 1959; Evans & O’Neill, 1988), nest
closure (Linsley & MacSwain, 1956; Evans, 1966b; Brockmann, 1985; Hager & Kurczewski, 1985;
Rosenheim, 1987a), temporary nest closure (Evans & Lin, 1959; Simonthomas & Simonthomas,
1972; Gess & Gess, 1975; Peckham, 1977; Brockmann, 1985; Spofford etal., 1986; Evans & O’Neill,
1988), levelled or conspicuous tumuli (Evans & Lin, 1959; Wcislo, 1984; Hager & Kurczewski, 1985;
Rosenheim, 1987a; Evans & O’Neill, 1988), daubing the outer surfaces of aerial mud nests with extra
mud (Freeman, 1982), building multicellular nests (Freeman & Ittyeipe, 1976; Evans, 1977),
differences in nest substrates (Taffe & Ittyeipe, 1975; Freeman, 1982), numbers of cells comprising
individual nests (Medler, 1964; Freeman & Ittyeipe, 1975; Evans, 1977; Freeman, 1977; Brooke,
1981; Evans & O’Neill 1988) and the inclusion of vestibular or empty cells (Evans & Eberhard, 1970;
Tepedino et ai, 1979; Coville & Griswold, 1984; Freeman & Ittyeipe, 1993).

The species included in this study differed markedly in their overall nest structures, the thickness
and materials used in construction of cell partitions and nest closures, the extent of daubing on mud
nests, use of temporary closures and the inclusion of vestibular cells. However, because of the lack
of significant interspecific differences in nesting success, there are no indications of links between
nest structure and success except with regard to numbers of cells comprising multi-celled nests.
Results indicate that there may be an optimum size at which success is maximised; for S. spirifex this
falls within the range of 11-22 cells per nest. However, the fact that the single- and two-celled nests
of A. aethiopicus and D. hottentottum respectively achieved higher levels of success suggests that
the optimum number of cells in aerial mud-nests varies with species. It is also likely that such an
optimum will vary even within a species according to population and types of parasites present, since
the effect of nest size in S. spirifex differed with different parasites.

Hostiparasite interactions. Significant differences in susceptibility to particular nest parasites
were recorded for certain species (Tables 14-18). However, in most cases these differences cannot
be definitely attributed to specific structures or behaviours because of the lack of direct observations
of interactions between nesting habits and parasites. Furthermore, levels of parasitism are the outcome
of the combined effects of several factors most of whose individual contributions cannot be reliably
assessed from the available data.

Nevertheless, there were certain trends that should be followed up experimentally or by direct
observation. Thick cell partitions of A. vitripennis with their very hard surfaces were associated with
reduced miltogrammine parasitism; the much thinner cell partitions of C. laevigatum&Wowed invasion

216

WEAVING: A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

of neighbouring cells by maggots which did not occur in A. vitripennis nests. However, this difference
in susceptibility could equally well be ascribed to numbers of prey per cell. Cells of A. vitripennis
are supplied with only one prey and are then immediately sealed, reducing the opportunity for
larviposition by flies. In contrast, C. laevigatiini cells remain open and unguarded for extended periods
whilst the females hunt for the many prey required. Further observations are needed to determine
which interpretation is correct. Perhaps both factors play a role.

Progressive-provisioning accompanied by nest guarding in 5. analis is the most likely explanation
for this species’ lack of parasitism by Chrysididae. Yet T. giierinii, which also employs extensive
nest guarding, suffers high levels of attack. At the same time, both species similarly suffered low
levels of parasitism by Miltogramminae which could also be attributed to nest guarding. Detailed
observations of how such parasitism is achieved are required to explain these apparent anomalies.

Osprynchotiis parasitised cells by ovipositing through the end seals of trap-nests or mud walls
of aerial mud-nests. It was obvious in the case of trap-nests, as with nests of C. laevigatiini in which
parasitism was confined to terminal cells, that mortality from such parasites was due to the absence
of vestibular cells. It also seems likely that very thick wal Is in aerial mud-nests prevent such parasitism,
as demonstrated by nests of S. analis compared with the progressively thinner-walled nests of S.
spirifex, D. hottentottiim and A. aethiopiciis (Table 16). A different pattern of parasitism by the
apparently very similar Stenarella emphasises the importance of making observations on modes of
attack. None were obtained for this species during this study, but this ichneumonid clearly uses a
different strategy from that of Osprynchotiis.

While natural enemies are considered to have controlled the development of many aspects of
nest structure and nesting behaviour (Wcislo et al., 1985; Spofford et al, 1986; Rosenheim, 1987a
and 1989), including the early steps leading to sociality (Rosenheim, 1989), the overall direction of
evolution of nesting behaviour and nest structure is in the end determined by complex interactions
between numerous factors (Wcislo et al, 1985). This makes it very difficult, if not impossible, to
assess the roles of individual factors,. including, of course, individual types of parasites, in isolation.

A further complication that does not appear to have been considered in the literature is how
representative the results obtained from a particular site are in relation to the species’ overall
distribution. This is of particular importance in comparative studies such as the present one. The sites
used in this study represent only a minute portion of each species’ incompletely known overall
distribution. It is often not known whether the sites are in a marginal or more central part of each
species’ range, nor how nesting success varies in different parts.

In conclusion, the present results indicate that the activities of parasites are a major factor in
regulating nesting success in solitary wasps. Since no species is equally protected against each type
of parasite, the extensive variation in nesting success that does occur is a reflection of the size and
species composition of parasite populations present at each site at the time of nesting. A high level
of success will be due either to low populations of nest parasites at the locality in question, regardless
of nesting behaviour, or to the fact that the species concerned is well protected, by virtue of certain
nesting habits, against the dominant parasite that is present.

ACKNOWLEDGEMENTS

Thanks are due to the Director of Natal Parks, Game and Fish Preservation Board for issuing
permits for carrying out research in areas under his control; to Mr T.C. White for allowing free access
to his farm Hilton; to Prof. D. Brothers of University of Natal for identifying the Mutillidae; to Dr
Thomas Pape of the Zoological Museum, University of Copenhagen for identifying the

217

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

Miltogramminae; to Dr F.W. Gess of Albany Museum for confirming the identity of Proepipona
meadewaldoi and for identifying the Chrysididae; to Dr J.M. Carpenter of the American Museum of
Natural History and Dr A. Giordani Soika of Museo Civico di Storia Naturale di Venezia for
identifying some Eumenidae; to Dr Jay Rosenheim of University of California, Dr Chris Starr of
University of the West Indies and Dr Bill Wcislo of Cornell University for kindly criticizing early
drafts of the manuscript; to Mrs Sarah Radloff of Department of Mathematical Statistics of Rhodes
University for conducting the loglinear, logistic regression and stepwise discriminant analyses; to
Albany Museum for providing funds in support of this project.

REFERENCES

Alcock, J. 1974. The behaviour of Philanlhus crabroniformis (Hymenoptera: Sphecidae). Journal of Zoology, London 173:
233-246.

Alcock, J. 1975. The nesting behavior of PhUanthus nmltimacidatiis Cameron (Hymenoptera, Sphecidae)./lw7cncfl/; Midland
Naturalist 93: 222-226.

Alm, S.R. and Kurczewski, F.E. 1984. Ethology of Anoplius tcncbrosus (Cresson) (Hymenoptera: Pompilidae). Proceedings
of the Entomological Society of Washington 86: 110-119.

Brockmann, H.J. 1985. Tool use in digger wasps (Hymenoptera: Sphecinae). Psyche 92: 309-329.

Brooke, M de L. 1981. The nesting biology and population dynamics of the Seychelles potter wasp Eumcnes aUuaudi Perez.
Ecological Entomology’ 6: 365-377.

Chapman, R.F. 1959. Some observations on Pachyophthalnius africa Curran (Diptera: Calliphoridae), a parasite of Eumenes
maxiUosus De Geer (Hymenoptera: Eumenidae). Proceedings of the Royal Entomological Society of London 34: 1 -6.
CoviLLE, R.E. 1979. Biological observatoions on Trypoxylon (Trypargilum) orizabense Richards in Arizona (Hymenoptera:
Sphecidae). Journal of the Kansas Entomological Society 52: 613-620.

CoviLLE, R.E. and Coville, P.L. 1980. Nesting biology and male behavior of Trypoxylon (Trypargilum) tenoctitlan in Costa
Rica (Hymenoptera: Sphecidae). A/i/itt/s of the Entomological Society of America 73: 110-119.

Coville, R.E. and Griswold, C. 1983. Nesting biology of Trypoxylon xanthandrum in Costa Rica with observations on its
spider prey (Hymenoptera: Sphecidae; Araneae: Senoculidae). Jour/m/ of the Kansas Entomological Society 56:
205-216.

Coville, R.E. and Griswold, C. 1984. Biology of Trypoxylon (Trypargilum) superbum (Hymenoptera: Sphecidae), a spider-
hunting wasp with extended guarding of the brood by males. Journal of the Kansas Entomological Society 57: 365-316.
Danks, H.V. 1971. Nest mortality factors in stem-nesting aculeate Hymenopteta. Journal of Animal Ecology 40: 79-82.
Endo, a. 1980. On the host-cleptoparasite relationship between the spider wasp Episyron arrogans (Smith) (Hymenoptera,
Pompilidae) and the miltogrammine fly Metopla sauteri (Townsend) (Diptera, Sarcophagidae). Japanese Journal
of Ecology 30: 117-132.

Evans, H.E. 1957. Studies on the comparative ethology of digger wasps of the genus Bembix. Ithaca, New York: Comstock.
Evans, H.E. 1962. The evolution of prey-carrying mechanisms in wasps. Evolution 16: 468-483.

Evans, H.E. 1966a. The accessory burrows of digger wasps. Science 152: 4,5,471.

Evans, H.E. 1966b. The comparative ethology and evolution of the sand wasps. Cambridge, Massachusselts: Harvard
University Press.

Evans, H.E. 1966c. The behavior patterns of solitary wasps. Antmal Review of Entomology 11: 123-154.

Evans, H.E. 1971. Observations on the nesting behavior of wasps of the tribe Cercerini.your/m/ of the Kansas Entomological
Society 44: 500-523.

Evans, H.E. 1977. Extrinsic versus intrinsic factors in the evolution of insect sociality. BioScience 27: 613-617.

Evans, H.E. 1987. Observations on the prey and nests of Podalonia occidentalis Murray (Hymenoptera: Sphecidae). Pan-
Pacific Entomologist 63: 130-134.

218

WEAVING: A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

Evans, H.E. and Eberhard, MJ.W. 1970. The Wasps. Ann Arbor: University of Michigan Press.

Evans, H.E. and Hook, A.W. 1982a. Communal nesting in AustralianCerccm digger wasps. In: The Biology of Social Insects.

Proc Ninth Congr Internat Union Stud Soc Insects; Boulder Colo; Aug 1982. 159-163.

Evans, H.E. and Hook, A.W. 1982b. Communal nesting in the digger wasp Cerceris australis (Hymenoptera: Sphecidae).
Australian Journal of Zoology 30: 557-568.

Evans, H.E., Kurczewski, F.E. and Alcock, J. 1980. Observations on the nesting behaviour of seven species of Crabro
(Hymenoptera, Sphecidae). Journal of Natural llistor)’ 14: 865-882.

Evans, H.E. and Lin, C.S. 1959. Biological observations on digger wasps of the genus Philantlius (Hymenoptera: Sphecidae).
The Wasmann Journal of Biology 17: 115-132.

Evans, H.E. and O’Neill, K.M. 1988. The natural history and behavior of North American beewolves. Ithaca, New York:
Comstock.

Freeman, B.E. 1973. Preliminary studies on the population dynamics of Sccliphron assimile (Dahlbom) (Hymenoptera:
Sphecidae) in Jamaica. Journal of Animal Ecology 42: 173-182.

Freeman, B.E. 1977. Aspects of the regulation of size of the Jamaican population of Sceliphron assimile Dahlbom.

Journal of Animal Ecology 46: 231-247.

Freeman, B.E. 1982. The comparative distribution and population dynamics in Trinidad of Sccliphron fistulariurn (Dahlbom)
and S. asiaticum (L.) (Hymenoptera: Sphecidae). Biological Journal of the Linnean Society 17: 343-360.
Freeman, B.E. and Ittyeipe, K. 1976. Field studies on the cumulative response of Melittobia sp. (Hawaiiensis complex)
(Eulophidae) to varying host densities. Journal of Animal Ecology 45: 415-423.

Freeman, B.E. and Ittyeipe, K. 1993. The natural dynamics of the eulophid parasite Melittobia australica. Ecological
Entomology 18: 129-140.

Freeman, B.E. and Parnell, J.R. 1973. Mortality of Sceliphron assimile Dahlbom (Sphecidae) caused by the eulophid
Melittobia chalybii Ashmead. Journal of Animal Ecology 42: 779-784.

Gess, F.W. 1980. Some aspects of the ethology of Dasyproctus westermanni (Dahlbom) (Hymenoptera: Sphecidae:
Crabroninae) in the Eastern Cape Province of South Africa. Annals of the Cape Provincial Museums (Natural
History) 13: 95-106.

Gess, F.W. and Gess, S.K. 1975. Ethological studies of Bembecinus cinguliger (Smith) and B. o.xydorcus (Handl.)

(Hymenoptera: Sphecidae), two southern African turret-building wasps. Annals of the Cape Provincial Museums
(Natural History) 11: 21-46.

Gess, F.W. and Gess, S.K. 1976. An ethological study of Parachilus insignis (Saussure) (Hymenoptera: Eumenidae) in the
Eastern Cape Province of South Africa. Annals of the Cape Provincial Museums (Natural History) 11: 83-102.
Gess, F.W. and Gess, S.K. 1982. Ethological studies of Isodontia simoni (du Buysson), I. pelopociformis (Dahlbom) and 1.

Stanley) (Kohl) (Hymenoptera: Sphecidae: Sphecinae) in the Eastern Cape Province of South Africa. An/w/s of the
Cape Provincial Museums (Natural History) 14: 151-171.

Gess, S.K. and Gess, F.W. 1989. Notes on nesting behaviour in Bembix bubalus Handlirsch in southern Africa with the
emphasis on nest sharing and reaction to nest parasites (Hymenoptera: Sphecidae). Annals of the Cape Provincial
Museums (Natural History) 18: 151-160.

G WYNNE, D.T. 1981. Nesting biology of the bumblebee wolf Philanthus bicinctus (Mickel) (Hymenoptera: Sphecidae).
American Midland Naturalist 105: 130-138.

Hager, B.J. and Kurczewski, F.E. 1985. Cleptoparasitism of Ammophila hard (Fernald) (Hymenoptera: Sphecidae) by
Senotainia vigilans Allen, with observations on Phrosinella aurifacies Downes (Diptera: Sarcophagidae). Psyche
92: 451-462.

Hook, A.W. and Matthews, R.W. 1980. Nesting biology of Oxybelus sericeus with a discussion of nest guarding by male
sphecid wasps (Hymenoptera). Psyche 87: 21-37.

Hungerford, H.B. and Williams, F.X. 1912. Biological notes on some Kansas Hymenoptera. Entomological News 23: 241-
260.

219

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

Kimsey, L.S. 1980. Notes on the biology of some Panamanian Pompilidae, with a description of a communal nest
(Hymenoptera). Pan-Pacific Entomologist 56: 98-100.

Krombein, K.V. 1967. Trap-nesting wasps and bees: life histories, nests, and associates. Washington, D.C.: Smithsonian Press.

Krombein, K.V. 1978. Biosystematic studies of Ceylonese wasps 111. Life history, nest and associates of Paraleptomenes
mephitis (Cameron) (Hymenoptera: Eumenidae). Journal of the Kansas Entomological Society 51: 721-734.

Krombein, K.V. 1979. Sphecoidea. In: Krombein, K.V., Hurd, P.D., Smith, D.R. and Burks, B.D. eds. Catalog of Hymenoptera
in America north of Mexico. Washington, D.C.: Smithsonian Institution Press, pp. 1573-1740.

Kurczewsk.1, F.E. 1989. Nesting behaviour of Tachysphex acutus, with a discussion of multicellular nest excavation in the
genus (Hymenoptera; Specidae). Journal of the Kansas Entomological Society 62: 44-58.

Kurczewskj, F.E. and Peckham, D.J. 1982. Nesting behavior of Z-vTor/asMb/rn (Say) (Hymenoptera: Sphecidae).Proceet/bigx
of the Entomological Society of Washington 84: 149-156.

Kurczewski, F.E. and Spofford, M.G. 1986. Observations on the nesting behaviors of Tachytes parvus Fox and T. obducUis
Fox (Hymenoptera: Sphecidae). Proceedings of the Entomological Society of Washington 88: 13-24.

Linsley, E.G. and MacSwain, J.W. 1956. Some observations on the nesting habits and prey of Cerccris californica Cresson
(Hymenoptera, Sphecidae). Annals of the Entomological Society of America 49: 71-84.

Matthews, R.W., Hook, A. and Krispyn, J.W. 1979. Nesting behavior of Crabro argusinus and C. hilaris (Hymenoptera:
Sphecidae). Psyche 86: 149-166.

McCorquodale, D.B. 1986. Digger wasp (Hymenoptera: Sphecidae) provisioning flights as a defence against a nest parasite,
Senotainia trilineata (Diptera: Sarcophagidae). Canadian Journal of Zoology 64: 1620-1627.

Medler, J.T. 1964. Parasitism of Eumeninae by cuckoo wasps in trap-nests in Wisconsin (Hymenoptera: Chrysididae).
Proceedings of the Entomological Society of Washington 66: 209-215.

O’Brien, M.F. and Kurczewski, F.E. 1982. Ethology and overwintering of Podalonia luctuosa (Hymenoptera: Sphecidae).
The Great Lakes Entomologist 15: 261-275.

Parker, F.D. 1984. Nests and nest associates of two “red” Od)7ien« wasps (Hymenoptera: Eumenidae). Jonrna/ of the Kansas
Entomological Society 57: 521-525.

Peckham, D.J. 1977. Reduction of miltogrammine cleptoparasitism by male Oxybclus subulatus (Hymenoptera: Sphecidae).
Annals of the Entomological Society of America 70: 823-828.

Peckham, D.J. , Kurczewski, F.E. and Peckham, D.B. 1973. Nesting behaviorofNearctic species of Ox>’bc/«s (Hymenoptera:
Sphecidae). Annals of the Entomological Society of America 66: 647-661.

Rau, P. and Rau, N. 1916. The biology of the mud-daubing wasps as revealed by the contents of their nests. Journal of Animal
Behaviour 6: 27-63.

Rosenheim, J.A. 1987a. Nesting behavior and bionomics of a solitary ground-nesting wasp, Ammophila dysmica
(Hymenoptera: Sphecidae): influence of parasite \itess\ite. Annals of the Entomological Society of America 80: 739-
749.

Rosenheim, J.A. 1987b. Host location and exploitation by the cleptoparasitic wasp Argochrysis armilla: the role of learning
(Hymenoptera: Chrysididae). Behavioral Ecology and Sociobiology 21: 401-406.

Rosenheim, J.A. 1988. Parasite presence acts as a proximate cue in the nest-site selection process of the solitary digger wasp,
Ammophila dysmica (Hymenoptera: Sphecidae). Jour/ta/ of Insect Behavior 1: 333-342.

Rosenheim, J.A. 1989. Behaviorally mediated spatial and temporal refuges from a cleptoparasite, Argochrysis armilla
(Hymenoptera: Chrysididae), attacking a ground-nesting wasp, Ammophila dysmica (Hymenoptera: Sphecidae).
Behavioral Ecology and Sociobiology 25: 335-348.

Rosenheim, J.A. and Grace, J.K. 1987. Biology of a wood-nesting wasp, Mimumesa mixta (W. Fox) (Hymenoptera:
Sphecidae), and its parasite,, Elampus viridicyaneus Norton (Hymenoptera: Chrysididae). Proceedings of the
Entomological Society of Washington 89: 351-355.

Sheehan, W. 1984. Nesting biology of the sand wasp Stictia heros (Hymenoptera: Sphecidae: Nyssoninae) in Costa Rica.
Journal of the Kansas Entomological Society 57: 377-386.

220

WEAVING: A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

SiMONTHOMAS, R.T. and Simonthomas, A.M.J. 1972. Some observations on the behaviour of females of Philanlhus thangidum
(F.) (Hymenoptera, Sphecidae). Tijdschrift voor Entomologie 115: 123-139.

Smith, A.P. 1978. An investigation of the mechanisms underlying nest construction in the mud wasp Paralastor sp.
(Hymenoptera: Eumenidae). Behaviour 26: 232-240.

SMiTtt, A. 1979. Life strategy and mortality factors of Sccliphron laetum (Smith) (Hymenoptera: Sphecidae) in Australia.
Australian Journal of Ecology 4: 181-186.

Snedecor, G.W. and Cochran, W.G. 1967. Statistical Methods. Ames: Iowa State University Press.

Spofford, M.G., Kurczewski, F.E. and Peckham, D.J. 1986. Cleptoparasitism of Tachysphc.x terminatus (Hymenoptera:
Sphecidae) by three species of Miltogrammini (Diptera: Sarcophagidae). Annals of the Entomological Society of
America 79: 350-358.

Spofford, M.G. and Kurczewski, F.E. 1992. Counter-cleptoparasitic behaviours of species of Sphecidae (Hymenoptera) in
response to Miltogrammini larviposition (Diptera: Sarcophagidae). yottrna/ of Natural History 26: 993-1012.

Taffe, C.A. and Ittyeipe, K. 1976. The effect of nest substrata on the mortality of Eumenes colona Saussure (Hymenoptera)
and its inquiVmes. Journal of Animal Ecology’ 45: 303-311.

Tepedino, V.J., McDonald, L.L. and Rothweli., R. 1979. Defence against parasitization in mud-nesting Hymenoptera: can
empty cells increase net reproductive output? Behavioral Ecology and Sociobiology 6: 99-104.

Tsuneki, K. 1963. Comparative studies on the nesting biology of the genus 5p/tcx(s.l.) in east Asia (Hymenoptera: Sphecidae).
Memoirs of the Faculty of Liberal Arts, Fukui University 13: 13-78.

Vinson, S.B., Frankie, G.W. and Coville, R.E. 1987. Nesting habits of Ccntrisflavofasciata Friese (Hymenoptera: Apoidea:
Anthophoridae) in Costa Rica. Journal of the Kansas Entomological Society 60: 249-263.

Wcislo,W.T. 1984. Gregarious nesting of a digger wasp as a “selfish herd” response to a parasitic fly (Hymenoptera: Sphecidae;
Diptera: Sarcophagidae). Behavioral Ecology and Sociobiology 15: 157-160.

WeiSLO, W.T., Low, B.S. and Karr, C.J. 1985. Parasite pressure and repeated burrow use by different individuals of Crabro
(Hymenoptera: Sphecidae; Diptera: Sarcophagidae). Sociobiology 11: 1 15-125.

WeiSLO, W.T., West-eberhard, M.J. and Eberhard, W.G. 1988. Natural history and behavior of a primitively social wasp,
Auplopus scmialatus, and its parasite, Irenangelus eherhardi (Hymenoptera: Pompilidae). Journal of Insect
Behavior 1: 247-260.

WEAVtNG, A.J.S. 1989. Nesting strategies in some southern African species of Ammophila (Hymenoptera: Sphecidae).yot/r;m/
of Natural History 23: 1-16.

Weaving, A.J.S. 1994a. Nesting behaviour in three Afrotropical trap-nesti ng wasps, Chalybion laevigatum ( Kohl ), Proepipona
meadcwaldoi Bequaert and Tricarinodynerus guerinii (Saussure) (Hymenoptera: Sphecidae, Eumenidae). The
Entomologist \\3: 183-197.

Weaving, A.J.S. 1994b. Notes on nesting behaviour in two Afrotropical auplopine wasps, Auplopus vitripennis Smith and/\.
femoralis (Arnold) (Hymenoptera: Pompilidae). The Entomologist 113: 140-153.

221

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

Appendix A. Success rates and mortality factors extracted from published accounts.

Per cent success and parasitism

Sp

SE

EM

DP

CH

MU

OS

MT

AERIAL MUD-NESTS
Auplopus esmeralda (Kimsey, 1980)

(72.5)

Delta maxillosum (Chapman, 1959)

(62.0) 38,0(22-51)
Eumenes alluaudi (Brooke, 1981)

65.9 8.2

73.0 (sealed cells)

5.0 (unsealed cells)

62.0 (c.3 cells/nest)

90.0 (16-26 cells/nest)

Eumenes colona (Taffe & Ittyeipe, 1976)

57.7 10.3

70.0 14.1
Pachodynerus nasidens (Jayasingh, in Taffe

& Ittyeipe, 1976)

Paraleptomenes mephitis (Krombein, 1978)

(85.0) 11.5 DP+HY

Sceliphron asiaticum (Freeman, 1982)

1.5

TRAP-NESTERS

Ancistrocerus antilope (Medler, 1964)

(39.5)

Euodynerus foraminatus (Medler, 1964)

(55.7)

Isodontia pelopoeiformis (Gess & Gess, 1982)

(75.0) 16.7 5.5

/. stanleyi (Gess & Gess, 1982)

(82.3) 11.8

Ody nevus erythrogaster 1984)

(78.0) 7.1 0.1

95(1 nest)
234

2.7 large

1268

21.1 rocks
0.4 roots
27.5 substrate
0.0 hanging

181

56.2 4.3 1.5

0,1

2.6

32.6

1280

S. assimile (Freeman, 1977 and 1973)

61.1

28.6

17059

59.9 8.3 0.1

30.6

2290

S.fistularium (Freeman, 1982)

64.4 6.6 5.4

0.3

1.5

19.7

1966

S. laetum (Smith, 1979)

61.8 21.0

7.9

7.9

1006

Trypoxylon albitarsis (Rau & Rau, 1916)
76.0 17.0

2.0

1282

159

260

144

17

222

2.2

712

WEAVING: A COMPARISON OF NESTING SUCCESS IN SOME AFROTROPICAL ACULEATE WASPS

Appendix A. (continued)

Sp SE EM DP CH MU OS MT

Trypargilum oryzabense (Coville, 1979)

(58.0)

23.0 17

T. superbum (Coville & Griswold, 1984)

57.1 29.0

91

T. xanthandrum (Coville & Griswold, 1983)

77.7 11.1 5.5

36

Trypoxylon tenoctitlau (Coville & Coville, 1980)

56.0 7.0 36.0

144

GROUND-NESTERS

Ammophila dysmica (Rosenheim, 1987a)

48.0 9.1 3.3 25.1

0.0

275

A. ferrugiueipes (Weaving, 1989)

(93.6) 3.2

37

A. insignis (Weaving, 1989)

(69.3) 3.1

65

A. harti (Hager & Kurczewski, 1985)

(80.0-96.0) 4.0

245

Bembecinus cinguUger (Gess & Gess, 1 975)
(97.4)

39

B. oxydorcus (Gess & Gess, 1975)

(100.0)

16

Centris flavofasciata (Vinson et ai, 1987)

(41.0)

22

Cerceris fumipennis (Evans, 1971)

(78.8) 9.6 11.5

52

Crabro advena Evans et al., 1980)

(14.0-100)

7/26/31 3 nests

C. cribrellifer (Evans et al., 1980)

(48.6) 51.4

72

C. monticola (Evans et al., 1980)

(90.7) 1.9 9.3

107

C. argusinus (Matthews et al., 1979)
(83.3-100)

?

C. hilaris (Matthews et al., 1979)

(73.3) 26.7

<40

C. cribrellifer (Wcislo et al., 1985; Wcislo, 1984)
(69.2) 30.8

C.155

(74.9) 25.1

20

Lyroda subita (Kurczewsi & Peckham, 1982)
(38.4) 4.5 47.6

21

Oxybelus sericeus (Hook & Matthews, 1980)
(72.0) 14.5

75

(60.3) 36.0 3.7

107

223

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 4 28 APRIL 1995

Appendix A. (continued)

Sp SE EM DP CH MU OS MT

O. subulatus (Peckham, 1977)

(57.1) 0.5

19.4 - 33(5-47) no males

361

18(4-33) -f males

Parachilus insigitis (Gess & Gess,

1976)

(73.0)

8.0

38

Philanthus bicinctus (Gwynne, 1981)

(92.5)

l.S

40

P. gibbosus (Evans & Lin, 1959)

(100.0)

12

P. politus (Evans & Lin, 1959)

(100.0)

18

P. sotivagus (Evans & Lin, 1959)

(97.5)

2.5

40

Podalonia luctuosa (O’Brien & Kurczewski, 1982)

(50.0)

10

P. occidentalis (Evans, 1987)

(25.1)

74.9

12

Tachysphex terminatus (Spofford et al., 1986)

(42.1-

57.9-

76

69.4)

30.6

98

T. acutus (Kurczewski, 1989)

(79.0)

(21.0)

14/4/1

Tflc/ijto parvMS (Kurczewski & Spofford, 1986)

(89.5)

10.5

19

PLANT/WOOD-BURROWERS

Dasyproctus westermanni (Gess, 1980)

(39.0) 38.0

100

Mimumesa mixta (Rosenheim & Grace, 1987)

(41.0)

(50.0)

548

Abbreviations for column headings:-

SE - successful cells; EM - endogenous mortality; DP - miltogramminae; CH - Chrysididae;
MU - Mutillidae; OS - Osprynchotus sp.; MT - Melittobia.

224

INSTRUCTIONS TO AUTHORS

LAYOUT of recent issues should be examined for details of acceptable style and format.

MANUSCRIPTS should be submitted on floppy disk accompanied by three print-outs
(spacing lYz or 2). In exceptional cases typed manuscripts will be accepted. These should be
submitted in triplicate, typed on A4 size paper, double-spaced, on one side of the paper only
and with wide margins all round.

Only generic and specific names should be underlined to indicate italics. All other marking
up should be left to the editor.

Footnotes should be avoided as far as possible.

All measurements should be metric.

Systematic papers should follow the rules of the International code of zoological
nomenclature or the International code of botanical nomenclature, as applicable.

ILLUSTRATIONS should bear the author’s name, title of paper and number in soft pencil on
the reverse side. Diagrams, line drawings and photographs are all regarded as figures, should
be termed as such and should be numbered in a single series.

In addition to the original illustrations a photocopy of each should be submitted.

Diagrams and line drawings should be reducible to a size not exceeding 14 x 16 cm (17,5 cm
including captions). Photographs for half-tone blocks should be final size, of high contrast
and printed on glossy paper. Colour illustrations are only printed under special circumstances
and their financing must be arranged by the author.

A scale should appear with all illustrations or the magnification should be given in the
caption.

A list of captions should be submitted on a separate sheet.

TABLES should be submitted on separate sheets. Foldouts and tables printed sideways should
be avoided.

REFERENCES should be listed alphabetically at the end of the paper. Abbreviations of
periodical titles should conform with those in the World list of periodicals (4th ed.). Layouts
must be as follows;

Alexander, J. 1969. The indirect evidence for domestication. In: Ucko, P. J. and G. W.
Dimbleby eds, The domestication and exploitation of plants and animals. London:
Duckworth, pp. 123-129.

De Winter, B. 1969. Plant taxonomy today. S.AfrJ.Sci. 65 (8): 229-242.

JuBB, R. A. 1967. Freshwater fishes of southern Africa. Cape Town: Balkema.

GALLEY PROOFS are sent (when practicable) to authors for checking printer’s errors.
Adjustments of language, style and punctuation and any other amendments of the original
manuscript, if made at this stage, may be charged to authors.

AUTHOR’S COPIES (50) are issued free of charge. When more than one author is involved
the fifty free copies will be sent to the senior author. Additional copies ordered before printing
are supplied at a reduced cost.

CADAR. P.E.-KS87

» /ft

'u.-f ;lj' ■ f \ *. ' > ' Jl'i.

r 1.| t >t :'>/p illlVltOn

•' ■• ••hh ViJ'Mi

. g. S .Q'il qt.

‘ Iiiui'lt'?

.‘V’ ■■»►' '■ '

^ ■, I, ill

iij) . Ill 'll >i)

"■tlbi'iil

ii 1 i|-i ■'•^1 i( \

iw-i I'.li. ijiii
. .11. ..(..mI 'M1«

U'lUi <mn

M './i '1. /■

. , ' . ' u ; " ’ I A

It

' •I'UtVsT
: I 'I nd

,/ I'l.i i.m

'li ' ,.li •'!

, f

' ._ ’

- , ,f . 1 '

7V"‘»

’ ' i.\ ....:•

1 'MlllUr*

■ ilf '4' ■ ‘ U'

.1 'T.,-'

Annals of the

Cape Provincial Museums

Natural History

Ann. Cape Prov. Mus. (nat. Hist.)

Volume 19 Part 5 28th May 1995

\

' A preliminary catalogue of fossil algal, plant, arthropod, and fish remains from a Late
I Devonian black shale near Grahamstown, South Africa

Published jointly by the Cape Provincial Museums
at the Albany Museum, Grahamstown, South Africa

ANNALS OF THE CAPE PROVINCIAL MUSEUMS

These Annals are published in two series. Natural History and Human Sciences, the latter series
covering cultural history, ethnology, anthropology and archaeology. They are issued in parts at
inegular intervals as material becomes available.

The primary objective of these Annals is to disseminate the results of research work undertaken by
staff members of the Cape Provincial Museums. However, a manuscript by any author is eligible
for publication provided it is based wholly or partly on the collection/s of one or more of these
museums or if the material on which the work is based is wholly or partly deposited in one of
these museums.

Parts are obtainable from the Librarian of the Albany Museum. Correspondence in connection
with these Annals should be addressed to the Editor, Albany Museum, Grahamstown 6140.

Editor

Dr F. W. GESS

Assistant Editor
Dr S. K. GESS

A preliminary catalogue of fossil algal, plant, arthropod, and fish remains
from a Late Devonian black shale near Grahamstown, South Africa

by

ROBERT W. GESS and NORTON HILEER=i=

(Department of Geology, Rhodes University, Grahamstown, 6140 South Africa)

ABSTRACT

A black carbonaceous shale, exposed in road cuttings that intersected rocks of the Witpoort
Formation (Witteberg Group), Eastern Cape Province, South Africa has yielded a variety of fossil
remains that have added considerably to our knowledge of Late Devonian life in southern Africa.
The rocks are interpreted as having been deposited in a coastal setting with the black shale representing
a stagnant back-barrier lagoon. Among the life forms present are charophyte and possible phaeophyte
algae and a diverse array of plant types including members of the Rhyniopsida, Zosterophyllopsida,
Lycopsida and Progymnospermopsida. Arthropods are represented by the remains of Ostracoda,
Conchostraca, and Eurypterida; fish by antiarch and arthrodire placoderms, acanthodians,
chondrichthyans, crossopterygians and dipnoans. Several trace fossil types have also been noted.

CONTENTS

page

Introduction 226

Stratigraphic setting 226

Sedimentology and depositional environment 226

Age and preservation 229

Occurrence of the fossil remains 229

Fossils associated with the sandstones 229

Fossils associated with the black shale 231

Algae 23 1

Phaeophyta 23 1

Charophyta 235

Plants 242

cf. Rhyniopsida 242

Zosterophyllopsida 242

Lycopsida 249

Putative arborescent Lycopsida 249

cf. Trimerophytopsida 261

Class indeterminate c/i Calcimophyton 261

Progymnospermopsida 261

Incertae sedis 270

Animals 274

Arthropoda 274

Pisces 274

• Placoderms 274

Acanthodians 288

Chondrichthyans 290

*Presenl address; c/o Geology Department, University of Canterbury. Private Bag 4800. Christ Church, New Zealand.

225

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Crossopterygians 294

Dipnoans 296

Incertae sedis 297

Acknowledgements 299

References 299

Appendix 1 : List of figured specimens 300

Appendix 2: List of additional specimens 303

INTRODUCTION

The construction in 1985 of a road bypass to the immediate south of Grahamstown (Cape
Province, South Africa) exposed sedimentary rocks of the Dwyka and Witteberg groups. Cuttings to
the west of the junction with the Port Alfred road all contain rocks belonging to the Witpoort Formation
of the Witteberg Group (Fig. 1). The Witpoort Formation is a predominantly arenaceous unit with
minor interbedded shales. The new road cuttings (Fig. 2) exposed an unusually thick layer of black
shale containing plant fossils. These fossils were similar to those described from a much thinner, well
known shale exposed in a road cutting in Howison’s Poort, about 10 km south-west of Grahamstown.
However, the new material was clearly more diverse and less fragmentary.

Both algal and vascular plant material was recovered initially, including apiece of Archaeopteris
frond.

As collecting continued, further plant material, remains of three arthropod groups and five fish
groups, placoderms, acanthodians, chondrichthyans, crossopterygians, and dipnoans, were
discovered.

Stratigraphic setting

The Witpoort Formation, which constitutes the middle part of the Witteberg Group, the youngest
division of the tripartite Cape Supergroup, is a predominantly quartz-rich sandstone unit with minor
interbedded black, grey, and maroon shales. In the Grahamstown area it is underlain by the Weltevrede
Formation, consisting of shales, siltstones, and minor sandstones and overlain by shales of the Lake
Mentz Subgroup.

Within the typical pale grey to white quartz arenites, quartz wackes and subarkoses of the Witpoort
Formation several black carbonaceous shales up to 1 m in thickness occur at several horizons. These
are often associated with fine plant fragments and trace fossils, frequently of the ichnogenus
Spirophyton. The best known of these shale horizons is exposed at Howison’s Poort. From it a few
plant species have been described from fragmentary specimens (Plumstead, 1967; Anderson and
Anderson, 1985; Rayner, 1988).

The relative stratigraphic positions of the shales at Howison’s Poort and at the new bypass site
suggest that they are probably neither continuous nor precisely contemporaneous.

Sedimentology and depositional environment

The fossil locality occurs in a road cutting approximately 500 m long and 20 m deep, where the
beds dip gently ( 1 0°) northwards. The sequence has been interpreted as representing a brackish coastal
back-barrier lagoon system, with significant freshwater input, sheltered by a barrier island (Hiller
and Taylor, 1992; Taylor and Hiller, 1993).

226

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

The lowermost part of the sequence exposed in the cutting consists of alternating beds of brown-
grey quartz arenite separated by thin shales. The quartz arenites appear largely structureless but contain

WASHOVER

DUNES

Fig. lA-D: A. Distribution ot the outcrop of Witteberg Group rocks in South Africa; B. Simplified geological map of the
Grahamstown area, showing the position of the site discussed; C. Key to A and B showing correspondence with
international geological time scale; D. Proposed depositional environment at the site.

227

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

some trough cross-bedding and horizontal lamination. These are overlain by about 50 m of thickly
bedded, pale grey quartz arenites with occasional thin, dark grey shales and quartz wackes. Erosional
contacts are common and primary structures within this sequence include horizontal lamination, ripple
cross-lamination, and trough cross-bedding. This cross-bedding suggests a general westerly to south-
westerly transport direction. Some bioturbation is evident in the wackes. This portion of the sequence

Fig. 2A and B: A. The road cutting from which the fossil material has been recovered; B. Gess excavating the black shale
horizon.

228

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

has been interpreted as shoreface deposits passing upwards into washover, tidal channel, tidal delta,
and back-barrier tidal channel deposits (Hiller and Taylor, 1992).

The middle portion of the exposed sequence, from which most of the fossil material has been
recovered, consists of laminated black carbonaceous shales up to 6 m thick. These sediments represent
a stagnant back-barrier lagoonal environment.

The black shales are overlain by creamy-coloured mica-rich quartz wackes containing several
lenses of black shale. The top of the sequence consists of white quartz arenites displaying trough
cross-bedding, ripple cross-lamination and horizontal lamination, overlain by horizontally laminated
micaceous quartz wackes. This probably represents a return to a tidal delta depositional environment
(Hiller and Taylor, 1992).

Age and preservation

Plumstead ( 1967) concluded that fossil plant fragments from the Witteberg Group were of Upper
Devonian age. Anderson and Anderson (1983) agreed with this estimate at least for the Witpoort
Formation. It is supported by the Early to Middle Devonian age assigned to the underlying Bokkeveld
Group on the basis of marine invertebrate evidence (Boucot, Johnson and Talent, 1969). Cooper
( 1 986), in attempting to derive a correlation with Northern Hemisphere Devonian rocks using eustatic
sea-level changes, suggested that the Witpoort Formation was of Famennian age. This estimate is
borne out by the presence among the material excavated during this research, of Archaeopteris, a
characteristic Late Devonian plant, as well as Groenlamiaspis and Bothriolepis placoderm fishes
characteristic of this age (Young, 1989).

The fossil material from the black shale is fairly poorly preserved, taking the form of silvery
white, near two-dimensional compressions. During low grade metamorphism, phyllosilicate minerals,
such as mica and possibly chlorite, grew in the cavities remaining after reduction of the organic matter.
These minerals subsequently altered to kaolinite during uplift to present levels. The condition of the
fossils has precluded excavation in the round and digestion of material from the rock. It is also not
possible to study fine details with the scanning electron microscope or high powered optical
microscope.

Much of the organic remains appear to have been in a state of decay at the time of burial and the
skeletal elements of the fish are partially, or completely dissociated. Water current activity in the
area of greatest concentration is likely to have been very slight, suggesting that dissociation was
aided by the gases of decay and the activities of scavengers.

Occurrence of the fossil remains

The majority of the fossils described in this catalogue were recovered from the carbonaceous
lagoonal black shale though several robust plant structures and trace fossils were discovered in the
sandstones of bander origin that stratigraphically underlie the shale.

FOSSILS ASSOCIATED WITH THE SANDSTONES

A variety of trace fossils was discovered as sole marks on the underside of sandstone layers
interbedded with thin reddish shales. The lower surfaces of the sandstones were prepared by exposing
them to the elements for a few months. By far the most common types present belong to the ichnogenera
Rusophycos and Cruziana, the former often associated with the end of several centimetre-long wavy

229

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Cruziaua tracks that are generally about 5 mm wide (Fig. 3A). Another interesting trace is a
presumably endogenic helical structure (Fig. 3B) that may have been produced by an annelid.

Fig. .tA-E: A. Riisoplixco.'i and Cruziaua troin sandstone interbedded with reddish .shale; B. Helical, prcsunied endogenic trace
fia)in sandstone interbedded with reddish shale; C-E. ‘A'/nVo/Av/o//" front near the top of the black shale.

230

GESS AND HILLER: CATALOGUE OE EOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Rusophycos and Cniziana are generally taken to represent the resting and feeding traces respectively
of arthropods, often trilobites, and to indicate a shallow marine environment. This accords with our
interpretation of the area as being close to the lagoon mouth.

Within the same set of interbedded sandstones an internal cast of the stem of the lycopod
Leptophloeum was found. This was probably \vashed down into the lagoon where it became stranded
on the sand flats along with smaller fragments of plants that are better preserved in the black shale.
Further Leptophloeum material, including young stems and root bases, was recovered from a mud-
chip conglomerate lens where it occurred with Haplostigma irregularis remains (Figs 22A, D; 23A-
C; 24; 27B-E).

A thin (50 mm) black shale interbedded with the sandstones has yielded several relatively well-
preserved pieces of the fronds of the progymnospcrmArchaeopteris (Figs 31, 32B). This occurrence
lies below the main black shale, which yielded other Archaeopteris material.

FOSSILS ASSOCIATED WITH THE BLACK SHALE

Trace fossils resembling small examples of the ichnogenus Spirophyton occur near the top of
the black shale. The specimens (Eig. 3C-E) are circular in plan view with an average diameter of 25
mm and a central shaft 6 mm across. If our identification is correct then this may represent an
opportunistic occurrence in an oxygen-depleted environment (Bromley, 1990).

Plant and algal remains of both terrestrial and aquatic origin are by far the most common fossils
recovered from the site. Below we document the diversity of plant types represented in the collection
and offer provisional identifications where possible.

ALGAE

Phaeophyta (Eigs 4-6)

A large amount of the material recovered can probably be assigned to the brown algae. The most
distinctive and among the most common were provisionally reported as “terminally-lobed algae”
(Taylor and Hiller, 1993), and are the subject of a separate paper (Hiller and Gess, in press).

The structure of this form is extremely variable. Typical specimens consist of thallae (or pieces
thereof) with an average length of about 60 mm, but reaching up to about 1 20 mm in more elongate
individuals (Fig. 5B). These thallae are on average 3-4 mm wide with bifurcations at irregular intervals,
typically about 15 mm apart (but sometimes at negligible intervals or far greater ones). Up to seven
bifurcations occur in some specimens. Micro-environmental influences such as water depth probably
account for this variability. Some specimens suggest a four-lobed overall structure, perhaps arising
from two very closely spaced initial bifurcations (Fig. 4A). In less than one-third of the specimens,
small, round, bud-like structures occur along the edges of thalloid strands (Figs 4D-E; 5A). These
may be reproductive. Two atypical specimens are also illustrated (Figs 5D; 6A). The first of these,
(Fig. 5D) undoubtedly represents the development of an adventitious branch.

The specimens strikingly resemble the more fragmentary remains described as Biithotrephis
trichotoma Douglas and Jell (1985) from the Early Devonian of central Victoria (Australia) and may
represent the same form. Individuals with a negligible distance between two main bifurcations (Fig. 5C)
can resemble Hungerfordia dichotoma Fry and Banks ( 1955), described from a single specimen from
the Upper Devonian of New York State. We believe that the American and Australian material is of the
same species. These specimens also bear a strong resemblance to the modern brown algal genus Dictyota.

231

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Other more elongated dichotomously branching structures from the locality similarly lack a
central strand. One specimen (Fig. 6B) calls to mind Yeaia flexuosa Douglas (1983) from the

Fig. 4 A-E. Hmifierfoniui: A. Showing po.s.sible four-lobed strucUire; B-C. Showing well preserved dicholoniising; D-E.
Showing bud-like structures.

GESS AND HILLER: CATALOGUE OE FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Barai’W'cincithia flora of central Victoria. Our specimen consists of a strap-like thalliis about 8-9 mm
wide, showing two bifurcations 50 mm apart. The exterior is irregularly covered in dots that are
presumed to be reproductive structures. Other, less complete pieces of similar but narrower straps
have also been found (Fig. 6C).

Fig. 5 A-D. HimgerfonUa: A. Showing unusual lobes; B. Large specimen; C. Resembling type specimen of Hiiiigeii'onlia
dichotoma Fry and Banks; D. With adventitious branch.

233

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

We believe that other algal material may be represented among the incertae sedis (Figs 39-41 ).

Fig. 6A-C: A. Hiiiif^crfordia. unusual specimen; B-C. Yeaia sp.

234

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE. S. AFRICA

Charophyta (Figs 7-11)

During the initial investigation only a few incomplete whorls of what the authors now interpret
as charophytes came to light. Their affinities were not recognised, so they were reported as “daisy-
wheels” (Taylor and Hiller, 1 993). However, many more isolated whorls, as well as short axes bearing
whorls at the nodes, have now been found. Fossil charophytes are known largely from their
reproductive structures (Hill and El-Khayal, 1983), so the discovery of associated vegetative structures
afforded us a rare opportunity to suggest reconstructions for these species (Gess and Hiller, in press).

Among the forms that we confidently assign to the charophytes we recognise four species, all
represented by both vegetative and reproductive structures. Two of these possess an octaradial
symmetry and the other two hexaradial symmetry, and on this basis we have assigned them to two
genera (Gess and Hiller, in press). The inclusion of these species in the Eocharaceae is suggested not
only by their age but also by their sinistrally spiral oogonia. Direct comparison with Eochara, which
is only known from three-dimensionally preserved isolated oogonia, is precluded because the two-
dimensional preservation of our specimens does not permit examination of the crown cell and
determination of the number of spiral cells (their high number being the diagnostic character of the
family). All four species consist of intemodes with regularly arranged fertile nodes.

The larger octaradial form (new genus 1, new species 1 ), with whorls of up to 14 mm in diameter,
has branches that are relatively broad with rounded terminations (Figs. 8A; 9B; lOA). Each nodal
whorl bears eight radially arranged branches that quadrifurcate after about one-third of their length.
Oogonia, which are attached within the junction of these four sub-branches, are almost spherical,
about 1.7 mm long and 1.6 mm wide at widest but tapering towards the attachment. On the upper
surface of each radial branch, just before the point of quadrification, a paired oval structure may
represent the antheridium (Fig. 9B).

The other octaradial form (new genus 1, new species 2), has a similar general arrangement, but
with branches that are narrow with slender tapering terminations, and whorls up to 10 mm in diameter
(Figs 7A-E; 9A; lOB). Oogonia of this species are ellipsoidal, about 1.5 mm long and 0.9 mm wide
at widest. In one specimen the sub-branches are seen to divide again (Fig. 7E). The internodes appear
uncorticated (Fig. 7A).

The intemodes of the hexaradial forms (new genus 2) also appear uncorticated (Figs 8C; 8E;
1 lA-E). Nodes in one species (new genus 2, new species 1) bear whorls about 6 mm in diameter,
consisting of six radial branches. Each branch consists of a short stalk that gives rise to a slender
pointed branchlet and carries an oogonium (Figs 8B-E; 9C; IOC). Oogonia are apparently attached
to the underside of branches, at the position from which the branchlet arises. The oogonia are pear
shaped, about 1.6 mm long and 1.2 mm wide at the widest, tapering towards the attachment.

The structure of the other species in this genus (new genus 2, new species 2) is less well known.
It has whorls of about 8.5 mm in diameter that consist of six branches. At about half their radius these
branches redivide, each giving rise to a smaller whorl, bearing six relatively small oogonia, couched
between branchlets arising from a further division (Fig. 1 1 ).

Although differing in superficial appearance, the architectures of these species are reminiscent
of the modern genus Nitella.

We believe that these four charophyte species probably lived in the shallow, still, brackish water
near the edge of the lagoon, close to their place of burial. Transport over any great distance would
probably have led to greater damage to, or destruction of, the vegetative material.

Fragmentary remains have been discovered, suggesting the presence of a fifth species of
charophyte.

235

ANN. CAPE PROV. MUS, (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Fig. 7A-E. Charophyta, new genus I , new species 2: A-B. Whorls on axis; C-D. Associated whorls; E. Showing sub-branches
redividing; F. Possibly related specimen.

236

I I I i

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE. S. AFRICA

Fig. 8A-E: A. Charophyta, new genus I . new .species I .. two whorls (scale bar = 10 mm); B-E. Charophyta, new genus 2. new
species I; B. Isolated whorl; C. Growing tip; D. Chain of associated whorls; E. Whorls on axis.

237

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Fig. 9 A-C. Charophyte nodal whorls: A. New genus I , new species 2; B. New genus I , new species I ; C. New genus 2, new
species 1. (Scale bar = 10 mm)

238

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Fig. 1 0 A-C. Charophyte axial lecoiistmction.s: A. New genus 1 . new species I ; B. New genus I , new species 2; C. New genus
2. new species I . (Scale bar = l(J mm)

239

ANN, CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Fig. I I A-H. Cliarophyta. new genus 2. new species 2: A. Specimen showing 4 whorls; B-C. Side view of whor! in A attached
to axis and containing oogonia; D-E. Side view of whor! in A attached to axis and lacking most oogonia; F. Oogonia
from three associated whorls; G. Oogonia attached to whorl in plan view; H. Plan view of whorl in A (counter
specimen).

240

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Fig. 1 2A-C. Presumed ac|iialie plant: A. Brandling axes bearing leallets and fertile nodes; B. Fertile node with well preserved
bracts; C. Fertile node.

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Another vegetative form bearing a superficial resemblance to the Charophyta cannot be
unequivocally assigned to any group at this time, but seemingly also consists of internodes and fertile
nodes (Fig. 12). Nodes occur at intervals of about 15 to 25 mm, with the fertile region having a
diameter of 5 mm, and the nodal bracts circumscribing a diameter of about 15 mm. The internodes
are branched and sometimes bear fine elongate “leaflets” about 5 mm long. This form is also assumed
to have been aquatic.

Nodes and internodes are also observed in another presumed aquatic form (Fig. 40A-B).
PLANTS

cf. Rhyniopsida (Figs 13, 14, 15A)

A single specimen has been found that the authors believe to be a gametophyte of this group
(Fig. 15A) as defined by Stewart and Rothwell (1993). It has a smooth axis 48 mm long, terminating
at one end in a thin putative sub-aerial axis, of which the proximal 16 mm is preserved. The other
end flares into a cup 16 mm wide at the lip. This specimen bears a striking resemblance to the
gametophyte genus Lyouophytou, though seemingly with fewer lobes around the lip of the cup.

We note that the South African species Dutoitia pulchra Hoeg ( 1 930) is very similar to the genus
of rhyniopsid gametophyte, Sciadopliyton. The only significant difference between these two genera
is the existence of small enations on the axis of Dutoitia pulchra. The authors believe the new specimen
described here represents a separate species of Dutoitia, a genus that should be restricted for South
African gametophyte rhyniopsids. The other two species added to this genus by Plumstead (1967)
are apparently sporophyte material of unrelated types and should therefore be removed from Dutoitia.

Note that Schweitzer (1981) proposed that Sciadophyton is the gametophyte correlate of
sporophytes of the Zosterophyllum and Taonocrada types. Zosterophyllum from the Grahamstown
site is discussed below.

Also present at the site are numerous specimens referable to Dutoitia alfreda Plumstead (1967)
(Fig. 13). These plants consist of slender, dichotomously branched naked axes about 0.5 mm wide,
terminating in club-shaped sporangia up to 7 mm long and 1 mm wide. As pointed out above, we
believe that Plumstead ( 1967) was wrong to assign this species to the genus Dutoitia and suggest the
creation of a new genus.

Closely associated on the same bedding planes with the “D. ” alfreda material are other, possibly
related, specimens with round sporangia 2 mm in diameter ( Fig. 14). The sporangia are found attached
to masses of dichotomously branched naked axes that vary from 1 mm wide at the thickest to less
than 0.25 mm. The thicker axes bear sporangia on short lateral branches (Fig. 14C) whereas the
thinner distal axes are often seen to terminate in sporangia (Fig. 14D).

Zosterophyllopsida (Figs 15B-D, 16-18)

A wealth of vegetative material attributable to the genus Zosterophyllum has been collected at
the site, although only one fertile spike is known. It is 23 mm long with closely packed sporangia
along the distal 13 mm (Fig. I5B). Unfortunately the state of preservation precludes detailed
examination of the fine structures of this spike. Far more frequently found are fragmentary "leaves”
and rhizomes, some showing the unusual branching forms that typify this genus (Figs 15C-D; 16A).
The “leaves” reach up to 1 10 mm in length, and fork towards the end in typical zosterophyll fashion
(Fig. 17, especially 17C). Frequently as.sociated with these axes are spheroid structures up to 15 mm

242

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S, AFRICA

Fig. 13. Dutoiliu cilfrcda Pluni.slead, a possible rhyniopsid.

243

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Fig. 14 A-D. Possible rhyiiiopsid: A. Seemingly near complete specimen; B. Detail of A; C. Sporungitim attached to side of
fairly proximal axis (detail of A); D. Sporangium attached to end of distal axis.

244

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Fig. I5A-D: A. Gametophyte rhyniopsid; B-D. Zostemphyllum: B. Fertile spike; C. H-branchiiig; D. K-branehing.

245

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Fig. 16A-C. ZosierophyUum: A. H-branching; B. "Spheioitr laterally attached to rhizome; C. Two "spheroid.s" laterally
attached to rhizome.

246

GESS AND HILLER: CATALOGUE OL LOSSILS LROM A LATE DEVONIAN BLACK SHALE, S. ALRICA

Fig. I7A-F. Zosterophylliim: A. ? “leaves" attaclied to ?rhizonie; B. Rhizome, “spheroid", “leaves" and “rootlets"; C.

“Spheroid" with “leaves"; D. “Spheroid" with “leal" bases; E. "Spheroid" with “leaf’ and “rootlets"; F. ? Rhizome,
“spheroid”, "leaves" and “rootlets".

247

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995 '

Fig. I8A-F: A. ? Zosterophyllwn: B-C. Zostcropinllwn "spheroids" apparently connecting two rhizomes; D-E. Unusual ?
Zostewphyllwu specimens; F. ? Zosterophyllwn rhizome and "rootlets" from which "spheroid" has been broken.

248

GESS AND HILLER: CATALOGUE OE FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

in diameter that apparently emit “leaves” in tufts (Fig. 1 7B-E); they may act as anchors and/or storage
organs. These “spheroids” also emit finer emanations that may anchor the tufts in the manner of
rootlets (Fig. 17B). These “spheroids” were apparently connected by rhizomes as they often appear
to terminate the latter (Fig. 17B-C). In some specimens they are laterally connected to the rhizomes,
and probably anchored them at intervals (Fig. 16B-C). Sometimes two rhizomes are found apparently
connected by the “spheroid” structures (Fig. 18B-C). These structures are reminiscent of the “corm”
of the extant genus Isoetes.

Other material that superficially resembles rhizome material is probably unrelated. Most
important is material bearing very small forked appendages and covered in “spots” (Fig. 40C), and
other material displaying branching of a type uncharacteristic of Zosterophyllum (Fig. 40F).

Among our incertae sedis is a circinate specimen that may also belong to this class (Fig. 40G).

Lycopsida (Figs 19-28, 29A-D)

Two putative herbaceous lycopsids from the site cannot be assigned with confidence to known
orders. One of these is represented by fertile and infertile axes (Fig. 19). The infertile axes reach at
least 80 mm in length and bear curved microphylls up to 13 mm long. The fertile axes are not
completely preserved, the largest specimen being only 30 mm long. Their microphylls are very similar
to those on infertile axes but they bear sporangia cradled in elongate boat-shaped cups on their adaxial
surfaces. The cups extend for about one-quarter of the length of the leaves.

Much larger, though structurally similar, fertile leaves have also been collected from the site
(Fig. 20). Although the full length is not preserved in any specimen, some leaves exceed 40 mm. The
sporophylls are apparently unbent and thus probably do not form part of a cone. They have a narrow
tapering attachment from which a midrib extends the full preserved length of each specimen. Distal
to the attachment, the leaf broadens to about 6 mm, producing a protective structure for a sporangium.
An extended lamina tapers naiTowly towards the leaf tip. Fine hair-like fimbriations along the margins
of the lamina are longest near the sporangium and extend for about one-half of the length of the leaf.

A further type of vegetative material present is not directly associated with reproductive material,
and may represent herbaceous, or arborescent material. It is represented by axes that dichotomise and
bear slender pointed microphylls (Fig. 21 ). The proximal axis, 40-100 mm long and up to 4 mm wide,
divides as many as three times, giving rise to thinner axes typically extending another 100 mm. The
microphylls average 3 mm in length.

Putative arborescent Lycopsida (Figs 22-28, 29A-D)

The arborescent lycopsid Leptophloeiim austnile (McCoy (1874)) Walton (1926) is well
represented in our collections as fragments of stems (Figs 22-24), some with attached roots, proximal
branch segments (Fig. 22D), and possible fertile structures (Fig. 29A-C). Stems bearing roots are
about 600 mm long and 45 mm wide. Above the roots (Fig. 24C-F) the epidermis is smooth for 220-
270 mm; beyond this rhomboid leaf scars become visible (Fig. 24B). At about two-thirds of the way
to this point the stems narrow to 30 mm. Fig. 23A-G illustrates the variation in morphology and
preservation of the leaf bases of L. austnile.

Fig. 23H shows an example of possible decortication in L. austnile. Further material exhibiting
this phenomenon is illustrated in Fig. 25, with an associated structure that may represent a shoot or
fertile body (Fig. 25D,E).

249

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Fig. I9A-I; A Putative herbticeous lycopsid (type one) (all to same scale): A-B. Infertile axes; C-D. Apices of fertile axes; E.

Microphylls from fertile axis; F. Micropliylls associated with small trilete spores; G. Infertile niicrophyll; H. Fertile
microphyll without sporangium; I. Fertile niicrophyll with sporangium.

250

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Fig, 20A-E. Putative herbaceous lycopsid (type two) (all to same scale) fertile microphylls.

251

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

252

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Fig. 22A-E. Leptophloewn austnile (McCoy) Walton: A. Stem from mud chip conglomerate lens; B. Stem from sandstone
displaying leaf impressions; C. Stem from black shale; D. Portion of branch from mud chip conglomerate lens; E.
Stem from black shale displaying preferential preservation of leaf impressions.

253

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Material displaying spirally arranged, closely spaced, elongate leaf scars (Fig. 26 A-C) is
tentatively assigned to the form genus Longicatrix Anderson and Anderson (1985). The authors do
not, however, exclude the possibility that the material could represent the decorticated inner core of
an arborescent lycopsid such as L. aiistrale.

A single specimen displays the distinctive hexagonal leaf scars characteristic of Archaeosigillaria
Kidston ( 1901 ) (Fig. 26D), most closely resembling Archaeosigillaria caespitosa (Schwarz (1906))
Plumstead (1967).

Fig. 23A-H: A-G. Lc/iiopliloc'inii aKStrale {McCoy )'Wi\hon displaying variation in rhombic leaf bases, and preservation thereof:
A-C. From mnd chip conglomerate lens; D-G. From black shale; H. ? Decorticating Lcptopliloeum from black shale.

254

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Fig, 24A-F. Leptopliloeum austnile (McCoy) Walton from mud chip congiomerale lens: A. In .situ excavation of stems: B.

Two stems bearing root bases (external motild in top specimen): C-D. Close up of root bases; E-F. Clarification of
C and D.

255

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Fig. 23A-E. '? Lcptopliloeiiin uiistrale (McCoy) Walton: A-C. F-H. ? Outer cortex: D-E. ? Shoot or fertile structure.

256

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S, AFRICA

Fig. 26A-D; A-C. Limf>icatri.\ Anderson and Anderson; D. Arcliacosi^illaria Kidson.

257

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Fig. 27A-E. Haplostigina irregularis (Schwarz) Seward: A. From black shale; B-E. From mud chip conglomerate lens.

258

GESS AND HILLER: CATALOGUE OE FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Fig. 28A-G; A-F. Palaeostigma rohusta Anderson and Anderson; G. ? Palacostigma

259

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Haplostigma irregularis (Schwarz (1906)) Seward (1932) has been recovered from the black
shale (Fig. 27A) as well as from the mud-chip lens (Fig. 27B-E). In the lens it is associated closely,
and exclusively, with the stems and subaerial portions of young Leptopliloeum australe. Plumstead

Fig. 29A-E: A-C. ? Leptopliloeum australe (McCoy) Walton cones; D. ? AiTorescent lycopsid cone; E. ? Trinierophyte
reproductive material.

260

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

( 1 967) commented on the frequent association of these two axial types and suggested that Haplostigma
may represent younger twigs of Leplophloeum. However, the material described here contains a
rhombically scarred axis narrower than many of the Haplostigma axes associated with it (Fig. 22D).
In addition, fine twigs of L. australe reported from Australia (White 1990, p.76), do not possess round
scars, but rather resemble the specimen figured in Plumstead ( 1967, pi. XIV-8). The authors believe
that the Haplostigma material in their collection probably represents the roots of L. australe. The
authors also assume that their associated Haplostigma material all belongs to the same species, despite
the similarity of some specimens to H. A'OH’/envA (Plumstead (1967)) Anderson and Anderson (1985).

Palaeostigma robusta Anderson and Anderson (1985) from the site (Fig. 28A-F) is nearly
identical in morphology and preservation to material recorded from the Howison’s Poort site.

Various long, narrow, cone-like structures (Fig. 29A-C) are presumed to represent the
reproductive structures of an arborescent lycopsid, possibly L. australe.

cf Trimerophytopsida (Fig. 29E)

A single, possibly reproductive specimen has a gross morphology that suggests trimerophyte
affinities (Fig. 29E). Seven centimetres of a terminating axis are preserved displaying alternate
branching. The first branch arises after 3.2 cm and the next after a further 1 cm. The branches divide
a further two to three times before terminating in expansions.

Class indeterminate cf. Calamophyton (Fig. 30A-C)

A reproductive truss believed to be from a plant of the Calamophyton type (Fig. 30A) has been
recovered. It is not associated with any sterile material, but we believe that some axes from the site
that display short lateral extensions could belong to this plant (Fig. 30B-C).

Progymnospermopsida (Figs 30D-H, 31-38)

Of the large amount of material collected that probably is assignable to this group, most can be
assigned to four species. Additional material of note includes arborescent axes (Fig. 30H) that as yet
have not been unequivocally correlated with finer axes. Some of this material possesses outgrowths
containing a central vascular strand that may represent roots (Fig. 41F). Various isolated leaf-like
structures have also been collected (Fig. 30D-G).

Archaeopteris sp. (species description by Anderson etai, in press) occurs in both the main shale
horizon and a thin shale interbedded with the sandstone a little lower in the sequence (Fig. 2A). It is
noteworthy as the first confirmed Archaeopteris from southern Africa. The material (Figs 31-33)
consists predominantly of infertile axes, fronds and frond fragments, but one fertile specimen has
been found (Fig. 32C). “Pinnules” are quite variable (Fig. 33), and occur attached to both “rachises”
(Fig. 32B) and “pinnae” (Fig. 33D). Towards the base of the “rachis” “rachial pinnules” are modified
into small bract-like structures (Fig. 31, top right).

Material that may represent another species o^ Archaeopteris is present, but consists purely of
non-fertile material (Fig. 34). This consists of specimens interpreted as “pinnae” and one (Fig. 34E)
in which putative “pinnae”, up to 90 mm long and 2 mm wide, are connected to a portion of an
apparent “rachis”, 65 mm long and 4mm wide. The putative “pinnules” are up to 10 mm long, are
unwebbed and divide as many as three times, once proximally, again at about mid-length, and finally
near the tip. A possible third species is represented by only two specimens (Fig. 41 A).

261

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Fig. 30A-H: A. cf. Cakunophxton reproductive head; B-C. cf. CaUmwphyton axes; D-G. Lsolated leaves; H. Progymnosperm
type arborescent axis.

The second well represented species (Figs 35-37) has fertile material which (Figs 36A-B, F-G;
37A) is very like that of Chaiileria. It consi.sts of small branches that bifurcate twice to produce four
recurved terminal axes, each bearing a small double pendulous structure. These structures were
described from the Howison's Poort site as Dutoitia luaraisia by Plumstead (1967), who believed
them to be psilophytes. The discovery of these structures associated with several orders of woody
branching has led the authors to believe that this species should be excluded from Dutoitia. The higher
order axes found (Figs 35; 36A) are generally 5-9 mm across and show preserved lengths up to 500

262

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AERICA

mm. An apparent striation may represent impressions of internal vascular material. The parallel
arrangement of these axes in our most complete specimen (Fig. 35) suggests that they branched off
a yet larger axis.

These axes give rise at irregular intervals to “secondary axes” that are borne in fans of at least
five arranged around a bud-like structure (Fig. 36C-E). Such fans may be spirally arranged on the
“primary axis”. The “secondary axes” are 2-5 mm across and show preserved lengths of up to 200
mm. “Tertiary axes” give rise to further branchlets that bear the fertile organs (Fig. 36F).

Associated with some specimens of this species we have found axes bearing tuft-like structures
that may represent terminal (Fig. 37B) or adaxial (Fig. 37C) growth points. Other branches carry
small leaf-like structures comprising very fine axes that bifurcate twice. These lack sporangia and
may represent the non-fertile material of this species (Fig. 37D).

Axes described by Anderson and Anderson (1985) from Howison's Poort as Praenmumciiliis
alteriKitirainus are almost certainly of the “secondary axis” type described above. McLoughlin and

Fig. 31. Specimen of Arcliaeopteris new species from a thin shale interhed, showing a "frond" tip, bract-like basal "rachial
pinnae" (top right), and normal “rachial pinnae" (bottom right).

263

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Long ( 1 994) have recorded this form species from Antarctica, in association with Haplostigma li)ieare,
Mahmzniiia sp. and Arcluieosigillaria cf. A. caespitosa. Comparison with their fragment would,
however, be liazardous.

F5g. ,t2A-C. Archaeopteris new species: A. Near complete ''frond" from the main black shale; B. ''Pinnae" and "piniinles"
attached to a "rachls", from the thin shale interbed; C. Piece of "rachis" with "rachial pinnules" and fertile material
from the main black shale.

264

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

The final type of putative progymnospermous material is less well represented than the previous
two. It consists of thin axes, about 1 mm across, that alternately bifurcate unequally at intervals of
about 40 mm (Fig. 38A). The thinner of the two daughter branches averages about 50 mm in length
and subdivides profusely before terminating in small expanded tips. Arranged around the point of
unequal bifurcation are other fine axes that terminate in the same way (Fig. 38A. B, D).

Fig. 33A-D. Anhaeopteris new species: A-C. Isolated "pinnae” from the main black shale; D. Isolated “pinnule" from the
main black shale.

265

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Fig. 34A-E. 1 Avchaeopteris sp.: A-D. ? “Pinnules”; E. ? “Pinnules" associated with portion of “rachis".

266

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Fig. 35. "Diiloitia" maniisia.

267

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Fig. 36A-G. "Dutoitia" mamisia: A. Reconstruction of branching system; B. Reconstruction of fertile structure; C-E.

Examples of "fan-like" branching; F. Fertile material attached to "quaternary branch"; G. Detail of a fertile structure
in F.

268

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Fig. 37A-D: A-C. "Dutoitia" maniisiu: A. Detail of fertile structLire. B. Terminal tuft-like growth point, C. Adaxial tuft-like
growth point; D. Probable leaves of "Dutoitia" maniisiu.

269

ANN. CAPE PROV, MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Incertae sedis

A large number of other reproduetive and vegetative specimens from the site cannot be

Fig. ,t8A-D. Pmbable progyiniiospenn: A. Axis with two unequal bif urcations alternately arranged; B. Detail of A showing
fine axes terminating in expanded tips; C. Thinner branch arising from point ol bifurcation; D. Axis showing unequal
bifurcation.

270

GESS AND HILLER; CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S, AFRICA

unequivocally assigned to any particular algal or plant group at present. A selection of these is
illustrated in Figs 39-4 1 .

Fig. 39. Incertae sedi.s.

271

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19. PT. 5 28 MAY 1995

Fig. 40. Incertae sedis.

272

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Fig. 41. Incertae sedis.

273

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

ANIMALS

Arthropoda (Figs 42-44)

The largest arthropod remains that have been recovered consist of occasional fragments of
eurypterid. The fragments were seemingly in a poor state prior to burial, perhaps reflecting the action
of scavengers and/or the relatively thin exoskeleton of eurypterids. Specimens include a possible
posterior limb (Fig. 42C), other limb fragments (Fig. 42D-E), a gnathostome (Fig. 42F), and several
fragmentary remains of body segments (Figs 43A, B). The only previously-reported eurypterids from
the Witteberg Group are Hastimima sp. from the Weltevrede Formation (Seward, 1909; Woodward,
1909) and Cyrtoctemis wittebergensis Waterston et ai, 1985 from the Waaipoort Formation. Given
the state of preservation, it is very difficult to say to which, if either, of these forms the present material
belongs. The former comes from older strata while the latter was found in younger rocks and it may
be that the new Witpoort Formation remains represent a different taxon.

Fairly common at the site, although often poorly preserved, are the remains of a large
conchostracan (Fig. 43D-G), which reaches up to 15 mm in width and 25 mm in length. Very small
specimens may represent a separate species. The material may belong to Palaeestlieria sp., described
by Rennie (1934), in association with plant fragments including Haplostigmo, from the somewhat
older black shale at Port Alfred. The type material of this species is, however, not well enough
preserved to allow thorough comparison.

Several species of ostracod have also been found. Most of the specimens have unornamented
carapaces (Fig. 44) but a single much smaller specimen, a mere 0.8 mm long, displays an interestingly
sculptured shell with two protrusions on each valve, a short distance from the hinge line (Fig. 43C).

Pisces

Fish material from the Grahamstown site can be assigned to several groups including
acanthodians, placoderrns, chondrichthyans, crossopterygians, and dipnoans (Anderson etal., 1994).
Of these, the placoderrns are the most common.

A series of more in depth papers on the fish material is to be prepared in conjunction with Dr
J.A. Long and Dr M.E. Anderson.

Placoderms (Pigs 45-55)

The placoderms are represented by arthrodires and antiarchs. The arthrodire material includes
three main types, the most common of which (Figs 45C, 46-49) is known from small juveniles (Fig.
49A) to large adults (Fig. 48A-B), and is assignable to the genus Groeiilandaspis (pers. comm. A.
Ritchie, September 1993). It is characterised by an unusually high anterior lateral plate (Figs 47G,
48C,E, 49D). Most plates of this species have been found dissociated (Fig. 49B-D). Another possible
Groeiilandaspis (Figs 50A-B, E-G, I-J, 5 1 ) has a lower anterior lateral plate (Pig. 50E) and a longer,
more tapering spinal plate (Figs 50A, B, I, 5 ID). A third groenlandaspidid arthrodire (Fig. 52A-D)
that may belong to a different genus was recognised among our specimens by Dr J.A. Long (September
1994). It has an extremely high median dorsal plate (Fig. 52A-D) but is represented at present by
only a few Isolated plates.

Other arthrodire material (Fig. 45 A, B) consisting of two plates bearing a distinctive ornament
cannot at present be further identified.

274

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

The only antiarch as yet recovered belongs to the genus Botin iolepis and is well represented by
both juvenile (Fig. 53) and large adult (Figs 54, 55A-D) specimens. This is probably a new species
(pers. comm. J.A. Long. September 1994). As with the arthrodires, small individuals are fairly well
articulated but the larger adult forms are mostly disarticulated.

Fig. 42A-F. Eurypterida: A-B. Body pails (to same scale); C. Possible posterior limb; D. Limb tragmeiU; E. Portion of limb;
F. Gnathostome.

275

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Fig. 43A-G: A-B. Pieces of eurypterid body segments; C. Ornamented ostracod; D-G. Conchostracans (? Palaeestlieria sp.)
(to same scale).

276

GESS AND HILLER: CATALOGUE OE EOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Fig. 44A-D. Various imornamented ostracods (A. 1.83mm long, B. 2.08mm long, C. 1.83mm long).

277

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Fig. 45A-C. Artlirodire placodenii I'isli remains: B. Possible Anterior Ventrolateral plate; C. Articulated Crocnhuukispis sp.
head and trunk plates (pers. comm. John Long. Septemher 1994).

278

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Fig. 46 A-E. Groenlandaspis sp.: A. Dissociated Groenlandaspis sp. head and trunk plates, Anterior Ventrolateral (trunk)
plate at bottom right; B-E. Details of specimen in A (to scale): B. Nuchal at left, right Paranuchal at bottom right,
right Central at top right (head plates), C. Pineal (head) plate at bottom, D. Suborbital (head) plate, E. Postorbital
(head) plate.

279

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Fig. 47 A-H. Groenlandaspis sp. : A. Dissociated Groenlandaspis sp. head and trunk plates (B-D to scale, E-H to scale); B-
H. Details of specimen in A: B. Paranuchal (head) plate at top. Nuchal (head) plate at bottom, C. Central (head) plate,
D. Possible superagnathal bone, E. Median Dorsal (trunk) plate, F. Posterior Dorsolateral (trunk) plate, G. Anterior
Dorsolateral (trunk) plate, H. Spinal plate.

280

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Fig. 48A-E. Groenlandaspis sp.: A. Semiarticulated head and trunk plates; head at top; B. Anterior Dorsolateral (trunk) plate
of specimen in A (most anterior ventral point at top right); C-D. Anterior Lateral, and Posterior Median Ventral
(trunk) plates of largest specimen found; E. Anterior Lateral (trunk) plate of small individual.

281

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 3 28 MAY 1995

Fig. 49A-D. Gi'oentandaspis sp.: A. Partially articulated juvenile left trunk; B. Preliminary reconstruction of right central head
shield plates, clockwise from top left - Nuchal plate. Pineal plate. Central plate and Paranuchal plate; C. Preliminary
reconstruction of ventral trunk plates, clockwise from top - Posterior Median Ventral plate, left Interolateral plate,
left Anterior Ventrolateral plate; D. Preliminary reconstruction of right dorsolateral trunk plates, clockwise from top
- Median Dorsal plate. Anterior Dorsolateral plate. Anterior Lateral plate. Spinal plate and Posterior Dorsolateral
plate. (B-D as in Anderson et al. 1994).

282

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE. S, AFRICA

Fig. 50A-J: A-B. Head and trunk plates of small articulated type two groenlandaspidids; C. Groenlaiidaspidid Anterior
Dorsolateral (trunk) plate; D. '? Type three groenlandaspidid Posterior Dorsolateral (trunk) plate; E. Type two
groenlandaspidid Anterior Lateral (trunk) plate; F. Type two groenlandaspidid plate; G. Type two groenlandaspidid
? Median Dorsal (trunk) plate; H. Groenlandaspidid trunk plate; I. Type two groenlandaspidid Spinal (trunk) plate;
J. Type two groenlandaspidid ? Rostral (head) plate.

283

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Fig. 5 1 A-D. Specimen of type two groenlandaspidid: A. Line drawing of head shield showing canals (Median Dorsal (trunk)
plate obscuring anterior margin); B-C. Specimen and counter specimen of head (Median Dorsal plate obscuring
anterior in C); D. Head, Anterior Lateral (trunk) plate, and Spinal (trunk) plate.

284

021 Oil 001

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Fig. 52A-F: A. High Median Dorsal (trunk) plate with associated Anterior Dorsolateral (trunk) plate (right) of type three
groenlandaspidid; B. Detail of Anterior Dorsolateral plate in A; C. Semi-articulated type three groenlandaspidid
trunk plates. Median Dorsal plate (top). Anterior Dorsolateral plate (bottom right). Posterior Dorsolateral plate
(bottom left); D. Counter specimen to specimen in C; E-F. Possible other high groenlandaspidid Median Dorsal
(trunk) plates.

285

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 3 28 MAY 1995

Fig. 53A-E. Botkriolepis sp. juveniles: A. Articulated ventral trunk plates; B. Specimen with both pectoral appendages; C.

Part ol'dorsal trunk. Anterior Dorsolateral plate (top left). Anterior Median Dorsal plate (top centre). Posterior Median
Dorsal plate (bottom centre); D-E. Pectoral appendages.

286

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Fig. 54A-C. Bothriolepis sp., assemblage of adiill plates: A. Anterior Dorsolateral (trunk) plate (top left), Mixilateral (trunk)
plate (top centre), upper pectoral appendage (top right). Posterior Median Ventral (trunk) plate (centre); B. Near
complete head, (piece of trunk at top left); C. Counter specimen of Mixilateral (top) and Anterior Dorsolateral (trunk)
plates.

287

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Acanthodians (Fig. 56)

The acanthodian material in our collection comprises a single isolated spine (Fig. 56E) and an

Fig. .55A-E: A-D. Bothriolepis sp. adult trunk plates; A-B. Pectoral appendages attached to Anterior Ventrolateral plates; C.
Posterior Ventrolateral plate (at left); D. Counter speciman of A; E. Indeterminate piscean body scales.

288

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

assemblage of about 12 closely associated spines (Fig. 56A-D). The assemblage includes left and
right spines and may well be derived from a single individual, possibly belonging to the Gyracanthidae
(Anderson cf «/., 1994).

Fig. 56A-E: Acanthodian spines: A-D. Part of assemblage of twelve spines (to scale); E. Isolated spine with counter specimen.

289

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Chondrichthyans (Figs 57,58)

Several specimens attributable to this group of fishes have been found at the Grahamstown site.

Pig. 57A and B: A. Whole (lodied puraseiachioinorpli speeimen showing (at left, top to bottom) high dorsal tin spine, spinal
eoltimn (extending right), scapnlocoracoid with attached pectoral tin. ;ind displaced toothless lower jaws. A partial
caudal fin impression is visible at the far right; B. Counter specimen to speciman in A. in which an apparent synarcual
is visible at the base of the dorsal fin spine.

290

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Fig. 58A-D: A. Enlargement of anterior of specimen in Fig. 57B; B. Possible reconstruction of the paraselachiomorph; C.

Enlargement of tail denticles of specimen in Fig. 57A; D. Dorsal fin spine of another individual with partial fin
impression.

291

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Three of these belong to a single previously unknown species. One specimen consists of a long dorsal
spine with associated fin impressions (Fig. 58D), a second consists of an isolated lower jaw, and the

Fig. 59A-E: A. Whole bodied specimen of a juvenile coelocanih; B. Provisional interpretation of specimen in A (outline
shortened and broadened through decay); C-D. Dorsal and ventral aspects of associated urohyal bones; E. Specimen
in D.

292

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

third is an almost complete individual (Figs 57, 58A-C). This possibly represents the oldest known
whole-bodied specimen of the group leading to holocephalomorphs (the paraselachomorphs)
(pers. comm. J.A. Long, September 1994).

Fig. 60A-E. Juvenile coelocanth remains (.scale in mm): A. Probable gular plate; B. Scale; C. Cheek plate; D. Detail of caudal
fin of specimen in .S9.A; E. First dorsal fin.

293

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Crossopterygians (Figs 59-62, 63A-C)

The site has so far yielded the remains of about a dozen juvenile coelacanths (Figs 59, 60) (Anderson
et cil.. 1994). Most are complete although faintly preserved (Fig. 59A) but others are repre,sented by
disarticulated skull elements (Figs 59C-D, 60A, C). In addition, several isolated plates belonging to
large individuals (pers. comm. J.A. Long, September 1994) have come to light (Fig. 62E-F).

Fig. 6IC-E. Rliipidistian fin remains: A. Ulnare; B. Fin basal; C-E. Supports (note body scale at top in E).

294

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Fig. 62A-F: A. Rhipidistian lower jaw: B-C. Associated material; B. Crushed rhipidistiaii maxillary and possible palatoqtiadrate
bones (anterior at top) (pers. comm, .loim Long); C. Rhipidistian ? cheek bone; D. Rhipidistian ? cleithriim part; E-
F. Associated material of probable large coelocantb; E. Skull elements; F. ? Gular plate.

The highly disarticulated remains of a large osteolepiform rhipidistian, probably a eusthenopterid,
have been recovered from a number of horizons. These include fin bones and rays (Fig. 6 1 ) and skull
components (Fig. 62A-D); other isolated elements (Fig. 63A-C) may belong with this species.

295

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Dipnoans (Fig. 63D)

A single partial parasphenoid of a large lungfish is the sole representative of this group so far
recovered from the site (Fig. 63D). It most closely resembles And reyeviclithys from Russia (Anderson
etcd., 1994).

Fig. 63A-D: A. ?Rliipidistian operculum; B. ?Rhipidi,stian bone; C. ?Rhipidistian quadrato-jugal (pers. comm. John Long);
D. Lungfish parasphenoid.

296

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S, AFRICA

Incertae sedis (Fig. 64)

Among the other fish remains are elements that the authors cannot place within any of the above

Fig. 64A-D. Incertae sedis; A. Poorly preserved specimen showing possible fin impression at left; B. Counter specimen of
that in A showing possible dorsal spine; C. Outline of specimen in A; D. Isolated possible fin impression.

297

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

groups, including a poorly preserved specimen featuring a possible dorsal spine (Fig. 64A-C) and a
possible isolated fin (Fig. 64D).

The presence of adult and juvenile fish material at the site suggests the possible use of the area

Fig. 65A-D. Possible pisceaii and/or eiirypterid coprolites: A. Elongate specimen; B-C. Typical specimens; D. Specimen with
carbonised material removed.

298

GESS AND HILLER: CATALOGUE OE EOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

as a nursery by the fish in the manner of many extant fish species. Abundant carbonized helically
spiral trace fossils (Fig. 65), some associated with fragmentary conchostracan and phaeophyte
remains, are interpreted as the coprolites of fishes of a range of sizes. This would imply that at least
some of the fish were living in the water column above the sediments and were not washed in after
death from some other locality. Eurypterids are also known to have had spiral alimentary canals, and
so it is possible that they may have been responsible for at least some of the apparent coprolites.

ACKNOWLEDGEMENTS

We would like to acknowledge the input of, amongst others: Mary White (Sydney), Alexander
Ritchie (The Australian Museum, Sydney) and Steve McLoughlin (University of Western Australia)
for their useful discussions with Gess in Australia ( 1993); John Long (Western Australian Museum)
for his input in Australia (1993) and whilst studying the material in South Africa (1994); Nicole
Grambast (Laboratoire de Paleobotanique, Montpellier) and Richard Bateman (Royal Botanic
Garden, Edinburgh) for their helpful correspondence; and Eric Anderson (J.L.B. Smith Institute,
Grahamstown) and Fleidi Anderson (Botanical Research Institute, Pretoria) for their co-operation.
We are grateful to Mary White for lending Gess a considerable collection of literature from her
personal library and to Alex Ritchie for a gift to Gess of casts of GroenUiiiclaspis material from Mount
Howitt, Australia.

The referees Richard Bateman, Nichole Grambast, John Long, Alexander Ritchie, Mary White
and a sixth anonymous referee are thanked for the time they spent in reading an early draft of this
paper, for their criticisms and for their constructive suggestions.

Elsevier Press is thanked for permission to reproduce the photographs here presented as Pig. 4
A, B, C, D and E, Pig. 5 A-D, and Pig. 6 A and B appearing in Hiller and Gess (in press) and Pig. I ,
Fig. 7 A-C and E, Pig. 8 A-E, Pigs 9 and 10, Pig. 1 I C and H, and Pig. 12 appearing in Gess and
Hiller (in press).

We are grateful to the Council of Rhodes University, Grahamstown for making funds available
for this project.

REPERENCES

Anderson, H.M.. Hiller, N, and Gess, R.W. (in pre.ss). /t/r/RK'o/^rf-n.v (Progymnosperniopsida) from the Devonian of southern
Africa. Botanical Journal of the LInnean Society 117:

Anderson, J.M. and Anderson, H.M. 1 985. The Palaeoflora of Africa: Prodronms of Southern African inegafloras, Devonian
to Lower Cretaceous. Rotterdam: Balkema.

Anderson, M.E.. Hiller, N. and Gess. R.W. 1994. The first /for/u /o/c/u'.v-associated Devonian fish fauna from Africa. South
African Journal of Science 90: 397-403.

Boucot, A.J., Johnson, J.G. and Talent, J. A. 1969. Early Devonian brachiopod zoogeography. Special Papers of the
Geological Society of America 199: 1-133

Bromley, R.G. 1990. Trace fossils: biology and taphononiy. London: Unwin Hyman.

Cooper, M.R. 1986. Facies shifts, sea-level changes and event stratigraphy in the Devonian of South Africa. South African
Journal of Science 82: 255-258.

Douglas, J.G. 1983. A new Late Silurian Thallophyte from central Victoria. Proceedings of the Royal Society of Victoria 95:
275-277.

Douglas, J.G. and Jell, P.A. 1985. Two thalloid (probably algal) species from the Early Devonian of central Victoria.
Proceedings of the Royal Society of Victoria 97: 157-162.

299

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Fry, W.L. and Banks, H.P. 1955. Three new genera of algae from The Upper Devonian of New York. Journal of Paleontology
29: 37-44.

Gess. R.W. and Hiller, N. (in press). Late Devonian Charophytes from the Witteberg Group, South Africa. Review of
Pakieobotany and Palynology.

Hill, C.R. and El-Khayal, A, A. 1983. Late Permian plants including Charophytes from the Khuff Formation of Saudi Arabia.
Bulletin of the British Museum (Natural History). (Geology) 37: 105-1 12.

Hiller, N. and Gess, R.W. (in press). Marine algal remains from the Upper Devonian of South Africa. Review of Palaeobotany
and Palynology.

Hiller, N. and Taylor, F.F. 1992. Late Devonian shoreline changes; an analysis of Witteberg Group stratigraphy in the
Grahamstown area. South African Journal of Geology 95: 203-2 1 2.

Hoeg, O.A., 1930. A psilophyte in South Africa. Kongelige Norske videnskabernes selskabs forhandlinger 3(24): 92-94,

Kidston, R. 1901. Carboniferous lycopods and sphenophylls. Transactions of the Natural Hi.story Society of Glasgow 6 (new
series): 25-140.

McCoy, F. 1874. Prodromus of Pal. of Victoria Dec. 1-5. Memoirs of the Geological Survey of Victoria.

McLoughlin, S. and Long, J.A. 1994. New records of Devonian plants from southern Victoria Land, Antarctica. Geological
Magazine 131: 81-90.

Plumstead, E.P. 1 967. A general review of the Devonian fossil plants found in the Cape System of South Africa. Palaeontologia
africana 10: 1-83.

Rayner, R.J. 1988. Early land plants from South Africa. Botanical Journal of the Linnean Society 97: 229-231 .

Rennie, J.V.L. 1934. Note on an Estheria from the Witteberg Series. South African Journal of Science 31: 233-235.

Schwarz, E.H.L. 1906. South African Palaeozoic Fossils. Records of the Albany Museum 1: 348-360.

Schweitzer, H. 1981. Der Generationswechsel rheinischer Psilophyten. Bonner paldobotanische Mitteilungen 8: 1-19.

Seward, A.C. 1909. Notes on fossil plants from the Witteberg Series of Cape Colony. Geological Magazine 6: 482-485.

Seward, A.C. 1932. Fossil Plants from the Bokkeveld and Witteberg Beds of South Africa. Quarterly Journal of the Geological
Society of London 88: 358-369.

Stewart, W.N. and Rothwell, G.W. 1993. Paleobotany and the Evolution of Plants. New York: Cambridge University
Press.

Taylor, F.F. and Hiller, N. 1993. A new Devonian fossil plant locality in the eastern Cape. South African Journal of Science
89: 565-568.

Walton, ,I . 1 926. On some Australian Fossil Plants Referable to the Genus Leptophloeum Dawson. Memoirs of the Manchester
Literaiy and Philosophical Society 70: 1 13-123.

Waterston, C.D., Oelofsen, B.W. and Oosthuizen, R.D.F. 1985. Cyrtoctenus wittebergensis sp. nov. (Chelicerata:
Furypterida), a large sweep-feeder from the Carboniferous of South Africa. Tran.sactions of the Royal Society of
Edinburgh If,: 339-358.

White. M.E. 1990. The Elowering of Gondwana. Princeton, New Jersey: Princeton University Press.

Woodward, H. 1909. Note on the genus Hastimima from Brazil and the Cape. Geological Magazine 6: 485-488.

Young, G.C. 1989. The Aztec fish fauna (Devonian) of southern Victoria Land: evolutionary and birigeographic significance. In:
Crame, J.A. ed.. Origins and Evolution of the Antarctic Biota. Geological Society Special Publication 47, pp.43-62.

APPENDIX 1 - LIST OF FIGURED SPECIMENS

All material illustrated was collected from the site on the N2 bypass outside Grahamstown and
is being deposited in the Albany Museum, Grahamstown. Most specimens are accompanied by counter
specimens.

Fig. 3A-E: A. AM5264. R.W. Gess, 1993; B. AM5265, R.W. Gess, 1993; C. AM5266, R.W. Gess, 1993; D. AM5267, R.W.

Gess, 1993; E. AM5292, R W. Gess, 1994.

300

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Fig. 4A-E: A. AM5224, R.W. Gess, 1 992; B. AM5225, R.W. Gess, 1 993; C. AM523 1 , R.W. Gess. 1 993; D. AM5229, R.W.
Gess. 1993; E. AM5230. R.W. Gess, 1993.

Fig. 5A-D; A. AM5227, R.W. Gess, 1994; B. AM4843, F. Taylor, 1991; C, AM5226, R.W. Gess, 1994; D. AM5232, R.W.
Gess, 1993.

Fig. 6A-C: A. AM5233, R.W. Gess, 1993; B, AM5223, R.W. Gess. 1993; C. AM5268, R.W. Gess. 1993.

Fig. 7A-F: A. AM5218, R.W. Gess, 1993; B. AM5219, R.W. Gess, 1994; C. AM5200, R.W. Gess, 1993; D. AM5201, R.W.

Gess, 1993; E. AM5220, R.W. Gess, 1994; F. AM5269, R.W. Gess, 1993.

Fig. 8A-E: A. AM4849, R.W. Gess and F. Taylor, 1991; B. AM5206, R.W. Gess, 1993; C. AM5209, R.W. Gess, 1993; D.

AM5207, R.W. Gess, 1993; E. AM5208, R.W. Gess, 1993.

Fig. IIA-G: A-E,H. AM5212, R.W. Gess, 1994; F. AM52I4, R.W. Gess, 1993; G. AM5215, R.W. Gess, 1993.

Fig. 12A-C: A. AM5270, R.W. Gess, 1993; B. AM5271, R.W. Gess, 1993; C. AM5272, R.W. Gess, 1993.

Fig. 13A-E: A. AM5273, R.W. Gess, 1993; B. AM5274, R.W. Gess, 1993; C. AM5277, R.W. Gess, 1993; D. AM5276, R.W.

Gess, 1993; E. AM5275, R.W. Gess, 1993.

Fig. 14A-D: A-C. AM5278, R.W. Gess, 1993; D. AM5279, R.W. Gess, 1993.

Fig. 15A-D: A. AM5280, R.W. Gess, 1993; B. AM5234, R.W. Gess, 1993; C. AM5254, R.W. Gess, 1993; D. AM5255, R.W.
Gess, 1993.

Fig. 16A-C: A. AM5253, R.W. Gess, 1993; B. AM4854, F. Taylor, 1991; C. AM5281, R.W. Gess, 1994.

Fig. 17A-F: A. AM5282, R.W. Gess, 1993; B. AM5252, R.W. Gess, 1993; C. AM525 1 , R.W. Gess, 1993; D. AM5283, R.W.

Gess, 1993; E. AM5284, R.W. Gess, 1993; F. AM5285, R.W. Gess, 1993.

Fig. 18A-F: A. AM5286, R.W. Gess, 1993; B. AM5287, R.W. Gess, 1993; C. AM5288, R.W. Gess, 1993; D. AM5289, R.W.

Gess, 1994; E. AM5290, R.W. Gess, 1993; F. AM5291, R.W. Gess, 1993.

Fig. 19A-I: A. AM5260. R.W. Gess, 1993; B. AM5300, N. Hiller, 1992; C. AM5299, R.W. Gess, 1989; D. AM5297, R.W.
Gess, 1994; E.AM5298, R.W. Gess, 1994; F. AM5301, R.W. Gess, 1993; G, AM5302, R.W. Gess, 1993; H. AM5303,
R.W. Gess, 1993; I. AM5304. R.W. Gess, 1993.

Fig. 20A-E: A-B. AM5305, R.W. Ge.ss, 1993; C. AM5306, R.W. Gess, 1993; D. AM5307, R.W. Gess, 1993; E, AM5308.
R.W. Gess, 1993.

Fig. 21A-D: A. AM5257, R.W. Gess, 1993; B. AM487I , G. Englebrecht, 1992; C. AM5258, R.W. Gess, 1993; D. AM5259,
R.W. Gess, 1993.

Fig. 22A-E: A. AM53 1 0. R.W. Gess, 1 994; B. AM53 1 2, A. Absalon, 1 992; C. AM4828, F. Taylor, 1 99 1 ; D. AM53 1 1 , R.W.
Gess, 1994; E. AM4832, R.W. Ge.ss, 1988;

Fig. 23A-H: A. AM5314, R.W. Gess, 1994; B. AM5315, R.W. Gess, 1994; C. AM5316, R.W. Gess, 1994; D. AM4828, F.
Taylor, 1991;E. AM5313, R.W. Gess, 1994; F. AM5317, N. Hiller. 1992; G. AM53 18. R.W. Gess, 1993; H. AM4826.
R.W. Gess, 1986.

Fig. 24A-D: AM5310. R.W. Gess, 1994.

Fig. 25A-H: A. AM4894(WMi029), R.W. Gess. 1993; B. AM53I9, R.W. Gess, 1994; C. AM4825. R.W. Gess, 1991; D-E.
AM5320, R.W. Gess. 1994; F. AM4894(WM 1027), R.W. Gess. 1993; G. AM4894(WM 1025), R.W. Gess. 1993; H.
AM4894, R.W. Gess, 1993.

Fig. 26 A-D: A. AM532!, R.W. Gess, 1993; B. AM5322, R.W. Gess, 1994; C. AM4833, F, Taylor, 1991 ; I). AM5323, R.W.
Gess. 1993.

Fig. 27A-E: A. AM5324, R.W. Gess, 1993; B. AM5325. R.W. Gess, 1994; C-D. AM5326. R.W. Gess, 1994; E. AM53IO,
R.W. Gess. 1994.

Fig. 28A-F: A. AM5327, R.W. Gess, 1993; B. AM5328, R.W. Gess, 1994; C. AM5329, R.W. Gess, 1993; I). AM5330. R.W.

Gess, 1993; E. AM533I. F, Taylor, 1991; F. AM5332, R.W. Gess. 1994.

Fig. 29A-E: A. AM5262, R.W, Gess, 1993; B. AM4834. F. Taylor, 1991; C. AM5334, R.W. Gess, 1993; D. AM5335, R.W.
Gess, 1993; E. AM4837, R.W. Gess, 1991.

Fig.30A-H: A. AM5263, R.W. Gess, 1993; B. AM5336, R.W. Gess, 1993; C. AM5337, R.W. Gess, 1993; I). AM5338, R.W.

301

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Gess, 1993; E. AM5339. R.W. Gess, 1993; F. AM5340, R.W. Gess. 1993; G. AM5341, R.W. Gess. 1993; H. AM5342,
N. Hiller, 1992.

Fig. 31; AM49 1 3. R.W. Gess, 1993.

Fig. 32A-C: A. AM49l4-b. R.W. Gess. 1993; B. AM49I6, R.W. Gess. 1993; C. AM49I8, R.W. Ge.ss. 1993.

Fig. 33A-D: A. AIVI492 1 -3a, R.W. Gess, 1 993; H. AM492 1 -4a. R.W, Gess, 1993; C. AM492 1 -2, R.W. Gess, 1 993; D. AM4923-
I, R.W. Gess, 1993.

Fig. 34A-E: A. AM5294, R.W. Gess, 1993; B. AM5295, R.W. Gess, 1993; C. AM4842, F. Taylor, 199 1 ; I). AM5296; R.W.

Ge.ss, 1993; E. AM.3293, R.W. Gess. 1994.

Fig. 35: AM5343, R.W. Gess and N. Hiller, 1993.

Fig. 36C-G: AM5343, R.W. Gess and N. Hiller, 1993.

Fig. 37A-I): A-C. AM5343. R.W. Gess and N. Hiller. 1993; D. AM.5.344. R.W. Gess, 1993.

Fig. 38A-D: A-B. AM5345. R.W. Gess, 1994; C. AM5346, R.W. Gess, 1993; 1). AM5347. R.W. Gess. 1993.

Fig. 39A-F: A. AM5.348. R.W. Gess, 1 993; B. AM5,349. R.W. Gess, 1993; C. AM.535(). R.W. Gess, 1 993; 1). AM535 1 . R.W.

Gess. 1993; E. AM53.52, R.W. Gess, 1993; F. AM.5353. R.W. Gess, 1988.

Fig. 40A-G: A. AM.5354. R.W. Gess, 1993; B. AM.5355. R.W. Gess, 1993; C. AM.5356, R.W. Gess, 1993; D. AM5358. R.W.

Gess. 1993. E. AM53.57, R.W. Gess. 1993; F. AM5359, R.W. Gess. 1993; G. AM536(I, R.W. Gess, 1993.

Fig. 41A-F: A. AM,S36I, R.W. Gess, 1993; B. AM4837, F. Taylor, 1991 ; C. AM.5363. R.W. Gess, 1993; 1). AM5364. R.W.

Gess. 1993; E. AM536.5. R.W. Gess. 1993; F. AM.5366, F. Taylor. 1991.

Fig. 42A-F: A. AM5367, R.W. Gess, 1993; B. AM5368, N. Hiller, 1992; C. AM,‘i26l , R.W. Gess, 1993; D. AM5369, R.W.

Gess. 1993; E. AM.5370. R.W. Gess. 1993; F. AM.5371, R.W. Gess. 1993.

Fig. 43A-G: A. AM5373, R.W. Gess, 1993; B. AM54()ft, R.W. Gess. 1993; C. AM5374, R.W. Gess. 1993; D. AMS379, F.

Taylor. 1991; F. AM.538(). R.W. Gess. 1993; F. AM.S381. R.W. Gess. 1993; G. AM5382, R.W. Gess. 1993.

Fig. 44A-D: A. AM.5375, R.W. Gess. 1993; B. AM,5376, R.W. Gess, 1993; C. AM.5377, R.W. Gess, 1993; I). AM3378, R.W.
Gess, 1993.

Fig. 45A-C: A-B. AM5383. R.W. Ge.ss. 1993; C. AM48I8. R.W. Gess, 1989.

Fig. 46A-E: AM4883, R.W. Gess. 1993.

Fig. 47A-H: AM49()2, R.W. Gess. 1993.

Fig. 48A-E: A-B. AM4898, R.W. Gess, 1993; C. AM4893, R.W. Gess, 1993; D. AIV14886. R.W. Gess, 1993; E. AM4895,
R.W. Gess. 1993.

Fig. 49A: AM4875. R.W. Gess. 1993.

Fig. 50A-J: A. AM48 1 9, R.W. Gess, 1991; B. AM4908. R.W. Ge.ss, 1 993; C. AM5387i, R.W. Gess, 1 994; D. AM5388, R.W.
Gess. l994;E.AM3384.R,W.Gess, 1994; F. AM5385b. R.W. Gess, I994;G. AM5385a. R.W. Gess. 1994; H. AM4238,
R.W. Ge.ss. 1994; I. AM4867, F. Taylor. R. Gess and M. Anderson, 1991 ; ,F. AM5386, R.W. Gess, 1994.

Fig.5IB-D: AM4907. R.W. Gess. 1993.

Fig. 52A-F; A-B. AM3247, R.W. Gess, 1993; C-D. AM.3246, R.W. Gess, 1993; E. AM5242; R.W. Gess, 1993; F. AM5240,
R.W. Gess, 1993.

Fig. 53A-E: A. AM490I , R.W. Gess, 1993; B. AM488.3, R.W. Gess, 1 993; C. AM488 1 , R.W. Gess, 1993; D. AM4897. R.W.

Gess. 1993; E. AM4900, R.W. Gess, 1993.

Fdg. 54A-C: AM4816. F. Taylor, M. Anderson and R. Gess, 1991.

Fig. 55A-E: A,D. AM4882, R.W. Gess. 1993; B. AM4888, R.W. Gess. 1993; C. AM4891, R.W. Gess, 1993; E. AM4889,
R.W. Gess. 1993.

Fig. 56A-E: A-D. AM4892, R.W. Gess, 1993; E. AM4880. R.W. Gess, 1993.

F4g. 57A-B: AM48I7. R. Gess and M. Stonestreet, 1990.

F4g. 58A, C and D: A and C. AM48I7. R. Gess and M. Stonestreet, 1990; D. AM4866, R.W. Gess, 1988.

Fig. 59A, C and D: A. AM49l2(BPCrl005a), R.W. Gess, 1993; C. AM491 2(BPCrl00l ). R.W. Gess, 1993; D.
AM49l2(BPCrl045), R.W. Gess, 1993.

Fig. 60A-E: A-C and E. AM4912(BPCrl00l ), R.W. Gess. 1993; D. AM49l2(BPCrl005a), R.W. Gess, 1993.

302

GESS AND HILLER: CATALOGUE OF FOSSILS FROM A LATE DEVONIAN BLACK SHALE, S. AFRICA

Fig. 61A-E: A. AM4868, R.W. Gess, 1989; B. AM5389b, R.W. Gess, 1994; C. AM5390, R.W. Gess, 1994; D. AM5389d.
R.W. Gess, 1994, E. AM5389ei, R.W. Gess, 1994.

Fig. 62A-F: A. AM5221, R.W. Gess, 1993; B. AM5391ai, R.W. Gess, 1993; C. AM5391bi, R.W. Gess, 1993; D. AM5389a,

R. W. Gess, 1993; E. AM5392aii, r!w. Gess, 1993; F. AM5392b, R.W. Gess, 1993.

Fig. 63A-D: A. AM5393a, R.W. Gess, 1993; B. AM5249, R.W. Gess, 1993; C. AM5394a, R.W. Gess, 1993; D. AM4821,

S. Coutouvides, 1989.

Fig. 64A-D: A. AM5395a. R.W. Gess, 1993; B. AM5395b, R.W. Gess, 1993; D. AM5396, R.W. Gess. 1987.

Fig. 65A-D: A. AM5397b, R.W. Gess, 1993; B. AM5398, R.W. Gess, 1993; C. AM5400. R.W. Gess, 1994; D. AM5399a,
R.W. Gess. 1994.

APPENDIX 2 - LIST OF ADDITIONAL SPECIMENS

Rusophycos and Cruziana: AM5402A-Z, R.W. Gess, 1993; AM3403A-B, R.W. Gess, 1993.

Spirophyton-Wke traces: AM5404A-E, R.W. Gess, 1993-1994

Hungerfordia sp.: AM4844, F. Taylor, 1991 ; AM4846, F. Taylor, 1991 ; AM5228, R.W. Gess, 1993; AM5372AA-AZ, BA-
BZ, CA-CU. R.W. Gess, 1985-1994; AM5405A-C, F. Taylor. 1991.
cf. Yeaia sp.: AM5268. R.W. Gess, 1993; AM561 1-5612, R.W. Gess, 1993.

Charophytes: new genus 1, new species 1: AM5193-5199, R.W. Gess, 1993-1994; AM52()5( 1 )-( 10). R.W. Gess, 1993-
1994; AM5346. R.W. Gess. 1994; AM5423, Hodgen Family, 1993; AM5424, R.W. Gess, 1993.

Charophytes: new genus 1, new species 2 : AM5202-5203, R.W. Gess, 1993; AM5204( 1 )-(4), R.W. Gess, 1993; AM5417-
5418, R.W. Gess. 1994; cf. AM5407-5416, R.W. Gess. 1992-1994.

Charophytes: new genus 2, new species 1: AM5210, R.W. Gess, 1993; AM52 11(1 )-(20), R.W. Gess, 1992-1993; AM5419A-
Z. R.W. Gess, 1 992- 1 994; AM5420A-N. R.W. Gess, 1 992- 1 994; AM542 1 A-D, F. Taylor. 1991; cf AM5422A-L. R.W.
Gess, 1992-1994

Charophytes; new genus 2, new species 2: AM521 3, R.W. Gess. 1993; AM52 16, R.W. Gess. 1993; AM521 7( 1 )-(3). R.W.
Gess. 1993.

Waterweed: (cf Fig. 12): AM5425-5426, R.W. Gess, 1993-1994; AM5600, R.W. Ge.ss, 1994.

Diitoitia alfreda: AM5427-5432. R.W. Gess. 1993-1994; AM5346, R.W, Gess, 1993.

cf Fig. 14: AM5433-5437, R.W. Gess, 1993-1994; AM5438, F. Taylor, 1991 ; AM56 13-56 14, R.W. Gess, 1993.
Zosterophyllopsida: with spheroids: AM5439-548 1 , R.W. Gess. 1993-1994; AM5476, F. Taylor, 1991 ; AM5482-5483. F.

Taylor, 1991 ; AM550 1-5502, R.W. Gess, 1993.

Zosterophyllopsida: cf Fig. 18F: AM5484-5486, R.W. Gess, 1993.

? Zosterophyllopsida: cf rhizomes: AM483 1 . F. Taylor, 1991; AM5488, F. Taylor, 1991; AM5489-5490. R.W. Gess, 1 993;

AM549l(c;/:5253), R.W. Gess, 1994; AM5492-5500, R.W. Gess, 1993-1994.

Lycopsida: cf Fig. 19: AM5503-5504. R.W. Gess, 1994; AM553I-5532, R.W. Gess, 1994.

Lycopsida: cf Fig. 20: AM5309, R.W. Gess. 1993; AM5505-5507, R.W. Gess, 1993-1994,

Lycopsida: cf Fig. 21: AM4838, F, Taylor, 1991; AM5508-55I6, R.W. Gess, 1993-1994; cf AM5517, R.W. Gess. 1993.
Leptophloeiim aiistrale: steins and hark: AM53I3. R.W. Gess, 1993; AM55I8A-H, R.W. Gess, 1993; AM55I9A-F, R.W.
Gess. 1993; AM5520A-B. R.W. Gess, 1993; AM5524-5525, R.W. Gess. 1993; AM5526A-G. R.W. Gess. 1993-1994;
AM5527-5529, R.W. Gess, 1993-1994.

Leptophloeiim austale: rooting base: AM552I, R.W. Gess, 1993.

Leptophloeiim aiistrale: ?rootlets: AM5522A-D, R.W. Gess, 1993.

? Leptophloeiim aiistrale - decortication rhombs: AM4894(BPWM1()09, R.W. Gess, 1993; BPI0I3-BPI0I6, BPI026,
BP 1028. BPI030-I039). AM5530A-Q, R.W. Gess, 1994; AM5533. R.W, Gess, 1993; AM5615-56I6. R.W. Gess, 1994.
Loiigicatrix sp.: AM5534, F. Taylor, 1991; AM5535, R.W. Gess, 1993.

Haplostigma irregularis: AM5523A-C. R.W, Gess, 1993.

303

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 5 28 MAY 1995

Axes with leaf/root bases: AM5536. R.W. Gess. 1993; AM5537A-D, R.W. Gess, 1993; AM5538A-C, R.W. Gess, 1993;

AM5539, H.M. Anderson, 1993; AM3617-5619, R.W. Gess, 1993-1994.

Striated axes: cf. Fig. 30H, 41F: AM4856, F. Taylor. 1991 ; AM5543, F. Taylor, 1991 ; AM5544-5546, R.W. Gess; 1993.
Archaeopteris new species: AM492 1 : 1 , 3-6. R.W. Gess, 1993; AM4923;2-3. R.W. Gess, 1993; AM4915, R.W. Gess, 1993;
AM49I7. R.W. Gess, 1993; AM49 1 9-4922. R.W. Gess, 1993; AM5547A-E. R.W. Gess, 1993; AM5548A-H, R.W.
Ge.ss, 1993; AM3349A-M. R.W. Gess, 1993; AM5392, R.W. Gess, 1994; AM3620, R.W. Gess, 1993.

? Archaeopteris new species 2: cf. Fig. 34: AM484 1 . F. Taylor, 1991; AM3350, R.W. Gess, 1 993.

Dutoitia maraisia: AM4836. F. Taylor, 1991; AM4875. F. Taylor, 1991 ; AM5343B-C. F-K. M-Z, AA-AD, R.W. Gess and
N. Hiller, 1993; AM555I-3536. R.W. Gess. 1993; AM3367, R.W. Gess, 1993.

? Progymnospermopsida: cf. Fig. 38: AM4836. F. Taylor, 1991; AM5566, F. Taylor, 1991; AIVI5337-3563, R.W. Gess,
1993-1994; AM3568. R.W. Gess, 1993.

Plant axis: cf Fig. 39C: AM5577, R.W. Gess, 1993.

Plant axes: cf. Fig. 39F: AM3369-5576. R.W. Gess. 1993.

Plant axes: cf Fig. 4IIC: AM5382-3583. R.W. Gess. 1993; AM5584, F. Taylor, 1991.

Plant axes: cf Fig. 40F: AM3394-5597. R.W. Gess. 1993-1994.

Plant axes: cf Fig. 40G: AM3398. F. Taylor. 1991 .

Plant axes: cf Fig. 41 E: AM3601-36I0. R.W. Gess. 1993-1994.

Plant axes with whorls: cf Fig. 40A-B: AM5378-5380, R.W. Ge.ss, 1993-1994; AM558I. F. Taylor, 1991.

Plant axis: cf Fig. 41A: AM5399. R.W. Gess, 1993.

Trilete spores: cf Fig. 40D: AM485I, F. Taylor. 1991; AM4853, F. Taylor, 1991; AM5386A-Z, R.W. Gess, 1993-1994;
AM5387A-1, R.W. Gess. 1993-1994.

Fertile capsules: cf. Fig. 40E: in whorls: AM5588-3589, R.W. Gess. 1993.

Fertile capsules: cf Fig. 40E: AM5590-5593, R.W. Gess. 1993-1994.

Eurypterid: limb fragments: AM562I -5622. R.W. Gess. 1993-1994.

Eurypterid fragments: AIVI5623-5641, R.W. Gess. 1993-1994.

Concho.stracon: AM3647A-Z, R.W. Gess. 1993- 1 994; AM5648A-Z, R.W. Gess. 1993- 1 994; AM5649A-Z, R.W. Gess. 1993-
1994. ^

Ostracod: AM5642-AM5646, R.W. Gess. 1993-1994; AM569I. R.W. Gess, 1994.

Arthrodire: AM4879. R.W, Gess, 1993; AM4884, R.W. Gess, 1993; AM4886=4893, R.W. Gess, 1993; AM4890, R.W. Gess,
1993; AM4896, R.W. Gess, 1993; AM4899. R.W. Gess, 1993; AM4903-4906, R.W. Gess, 1993; AM5235=5237, R.W.
Gess. l993;AM5236,M.Stonestreet, l993;AM.524l=AM5248,R.W.Gess, 1993; AM5244, R.W. Gess. 1993; AM.S650-
5667, R.W. Gess. 1993-1994; AM5670. R.W. Gess, 1994; AM5686-5688, R.W. Gess, 1993-1994.

Bothriolepis sp.: AM49I I. R.W. Gess, 1993; AM5239, N. Hiller. 1992; AM5668, R.W. Gess. 1994; AM5669, N. Hiller,
1992.

Piscean spines: AM568I-5683. R.W. Gess. 1993-1994.

Chondrichthyes: AM5243, R.W. Gess, 1993; AM5245. R.W. Gess, 1994.

Actinistia: Juveniles: AM4912(BPCrlO()2-BPCrlO()4. BPCrl006, AMBPCrlOlO), R.W.Gess, 1993.

Actinistia: adults: AM4909i(scale), R.W. Gess. 1989; AM5675-5676, R.W, Gess, 1994; AM5697A-0(,scales), R.W.Gess,
1993-1994.

Rhipidistia: AM4909a-h, k-l (scales). R.W. Gess, 1989; AM5389f-z. aa-ag, R.W. Gess, 1992-1994; AM567I, N. Hiller,
1992; AM5672-3. R.W. Gess. 1993; AM5674A-L(scales), R.W. Gess, 1985-1994.

Pisces: AM5677-5679, R.W. Gess, 1993; AM568()A-Q(.scales), R.W. Gess, 1993-1994; AM5684-5685, R.W. Gess, 1994;
AM5689. R.W. Gess. 1994.

?Piscean coprolites: AM540IA-O, R.W. Gess. 1993-1994; AM569(). R.W. Gess, 1994.

Other ?fecal material: AM5642-3643, R.W. Gess, 1993; AM569I-5695, R.W, Gess, 1994.

304

INSTRUCTIONS TO AUTHORS

LAYOUT of recent issues should be examined for details of acceptable style and format.

MANUSCRIPTS should be submitted on floppy disk accompanied by three print-outs
(spacing I 'A or 2). In exceptional cases typed manuscripts will be accepted. These should be
submitted in triplicate, typed on A4 size paper, double-spaced, on one side of the paper only
and with wide margins all round.

Only generic and specific names should be underlined to indicate italics. All other marking
up should be left to the editor.

Footnotes should be avoided as far as possible.

All measurements should be metric.

Systematic papers should follow the rules of the International code of zoological
nomenclature or the International code of botanical nomenclature, as applicable.

ILLUSTRATIONS should bear the author’s name, title of paper and number in soft pencil on
the reverse side. Diagrams, line drawings and photographs are all regarded as figures, should
be termed as such and should be numbered in a single series.

In addition to the original illustrations a photocopy of each should be submitted.

Diagrams and line drawings should be reducible to a size not exceeding 14 x 16 cm ( 17,5 cm
including captions). Photographs for half-tone blocks should be final size, of high contrast
and printed on glossy paper. Colour illustrations are only printed under special circumstances
and their financing must be arranged by the author.

A scale should appear with all illustrations or the magnification should be given in the
caption.

A list of captions should be submitted on a separate sheet.

TABLES should be submitted on separate sheets. Foldouts and tables printed sideways should
be avoided.

REFERENCES should be listed alphabetically at the end of the paper. Abbreviations of
periodical titles should conform with those in the World list of periodicals (4th ed.). Layouts
must be as follows:

Alexander, J. 1969. The indirect evidence for domestication. In: Ucko, P. J. and G. W.
Dimbleby eds, The domestication and exploitation of plants and animals. London:
Duckworth, pp. 123-129.

De Winter, B. 1969. Plant taxonomy today. S.Afr.J.Sci. 65 (8): 229-242.

JuBB, R. A. 1967. Freshwater fishes of southern Africa. Cape Town: Balkema.

GALLEY PROOES are sent (when practicable) to authors for checking printer’s errors.
Adjustments of language, style and punctuation and any other amendments of the original
manuscript, if made at this stage, may be charged to authors.

AUTHOR’S COPIES (50) are issued free of charge. When more than one author is involved
the fifty free copies will be sent to the senior author. Additional copies ordered before printing
are supplied at a reduced cost.

CADAR, P.E.-K887

ISSN 0570-1880

0^ H
I

A(p\S
J\j fl

Annals of the

Cape Provincial Museums

Natural History

Ann. Cape Prov. Mus. (nat. Hist.)

Volume 19 Part 6 29th April 1996

M4Y 3 0 1997

Alien and translocated aquatic animals in southern Africa (excluding Zimbabwe and
Mozambique) - revised checklist and analysis of distribution on a catchment basis

I.J. DE MOOR and M.N. BRUTON

Published jointly by the Cape Provincial Museums
at the Albany Museum, Grahamstown, South Africa

ANNALS OF THE CAPE PROVINCIAL MUSEUMS

These Annals are published in two series. Natural History and Human Sciences, the latter series
covering cultural history, ethnology, anthropology and archaeology. They are issued in parts at
irregular intervals as material becomes available.

The primary objective of these Annals is to disseminate the results of research work undertaken by
staff members of the Cape Provincial Museums. However, a manuscript by any author is eligible
for publication provided it is based wholly or partly on the coliection/s of one or more of these
museums or if the material on which the work is based is wholly or partly deposited in one of
these museums.

Parts are obtainable from the Librarian of the Albany Museum. Correspondence in connection
with these Annals should be addressed to the Editor, Albany Museum, Grahamstown 6140.

Editor

Dr F. W. GESS

Assistant Editor
Dr S. K. GESS

Alien and translocated aquatic animals in southern Africa (excluding
Zimbabwe and Mozambique) - revised checklist and analysis of distribution

on a catchment basis

by

I.J. DE MOOR and M.N. BRUTON*

(J.L.B. Smith Institute of Ichthyology, Grahamstown, 6140 South Africa)

ABSTRACT

The distribution patterns of 44 alien and translocated freshwater aquatic animals in southern
Africa (excluding Zimbabwe and Mozambique) are updated and re-assessed. A comparison with
1988 records indicates the presence of two alien species not previously recorded in natural waters.
The distributions of five species, previously recorded in catchments outside their native ranges, have
been extended into new catchments. The common carp Cypriuus carpio Linnaeus 1758 has invaded
more catchments than any other species and its range has recently been extended into the Phongolo
and upper Mkuze systems.

The composition of alien species in each major catchment and the catchments invaded by
individual alien species are documented. Results indicate a positive correlation between the number
of alien species and water availability in catchments. Regions with the highest number of alien species
are associated with major urban centres. Records suggest that high numbers of alien species are found
in regions with a low diversity of indigenous species, however, this trend could not be statistically
validated. In eight out of 19 catchments/waterbodies investigated more than half of the fish species
were aliens. In the Kei catchment 1 1 out of 13 fish species are aliens.

Certain waterbodies ofhigh conservation status (Okavango, Lake Sibaya and the Mkuze Swamps)
in sub-tropical regions are free of alien fish. Temperatures in these waterbodies are likely to be within
the tolerance range of most tropical species associated with the aquarium trade. It is recommended
that special measures be taken to prevent introductions of alien species into these systems. The presence
of Cypriuus carpio, Micropterus salinoides (Lacepede 1802) and Oreochromis mossainbicus ( Peters
1852) in the Omatako drainage raises the possibility of the future introduction of these species into
the Okavango system. Potential introductions through interbasin transfer schemes are also discussed.

CONTENTS

page

Introduction 306

Methods 308

Results 309

Discussion 3 1 1

Reclassification of distribution records 31 1

New species and range extensions of naturalised species 316

Degree of invasion of catchments 317

Invasive species 322

Future introductions 324

Interbasin transfers (IBTs) 324

Possible future introductions into the Okavango region 325

Conclusion 326

* Pre.sent address: Two Oceans Aquarium, P O Box 50603, Wateifront, 8002 South Africa.

305

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 6 29 APRIL 1996

Acknowledgements 326

References 326

Appendix 1 : Major catchments and sub-catchments in the study area 333

Appendix 2: Checklist of alien species introduced into southern Africa 336

Appendix 3: Checklist of translocated indigenous species within the study area 339

Appendix 4: References associated with Appendices 2 and 3 341

Appendix 5: Existing and future interbasin transfer schemes 343

INTRODUCTION

Numerous ecological problems have resulted from the introduction of alien aquatic animals into
inland waters in many parts of the world (Welcomme, 1988). These include the degradation of the
environment due to habitat alteration, the disruption of the host community through competition with
and predation on indigenous species, deterioration of fish stocks (through stunting of the introduced
population), genetic contamination of the host community, and the introduction of parasites and
diseases.

Serious economic and health problems have arisen as a result of some introductions. The recent
introduction of the zebra mussel Dreisseiia polymorpha ( Pallas 1771) into the Laurentian Great Lakes
in North America has caused widespread economic damage as a result of its prolific growth in the
water pipes of electricity-generation and water-treatment facilities (Griffith et ah, 1991). The
introduction of the malaria-vector mosquito Anopheles gambiae sensu lato Giles 1920 into South
America resulted in a malaria epidemic causing the death of over 20 000 people (Soper and Wilson,
1943).

Bruton ( 1986) noted that the life-history style adopted by introduced species was an important
factor in determining the success or failure of the introduction. Preliminary distribution records
suggested that altricial fish (which produce small, incompletely developed young and are generalists
capable of surviving in an unstable, uncrowded environment), such as rainbow trout Oncorhynchiis
mykiss (Walbaum 1792) and the common carp Cyprinus carpio, had successfully invaded species-
poor, abiotically harsh environments. Precocial species (which produce large, well- developed young
and are specialists adapted to survive in a stable, crowded environment), such as the largemouth bass
Micropteriis sahnoides, swordtail Xiphophorns helleri Heckel 1 848, and the guppy Poecilia reticulata
Peters 1859, tended to invade species-rich, biotically harsh environments. The present paper will
attempt to evaluate some of these hypotheses.

De Moor and Bruton ( 1988) reviewed the problem in southern Africa and found that 58 alien
species had naturalised breeding populations in the southern African region (excluding Zimbabwe
and Mozambique). Of these, 33 were from other countries and 25 were transfers of species into new
catchments within southern Africa. Of the 58 species which were frequently recorded in natural
habitats, 37 were considered to be detrimental, three to be beneficial and 18 to be equivocal (i.e.
beneficial in some respects and detrimental in others). Detrimental impacts included the alteration
of the habitat, predation of and competition with indigenous species, the introduction of parasites,
and the threat of genetic contamination (de Moor and Bruton, 1988). Although such impacts can be
regarded as being significant in terms of environmental effects, alien aquatic animals in southern
Africa have not yet caused any major medical or economic impacts on the scale of other well-known
disasters mentioned above.

The aim of this paper is to provide an updated checklist of alien aquatic animals in southern
Africa and to revise distribution records in terms of catchment distributions. This will enable
conservation officials and other managers to rapidly assess the degree of invasion in each catchment

306

DE MOOR AND BRUTON: ALIEN AND TRANSLOCATED AQUATIC ANIMALS IN SOUTHERN AFRICA

and to identify catchments which are susceptible to further invasions. Information from previous
publications (Bruton, 1986; de Moor and Bruton, 1988) is also updated and re-assessed. It is hoped
that the synthesis of distribution records presented in this paper will serve as a basis for further
investigations into the biogeography and ecology of alien aquatic animals in southern Africa.

It should be noted that the invasion of freshwater ecosystems is an ongoing process and the
present review represents a “snapshot” picture of known distribution records at present. There are
notable gaps in distribution records, particularly with respect to fish parasites and certain popular
angling species (such as Cypriniis carpio, Microptenis salmoides, and Oreochromis mosscuuhicus)
which have been illegally translocated into many catchments in southern Africa.

With the exception of the grass carp Ctenophcuyjigodou /Jc/Zu (Valenciennes 1844) (a long-lived
species which does not breed in natural waters in southern Africa) only naturalised alien species are
included in this review. The regions covered include South Africa, Lesotho, Swaziland, Namibia and
Botswana. With the exception of the latter two countries, descriptions of major catchments (Fig. I )
follow the convention of the Department of Water Affairs (Anon, 1986). Appendix 1 describes the
catchments and sub-catchments illustrated in Fig. 1, as well as the major rivers, impoundments and
towns in each region.

Certain categories of species listed in de Moor and Bruton ( 1988) have been excluded from this
review. These include: marine species; threatened species translocated to regions close to their native
range; species translocated within the same major catchment; and biocontrol agents which have a
beneficial impact on the environment.

A number of terms used in this publication are defined below.

The study area. This is indicated in Figure 1. The area includes all catchments south of the
Limpopo and Cunene rivers as well as the Okavango region which forms part of the upper Zambezi
catchment. For political reasons distribution records from Zimbabwe and Mozambique were not
available during the time when work on this project was being carried out.

Natural and semi-natural habitats. For the purpose of this study this includes natural
watercourses, large impoundments and small earthem impoundments either of which may overflow
into natural waters during flood periods. Excluded from this definition are garden ponds, closed-
system aquaculture installations and concrete farm reservoirs.

Alien species. A species which has been translocated, either intentionally or unintentionally, by
man into catchments beyond its native range.

Translocated indigenous species. A species indigenous to the study area, which has been
translocated by man into new catchments within the study area. Translocated species can be regarded
as a special category of alien species.

Invasive species. An alien species which has spread unaided within freshwater ecosystems.

Naturalised species. An introduced species which has established self-sustaining populations in
areas of natural or semi-natural vegetation or habitat.

Detrimental impact. The introduction has had a harmful impact on the environment.

Beneficial impact. The introduction has apparently had an advantageous impact on the
environment. This term is confined to host-specific alien species introduced for the purpose of
biocontrol, which successfully controlled the targeted host species.

Major catchments. Catchments categorised by the Department of Water Affairs (Anon, 1986)
as major and here designated by letters of the alphabet (Fig. 1 ). In most instances these catchments
are discrete and separated from other catchments by watershed boundaries. Exceptions are the Olifants
and the Vaal (catchments B & C, Eig. I ) which represent major tributaries of the Limpopo and Orange

307

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 6 29 APRIL 1996

Fig. I : The study area, indicating major catchments and sub-catchments (refer to Appendix 1 for information on major rivers,
waterbodies and place names in each catchment).

rivers (catchments A & D, Fig. I ) respectively.

In some instances a number of small discrete coastal catchments (within major catchments G,
K, M, P, R, T, U and W: Fig. 1 ) are categorised together as a “major catchment area”.

Snh-catchments. Catchments of tributaries within major catchments, designated by numbers in
Fig. 1.

Successful iutrocluctiou. An introduction is described as successful where an alien species has
established a naturalised population in natural or semi-natural waters.

Failed iutrocluctiou. An introduction is regarded as unsuccessful or failed where an alien species
has not succeeded in establishing a naturalised population, despite repeated introductions into natural
waters.

METHODS

Distribution records were obtained from: the Albany Museum and the J.L.B. Smith Institute of

308

DE MOOR AND BRUTON: ALIEN AND TRANSLOCATED AQUATIC ANIMALS IN SOUTHERN AFRICA

Ichthyology, Grahamstown (Appendices 2 and 3); literature records of species from accredited
scientific journals and reports (Appendices 2, 3 and 4); and personal communications with nature
conservation officials and other experts in the field (Appendices 2, 3 and 4).

Distribution records were categorised as follows.

S - Self-sustaining populations (=breeding populations).

P - Probable presence. Where no formal record has been received of the presence of a widespread
species in a particular catchment sub-division but where such a species is widespread in an
adjacent catchment it is considered that the presence of the species is “probable” in the
catchment concerned.

R - Regularly stocked. Where the species is regularly stocked in a particular catchment but
populations are probably not self-sustaining.

M -Possibly stocked. Where there are no definite records of stocking but it is suspected that the
species may be stocked somewhere in the catchment. It is unlikely that self-sustaining
populations occur in these catchments.

H - Historical. Where only isolated distribution records prior to 1985 exist so that it is not known
whether self-sustaining populations are present.

B - Boundary. Where a self-sustaining population is found in a region close to the watershed
boundary and consequently the catchment to which the locality belongs is uncertain.

* - New distribution record. Not recorded in de Moor and Bruton (1988).

Latitude/longitude co-ordinates Vv'ere obtained from ordinance survey maps, the Times World
Atlas or Skead (1973). The catchment associated with each record was then identified by reference
to Fig. 1.

In the case of certain species it was possible to estimate abundance in selected catchments by
reference to literature or through personal communication with researchers. If only one record exists
of the species, then it was either classified as rare (if a recent record) or as an historical record. If
museum records indicated that the species had been collected in a number of localities in at least three
tributaries, or in more than four localities in the mainstream of rivers, then the species was classified
as widespread within the catchment or sub-catchment.

Based on the above information, abundance Vv'as noted within certain sub-catchments and
classified as follovv's (Tables 1 - 4).

1 - Very rare. Only a single record exists.

2 - Isolated populations in small restricted region/regions. These may, however, be thriving

populations.

3 - Widespread. Found throughout the upper and lower catchment or sub-catchment.

3u - Widespread in the upper catchment.

3 1 - Widespread in the lower catchment.

It must be emphasised that records of abundance are incomplete, and have only been noted when
the relevant information is available. This implies that the distribution of most alien species is greater
than indicated by records summarised in Tables 1-4.

RESULTS

The checklists of alien and translocated indigenous species (Appendices 2 and 3 respectively)
include brief notes on the native range, date of first introduction, impact, and abundance of each
species, as well as code numbers indicating selected references (listed in Appendix 4). For further
details see de Moor and Bruton (1988).

309

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 6 29 APRIL 1996

Table 1 . Distribution records of alien aquatic animals present in catchments and sub-catchments
A - G illustrated in Fig. 1 .

Species

Catchments

Al

A2

A3

B1

B2

B3

Cl

C2

C3

D1

D2

D3

D4

D5

D6

E

F

G

Ichthyophthirius inultifilis

S

S*

S

Bothriocephalus acheilognathi

S

S

Trichodina acuta

S

S

S

S

s

Craspedacusta sowerhyi

Argulus japonicits

S

S

s

Cherax tenidmanus

Procamharus clarkii

Phvsa acuta

s

s

S

S

S

3

3

3

3

Helisoma duryi

H

Lvmnaea columella

S

s

S

S

S

S

s

S

S

S

3

3

3

3

3

3

Oncorhvnclius mykiss

R

M

S

R

S/R

S/R

M

M

S

S

S

S

2

2

3u

3u

2

2

Salmo trutta

M

M

S

M

S

S

R

S

S

2

2

2

2

2

Carassius a u rat us

S

1

S

S

Ctenopharyugodon idellus

M

R

M

Cyprinus carpio

S

P

S

P

S

S

P

S

S

S

s

s

s

S

M

s

3

2

3

3

3

3

Hypophthalmichthys molitrix*

S*

Tinea tinea

S

!

Gambusia affinis

S*

H

S*

S

S*

Poecilia reticulata

s

Xiphophorus helleri

Lepomis macrochirus

S

S

S

P

P

s

p

S

S

3

3

Micropterus dolomieu

S

S

s

P

S

P

s

P

p

p

S

S

3

3

Micropteriis punctulatus

S*

1

H

Micropterus salinoides

s

P

s

s

S

s

s

s

S

s

s

S

S

S

2

3

3 I

310

DE MOOR AND BRUTON: ALIEN AND TRANSLOCATED AQUATIC ANIMALS IN SOUTHERN AERICA

Table 1. Continued.

Species

Catchments

Al

A2

A3

Bl

B2

B3

Cl

C2

C3

Dl

D2

D3

D4

D5

D6

E

F

G

Perea fluviaiilis

S

S

Oreochromis aureus

H

Oreochromis niloticus

H

Truchemys s. elegans

S

Code for distribution
S - Self-sustaining populations
P - Self-sustaining populations probable,
but no definite records

R - Regular stocking, probably not self-sustaining
M - Maybe stocked somewhere in catchment,
self-sustaining populations unlikely
H - Historical. May or may not still be present
U - Uncertain historical translocation
* - New distribution record since de Moor & Bruton ( 1988)

Code for abundance

1 - Very rare. Probably a single record

2 - Isolated population in small restricted region/s

3 - Widespread

3u- Widespread in upper catchment
3i - Widespread in lower catchment

B - Boundary: Close to watershed between two catchments,
so exact catchment is uncertain

Table.s 1 and 2 summarise the distribution of alien and translocated indigenous species
( respectively ) in catchments A-G. Tables 3 and 4 summarise the distributions of alien and translocated
indigenous species (respectively) in catchments H-Z. The number of species in each sub-catchment
and major catchment is also noted. Tables 3 and 4 also indicate the number of catchments (from A-
Z) invaded by each species. It should be noted that in calculating these totals, only the “S” (species
survives) and “R” (regularly stocked) categories of distribution records are considered. Historical
records are noted separately.

DISCUSSION

Reclassification of distribution records

Species for which there were previously single distribution records (de Moor and Bruton, 1988)
have now been classified as having “historical distributions” since no further reports have been
received of their presence in natural waters (Tables 1 - 4). These species include two alien cichiids
(Oreochromis aureus (Steindachner 1864) and O. nilolicus (Linnaeus 1758)) previously recorded in
Natal and the Western Cape, one alien crayfish (Clierax lenuimauus (Smith 1912)) previously
recorded in the Buffalo River (Eastern Cape), and four translocated indigenous cichiids ( Oreochromis
andersouii (Castelnau 1861), Sargochromis codringtoui (Boulenger 1908), Serranochromis
tlnimhergi (Castelnau 1861) and S. angusticeps (Boulenger 1907)) which were included in a group
of species translocated into the Shashi impoundment from the Okavango region (Gilmore, 1978).

It should be noted that the term “historical” does not imply that the species has definitely
disappeared from the region in question. In the context of this publication “historical” indicates that
no further records have been received in the past 10 years, and it is possible that the species may still
be present in the region.

311

ANN. CAPE PROV. MUS, (NAT. HIST.) VOL. 19, PT. 6 29 APRIL 1996

Table 2. Distribution records of translocated indigenous aquatic animals present in catchments
and sub-catchments A-G illustrated in Fig.l. Total numbers of alien species (including species listed
in Table 1 ) in each catchment are also given.

Species

Catchments

A1

A2

A3

B1

B2

B3

Cl

C2

C3

D1

D2

D3

D4

D5

D6

E

F

G

Barbus aeneus

Barbus uiioplus

Labeo capensis

Labeo uinbratus

S

Clarias gariepinus

S

Chetia flaviventris

S*

Oreochroniis amlersoiiii

H

Oreochromis macrochir

S

Oreochroniis mossambicus

S

S

3i

S

s

Sargocliroinis codringtoni

H

Serranochroinis tliuinbergl

H

Serranochromis r. jallae

H

H

Serranochromis angusticeps

H

Tilapia r. swierstrae

Tilapia sparnnanii

S

S*

3

Sandelia capensis

s

No. alien
species in s-c

11

1

+

5H

7

-1-

IH

9

10

7

3

1 1

5

2

6

3

2

3

3

9

0

15

+

5H

No. alien species in m-c’s

14

15

13

10

9

0

15

s-c = sub-catchment
m-c = major catchment

Code for distribution
S - Self-sustaining populations
P - Self-sustaining populations probable,
but no definite records

R - Regular stocking, probably not self-sustaining
M - Maybe stocked somewhere in catchment,
self-sustaining populations unlikely
H - Historical. May or may not still be present
U - Uncertain historical translocation
* - New distribution record since de Moor & Bruton ( 1988)

Code for abundance

1 - Very rare. Probably a single record

2 - Isolated population in small restricted region/s

3 - Widespread

3li - Widespread in upper catchment
3i - Widespread in lower catchment

B - Boundary: Close to watershed between two catchments,
so exact catchment is uncertain

312

Table 3. Distribution records of alien aquatic animals present in catchments and sub-catchments H - Z illustrated in Fig.l.
Total catchments (from A - Z) invaded by alien species is also given.

DE MOOR AND BRUTON: ALIEN AND TRANSLOCATED AQUATIC ANIMALS IN SOUTHERN AFRICA

313

Table 3. Continued.

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 6 29 APRIL 1996

Total No.
Catch. 's In-
vaded (A-Z)

m-c

m

m

X

OO

<N

K

O'!

X

<N

o\

s-c

m

ro

OS

OO

32

CA

Catchments

N

a

s

GO CQ

CO ro

Y2

s

B

(zi Da

CO r<S

X3

CO

CO

ZX

CO

CO

CO

*

CO

W2

lyy

*

CO

W1

X

CO

>

CO CO

CO

CO (N

CO

c/5

CO

CO ro

CO

CO CN

CO fo

ac

oa

CO

T3

*

CO

CO

CO

T2

*

CO

•vf

CO

CO

*

CO

H

CO

Cu

GO

CO

CO

CO

CO

od

CO

CO

*

CO

CO rt")

CO

o

an

*

CO

I

0-

*

CO

CO —

CO

z

a:

S

CO

CO

j

* _
CO

CO ro

CO

CO

CO

CO

-

4f

CO

CO

X

CO

CO ro

CO ro

Species

Poecilia reticulata

Xiphophorus helleri

Lepomis macrochirus

Micropterus dolomieu

Micropterus pimctulatus

Micropterus salmoides

1 Perea fliiviatilis

1 Oreochromis aureus

1 Oreochromis niloticus

II Trachemys s. elegans

M 2

.5 E

E E

D, ^

^ O- T3 "O ^

(U (U (U

cd *0 Cu O, G, i2

> s ^

<N m CO CO

c £
<u -G

E id

OD W) ^

O Q

c •=:

X) <u
e3 -C

O S
G- S
' o

bij c/5

-E "O

G O

^ I -H

S i: o

! « E

° -C C
^2-2

•S •- o

^ ^ ^ W)

(U (U 3

GO C/5 XI Oh

'u t: -TD

O <U :>

^ e ^

K D Z

a: S

I 5

314

Table 4. Distribution records of translocated indigenous aquatic animals present in catchments and sub-catchments H-Z
illustrated in Fig.l. Total numbers of alien species (including species listed in Table 3) and total numbers of catchments (from
A-Z) invaded by alien species are also given.

DE MOOR AND BRUTON: ALIEN AND TRANSLOCATED AQUATIC ANIMALS IN SOUTHERN AFRICA

Total No.
Catch. ’s In-
vaded (A-Z)

m-c

to

(N

rj

1

0-)

-

0\

X

o\

X

n

CD

CN

s-c

(N

(N

-

-

OS

0-)

ri

Catchments

N

00

CO TO

CO

lO

Y2

CO fO

3

2B

TO

O

X3

ir-i

-

cX

O

X

oo

W2

CO —

r-

OO

W1

X

4

2H

>

*

00

X

*

CO

*

X ^

O X

r^l

o

D

X

■X-

co

CO

oo X

rr-,

00

T3

*

oo

CO

*

CO

d-

HI

L

H

00

lO',

00

00

Cl

ri

CO

CO

00

Tt

't

a

OO

CO

CO

CO

7

2H

r-

Cu

00

CO

CO

OS

os

z

CO

00 ro

CO

5

IH

S

00

CO

00

00

CO

lO-j

-X-

co

OS

Ov

-

oo

CO

SO

SO

00

CO

O

o

Species

Barbus aeneus

II Barbus anoplus

II Labeo capensis

Labeo wnbratus

Clarias gariepinus

Chetia flaviventris

Oreochromis andersonii

Oreochromis macrochir

Oreochromis mossambicits

Sargochromis codringtoni

Serranochromis thimbergi

Serranochromis r. jallae

1 Serranochromis angusticeps

Tilapia r. swierstrae

Tilapia sparrmanii

1 Sandelia capensis

Total No. alien species in s-c

Total No. in m-c’s

315

Legend. As for Table 1

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 6 29 APRIL 1996

Table 5. Updated distribution records (compared to de Moor and Bruton, 1988).

Species

Range

No. catchments

Distribution

extensions

where new

code

(catchments)

distributions

(Fig. 1)

recorded

s-c

m-c

Species not previously recorded in study area

Chetia flaviveutris

B3

1

1

NR

HypophthaUnichthys molitrix

B2

1

1

NR

Procamharus clarkii

X2

1

1

NR

New distributions of species previously recorded in study area

Bothriocephaliis acheiloguatlii

U,X1

2

2

RE

Carassiiis aiiratus

P,T3,U

3

3

CE

Cteuopharyngodon idellus

T2,W2

2

2

CE

Cyprimis carpio

W2

1

1

RE

Poecilia reticulata

T3

1

1

CE

Lepomis macrochirus

L,P,T2,

4

4

CE

Micropteriis punctulatus

D2,Q,R,T2,

4

4

CE

Micropterus salmoides

W2,X1

2

2

CE

Perea fluviatilis

T2

1

1

CE

Barbus aeiieus

V

1

1

RE

Barbus anoplus

T3

1

1

CE

Labeo capensis

V

1

1

RE

Labeo umbratus

V*

1

1

RE

Tilapia r. swierstrae

T3, U

1

1

CE

Tilapia sparnuauii

G,K,S,T1

4

4

CE

s-c - sub-catchments

m-c - major catchments

NR - New records: alien species not previously recorded in natural waters in the study area.

CE - Records which are probably the result of greater collection effort. Unlikely to represent a

range extension of the species.

RE - New distribution records which represent probable range extensions of existing alien species.

* - Only a single specimen was collected: may not have established a population, so record is

classified as historical (H) (Table 2)

New species and range extensions of naturalised species.

Table 5 summarises new records (NR) of three species (two alien and one translocated indigenous
species) not previously recorded, as well as apparent range extensions (into new catchments or sub-
catchments) of 15 alien species which were listed in de Moor and Bruton ( 1988). The latter records
are classified as either new records as a result of greater collection effort (CE) or as probable range
extensions (RE).

316

DE MOOR AND BRUTON: ALIEN AND TRANSLOCATED AQUATIC ANIMALS IN SOUTHERN AERICA

The two recently-recorded alien species (Procambams clarkii (Girard 1852) and
Hypophthalmichthys molitrix (Valenciennes 1844)) were previously present in southern Africa but
kept in captivity. Procambams clarkii was illegally imported through the aquarium trade, reports
having been received of its presence in various retail outlets (Anon, 1987). It was first recorded from
the Crocodile River (Nkomati system) in 1993 (Schoonbee, 1993). Hypophthalmichthys molitrix was
widely distributed in aquaculture ponds but has not been previously recorded from natural waters.

The new record of a translocated indigenous species, Chetia flaviventris Trewavas 1961 (the
canary kurper) (Table 5), is probably the result of an unrecorded translocation. Its presence has recently
been noted in the B3 catchment (Fig. 1) (Hoffman, pers. comm.).

Of the new distribution records of previously-recorded alien species (Table 5), the apparent range
extensions of ten species are probably the result of greater collection effort. In the case of five species
{Bothriocephalus acheilognathi Yamaguthi 1934, Barbus aeneiis (Burchell 1822), Cyprinus carpio,
Labeo capensis (Smith 1841) and L. umbratiis (Smith 1841 )) the records probably represent true
range extensions (Table 5). Cyprinus carpio has recently been recorded for the first time in the Mkuze
system (Albany Museum records) in the W2 catchment. Its range within the W 1 catchment has also
been extended. Previously there was a single record of this species in the Phongolo floodplain at Tete
Pan (de Moor and Bruton, 1988). It has now been recorded at numerous localities in this region
(Merron, pers. comm.). Three other species (B. aeneiis, L. capensis and L. umbratiis) were recently
recorded in the Kilbum Dam (Tugela system), the outlet of the Vaal-Tugela Interbasin transfer (IBT)
scheme (Coke, pers. comm.).

The new distribution records of B. acheilognathi and C. carpio are cause for concern. The spread
of the former species, a fish parasite, is highly undesirable and its extension into four major catchments
less than 20 years after its initial introduction into southern Africa (Brandt et ah, 1981 ) indicates that
it is highly invasive. Cyprinus carpio has been in southern Africa for over 200 years (Bruton and van
As, 1986) and has invaded most major catchments. The lowland region of northern Zululand (W1
and W2 catchments) was one of the few areas not yet invaded by this species. Carp has now extended
its range into both of these catchments. Its presence in the upper Mkuze system (W2) is of particular
concern, since it raises the possibility that this species may invade the Mkuze Swamps region which
is a relatively pristine area (river conservation map associated with O’Keeffe, 1986).

Degree of invasion of catchments

The population size of alien species in comparison with that of indigenous species is probably
the best estimate of the degree of invasion. The presence or absence of the alien species does not give
a true reflection of the degree of invasion, since a small isolated population in a single locality would
carry the same weighting as a species with a high population density throughout the catchment. Since
information on population densities of alien species is not readily available for all catchments in the
study area, discussion will be confined to certain trends.

The total number of alien species in sub-catchments and small major catchments (in sequence
from the highest to the lowest number) and the areas of the catchments are listed in Table 6. Based
on these figures, catchments were categorised into four classes according to the number of alien
species present (Fig. 2). Although Table 6 indicates that certain regions, such as the Okavango, are
free of alien aquatic animals, this must be seen in the context of this study (which excludes successful
biocontrol agents and threatened species translocated to sub-catchments close to their native range).
Certain biocontrol agents (i.e. two species of aquatic weevils, Cyrtobagous salviniae Calder & Sands
1985 and C. singularis (Mustache 1929)) have established populations in the Okavango (Schlettwein,

317

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 6 29 APRIL 1996

Table 6. Total number of alien species present in smaller catchments and in all sub-catchments
(listed in sequence from the highest to the lowest number). Where figures are available*, the mean
annual runoff (MAR), area of catchment, and MAR/unit area are also listed.

Catdiment

MAR p.a.
(I0'’x m’)

Area

(kmh

MAR/ unit area
(10'‘xmV100kmh

Total no. alien
species

U - Mgeni

3174

18 267

17.38

18

G - Berg - W. Cape

2158

25 415

8.49

15

R - Buffalo

601

7 984

7.53

14

S - Kei

1064

20 535

5.18

12

T3 - Natal S. coast

1932

8 876

21.77

1 1

A1 - Upper Limpopo

916

43 244

2.12

1 1

C2 - Vaal - middle

1494

69 504

2.15

11

V - Tugela

4589

28 966

15.84

10

B2 - Olifants (Limpopo) - Elands

409

20 939

1.95

10

H - Bree

1954

15 658

12.48

10

X2 - Nkomati - Crocodile

1238

10 455

1 1.84

10

Bl - Olifants (Limpopo)- upper

1438

2 1 354

6.73

9

E - Olifants (W. Cape)

1015

48 880

2.08

9

K - S. Coastal

1372

7 227

18.98

9

P - Bushmans/Kariega

102

5 569

1.83

9

M - Swartkops

129

2 536

5.09

8

A3 - Limpopo (Levuvhu)

1374

66 360

2.07

7

B3 - Olifants (Limpopo) - Letaba

1344

31 241

4.30

7

Q - Fish

529

30 245

1.75

7

T2 - Transkei

4008

23 522

17.04

7

W2 - Zululand

2349

27 912

8.42

7

D2 - Orange - Caledon & NE Cape

6465

64 931

9.96

6

J - Gouritz

539

45 028

1.20

6

C3 - Vaal - lower

705

88 234

0.80

5

L - Gamtoos

504

34 790

1.45

5

N - Sundays

200

21 056

0.95

5

Tl - Transkei - southern

2018

11 497

17.52

5

XI - Nkomati - Sabie

883

9 423

9.37

5

Z - Namibia - Swakop/Kuiseb

-

-

-

5

W1 - Swaziland - Usuthu, Phongolo

3545

32 360

10.95

4

X3 - Nkomati - Komati

1443

11 279

12.79

5

Cl - Vaal - upper

2288

38 609

5.93

3

D3 - Orange - middle

585

107 393

0.54

3

D5 - Orange - N. arid tributaries

202

111 111

0.18

3

D6 - Orange - lower

-

-

-

3

Y2 - Namibia - Omatako

-

-

-

3

D 1 - Orange - upper

-

-

-

2

D4 - Orange - s. tributaries

268

93 690

0.29

2

A2 - Limpopo - Botswana tributaries

-

-

-

1

F - West coast (W. Cape)

1 10

28 629

0.38

0

Y I - Okavango

-

-

-

0

* Department of Water Affairs statistics (Anon, 1986)

318

DE MOOR AND BRUTON: ALIEN AND TRANSLOCATED AQUATIC ANIMALS IN SOUTHERN AFRICA

Fig. 2: Map of catchment.s in the study area illustrating the total numbers of alien species in each catchment or sub-catchment.

1985; de Moor and Bruton, 1988).

Regions with the highest number of alien species are associated with the major urban centres of
the Pretoria-Witwatersrand region (catchments Al, C2), Cape Town (catchment G), Durban
(catchment U ) and East London ( R ) (Fig. 2). Low numbers of alien species are associated with sparsely
populated regions in the high-altitude upper catchments of the Vaal and Orange (Cl and D1 ) or with
the sparsely populated, arid regions (middle and lower Orange, West Cape coast, Botswana and
Namibia). There are two noticeable exceptions to this trend. The Kei catchment (S) and the Natal
south coast catchment (T3) have high numbers of alien species present (12 and 1 1 respectively) and
neither is associated with major urban centres. It could also be expected that the Port Elizabeth region
(catchment M) would have a higher number of alien species. The number of alien species in the
catchment (8) is, however, high in relation to the size of the catchment (Table 6).

To test the relationship between the number of alien species present and water availability, a
scattergram (based on data from Table 6 ) of the number of species present vs. mean annual runoff/unit
area in 35 sub-catchments was constructed (Fig. 3). CoiTelation analysis (using Spearmann’s Rank

319

ANN. CAPE PROV. MUS. (NAT, HIST.) VOL. 19, PT. 6 29 APRIL 1996

coiTelation test) indicates a positive relationship (n = 0.45) which is significant at the 0.01 level (P =
0.009; n = 35).

A number of experts in the field of invasive ecology (Elton, 1958; Fox and Fox, 1986) have noted
a general tendency for relatively pristine ecosystems with a high diversity of species to have a greater
resistance to invasion than is the case for less diverse or disturbed ecosystems. Since information on

Fig. 3: Scattergram analysis of the total ntimber of alien aquatic animal species pre.sent in individual catchments and sub-
catchments vs. mean annual runoff (MAR) per unit area (million m'/iOOkmO in 35 catchments in southern Africa.

the number of indigenous aquatic invertebrates is not readily available, analysis was confined to a
comparison of fish species present in the study area. Information on the species composition of
indigenous fish (excluding diadromous species) in 19 selected waterbodies and catchments was
obtained from a number of sources (Crass, 1966; Scott and Hamman, 1984; Skelton et al., 1985;
Skelton, 1988, 1993; Bruton, 1993; Skelton, pers. comm.) and compared to the number of alien fish
species present in the catchments concerned (Table 7). Results indicate that in eight of these
catchments/waterbodies alien species represented over 50% of the total number offish species present
(Table 7). The Kei catchment, in which only two indigenous non-diadromous fish species have been
found (Skelton, pers. comm.), had the greatest proportion of alien fish (84.6% of the total number of
species) (Table 7).

320

DE MOOR AND BRUTON: ALIEN AND TRANSLOCATED AQUATIC ANIMALS IN SOUTHERN AFRICA

Examination of Table 7 suggests a trend in which catchments or waterbodies with a high diversity
of indigenous species (e.g. the Okavango region, the Phongolo system, Mkuze and Lake Sibaya) have
fewer alien species than those v/ith a low diversity of indigenous species (e.g. Kei, Great Fish, Sundays
and Berg). In this context it should be noted that the two alien species recorded in the Phongolo system
{ Oncorhynchus mykiss and Cyprinus carpio) are relatively rare in the system. The former species is
only found in a few small tributaries in the upper catchment and C. carpio has only been recorded in
some isolated localities (Pongolapoort impoundment and in some regions of the Phongolo floodplain)
where the population size is reported to be low (de Moor and Bruton, 1988; Merron, pers. comm.).
Carp has, however, recently increased its range in this region (Merron, pers. comm.).

Correlation analysis of the number of alien fish species vs. number of indigenous fish species
within catchments indicates no significant relationship between these two variables (n = 0.28; P =
0.23; n = 19). These results are based on total numbers of species within major catchments, since
data on species diversity of sub-catchments were not available. Should this information become

Table 7. Comparison of the number of indigenous and alien fish species in selected catchments
and waterbodies.

River

No. alien

No. indigenous

Alien fish species

system

fish

species

fish species*

as % of total
fish fauna

A. & B. Limpopo

12

48

20.0%

(inch Olifants)

C/D. Orange/V aal

13

15

46.4%

X. Incomati

6

49

10.9%

Phongolo

2

43

4.4%

Mkuze system

2

25

7.4%

Lake Sibaya

0

19

0.0%

U. Umgeni

13

8

61.9%

V. Tugela

10

14

41.7%

R. Buffalo/

11

5

68.8%

Keiskamma

S. Kei

11

2

84.6%

Q. Great Fish

6

4

60.0%

N. Sundays

3

4

42.9%

L. Gamtoos

5

6

45.5%

J. Gouritz

6

6

50.0%

H. Bree

9

4

69.2%

Berg

9

4

69.2%

E. Olifants

8

10

44.4%

P. Bushmans/

9

6

60.0%

Kariega/Kowie

Okavango Delta

0

80

0.0%

* This figure excludes diadromous species

321

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 6 29 APRIL 1996

available, it would be possible to refine the resolution of the analysis. An increase in the number of
samples would also improve the statistical validity of the correlation test and might produce a more
significant result.

Many factors influence the degree of invasion of catchments, such as the opportunity of species
to invade, niche space and habitat availability in the target environment (Stauffer, 1984). The question
of suscepibility of catchments to invasion is therefore extremely complex. Further analysis of this
topic would require a multivariate analysis of the factors which influence the success or failure of
invasions. This is beyond the scope of the present study.

Invasive species

The number of major catchments invaded by alien and translocated indigenous species is
summarised in Tables 8 and 9 respectively. The number of sub-catchments where certain species are
reported to have a widespread distribution (Tables 8 and 9) as well as the number of catchments
within the study area which are beyond the native range of translocated indigenous species (Table 9)
have also been noted. The latter was calculated by subtracting the number of catchments in the native
range (after Skelton, 1993) from the total number of catchments in the study area.

The species listed as widespread in certain catchments (C carpio, Microptenis salmoides, M.
dolomieu Lacepede 1 802, Lymnaea columella Say 1817,0. mykiss, Lepomis macrochiriis Rafinesque
1819, Physa acuta Drapamaud 1805, Tilapia sparrmanii Smith 1840 and O. mossambicus) can all
be regarded as “invasive” species, with the possible exception of O. mykiss which is regularly stocked
in many catchments (Tables 8 and 9). Information on the abundance of species is incomplete and
species have only been recorded as being “widespread” when sufficient information is available.
Certain species, such as three of the fish parasites (Trichodina acuta Lorn 1961, Argulus japonicus
Thiele 1900 and Ichthyophthirius multifdis Fouquet 1876), are likely to have a far wider distribution
than those recorded in Tables 1 - 4. Other species (e.g. P. clarkii, B. acheilognathi and Clarias
gariepinus (Burchell 1822)) have been introduced into new catchments within the past 20 years and
have not yet had the opportunity to invade many catchments but are likely to be invasive species.

Among the fishes, the most widespread species include members of many reproductive guild
categories, i.e. non-guarders (e.g. C. carpio), guarders (e.g. T. sparrmanii, L. macrochiriis and M.
salmoides) and external bearers (O. mossambicus). This supports Bruton’s (1986) observation that
it is not only the altricial species which become invasive. A closer analysis of distribution records of
“widespread” species (Tables 1 - 4) indicates that C. carpio (a more “altricial” species) and M.
salmoides (a more “precocial” species) do not conform to the expected distribution patterns (discussed
in the introductory section) proposed by Bruton (1986). Both of these species are widespread in a
number of relatively species-poor catchments. In some cases (such as in catchments G, U, Y2 and
Z) both species are widespread within the same catchments. In other cases, there are differences in
the distribution patterns but these can usually be ascribed to differences in habitat requirements. For
example, C. carpio is widespread in certain turbid rivers such as the Vaal/Orange and the Fish River
(catchments C, D and Q) whereas M. salmoides is rare in these systems (Tables 1 and 3). M. salmoides
is widespread in the Olifants system (catchment E) where C. carpio has not been recorded. All of
these rivers are relatively species-poor with respect to fish. On the other hand, neither carp nor bass
have been successful in invading certain relatively species-rich catchments such as the Phongolo
(Wl) and northern Zululand (W2) (Tables 1 and 3) or in the more diverse lowveld regions of the
Incomati (X) and Limpopo systems (lower A3) (van der Waal, pers. comm.). M. salmoides has,
however, been more successful than C. carpio in invading the upper Incomati system (Table 3), which

322

DE MOOR AND BRUTON: ALIEN AND TRANSLOCATED AQUATIC ANIMALS IN SOUTHERN AFRICA

Table 8. The degree of invasion of alien species in different catchments and sub-catchments.

Species

Total catchment number

No. catchments
invaded

s-c m-c

41 24

No. sub-catchments
in which the
species is
widespread

Cyprinus carpio

32

22

12

Micropterus sahnoides

32

21

9

Lymnaea columella

24

16

9

Oncorhynchus mykiss

23

16

3

Lepomis macrochirus

19

16

6

Micropterus dolomieu

17

15

5

Salmo trutta

13

11

-

Micropterus punctulatus

8

8

-

Physa acuta

8

7

5

Carassius auratus

8

7

-

Ctenopharyngodon idellus

6

5

-

Ichthyophthirius rnultifilis

6

5

-

Gambusia ajfmis

5

5

-

Poecilia reticulata

5

5

-

Bothriocephalus acheilognathi

5

4

-

Trichodina acuta

6

4

-

Perea fluviatilis

4

4

-

Argulus japonicus

4

3

-

Tinea tinea

3

3

-

Xiphophorus helleri

3

3

-

Trachemys scripta

2

2

-

Hypophthalmichthys moUtrix

]

1

-

Craspedacusta sowerbyi

1

1

-

Procambarus clarkii

1

1

-

s-c = sub-catchment
m-c = major catchment

= information not available

lends some support to Bruton’ s ( 1 986) hypothesis. In order to further test this hypothesis, more records
of the distribution and abundance of all the species mentioned (i.e. O. mykiss, C. carpio, C. gariephms,
T. sparrmanii, M. sahnoides, L. macrochirus, O. mossambicus and P. reticulata) would have to be
obtained and analysed.

323

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 6 29 APRIL 1996

Table 9. The degree of invasion of translocated indigenous species. The number of catchments
which are beyond the native range of each species is also given.

Species

No. catchments
beyond native
range*

No.

catchments

invaded

No. catch-
ments where
widespread

Tilapia sparnnanii

17

15

5

Oreochromis mossambicus

11

9

4

Clarias gariepimis

16

5

-

Barbus aeneus

22

5

-

Laheo capeusis

22

2

-

Serranochromis r. jallae

23

2

-

Tilapia r. swierstrae

19

2

-

Sandelia capeusis

18

2

-

Barbus anopius

5

2

-

Laheo umbratus

17

2

-

Chetia fiaviveutris

23

1

-

Oreochromis macrochi r

23

1

-

* Information on native range obtained from Skelton (1993)
- = information not available

Future introductions

Interbasin transfers (IBTs)

There are at present seven completed interbasin transfer schemes in southern Africa (Pettijean
and Davies, 1988), and five of these cross major catchment boundaries. A further six such schemes
are either proposed or under construction in southern Africa (Pettijean and Davies, 1988) (Appendix
5). Although many hazards are encountered in most IBT’s, certain species are able to survive such
hazards. For example, in order to survive the journey from Teebus (Orange River) to the Grassridge
Dam (Fish River), fish (or their eggs) have to pass through grids of 6.5 cm width, baffles and pepper-
pot valves, as well as high pressures within the take-off chamber (Pettijean and Davies, 1988). Despite
this, seven species of Orange River fish survived the passage through the Orange-Fish tunnel and
have been recorded in the Fish River. Of these, Laheo capeusis and Clarias gariepinus, which were
previously not present in the Fish River, have established breeding populations (Cambray and Jubb,
1 977 a and b; Laurenson and Hocutt, 1986). This case illustrates the likelihood of future translocations
(via IBT’s) into a number of major catchments (see Appendix 5).

Following the completion of the Vaal-Tugela scheme, three Vaal River species {Barbus aeneiis,
Laheo capeusis and L. iimbratiis) were recorded in Kilburn Dam (Tugela system) (Coke, pers. comm.).
The two former species appear to have established populations in the recipient river system (Coke,
pers. comm.).

Skelton and Merron (1984) have completed a survey of the ichthyofauna in the donor (Okavango)
and recipient rivers (Swakop and Omatako) of the Eastern National Water Camer scheme (Appendix
5) which is still under construction in Namibia. Sixty-two species were recorded in the Okavango

324

DE MOOR AND BRUTON: ALIEN AND TRANSLOCATED AQUATIC ANIMALS IN SOUTHERN AFRICA

River whereas only five species were recorded in the von Bach Dam (Swakop system) and Omatako
Dam (Omatako River). It was considered that most of the Okavango River species would be unlikely
to survive in the recipient system. There were, however, twenty species with broader ecological
tolerances which they considered might not survive well in the canals and pipelines but could populate
the various systems along the route of the carrier.

Possible future introductions into the Okavango region

The widespread presence of three highly invasive alien species (C. carpio, M. salmoides and O.
mossambicus) in small farm impoundments in the arid Omatako-Omarumba region (catchment Y 1 )
in Namibia (Schrader, 1985; de Moor and Bruton, 1988), raises the possibility that these species may
find their way into the Okavango system in the future. During normal flow rates the water bodies in
the Omatako region are not connected to the Okavango system. In the event of rare flash floods these
rivers flow into the Okavango River providing the opportunity for these species to reach the Okavango
Swamps (Schrader, 1985). Water temperatures in the Okavango region range from approximately 1 8
- 29°C (Reavell, 1980). This is within the tolerance range of the three species mentioned above (de
Moor and Bruton, 1988) so it is probable that they will be able to survive in the region. The possible
negative impacts of invasions by these species into the Okavango Swamps area are described below.

Cyprimis carpio. Some workers have noted that this species uproots submerged vegetation
(Crivelli, 1983) and disturbs bottom sediments. This has the overall effect of increasing the turbidity
in the water. The altered environment then becomes less suitable for indigenous species (McCrimmon,
1968; Welcomme, 1984). There is, however, some dispute about the role of carp in causing these
effects. Fletcher et al. (1985) found that there was no association between high carp densities and
turbidity. They did however conclude that the presence of carp could cause a reduction in shallow-
rooted, soft-leaved aquatic vegetation. Should carp succeed in establishing breeding populations in
the Okavango, it could cause widespread ecological problems associated with the disturbance of
bottom sediments.

Oreochromis mo.ssambicus. This species could hybridise with indigenous cichlid species such
as O. andersonii and O. inacrochi r (Bou\enger 1912) in the Okavango region (Skelton, pers. comm.).
Such genetic contamination could decrease the biological fitness of indigenous fish stocks
(Welcomme, 1988).

Microptenis salmoides. This is a predatory species that has had major negative impacts in isolated
river systems in the southern Cape, which have a high degree of endemism. Since a number of
voracious indigenous predators (such as the tigerfish, Hydrocynus vittatus Castelnau 1861 and the
African pike, Hepsetiis odoe (Bloch 1794)) occur in the Okavango (Skelton et al., 1985), indigenous
species may not be so vulnerable to predation as was the case in the Cape rivers, where no large
specialist indigenous predatory species occurred prior to the introduction of trout and bass.

It is difficult to predict with certainty whether the three alien species mentioned above would be
successful in establishing populations in the Okavango which is a diverse ecosystem with
approximately 80 species of fish (Skelton et al., 1985). Given the evidence that carp and bass have
not succeeded in invading species-rich catchments in the eastern Transvaal lowveld, it seems unlikely
that these species will succeed in the species-rich Okavango system so long as the condition of this
system remains in a relatively pristine state. O. mossambicus, a “tropical” species with a precocial
life-history style, seems more likely to succeed in invading the Okavango.

The Okavango system, although still relatively pristine compared to many rivers in southern
Africa, has been subjected to some major disturbances such as water abstraction, poor catchment

325

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 6 29 APRIL 1996

management, insecticide spraying, and channelisation (Skelton et al., 1985; Merron and Bruton,
1990). Fox and Fox (1986) found a direct correlation between the degree of disturbance and the
susceptibility of a biotope to invasion by alien species. An increase in the levels of disturbance is
therefore likely to increase the vulnerability of the Okavango region to invasion. For this reason, it
would be unwise to adopt a complacent attitude towards the potential of alien species to invade the
system.

Conclusion

Invasive aquatic animals disrupt ecological processes, threaten the genetic integrity of indigenous
species, and alienate aquatic habitats. They therefore constitute a severe threat to aquatic ecosystems
and need to be monitored and controlled. Distribution records of alien species are not complete, but
these do indicate that the problem has reached alarming proportions. The threat posed by invasive
species needs to receive as much attention in South Africa as water pollution and habitat deterioration
because it contributes to both and causes additional negative repercussions. The importation and
translocation of invasive species needs to be strictly controlled and quantitative studies need to be
carried out on the impacts of widespread invasive species. It is recommended that special measures
be taken to minimise the possibility of the introduction of alien species into certain waterbodies of
high conservation status (the Okavango, Mkuze Swamps and Lake Sibaya) which have not as yet
been invaded by alien fish. It is noted that these waterbodies are in tropical or sub-tropical regions
where water temperatures are likely to be within the tolerance range of many species associated with
the aquarium trade.

ACKNOWLEDGEMENTS

The authors would like to thank Dr P.H. Skelton, Dr J.A. Cambray, Mrs P. Black and Mrs J.
Holiday for allowing them access to comprehensive museum distribution records; Dr Skelton for
providing information on native ranges and fish species assemblages; Mrs S. Abraham and Mr O.
West for their assistance in cartographic work; Mrs S. Radloff for her advice on the statistical analysis;
and Mr J. Alletson, Mr M. Coke, Mr R. Crass, Mr J. Engelbrecht, Dr G. Merron, Mr A. Smith, Mr
S. Thome, Dr B. van der Waal and numerous other workers in nature conservation departments for
their help in compiling the distribution records.

REFERENCES

Alletson, D.J. 1985. Observations on some piscivorous birds in a trout fishing area of Natal. Lainmergeyer 35: 41-46.
Anon 1944. Cape Provincial Administration. Inland Fisheries Department Report No. 1 : 1-5.

Anon 1958. Large carp caught in Liesbeeck Lake. Piscator 43: 80.

Anon 1959. Importation of carp. Piscator 46: 47.

Anon 1960. Survey of dams in the northern Cape. CPA Department of Nature Conservation Report No. 17: 55-56.

Anon 1962/1963. From the Cape Agricultural Journal, July 26, 1894: acclimation of trout, Piscator 56: 109-1 10.

Anon 1974. A strange find. Fauna and Flora 25: 17.

Anon 1975/1976. Large tench found dead at Paardevlei, Somerset West. Piscator 95: 141.

Anon 1 982. The Olifants River system - unicpte habitat for rare Cape fishes. Cape Conservation Series.

Anon 1986. Management of the water resources of the Republic of South Africa. Compiled, edited and published by the
Department of Water Affairs, Cape Town.

326

DE MOOR AND BRUTON: ALIEN AND TRANSLOCATED AQUATIC ANIMALS IN SOUTHERN AFRICA

Anon 1987. Wildlife Newsletter. Exotic freshwater crayfish in the Cape Province. African Wildlife 41(2): 89.

Appleton, C.C. 1974. Checklist of the flora and fauna of the Gladdespruit, Nelspruit district. Eastern Transvaal. Limnological
Society of Southern Africa Newsletter 22: 49-58.

Appleton, C.C. 1977. The exotic freshwater snail Helisoma duryi (Weatherby 1879) (Planorbidae) in southern Africa.
Zoologische Mededeelingen 52: 125-135.

Balon, E.K. 1974. Domestication of the carp Cyprinus carpio L. Miscellaneous Publications of the Royal Ontario Museum
of Life Sciences: 1-37.

Barrow, I. 1971. Freshwater fish. In: The Swartkops estuary: an ecological sun’ey. Port Elizabeth: The Zwartkops Trust, pp.
33-34.

Basson, L., van As, J.G. and Paperna, I. 1983. Trichodinid ecotoparasites of cichlid and cyprinid fishes in South Africa and
Israel. Systematic parasitology 5: 245-257.

Batchelor, G.R. 1974. An ecological investigation of the Doorndraai Dam, Sterk River, Transvaal, with .special reference
to fish management. MSc. dissertation, Rand Afrikaans University, Johannesburg,

Batchelor, G.R. 1978. Aspects of the biology of Tilapia rendalli in the Doorndraai Dam, Transvaal. Journal of the
Limnological Society of Southern Africa 4(1): 65-68.

Begg, G.W. 1976. Some notes on the Sandvlei fish fauna, Muizenberg, Cape. Pi.scator 96: 5-14.

Boomker, J., Huchzermeyer, F.W. and Naude, T.W. 1980. Bothriocephalus in the common carp in the eastern Transvaal.
Journal of the South African Veterinaty Association 51: 263-264.

Bourquin, O. 1985. Conservation aspects related to the development of aquaculture in South Africa. In: Hecht, T, Bruton,
M.N. and Safriel, O. eds. Aquaculture South Africa. Occasional Report Series No. 1 . Foundation for Research
Development. Pretoria: CSIR, pp. 60-63.

Bourquin, O. and Mathias, I. 1984. The vertebrates of Oribi Gorge Nature Reserve: I . Lammergeyer 33: 35-44.

Bourquin. O., Pike, T., Johnson, D., Rowe-Rowe, D. and Appleton, C.C. 1984. Alien animal species. Internal report of the
Natal Parks, Game and Fish Preservation Board, Pietermaritzburg.

Bourquin, O. and Sowler, S.G. 1 980. The vertebrates of the Vernon Crookes Nature Reserve: 1 . Lammergeyer 28: 20-32.

Bourquin, O. and van Rensberg, J. 1984. Vertebrates of Vernon Crookes Nature Reserve: additional and confirming records.
Lammergeyer 34: 59-6 1 .

Braack, H.H. 1981. Lower vertebrates of the Bontebok National Park, Koedoe 24: 67-77.

Brandt, T. de W., van As, J.G. Schoonbee, H.J. and Hamilton- Attwell, V.L. 1981. The occurrence and treatment of
Bothriocephalosis in the common carp, Cyprinus carpio. in fish ponds with notes on its presence in the largemouth
yellowfish Barbus kimberleyensis from the Vaal Dam, Transvaal. Water SA 7( I ): 35-42.

Brown, D.S. 1967. A review of the freshwater Mollusca of Natal and their distribution. Annals of the Natal Museum 18: 447-
494.

Brown, D.S. 1978. Freshwater molluscs. In: Werger, M.J.A. and van Bruggen, A.C. eds. Biogeography and ecology of .southern
Africa. The Hague: W. Junk, pp. 1 153-1 180.

Brown, D.S. 1980. Freshwater snails of Africa and their medical importance. London: Taylor and Francis.

Bruton, M.N. 1986, Life history styles of invasive fish in southern Africa. In Macdonald, I.A.W., Kruger, F.J. and Ferrar,
A. A. eds. The ecology and management of biological invasions in southern Africa. Cape Town: Oxford University
Press, pp. 201-208.

Bruton, M.N. 1993, First record of Ambassis productus from endorheic Lake Sibaya. Lammergeyer A2: 18-20.

Bruton, M.N. and Appleton, C,C, 1975, Survey of the Mgobezeleni lake-system in Zululand, with a note on the effect of a
bridge on the mangrove swamp. Transactions of the Royal Society of South Africa 41: 282-294,

Bruton, M,N, and Kok, H,M, 1980, The freshwater fishes of Maputaland, In: Bruton, M,N. and Cooper, K.H. eds, Studies
on the ecology of Maputaland. Grahamstown: Rhodes University and Durban: Natal Branch of the Wildlife Society
of Southern Africa, pp. 210- 244.

Bruton, M.N. and van As, J,G, 1986, Faunal invasions of aquatic ecosystems in southern Africa, with suggestions for their

327

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 6 29 APRIL 1996

management. In: Macdonald, I.A.W., Kruger, F.J. and Ferrar, A. A. eds. The ecology and management of biological
invasions in southern Africa. Cape Town: Oxford University Press, pp. 47-61 .

Cambray, J.A., Hahndsek, S. and Hahndiek, Q. 1977. A survey of fishes in Grassridge Dame. Research Report of the
Department of Nature and Environmental Consen’ation. Cape Provincial Administration: 107-124.

Cambray, J.A. and Jubb, R.A. 1977a. The Orange-Fish tunnel. Piscator99: 4-6.

Cambray, J.A. and Jubb, R.A. 1977b. Disperal of fishes via the Orange-Fish tunnel. South Africa. Journal of the Limnological
Society of Southern Africa 3( 1 ): 33-35.

Cambray, J.A. and Meyer, K. 1988. Early ontogeny of an endangered relict, cold-water cyprinid from Lesotho, Oreodaimon
quathlambe (Barnard, 1938). Revue d'Hydrobiologie Tropicale 21(4): 309-333.

Cambray, J.A. and Stuart, C.T. 1985. Aspects of the biology of a rare redfin minnow, Barbus burchelli (Pisces: Cyprinidae),
from South Africa. South African Journal of Zoology 20( 3): 1 55- 1 65.

CoETZEE, P.W. 1977. The history and distribution of freshwater fish in Albany: smallmouth yellowfish. Albany Freshwater
Angling Association Newsletter 36: 3-4.

CoNANT, R. 1958. A field guide to reptiles and amphibians of the United States and Canada east of the 100th Meridian.
Cambridge: The Riverside Press.

Crass, R.S. 1964. Freshwater fishes of Natal. Pietermaritzburg: Schuter and Shooter.

Crass, R.S. 1 966. Features of freshwater fish distribution in Natal and a discussion on controlling factors. Limnological Society
of Southern Africa Newsletter 7: 3 1 -35.

Crass, R.S. 1 969a. The effects of landuse on freshwater fish in South Africa, with particular reference to Natal. Hydrobiologia
34(1): 38-56.

Crass, R.S. 1969b. Trout in the Bushmans River. Limnological Society of Southern Africa Newsletter 12: 17-20.

Crivelli, A.J. 1983. The destruction of aquatic vegetation by carp. Hydrobiologia 106: 37-41.

Day, L.A. 1932. The introduction of Salmonidae into Natal rivers, with special reference to Sahno irideus. South African
Journal of Science 29: 473-479.

De Moor, I.J. and Bruton, M.N. 1 988. Atlas of alien and translocated indigenous aquatic animals in southern Africa. South
African National Scientific Programmes Report No. 144. Pretoria: CSIR.

Dixon, J.E.W. and Blom, M.J. 1974. Some aquatic vertebrates from the Namib Desert, South West Africa. Madoqua 2(3):
31-32.

Donnelly, B.G. 1965. The first record of rainbow trout breeding in the Port Elizabeth area. Piscator 64: 91-92.

Du Plessis, S.S. 1961. Trout pioneer F.C. Braun ( 1875-1959). foinw ant/ F/ora 12: 17-23.

Du Plessis, S.S. and le Roux, P.J. 1965. Sport fisheries in river development with references to the Orange River scheme.
South African Journal of Science 61(3): 137-146.

Elton, C.S. 1958. The ecology of invasions by animals and plants. London: Methuen Books.

Fletcher, A.R. , Morrison, A.K. and Hume, D. J. 1 985 . Effect of carp, Cyprinus carpio L., on communities of aquatic vegetation
and turbidity of waterbodies in the lower Goulbum River Basin. Australian Journal of Marine and Freshwater
Research 36: 3 1 1 -321 .

Fogarty, A. 1980. New fish species in eastern Cape waters. The Eastern Cape Naturalist 69: 24.

Fox, M.D. and Fox, B.J. 1986. The susceptibility of natural communities to invasion. In: Groves, R.H. and Burdon, J.J. eds.
Ecology of biological Invasions: An Australian perspective. Canberra: Australian Academy of Science and
Cambridge: Cambridge University Press, pp. 57-66.

Gaigher, C.M. 1981. Tiny jewels of the Cedarberg. African Wildlife 35(3): 25-29.

Gaigher, I.G. 1975a. The ecology of a minnow, Barbus trevelyani (Pisces: Cyprinidae) in the Tyume River, Eastern Cape.
Annals of the Cape Provincial Museums 11 (1): 1-19.

Gaigher, I.G. 1975b. The Hardap Dam: an angler’s paradise in South West Africa. Piscator 93: 4-8.

Gaigher, I.G., Hamman, K.C.D. and Thorne, S.C. 1 980. The distribution, conservation status and factors affecting the survival
of indigenous freshwater fishes in the Cape Province. Koedoe 23: 57- 88.

328

DE MOOR AND BRUTON: ALIEN AND TRANSLOCATED AQUATIC ANIMALS IN SOUTHERN AFRICA

George, T.T. 1976. Introduction and transplantation of cultivable species into Africa. Reviews ami Experience Papers. CIFA
Technical Papers No. 4, Supplement I: 408-432.

Gephard, S.R. 1977. Status of trout populations of Lesotho. Pi.'icator 100; 71-74.

Gilmore, K.S. 1978. A fisheiy survey of the Shashi Dam. Unpublished report of the Department of Agriculture and Fisheries.
Botswana: Ministry of Agriculture.

Goldner, H.J. 1967. 'n Popukisiestudie van varswatervisse in Barherspan, wes-Transvaal. MSc. thesis, Potchefstroom
University for CHO, Potchefstroom.

Griffiths, R.W., Schloesser, D.W., Leach, J.H. and Kovalak, W.P. 1991. Distribution and dispersal of the zebra mussel
{Dreissemi pohmorpha) in the Great Lakes Region. Canadian Journal of Fisheries and Aquatic Science 48: 1381-
1388.

Hamilton-Attwell, V.L., de Kock. K.N. and van Eeden, J.A. 1970. The occurrence and distribution of Physa acuta
Drapernaiid in the Republic of South Africa. Scientific contributions of Potchefstroom University for CHO. Series
B: Natural Sciences 26: 1-11.

Hamman. K.C.D. 1980. Post-impoundment trends in the fish populations of the Hendrik Verwoerd Dam, South Africa. Journal
of the Limnological Society' of Southern Africa 6(2): 101-108.

Hamman, K.C.D., Thorne, S.C. and Skelton, P.H. 1984. The discovery of the Cape kurper, Sandelia capensis (Cuview in
C & V. 1831 ) in the Olifants River System (Western Cape Province). The Naturalist 28: 24-26.

Harrison, A.C. 1 938. Fish of the rivers of the Langeberge, south- western Cape. Cape Piscatorial Society Freshwater Fisheries
Circular No. 30: 2-5.

Harrison, A.C. 1 939. Report of the inland Fisheries Advisory Officer for the year 1 938. Province of the Cape of Good Hope.

Harrison. A.C. 1948. Report on the rivers of Maclear and East Griqualand, April and May 1948. Inland Fisheries Department.
Cape Provincial Administration Report No. 5: 15-19.

Harrison, A.C. 1953. Introduction of exotic fishes to the Cape Province. Section III: Black bass and bluegills (continued).
Piscatorli: 12-15.

Harrison, A.C. 1954. Trout pioneering in the Eastern Province. No. 9; Drought, sea trout and farewell to Stenning. Piscator
30: 47-53.

Harrison, A.C. 1958. Sea-run trout of the Eerste River. Piscator Ai: 44-53.

Harrison, A.C. 1960/1961 . Tiger trout (Salino trutta female x Salvelinus foniinalis male). Piscator 50: 85-93.

Harrison, A.C. 1964. Steenbras reservoir: trout plantings and recoveries. Pi.scator 6(i: 29-46.

Harrison, A.C. 1964/65. Spotted bass. Piscator 62: 127-128.

Harrison, A.C. 1966. Trout in the peat-stained waters of the southern Cape. Piscator 66: 22-23.

Harrison, A.C. 1966/1967. Rainbow trout in Wemmershoek Reservoir. Piscator 6S: 126-134.

Harrison, A.C. 1972/1973. Early records of trout stocking in the vleis of the Cape Flats. Piscator ^6: 1 14.

Harrison, A.C, and Lewis, D.C. 1968/1969. Twenty-one years of trout fishing in Liesbeek. Piscator 74: 128-144.

Heard, H.W. and King, M. 1981, A report on the fish populations and ecology of the Baakens River. Internal Report, Cape
Department of Nature and Environmental Conservation.

Heard, H.W. and King, M. 1982. The fish populations and ecology of the Baakens River. The Naturalist 26(2): 21-24.

Heeg, J. 1983. Secondary production - utilization of invertebrates. In: Breen, C.M. ed.. Limnology of Lake Midinar. South
African National Scientific Programmes Report No. 78. Pretoria: CSIR. pp. 113-114.

Hey. S.A. 1926. Fisheries survey 1925-1926: inland waters. Department of Mines and Industries Report: No. 2.

Hocutt, C.H. and Skelton. P.H. 1983. Fishes of the Sak River, South Africa, with comments on the nomenclature of the
smallmouth yellowfish. Barbus aetieus (Burchell, 1822). JLB Smith institute of Ichthyology Special Publication
No. 32.

Hoffman, G.L. and Schubert, G. 1984. Some parasites of exotic fishes. In: Courtenay, W.R. and Stauffer, J.R. eds.

Distribution, biology and management of exotic fishes. Baltimore: Johns Hopkins University Press, pp. 233-261 .

Holcik, J. and Mihalik, J. 1968. Freshwater fishes. London: Hamlyn.

329

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 6 29 APRIL 1996

Jackson, P.B.N. 1982, Fish in the Buffalo River catchment system. In: Hart, R.C. ed„ Water quality in the Bujfalo River
catchment: a synthesis. Rhodes University, Grahamstown, pp. 1 19-132.

Jackson, P.B.N., Cambray, J.A., Eccles, D.H., Hamman, K.C.D., Tomasson, T. and White, P.N. 1 983. Distribution, structure
and relative abundance of fish populations. In: Allanson, B.R, and Jackson, P.B.N. eds. Limnology and fisheries
potential of Lake le Roux. South African National Scientific Programmes Report No. 77. Pretoria: CSIR, pp. 77-
107.

Jacobs, K. 1971. Livehearing aquarium fishes. London: Studio Vista.

Jayaram, K.C. 1981. The freshwater fishes of India. A handbook. Calcutta: Zoological Survey of India.

JouBERT, P. 1984. Daar’s baars in die Vaal. Stywe Lyne January: 5-7.

JuBB. R.A. 1959. Carp, mud-mullet and eels in Eastern Cape Rivers. Piscator 46: 56-60.

Jlibb, R.A. 1963. Transportation of Orange/Vaal yellowfish and the proposed eel fishery. Piscator 51: 22.

JuBB. R.A. 1965. Freshwater fishes of the Cape Province. Annals of the Cape Provincial Museums 4: 1-72.

Jubb, R.A. 1966. Some notes on Labeo or mudsuckers. Piscator 66: 25- 29.

JuBB, R.A. 1966/67. Notes on the popular angling yellowfishes and distribution outside their native ranges. Piscator 6^: 120-
125.

Jubb, R.A. 1967. Freshwater fishes of southern Africa. Cape Town: Balkema.

Jubb, R.A. 1968. North End Lake, Port Elizabeth. Piscator li: 53-58.

Jubb, R.A. 1971a. Freshwater fishes. In: Potgieter, D.J., dii Plessis, P.C. and Skaife, S.H. eds. Animal life in southern Africa.
Pretoria: NASOU, pp. 256-261.

Jubb, R.A, 1971b. Some interesting Eastern Cape freshwater fishes. The Eastern Cape Naturalist 45: 7-9.

Jubb, R.A. 1972. The Hendrik Verwoerd Dam and Orange River fishes. Piscator 84: 22-26.

Jubb, R.A. 1973. Notes on exotic fishes introduced into South African inland waters. Piscator 81: 9-12.

Jubb, R.A. 1976/1977. Notes on exotic fishes introduced into South African waters. Part II: live-bearing sharptooth carps.
Piscator 98: 132-134.

Jubb, R.A. 1978. The distribution of the common caip, Cyprinus carpio. in southern Africa. Piscator 88: 63-65.

Jubb, R.A. 1979. Some endangered freshwaters fish species. The Eastern Cape Naturalist 68: 24-26.

J UNOR, E.J .R. 1 969. Tilapia melanopleura Dum, in artificial lakes and dams in Rhodesia, with special reference to its undesirable
effects. Rhodesia Agricultural Research 1: 61-69.

KiNG.M.J.andBoK, A.H. \ 98A. Report on the ichthyofauna and ecology oftheBaakens River. Internal Report, Cape Department
of Nature and Environmental Conservation.

Kleynhans, C.J. 1983. A checklist of the fish species of the Mogol and Palala Rivers (Limpopo system) of the Transvaal.
Journal of the Limnological Society of Southern Africa 5(1): 27-29.

Kleynhans, C.J. 1984. Die verspreiding en status van sekere seldsame vissoorte van die Transvaal en die ekologie van
sommige spesies. D.Sc. thesis. University of Pretoria, Pretoria.

Koch, B.S. and Schoonbee, H.J. 1980. A fish mark-recapture study in Boskop Dam, Western Transvaal. Water SA 6(3): 149-
155.

Kruger, E.J. 1971. 7; Ondersoek na die ekologiese voedselstruktuur van .sekere visspesies in Loskopdam. D.Sc. thesis,
Potchefstroom University for CHO. Potchefstroom.

Kruger, I. van As, J.G. and Saayman, J.E. 1983. Observations on the occurrence of the fish louse Argulus japonicus Thiele,
1900, in the western Transvaal. South African Journal of Zoology 18(4): 408-410.

La Hausse de Lalouviere. P. 1 986. Notes on unusual fishes from the Pongolo River system, northern Zululand. Lammergeyer
37: 50-54.

Laurenson. L.J.B. and Hocutt, C. H. 1986. Colonization theory and invasive biota: the Great Fish River, a case history.
Environmental Monitoring and Assessment 6( 1 ): 71-90.

Lee, D.S., Gilbert, C.R., Hocutt, C.H., Jenkins, R.E., McAllister, D.E. and Stauffer, J.R. 1980. Atlas of North American
freshwater fishes. North Carolina State Museum of Natural History.

330

DE MOOR AND BRUTON: ALIEN AND TRANSLOCATED AQUATIC ANIMALS IN SOUTHERN AFRICA

Marshall, R.D. 1972. A preliminary fish survey of the Caledon River system. The Civil Engineer in South Africa 14(2): 94-95

Mayekiso, M. 1986. Some aspects of the ecology of the Eastern Cape rocky Sandelia bainsii (Pisces: Anabantidae) in the
Tyume River, Eastern Cape, South Africa. M.Sc. thesis, Rhodes University, Grahamstown.

McCrimmon, H.R. 1968. Carp in Canada. Fisheries Research Board of Canada Bulletin No. 165.

McVeigh, S.J. 1984. The sunfish syndrome. Tight Lines 22 ( 12): 52- 53, 56.

Merron, G.S. and Bruton, M.N. 1990. The physiological and toxicological effects of aerial spraying with insecticides on the
fish stocks of the Okavango Delta, Botswana. JLB Smith Institute of Ichthyology, Investigational Report No. 3 1 .

Morrissy, N.M. 1978. The past and present distribution of marron, Cherax tenuimanus (Smith), in Western Australia. Fisheries
Research Bulletin, Fisheries Department, Western Australia 22: 1-38.

Mulder, P.F.S. 1986. Fish populations in the Vaal River. In: Walmsley, R.D. and Rogers, K. eds. The Vaal River Ecosystem:
status and problems. Occasional Report Series No. 5. FRD, Pretoria: CSIR, pp. 201-208.

Newbery, R. 1984. The law and the red-eared temapin. African Wildlife 38(5): 189.

Oberholzer, G. and van Eeden, J.A. 1967. The freshwater moliusca of the Kruger National Park. Koedoe 10: 1-42.

O’Keeffe, J.H.(ed.) 1986. The conservation of South African rivers. South African National Scientific Programmes Report
No. 131. Pretoria: CSIR.

Paperna, I. 1980. Parasites, infections and diseases of fish in Africa, CIFA Technical Paper No. 7.

Pettijean, M.O.G. and Davies, B.R. 1988. Ecological impact of inter- basin water transfers: some case studies, research
requirements and assessment procedures in southern Africa. South African Journal of Science 84: 819-828.

Philippart, J-C. and Ruwet, J-C. L. 1982. Ecology and distribution of tilapias. In: Pullin, R.S.V. and Lowe-McConnell, R.H.

eds. The biology and culture of tilapias. Proceedings of the International conference on the biology and culture of
tilapias, 2-5 September 1980, Italy, Manila: ICLARM, pp. 15-59.

Pienaar, U. de V. 1978. Undesirable immigrants in the Kruger National Park. Custos 7( 10), 6-7, 14-15.

Pike, T. 1980a. An historical review of freshwater fish hatcheries in Natal. Piscator 106: 49-53.

Pike, T. 1980b. Initial investigation of the grazing effects of Ctenopharyngodon idella on submerged aquatics in farm dams
in Natal. Interna! report of the Natal Parks, Game and Fish Preservation Board, Pietermaritzburg.

Pike, T. 1983. A review of freshwater fish and invertebrates which have been used for food production or angling purposes
throughout the world, and some effects caused by exotics on indigenous South African populations. Internal report
of the Natal Parks, Game and Fish Preservation Board, Pietermaritzburg.

Pike, T. 1986. Grass carp for biological control of aquatic plants in farm dams. Natal Parks Board. Research communication
No. 55. April.

Pike, T. and Tedder, A.J. 1973. Rediscovery of Oreodaimon quathlcimbae (Barnard). Lammergeyer 19: 9-15.

Plumstead, E.E., Prinsloo, J.F. and Schoonbee, H.J. 1985. A survey of the fish fauna of Transkei estuaries. Part 1: The Kei
River Estuary. South African Journal of Zoology 20(4): 213-220.

POTGIETER, F.J. 1974. 'n Ekologiese studie van die rooibors terperTilapia melanopleura Dumeril ( 1859} ( Pisces: Cichlidae)
in Transvaal met verwysing na geassossieerde varswatenissoorte. M.Sc. dissertation, Rand Afrikaans University,
Johannesburg.

Pott, R. Me C. 1981. The Treur River barb: a rare fish in good company. African Wildlife 35(6): 29-31.

Pretorius, S.J., DE Kock, K.N., JouBERT, P.H., Ryke, P.A.J. and van Eeden, J.A. 1980. Varsv/atermolluske in de
Olifantsrivierbekken. I. Die bekken stroomopwaarts vanaf die Marble Hail omgewing. Scientific Contributions of
Potchefstroom University for CHO. Series B: Natural Sciences 100: 1-50.

Pretorius, S.J., Jennings, A.C., Coertze, D.J. and van Eeden, J.A. 1975. Aspects of the freshwater Moliusca of the Pongolo
River floodplain pans. South African Journal of Science 71: 208-212.

Rayner, N.A. 1981. Studies on the zooplankton of Lake Midmar. M.Sc. thesis. University of Natal, Pietermaritzburg.

Rayner, N.A. 1988. First record of Craspedacusta sowerbyi Lankester (Cnidaria: Limnomedusae) from Africa. Hydrobiologia
162: 73-77.

Reavell, P.E. 1980. A study of the relationship between phytoplankton growth and nutrient concentrations in the waters of
the seasonally inundated Okavango delta, Botswana. M.Sc. thesis, University of the Witwatersrand, Johannesburg.

331

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 6 29 APRIL 1996

Rondorf. D.W. 1976. New locations of Oreodaimon qiiathlamhae (Barnard) (Pisces: Cyprinidae) populations. South African
Journal of Science 11(5): 150-15 1 .

Schrader, H.J. 1985. Invasive alien fishes in South West Africa/Namibia. In: Brown, C.J., Macdonald, I.A.W. and Brown,
S.E. eds. Invasive Alien orgcuiisms in South West Africa/Nainihia. South African National Scientific Programmes
Report No. 1 19. Pretoria: CSIR, pp. 35-40.

ScHOONBEE, H.J. 1 993. Occurrence of the red swamp crayfish Procamharus clarkii (Crustacea: Cambaridae) in the Crocodile
River at Dullstroom, Transvaal. Water SA 19(2): 163-166.

SCHLETTWEIN. C.H.G. 1985. The biological control of Salvinia molesta Mitchell in the Eastern Caprivi Zipfel. Report to the
Department of Water Affairs, Windhoek, SWA/Namibia.

ScHUTTE. C.H.J. and Frank, G.H. 1 964. Observations on the distribution of freshwater Mollusca and chemistry of the natural
waters in the south-eastern Transvaal and adjacent northern Swaziland. Bulletin of the World Health Organisation
30: 389-400.

Scott, H.A. and Hamman, K.C.D. 1984. Freshwater fishes of the Cape. Cape Conservation Series No. 5.

Scott, W.E, and Crossman, E.J. 1973. Freshwater fishes of Canada: Bulletin 184. Ottawa: Fisheries Research Board, Canada.

Siegfried, W.R. 1962. Introduced vertebrates in the Cape Province. Department of Nature Conseiyation, Cape Provincial
Administration. Report No. 19: 80-87.

Skead, C.J. 1973. Zoo-historical Gazetteer. Annals of the Cape Provincial Museums 10: 1-259.

Skelton, P.H. 1972. Smallmouth yellowfish (Barbus holuhi) in Settlers Dam, Grahamstown. Pl.scatorS4: 27-30.

Skelton, P.H. 1981 . Fishes of the Aughrabies Falls National Park and the lower Orange River. Research Report, Albany
Mirseum. Grahamstown.

Skelton. P.H. 1986. Fish of the Orange-Vaal system. In: Davies, B.R. and Walker, K.F. eds. The ecology of river systems.
Dordrecht: Kluwer Academic Publishers, pp. 143-161.

Skelton, P.H. 1987. South African Red Data Book. Fishes. South African National Scientific Programmes Report No. 137.
Pretoria: CSIR.

Skelton, P.H. 1988. Diversity and distribution of freshwater fishes of the eastern Cape. In: Bruton, M.N. and Gess, F.W. eds.
Towards an environmental plan for the eastern Cape. Grahamstown: Rhodes University, pp. 1 76-196.

Skelton P.H. 1 993. A complete guide to the freshwater fishes of southern Africa. Halfway Hou,se: Southern Book Publishers.

Skelton, P.H., Bruton. M.N., Merron, G.S. and van der Waal, B.C.W. 1985. The fishes of the Okavango drainage system
in Angola, South West Africa and Botswana: Taxonomy and distribution. Ichthyological Bulletin of the JLB Smith
Institute of Ichthyology 50: 1-21.

Skelton, P.H. and Merron, G.S. 1984. The fishes of the Okavango River in South West Africa with reference to the possible
impact of the Eastern National Water Carrier on fish distribution. JLB Smith Institute of Ichthyology, Investigational
Report No. 9.

Skelton, P.H. and Skead, J. 1 984, Early reports and paintings of freshwater fishes in the Cape Province. Africana Notes and
Records 26( I ): 29-34.

Smith. G.R. and Stearley, R.F. 1989. The classification and scientific names of rainbow and cutthroat trouts. Fisheries
14(1): 4-10.

Smith, W. 1983. Rietviei - pa.st and present. Tight Lines. October: 14-18.

Smith, W. 1984. Countdown for SA bass team. Tight Lines. March: 19- 23.

Soper, F.L. and Wilson, D.B. 1943. Anopheles gambiae in Brazil 1930- 1940. New York: The Rockefeller Foundation.

Stauffer, J.R. 1984. Colonization theory relative to introduced populations. In: Courtenay, W.R. and Stauffer, J.R. eds.
Distribution, biology and management of exotic fishes. Baltimore: Johns Hopkins University Press, pp. 8-21.

Sterba, G. 1962. Freshwater fishes of the world (translated and revised by D.W, Tucker). London: Vista Books.

Trewawas, E. 1 983. Tilapine fishes of the genera Sarotherodon, Oreochromis and Danakilia. London: Trustees of the British
Museum (Natural History).

Van As, J.G., Basson, L. and Theron, J. 1981. Further records of the occurrence of Bothriocephalus (Cestoda:

332

DE MOOR AND BRUTON: ALIEN AND TRANSLOCATED AQUATIC ANIMALS IN SOUTHERN AFRICA

Pseudophyllidae) in the Transvaal. South African Journal of Science 77: 343.

Van As, J.G. and Basson, L. 1984. Checklist of freshwater fish parasites from southern Africa. South African Journal of
Wildlife Research 14(2): 49-61 .

Van Bruggen, A.C. 1964. The distribution of introduced mollusc species in southern Africa. Beaufortia 11: 161-169.

Van Bruggen, A.C. 1966. Physa acuta, an introduced mollusc species in southern Africa. Journal of Conchology 26: 49-50.

Van der Merwe, C.V. 1970. A short description of the Goukamma Nature Reserve. Department of Nature Consenation.
Cape Provincial Admini.stration Investigational Report No. 16.

Van Eeden, J.A. and Brown, D.S. 1966. Colonisation of fresh waters in the Republic of South Africa by Lymnaea columella
Say (Mollusca: Gastropoda). Nature 210: 1 172-1 173.

Van Eeden, J.A., de Kock, K.N., Pretorius, S.J., Joubert, P.H. and Ryke, P.A.J. 1980. Die voorkoms en verspreiding van
varswatermolluske 'mAKSMoXozmy'iexbeCktn. Scientific contributions of Potchefstroom University for CHO. Series
B: Natural Sciences 99: 1 -27.

Van Rensburg, K.J. 1966. Die vis van die Olifantsrivier (weskus) met spesiale verwysings na die geelvis (Barbus capensis)
en saagvin (Barbus serra). Department of Nature Conservation, Cape Provincial Administration. Investigational
Report No. 1 0.

Van Schoor, D.J. 1966. Studies on the culture and acclimation of Tilapia in the Western Cape Province. Department of Nature
Consen’cition, Cape Provincial Administration Investigational Report No. 7.

Van Schoor, D.J. 1 969a. The growth of largemouth bass Micropterus salmoides at Jonkershoek Fish Hatchery without benefit
of forage fish. Department of Nature Conservation. Cape Provincial Administration Investigational Report No. 1 I .

Van Schoor, D.J. 1969b. The introduction of Serranochromis robustus jallae (Boulenger) to the Eerste River Basin, Western
Cape Province. Department of Nature Consenation. Cape Provincial Administration Investigational Report No. 1 2.

ViLJOEN, S. and van As, J.G. 1985. Sessile peritrichs (Ciliophora: Peritricha) from freshwater fish in the Transvaal, South
Africa. South African Journal of Zoology 20: 79-96.

Walmsley, R.D. 1988. Status of man on farming in Australia and South Africa. In: Walmsley, R.D. ed.. An overview of marron
farming and its potential in South Africa. Occasional Report Series No. 29. Pretoria: CSIR.

Welcomme, R.L. 1981 . Register of international transfers of inland fish species. FAO Fisheries Technical Paper No. 213: I-

120.

Welcomme, R.L. 1984. International transfers of inland fish species. In: Courtenay, W.R. and Stauffer, J.R. eds. Distribution,
biology, and management of exotic fishes. Baltimore: Johns Hopkins University Press, pp. 22-40.

Welcomme, R.L. 1988. International introductions of inland aquatic species. FAO Fisheries Technical Paper 294. FAO
Fisheries Department, Rome,

APPENDIX 1

Major catchments and sub-catchments in the study area (illustrated in Fig. 1) as well as the major
rivers, impoundments and towns in each region. DWA sub-catchments also listed where such

information is available*.
Region Major Rivers & tributaries

Major waterbodies/impoundments

Towns/places

A - Limpopo

Al

-Crocodile, Jukskei, Marico

Rietvlei Dam, Hartbeespoort Dam,

N-Johannesburg, Pretoria

(A10,A20,A30)

Roodeplaat Dam, Marico-Bosveld Dam

A2

Botswana tributaries:

Shashi Dam

Gaberone

-Lotsani, Shashi, Metsemothlaba

Gaberone Dam

A3

-Levuvhu

Nzhelele Dam, Albasina Dam,

Pietersburg, Potgietersrus

(A40,A50,A60,A70,A80,A90)

Glen Alpine Dam, Hans Strydom Dam

333

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 6 29 APRIL 1996

Appendix 1. Continued.

Region Major Rivers & tributaries

Major waterbodies/impoundments

Towns/places

B - Olifants

B1

-Blyde, Treur, Steelpoort

Bronkhorstspruit Dam,

Bronkhorstspruit

(B10,B20,B40,B60)

Blyderivierspoort Dam

B2

-Elands (B30.B50)

Rust de Winter, Loskop Dam

Marble Hall, Witbank

B3

-Letaba (B70.B80.B90)

Fanie Botha Dam, Ebeneezer Dam

C - Vaal

Cl

-Klip, Wilge (CIO, C80)

Grootdraai Dam, Sterkfontein Dam

Bethlehem, Standerton

C2

-Skoonspruit, Vet, Mooi, Vais

Allemanskraal Dam, Boskop Dam

S. Johannesburg, Klerksdorp,

(C20,C40,C60,C70)

Erfenis Dam, Vaal Dam

Parys, Potchefstroom,

C3

-Hartz, Modder, Riet

Barberspan, Bloemhof Dam, Kalk-

Bloemfontein, Kimberley

(C30,C50,C90)

fontein Dam, Spitzkop Dam

D - Orange

D1

- Malibamatso, Mokhotlong, Senqu

Lesotho highlands

D2

-Caledon, Kraai, Makeleng (D10,D20)

Maseru

D3

-Seekoei (D30,D60,D70)

PK le Roux Dam, Verwoerd Dam

Douglas, Upington

D4

-Hartbees, Sak (D50)

Rooidam

D5

-Kuruinan, Malopo (D40)

Kuruman Oog, Malopo Oog

Kuruman

D6

-Fish (D80-t-Fish R,)

Hardap Dam

Oranjemond

E

Olifants (Clanwilliam)

Bulshoek Dam, Clanwilliam Dam

Clanwilliam

- Tankwa, Dol ing, Twee, Driehoek

F

Buffalo, Swartdoring

G

Berg, Bot, Eerste, Heuningnes,

Paardevlei, Princessvlei,

Cape Town, Somerset W.,

Klein, Palmiet

Seekoeivlei

Stellenbosch

H

Breede

- Flex, Nuy, Riviersonderend

Brandviei Dam, Theewaterskloof

Riviersonderend

J

Gouritz

Gamkapoort Dam

Oudtshoorn

- Garnka, Groot, Touwsrivier

K

South coastal

Keurbooms, Kromme, Tsitsikama,

Groenvlei, Rondevlei, Swartvlei

Knysna, Plettenberg Bay

L

Gamtoos

- Baviaanskloof, Groot, Wit

Paul Sauer Dam

Patensie

M

Baakens, Swartkops,

Port Elizabeth, Uitenhage

N

Sundays

de Hoop Dam, Lake Mentz,

Kirkwood

P

Bushmans, Kariega, Kowie

Howiesonspoort Dam

Grahamstown, Port Alfred

334

DE MOOR AND BRUTON: ALIEN AND TRANSLOCATED AQUATIC ANIMALS IN SOUTHERN AFRICA

Appendix 1 . Continued.

Region Major Rivers & tributaries

Major waterbodies/impoundments

Towns/places

Q

Fish

Grassridge Darn, Lake Arthur

Cradock

R

Buffalo, Keiskamma, Nahoon

Bridle drift Dam, Maden Dam,

East London,

Rooikrans Dam

Kingwilliamstown

S

Kei

Gubu Dam, Lubisi Dam,

Stutterheim

- Kubusie, Swart Kei

Nqora Dam, Xonxa Dam,

T-

S. Natal/Transkei

TI

Mngazi to Kobongaba: Mbashe
Mngazi, Mtata (T10,T20,T70,T80)

Mtata Dam

Umtata

T2

Mzamba to Mzimvubu (T30,T60)

Mount Frere, Kokstad

Mzimvubu - Mzimhlava, Tina, Tinana

Port St. Johns

T3

Mzimkulu (T40,T50)

- Pholela, Mzimkulwana, Mtamvuna,

- Ngwangwana

Harding, Margate

U

Mgeni - Karkloof, Msindusi,

Albert Falls Dam, Midmar Dam,

Durban,

Mkomazi - Mquatsheni, Loteni

Nagle Dam

Pietermaritzburg

V

Tugela - Buffalo, Mooi

Chelmsford Dam, Spionkop Dam,
Wagendrift Dam, Woodstock Dam

Estcourt, Newcastle

w-

Zululand/Swaziland

W1

Ngwavuma, Phongolo, Usuthu

Jericho Dam, Kosi Bay,

Mbabane

(W40,W50,W60)

Pongolapoort Dam, Westoe Dam

Pongola

W2

Mfolozi, Mhlatuze, Mkuze,
(W10,W20,W30,W70)

Mkuze Swamps, Lake Sibaya

Richards Bay

X-

Nkomati

XI

Sabie (X30,X40)

Sabie

X2

Crocodile (X20)

Nelspruit

X3

Komati, Mbuluzi (XIO)

Kafferskraal, Nooitgedacht

N. Swaziland

Y-

Botswana/Namibia

Yl

Chobe, Okavango

Makgadkgadi Pans, Okavango Swamps

Maun

Y2

Omataka Omarumba

Z

Swakop, Kuiseb

von Bach Dam, Kuiseb Canyon

Windhoek, Swakopmund

- Indicates major tributaries of river indicated.

* Code for sub-catchments according to Department of Water Affairs statistics (Anon, 1986).

335

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 6 29 APRIL 1996

APPENDIX 2

Checklist of alien species introduced into southern Africa together with selected bibliography
(in numerical and alphabetical code, listed in Appendix 4).

Scientific classification

Native range

Date of

Impact'

Distribution

and common name

first

references

introduction

Ciliophora

Possibly

Uncertain

PF(116)

116,149,

Ichthyophthiriidae

Asia

(78)

major

1

Ichthyophthirius midtifilis
Fouquet 1 876
whitespot, ich

(78)

Oligohymenophorea

Uncertain

Possibly

PF(116)

14, 149

TrichocUna acuta Lorn 1961

(116)

18th C.

major

trichodina

(c)

Cnidaria

N. hemi-

late

LI

130

Limnomedusae

sphere

1970’s

(129,

Craspedacusta sowerhyi
Lankester 1880
Freshwater medusa

(129)

(130)

130)

Platyhelminthes

Asia

1975

PF(25)

18,25,148

Bothriocephalidae
Bothriocephaliis
achedognathi Yamaguthi 1934
fish tape worm

(78)

(25)

major

Arthropoda

Possibly

Prior to

PF

104,149,

Crustacea

Asia

1983

(104)

1

Argulidae
A rgidus japonicus
Thiele 1900
fish louse

(78)

(104)

major

Parastacidae

SW

c. 1982

LI

160,

Cherax tenidmanus

Australia

(23)

( 160)

r

(Smith 1912)

(112)

marron

Cambaridae

N. America

c. 1992

Possibly

134

Procambarus clarkii

(134)

(134)

HD

(Girard 1852)
red swamp crayfish

(134)

Mollusca

Probably

c. 1966

LI

27,57,128,154

Physidae

N. America

(57,151)

336

DE MOOR AND BRUTON: ALIEN AND TRANSLOCATED AQUATIC ANIMALS IN SOUTHERN AFRICA

Appendix 2. Continued.
Scientific classification
and common name

Native range

Date of
first

introduction

Impact'

Distribution

references

Physa acuta Draparnaud 1 805

26

physa snail

Planorbidae

N. America

c. 1969

LI

1 1

Helisoma diiryi Weatherby 1879

(b)

(b)

(11)

helisoma snail

Lymnaeidae

N. America

c. 1944

VPL

29,1 15,127,128,

Lymnaea columella Say 1817

(b)

(28)

27,28

133,150,153,154

lymnaea snail

major

Chordata

Western

C.1897

p

39,40,41,42,45,47,

Osteichthys

N. America

(2,43,46,

major in

50,54,62,70,87,

Salmonidae

& E. Asia

64,71,76)

some

108,123,139

Oncorhynchus mykiss

(108,145)

cases

a,e,g,h,i,j,k,l,

(Walbaum 1792)

(36,103)

m,n,o,p,x,y,z

rainbow trout

1000, 2000

Sabno trutta

Europe,

Late

P

40,41,50,54,65,66,

Linnaeus 1758

W. Asia

19th C.

(140)

67,69,75,80,81,94,

brown trout

& Morocco

(2,6,

major in

131,140,155

(161, J)

119)

some cases

h,k,t,y

(123)

1000, 2000

Cyprinidae

E. Europe,

c. 1728

IP

39,40,60,72,77,

Carassius aunitus

central

(142)

78

100,

(Linnaeus 1758)

Asia 8l

P

goldfish

China ( 163)

1000, 2000

Ctenopliaryngoclon idellus

Asia

1967

DAV,

P

(Valenciennes 1844)

(83)

(23)

IP

grass carp

(25,120,122)

Cyprinus carpio

Asia &

18th

IP,

3,5,9,15,17,24,

Linnaeus 1758

E. Europe

century

DAV, HD

31,35,36,40,44,47,

common carp

(12,85)

(4,142)

(2,78,85,

51,56,58,62,74,75,

109,162,

77,81,95,100,101,

164,165)

103,105,106,109,1

124,132,139,143,

i,q,s,w,y,aa

1000, 2000

Hypoph thahn icirthys

China &

1975

unknown

1

moUtrix

E. Siberia

(163)

2000

(Valenciennes 1844)

(163)

silver carp

337

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 6 29 APRIL 1996

Appendix 2. Continued.

Scientific classification
and common name

Native range

Date of
first

introduction

Impact'

Distribution

references

T’/ttcfl Linnaeus 1758

Europe &

1910

LI

8,24,97,

tench

W. Asia
(162)

(2)

o,w

Poeciliidae

SE&

1936

C

152.

Ganihusia afflnis
(Baird & Girard 1854)
gambusia, mosquitofish

central
N. America
(108)

(2)

l.p.u

PoecUia reticulata

S. America

1912

C,

f.v.w

(Peters 1859)
guppy

north of
Amazon R.
(82)

(61,136)

1000,2000

Xiphophorus helleri

S. Mexico

Before

C, PEL

10

Heckel 1840

& Guatemala

1974

(96)

w,x

swordtail

(146)

(7,96)

1000.2000

Centrarchidae

E & central

1938

C, PEL

20,22,35,39,40„5(),

Lcponiis nuicroclunis

N. America

(61)

major

58,81.94,1 1 1.139,

Rafinesque 1819
bluegill sunfish

(135)

(49,85,

87)

152,155,
d.ky.aa, 1000

Micropterus dohnnieu

E. Canada

1937

P major

24,39,80,87,93,98,

Lacepede 1 802

& USA

(63)

(9,35,49.

101,102,126,159

smallmoLith bass

(108)

102,126,

140)

e.k.y.ab

1000,2000

Micropterus pimctidatus

SE of USA

1939

unknown

e,g.m

(Rafinesque 1819)
spotted bass

(108)

(68)

1000,2000

Micropterus sahuoides

N. America

1928

P major

13,15,21,37.40,

(Lacepede 1802)
largemouth bass

(135)

(2)

2,13,23.

35,87,140

44,47.53.58.73,75,80,
81,84.90.94.1 13,132,
139,143,144,152.157,
e,g,h,y,z,aa,ab
1000,2000

Percidae

Europe &

1927

LI

P

Perea fluviatilis
Linnaeus 1758
perch

Asia (79)

(2)

(23)

1000

Cichlidae

Middle E

c, 1965

LI

157

Oreochroiu is aureus

& NE Africa

(157)

PH

338

DE MOOR AND BRUTON: ALIEN AND TRANSLOCATED AQUATIC ANIMALS IN SOUTHERN AERICA

Appendix 2. Continued.
Scientific classification

Native range

Date of

Impact'

Distribution

and common name

first

intro.

References

(Steindachner 1864)
Israeli tilapia

(147)

(31)

Oreochromis niloticus

Middle E.

c. 1959

LI

156

(Linnaeus 1758)

& NE Africa

(156)

PH

Nile tilapia

(147)

(156)

Reptilia

SE of USA

Before

Poss.

1 14

Testudinae

(38)

1984

VPM

Trachemys scripta elegans

(114)

(114)

(Schoepff 1792)
red-eared terrapin

Legend for impact of species.

C - Competition with indigenous species
DAV - Destruction of aquatic vegetation
HD - Habitat destruction

IP - Associated with the introduction of parasites

PH - Possibility of hybridisation with congeneric indigenous species

LI - Little impact

P - Predator of indigenous species

PF - Parasite of indigenous fish

PFL - Predator of fish larvae and eggs

VPL - Vector of parasites of livestock

Major - Indicates major impact of species on the aquatic environment

APPENDIX 3

Checklist of translocated indigenous species whose native range is within the study area together
with selected bibliography (in numerical and alphabetical code, listed in Appendix 4).

Scientific classification
and common name

Catchments* of
native range

Impact'

Distribution

references

Cyprinidae

C,D

uncertain,

32,33,34,86,

Barbus aeneus (Burchell 1822)

(141)

PH

92,107,137,

smallmouth yellowfish

(107)

e,i, 1000

Barbus anoplus Weber 1897
chubbyhead barb

A1,B1,C.D1,D2,
D3,D4,D5,E,J,
L,M,P,Q,R,S,T3=
U=,V,W,X (141)

uncertain

1,44

339

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 6 29 APRIL 1996

Appendix 3. Continued.
Scientific cla,ssification
and common name

Catchments* of
native range

Impact'

Distribution

references

Labeo capensis (Smith 1841 )
Orange River labeo

C,D(14I)

C,PH

32,33,34,48,i,

1000

Labeo umhmtiis (Smith 1841)
mud mullet

C,D,J,L,N,P,Q

(141)

HD

(88)

32,87,88,90,
!41,i, 1000, 2000

Clariidae

Clarias gariepinus
(Burchell 1822)
sharptooth catfish

A,B,C,D,V,W,X,Y

(141)

P.C

expect major

impact

(107,110)

33,34,52,107,

1 10, 1000,2000

Cichlidae
Chetia flaviventris
Trewavas 1961

A

(141)

uncertain

1

Oreochromis andersonii
(Castelnau 1861 )
three spot tilapia

Y

(1 17,141)

PH

55

Oreochromis macrochir
(Boulenger 1912)
greenhead tilapia

Y

(141)

PH

55,1

Oreochromis mossambicus
(Peters 1851)

Mozambique tilapia

A,B,P,Q,R,S,T,

U,V,W,X

(141)

C,PH,

expect major

impact

(41,52,132)

17,41,44,51,91,

132,138,141,158

1000

Sargochromis codringtoiii
(Boulenger 1908)
green happy

Y(117)

PH

55

Serranochromis angusiiceps
(Boulenger 1907)
thinface largemouth

Y (141)

PH

55

Serranochromis robustus
(Boulenger 1896)
nembwe

Y (141)

PH

55,119,121,

s

Serranochromis thunbergi
(Castelnau 1861)
brownspot largemouth

Y (141)

PH

55

Tilapia rendalli
Gilchrist & Thompson 1917
redbreast tilapia

A.B.W.X.Y

(141)

DAV

major impact
(16,99,125)

19,30,119,122,125

1000,2000

340

DE MOOR AND BRUTON: ALIEN AND TRANSLOCATED AQUATIC ANIMALS IN SOUTHERN AFRICA

Appendix 3. Continued.
Scientific classification

Catchments* of

Impact'

Distribution

and common name

native range

References

Tilapia sparnnanii
Smith 1 840
banded tilapia

Anabantidae
Sandelia capensis
(Cuvier 1831)

Cape rocky

A,B,C,D3.D4,D5,

D6,W,X,Y

(141)

G,H,J,K,L,M

(141)

C

(49,140)

C

(59)

* Ref. Fig. 1 & Appendix 1 for codes and description of catchments

Legend for the impact of species as in Appendix 2

24,35,50,74,87,

140,155,1000,2000

,59,x,1000

Although the native range includes most of the T and U catchments (Skelton 1993), Barbus anoplus is absent from the
Mkomazi and Mzimkulu catchments within U and Tj( respectively.

Although normally translocations within major catchments are beyond the scope of this paper, an exception is made here
since these catchments are completely separated from others within the major catchments of U and T3,

APPENDIX 4

References associated with Appendices 2 and 3

1.

Alletson, 1985

2.

Anon, 1944

3.

Anon, 1958

4.

Anon, 1959

5.

Anon, I960

6.

Anon. 1962/63

7.

Anon, 1974

8.

Anon, 1975/76

9.

Anon, 1982

10.

Appleton, 1974

II.

Appleton, 1977

12.

Balon, 1974

13.

Barrow. 1971

14.

Basson et al., 1983

15.

Batchelor. 1974

16.

Batchelor, 1978

17.

Begg, 1976

18.

Boomker et al., 1980

19.

Bourquin, 1985

20.

Bourquin and Mathias, 1984

21.

Bourquin and Sowler, 1980

22.

Bourquin and van Rensberg, 1984

23.

Bourquin et al., 1984

24.

Braack, 1981

25.

Brandt et al.. 1981

26.

Brown, 1967

27.

Brown, 1978

28.

Brown, 1980

29.

Bruton and Appleton. 1975

30.

Bruton and Kok, 1980

31.

Bruton and van As, 1986

32.

Cambray et al.. 1977

33.

Cambrey and Jubb, 1977a

34.

Cambray andJubb. 1977b

35.

Cambray and Stuart, 1985

36.

Cambray and Meyer, 1988

37.

Coetzee. 1977

38.

Conant. 1958

39.

Crass, 1964

40.

Crass, 1966

41.

Crass, 1969a

42.

Crass, 1969b

43.

Day, 1932

44.

Dixon and Blom, 1974

45.

Donnelly, 1965

46.

du Plessis, 1961

47.

du Plessis and le Roux, 1965

48.

Fogarty, 1980

49.

Gaigher (C.M.), 1981

50.

Gaigher (IG), 1975a

51.

Gaigher. 1975b

52.

Gaigher et al., 1980

53.

George, 1976

54.

Gephard, 1977

55.

Gilmore, 1978

56.

Goldner, 1967

57.

Hamiiton-Attwell et al., 1970

58.

Hamman. 1980

59.

Hamman et al., 1984

60.

Harrison, 1938

341

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 6 29 APRIL 1996

61. Harrison, 1939

64. Harrison, 1954

67. Harrison, 1964

70. Harrison, 1966/67
73, Heard and King, 1981

76. Hey, 1926

79. Holcik and Mihalik, 1968

82. Jacobs, 1971

85. Jubb. 1959

88. Jubb, 1966

91. Jubb. 1968

94. Jubb, 1972

97. Jubb, 1978

100. King and Bok, 1984

103. Kleynhans, 1984

106. La Hausse de Lalouviere. 1986
109. Marshall, 1972

112. Morrissy, 1978

1 15. Oberholzer and van Eeden. 1967

118. Pienaar, 1978

121. Pike, 1983

124. Plumstead et al.. 1985

127. Pretorius et al., 1975
130, Rayner, 1988

133. Schutte and Frank. 1964

136. Siegfried. 1962

139. Skelton, 1986

142. Skelton and Skead. 1984

145. Smith and Stearley, 1989

148. van Aset ah. 1981

151. van Bruggen. 1966

154. van Eeden et ah. 1980

157. van Schoor, 1969a

160. Walmsiey, 1988

163. Welcomme, 1988

Personal communication
a- Alletson, D.J.
b - Appleton, C.C.
c - Basson, L.
d - Benade, B.
e - Bok, A.
f - Bourquin, O.
g - Cambray, J.A.
h - Coetzee, A.
i - Coke, M.
j - Crass, R.

62. Harrison, 1948

65. Harrison. 1958

68. Harrison, 1964/65

71. Harrison. 1972/73

74. Heard and King, 1982

77. Hocutt and Skelton, 1983

80. Jackson, 1982

83. Jayaram. 1981

86. Jubb, 1963

89. Jubb, 1966/67

92. Jubb, 1971a

95. Jubb. 1973

98. Jubb, 1979

101. Koch and Schoonbee. 1980

104. Kruger, 1971

107. Laurenson and Hocutt, 1986

no. Mayekiso, 1986

113. Mulder, 1986

116. Paperna, 1980

119. Pike, 1980a

122. Pike, 1986

125. Potgieter, 1974

128. Pretorius et ah, 1980

131. Rondorf, 1976

134. Schoonbee. 1993

137. Skelton, 1972

140. Skelton, 1987

143. Smith. 1983

146. Sterba, 1962

149. van As and Basson, 1984

152. van der Merwe, 1970

155. van Rensburg. 1 966

158. van Schoor, 1969b

161. Welcomme, 1981

164. Crivelli, 1983

63. Harrison. 1953

66. Harrison. 1960/61

69. Harrison, 1966

72. Harrison and Lewis, 1968/69

75. Heeg, 1983

78. Hoffman and Schubert, 1984

8 1 . Jackson et ah. 1983

84. Joubert, 1984

87. Jubb. 1965

90. Jubb. 1967

93. Jubb, 1971b

96. Jubb, 1976/77

99. Junor, 1969

102. Kleynhans, 1983

105. Kruger et ah, 1983

108. Lee et ah, 1980

111. McVeigh. 1984

1 14. Newbery, 1984

1 17. Philippart and Ruwet. 1982

120, Pike. 1980b

123. Pike and Tedder, 1973

126. Pott, 1981

129. Rayner, 1981

132. Schrader, 1985

135. Scott and Crossman, 1973

138. Skelton, 1981

141. Skelton, 1993

144. Smith, 1984

147. Trewawas, 1983

150. van Bruggen, 1964

153. van Eeden and Brown, 1966

156. van Schoor, 1966

159. Viljoen and van As, 1985

162. Welcomme, 1984

165. Fletcher et ah, 1985

Natal Parks Board, Pietermaritzburg.

Department of Zoology, University of Natal, Pietermaritzburg.

Department of Zoology, University of the Orange Free State, Bloemfontein,
Cape Department of Nature Conservation, Kimberley.

Cape Department of Nature Conservation, Port Elizabeth.

Natal Parks Board, Pietermaritzburg.

Albany Museum, Grahamstown.

Fisheries Research Station, Marble Hall.

Natal Parks Board, Pietermaritzburg.

(formerly) Natal Parks Board, Pietermaritzburg.

342

DE MOOR AND BRUTON: ALIEN AND TRANSLOCATED AQUATIC ANIMALS IN SOUTHERN AFRICA

k - Davies. M.T.T.

I - Engelbreclit, J.
m - Hamman, K.C.D.
n - Hoffman, J.

0 - James, N.P.E.

p - Kleynhans, C.J.
q - Merron, G.S,
r - Palmer, C.
s - Pike, T.

1 - Pitt. R.

u - Ratte, T.

V - Reavell. P.

w - Ribbink. A

X - Skelton P.H.

y - Smith, A.

z - Thorne, S.

aa - van der Waal, B.

ab - van Loggerenberg,

Museum records

Department of Ichthyology and Fisheries Science, Rhodes University, Grahamstown.
Transvaal Nature Con.servation Division. Lydenberg.

Cape Department of Nature and Environmental Conservation, Bellevue, Cape Town.
Fisheries Research Slalion, Marble Hall.

J.L.B. Smith Institute of Ichthyology, Grahamstown.

Water Research Commission, Pretoria.

J.L.B. Smith Institute of Ichthyology, Grahamstown.

Institute for Water Research, Rhodes University, Grahamstown.

Natal Parks Board. Pietermaritzburg.

Federation of South African Fly Fishers, Eastern Cape Chapter.

Cape Department of Nature and Environmental Conservation, Sedgefield.

Lhiiversity of Zululand. Empangeni.

J.L.B. Smith Institute of Ichthyology, Grahamstown.

J.L.B. Smith Institute of Ichthyology, Grahamstown.

Cape Department of Nature and Environmental Conservation, Stellenbosch.

Cape Department of Nature and Environmental Conservation, Stellenbosch.
University of Venda, Sibasa.

Transvaal Nature Conservation Division, Lydenburg.

1000 - Albany Museum record.

2000 - J.L.B. Smith Institute of Ichthyology record.

APPENDIX 3

Existing and future interbasin transfer schemes between major catchments in southern Africa
which could result in further translocations (after Pettijean and Davies, 1988).

Scheme

Status

Donor catchment

Recipient river(i
& catchment

Usuthu-Vaal

Completed

Vaal

Steenkoolspruit

(Cl)

Olifants (B 1 )

Usuthu

Vaal

(Wl)

(Cl)

Tugela-Vaal

Completed

Tugela

Vaal

(V)

(Cl)

Orange project

Sections completed

Orange

Fish

(D3)

(Q)

Fish

Sundays

(Q)

(N)

Riviersonderend-

Completed

Breede

Eerste

Berg

(H)

(G)

343

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 6 29 APRIL 1996

Appendix 5. Continued.
Scheme

Status

Donor catchment

Recipient river(s)
& catchment

Ea.stern National

Under construction

Kavango

Swakop

Water Carrier

(Y2)

(Z)

Lesotho Highlands

Under construction

Upper

Vaal

Orange ( D 1 )

(C2)

Amatola

Under construction

Toise &

Nahoon &

Kubusie

Yellowoods

Kei (S)

(R)

Mooi-Mgeni

Under construction

Mooi (Tugela)

Mpofana (Mgeni)

(V)

(LI)

Mzinikulii-Mkomazi

Linder construction

Mkomazi

Mgeni & lllovo

(U)

(U)

Mzimkuiu

Mkomazi

(T3)

(U)

Zambezi ac|ueduct

Proposed

Zambezi

Botswana & Vaal

(YI.D5, C3)

Addendum

At the time of going to press the following two new distribution records were received:
Gambusici ciffinis in the Fish River (catchment Q) (Ter Morshuizen pers. comm.)
Oreochromis mossambicus in the Gouritz catchment (J) (Bok pers. comm.)

344

INSTRUCTIONS TO AUTHORS

LAYOUT of recent issues should be examined for details of acceptable style and format.

MANUSCRIPTS should be submitted on floppy disk accompanied by three print-outs
(spacing I/2 or 2). In exceptional cases typed manuscripts will be accepted. These should be
submitted in triplicate, typed on A4 size paper, double-spaced, on one side of the paper only
and with wide margins all round.

Only generic and specific names should be underlined to indicate italics. All other marking
up should be left to the editor.

Footnotes should be avoided as far as possible.

All measurements should be metric.

Systematic papers should follow the rules of the International code of zoological
nomenclature or the International code of botanical nomenclature, as applicable.

ILLUSTRATIONS should bear the author’s name, title of paper and number in soft pencil on
the reverse side. Diagrams, line drawings and photographs are all regarded as figures, should
be termed as such and should be numbered in a single series.

In addition to the original illustrations a photocopy of each should be submitted.

Diagrams and line drawings should be reducible to a size not exceeding 14x16 cm (17,5 cm
including captions). Photographs for half-tone blocks should be final size, of high contrast
and printed on glossy paper. Colour illustrations are only printed under special circumstances
and their financing must be arranged by the author.

A scale should appear with all illustrations or the magnification should be given in the
caption.

A list of captions should be submitted on a separate sheet.

TABLES should be submitted on separate sheets. Foldouts and tables printed sideways should
be avoided.

REFERENCES should be listed alphabetically at the end of the paper. Abbreviations of
periodical titles should conform with those in the World list of periodicals (4th ed.). Layouts
must be as follows:

Alexander, J. 1969. The indirect evidence for domestication. In: Ucko, P. J. and G. W.
Dimbleby eds. The domestication and exploitation of plants and animals. London:
Duckworth, pp. 123-129.

De Winter, B. 1969. Plant taxonomy today. S.Afr.J.Sci. 65 (8): 229-242.

Jlibb, R. a. 1967. Freshwater fishes of southern Africa. Cape Town: Balkema.

GALLEY PROOES are sent (when practicable) to authors for checking printer’s errors.
Adjustments of language, style and punctuation and any other amendments of the original
manuscript, if made at this stage, may be charged to authors.

AUTHOR’S COPIES (50) are issued free of charge. When more than one author is involved
the fifty free copies will be sent to the senior author. Additional copies ordered before printing
are supplied at a reduced cost.

CADAR, P.E.-K887

, ISSN 0570-1880

NH

Annals of the

Cape Provincial Museums

Natural History

Ann. Cape Prov. Mus. (nat. Hist.)

Volume 19 Part 7 29th April 1996

Nesting and flower visiting of some southern African Anthophorini
(Hymenoptera: Apoidea: Apidae: Apinae)

F.W. GESS and S.K. GESS

m 3 0 1997

tlBRARlE-^

Published jointly by the Cape Provincial Museums
at the Albany Museum, Grahamstown, South Africa

ANNALS OF THE CAPE PROVINCIAL MUSEUMS

These Annals are published in two series. Natural History and Human Sciences, the latter series
covering cultural history, ethnology, anthropology and archaeology. They are issued in parts at
irregular intervals as material becomes available.

The primary objective of these Annals is to disseminate the results of research work undertaken by
staff members of the Cape Provincial Museums. However, a manuscript by any author is eligible
for publication provided it is based wholly or partly on the collection/s of one or more of these
museums or if the material on which the work is based is wholly or partly deposited in one of
these museums.

Parts are obtainable from the Librarian of the Albany Museum. Correspondence in connection
with these Annals should be addressed to the Editor, Albany Museum, Grahamstown 6140.

Editor

Dr F. W. GESS

Assistant Editor
Dr S. K. GESS

Nesting and flower visiting of some southern African Anthophorini
(Hymenoptera: Apoidea: Apidae: Apinae)

by

F.W. GESS and S.K. GESS
(Albany Museum, Grahamstown, 6140 South Africa)

ABSTRACT

Accounts are given of nesting in horizontal or gently sloping ground by two species of Ainegilla
Friese, A. (Micrainegilla) atrocincta (Lepeletier) and A. (Zebramegilla) punclifrons (Walker). The
burrow of A. (M.) atrocincta is excavated using water and the entrance is surmounted by a turret.
Each shaft, of which there may be more than one per nest, is terminated by a single cell. Water is not
used in excavation by A. (Z.) pimctifrons. Soil particles are extracted using the mandibles and are
raked out of the burrow. Each shaft, of which there may be more than one per nest, is terminated by
several cells in linear series. Notes are given on the nesting in vertical earthen banks by two species
of Anthophora Latreille, A. ( Heliophila ) praecox Friese and A. ( ParamegiUa ) epichariformis Gribodo
neither of which constructs an entrance turret.

Evidence is given for possible associations between Thyreus alfkeni Brauns and Thyreus
delumbatus (Vachal) and A. (Z.) punctifrons, and between Thyreus calceatiis (Vachal) and A. (H.)
praecox.

Flower visiting records, totalling 502, are given for 16 species of Ainegilla and 10 species of
Anthophora. Analysis of these records shows that the differences in flower choice between genera
and also between subgenera (despite the lack of distinction between pollen and nectar collection)
indicate that the Anthophorini do not as a rule practise indiscriminate broad polylecty.

INTRODUCTION

Remarkably little has been published on the biology of the southern African species of
Anthophora Latreille and Ainegilla Friese (Apoidea: Apidae: Apinae: Anthophorini). Brauns (1913)
and Rozen (1969) gave life history notes on two species of Anthophora. Although some observations
were made by the present authors from time to time over the past twenty years the difficulty
experienced in determining the voucher specimens discouraged more intensive investigation. With
the recent taxonomic revisions of the southern African species of these genera (Eardley and Brooks
1989, Eardley 1994) the identification of these bees has been greatly facilitated. It seems useful to
present and discuss the available nesting observations and flower visiting records despite the fact that
on the whole they are of a somewhat fragmentary nature. This may lay the foundation and provide
the stimulus for further investigations.

NESTING

Ainegilla ( Micrainegilla ) atrocincta (Lepeletier)

Geographic distribution

Ainegilla (Micramegilla) atrocincta occurs throughout southern Africa, with the noticeable
exception of the region incorporating southern Namibia, and the northern, central and western Cape
Province (Eardley 1994).

347

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 7 29 APRIL 1996

Nesting areas, nesting sites and nesting density’

Two nesting areas have been located on the farms Hilton (27.15S, 32.2 IE) and Clifton (27.1 IS,
32.24E), respectively 1 8 and 19 kilometres WNW Grahamstown in the southeastern extension of the
Nama Karoo. The vegetation of Clifton, the more easterly farm, differs from that of Hilton in having
a higher succulent element. The nesting sites were in bare areas of non-friable clayey soil in low
scrub, all in close proximity to water. The surface of the ground chosen for nesting was in all instances
horizontal to subhorizontal.

During the eight years from 1973 to 1981 during which the authors were working intensively on
the farm Hilton Ainegilla atrocincta was very rarely sighted. Only two single nests were found, one
in 1973 and one in 1978.

In some years Ainegilla atrocincta has been relatively commonly encountered on forage plants
at Clifton; however, the nests have proved difficult to locate. No aggregations of nests were found,
only single widely scattered nests. During the summer 1985/1986, when a deliberate effort was made
to locate nests, only five nests were found.

Brauns ( l913)commented that, at Willowmore, he found A. atrocincta (as Antliopliora plumipes)
nesting in areas of hard level ground, the nest entrance generally covered by a stone. No nests so sited
have been found by the authors.

Flight period

Collecting dates (collected records from Eardley 1 994, additional records from label data Albany
Museum) throughout the range of A. atrocincta suggest that this bee is most commonly in flight from
mid- to late-summer, December to March in the Cape Province. Certainly in the vicinity of
Grahamstown, where the Gesses have collected throughout the year, this bee is most commonly
encountered in January and February. To the north in the Transvaal, Zimbabwe and Namibia the
flight period is more extended there being scattered records from September through to May.

Flower visiting

All plants in flower on Hilton and Clifton were sampled for flower visitors at intervals throughout
the summer. It was found that males and females of Ainegilla atrocincta commonly visited the white
flowers of Blepharis capensis (L.f.) Pers. ( Acanthaceae) at Clifton during January and February. An
additional two males were taken on flowers of B. capensis at Waterford (33.07S, 25.01E). Though
not sampled A. atrocincta was observed visiting the yellow flowers of Cotyledon campamilata
Marloth (Crassulaceae) in addition to those of B. capensis during January 1986 when the nesting
behaviour of the bees was being investigated on Clifton. No forage plants of these bees were located
at Hilton. The only other flower visiting records available are of two females taken on the yellow
flowers of Berkheya heterophylla (Th.) O.Hoffm. (Asteraceae) between Lynton and Vlakwater
(27.09S, 32.18E). Voucher specimens are listed in Appendix 1 under Grahamstown and Waterford.

Provision

The provision is in the form of a syrup on which the egg or larva floats. Pollen from the cells of
two nests at Clifton was examined microscopically. Though mixed, the bulk of the pollen from both
nests matched that of C. campannlata. No pollen matching that of B. capensis was found suggesting
that this plant was principally visited for nectar. However, provision from a third nest, investigated

348

GESS AND GESS: NESTING AND FLOWER VISITING OF SOME SOUTHERN AFRICAN ANTHOPHORINI

when the flowers of C. campamdata were almost over, contained an appreciable proportion of pollen
matching that from B. capensis and Peristrophe cermia Nees (Acanthaceae) but very little pollen
matching that from C. campamdata.

The cells when opened gave off a marked odour which could be described as “meady” or “cheesy”.
It was probably a combination of odours derived from the provision and the waxy Dufour’s gland
secretion.

Male behaviour

At Clifton, in addition to feeding at the flowers of B. capensis males were frequently observed
patrolling a patch of this plant growing in close proximity to nest sites. None was observed in the
vicinity of nests.

Sheltering and sleeping

A nesting female was observed by the present authors sheltering head downwards in her nest.
Males and un-mated, and therefore nestless, females were observed by Brauns (1913) spending the
night on low plants, up to 10 bees in a row on a single twig, holding on firmly with their mandibles,
and in smaller numbers under stones.

Description of nest

The nest consists of a subterranean burrow surmounted by a cylindrical erect or inclined tun-et
constructed from mud and having an inner diameter of 9 mm and reaching lengths of up to 80 mm
(Fig. 1). Individual applications of mud are visible on the outside of the turret but the inside is
smoothed.

The burrows of six of the seven nests investigated consisted of a single subvertical shaft, 8-9
mm in diameter. Four terminated in a single cell and two had not reached the stage of cell construction.
The burrow of the sixth nest bifurcated just below the surface of the ground (Fig. 2). One shaft,
apparently the first excavated, led directly downwards whereas the other at first sloped gently before
continuing downwards parallel with the first shaft. Each shaft terminated in a single cell at an average
depth of 172.4 mm (sample of five, range 130-208 mm). The sides of the cells were parallel and the
base was rounded. The diameter of the cells was the same or marginally greater than that of the shafts.
The lengths of the two cells measured were 20 and 23 mm. The walls were smooth and lined with
v/hite wax. Each cell was closed with a mud and wax plug which was somewhat smoothed above but
not below and which was slightly concave above and was either flat or somewhat concave below as
the plug was sometimes deeper around its circumference than at its centre. Above each cell plug was
a capping of mud. Above this a portion of shaft, slightly shorter than a cell, with smoothed walls
formed a chamber. This chamber v/as full of mud pellets, and was sealed with a mud plate. The shaft
above this chamber was full of loose earth and there was an additional mud plate at a variable height
along its length. In one nest a mud plate sealed the shaft at ground level within the turret.

Method of construction

Water is required for the excavation of the burrow, construction of the tuiTet, smoothing of the
cell walls and construction of the mud seals. Water is collected by the female from a nearby pool.
Whilst taking in water she stands on the mud at the water’s edge.

349

ANN. CAPE PROV. MUS, (NAT. HIST.) VOL. 19, PT. 7 29 APRIL 1996

•V

¥

350

GESS AND GESS: NESTING AND FLOWER VISITING OF SOME SOUTHERN AFRICAN ANTHOPHORINI

Fig. 2. Vertical plan of one of the nests of Amegilia (Micramegilla) atrocincta (Lepeletier) investigated on the farm Clifton
(27.1 IS, 32.24E). Scale bar = 10 cm.

351

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 7 29 APRIL 1996

Having selected a suitable nesting site and returned with a load of water the female initiates
excavation. She regurgitates the water and mixes it with the soil to form mud which she passes back
beneath herself, rotating steadily. The mud is positioned around the rim of the shaft initial to form
the base of the turret. Further extracted mud is passed hack in the same manner and is positioned by
the tip of the abdomen. The lengths of time spent away from the nest collecting water, actually
imbibing water and working at the nest excavating mud and working on the construction of the nest
were measured for one of the builders. She was found to take 20-34 seconds away from the nest to
fetch water. 13-14 seconds to tank up, and for each water load 2-3 minutes excavating, adding two
loads of mud to the tuiret and smoothing it on the inside. On her return flights to the nest she circled
above the nest and then flew straight in.

In one instance a shaft passed through a patch of weakly structured soil. The walls of this section
of the shaft had been shored up by cementing with mud.

The excavated cell walls were smoothed apparently with the use of water. There was no
constructed mud cell within the excavated cell. The wax lining of the cell forms a distinct layer which
can be separated from the earthen wall.

The wax coating of the cell wall extends into the mouth of the cell. The mud and wax plug sealing
the ceil is constructed within the mouth of the cell fitting like a stopper into the neck of ajar. The
mud is laid down spirally; however, the spiral is not continuous as each load of mud is added without
matching the end of the previous spiral which is little more than a ring with overlapping ends. Five
such broken rings could be counted on one of the plugs. The central remaining aperture is crudely
plugged with a final load of mud which may project on the lower surface as a rough knob. The manner
in which the wax is applied is not clear. As the lower surface of the plug including the knob has a
visible white coating it is thought that the wax is probably added with the mud. The upper surface is
roughly smoothed and given a light wax coating. An additional mud capping is added above the
stopper and this bonds with the walls of the shaft above the cell.

The mud capping added above the cell plug, unlike the plug shows no patterning. It is, apparently,
simply a layer of mud spread over the plug and onto the walls at the base of the chamber above the
cell. Its function appears to be to completely seal the cell, as the plug is not firmly attached to the cell
wails there being a layer of wax between the earthen walls and the plug. No wax is added to this
capping or to the walls of the chamber above the cell. The function of the wax appears to be to
waterproof the cell, the provision being fluid.

Associated insects

Of the seven nests two were found to have been “parasitised”.

One nest had been usurped by a megachilid. This nest contained one A. atrocincta cell which
had been lined with wax and partly provisioned. The cell and the shaft above it had been filled with
gravel to a height of 30 mm above the cell. The megachilid bee had then constructed a petal cell
sealed above with a petal disc. The sealing disc was identified as being cut from a white petal of
B. capensis. The remainder of the shaft had been filled with gravel. Pollen from the provision was
examined and found to be of two forms, one of which matched that of P. cernua (Acanthaceae) and
the other Pentzia incana (Thunb.) Kuntze (Asteraceae).

In the other nest eight non-phoretic triungulin larvae of a meloid beetle were found in the single
sealed cell.

352

GESS AND GESS: NESTING AND FLOWER VISITING OF SOME SOUTHERN AFRICAN ANTHOPHORINI

Amegilla (Zebramegilla) piinctifrom (Walker)

Geographic distribution

Amegilla {Zebramegilla} piinctifrons is widespread in subsaharan Africa, and apparently occurs
throughout the subcontinent, except that it is noticeably absent from the western, southwestern, central
and northern Cape Province, and from the southern half of Namibia (Eardley 1994).

Nesting areas, nesting sites and nesting density

Brauns (1913) noted that A. (Z.) piinctifrons (as Anthophora circulata) nests in extensive
aggregations in bare level places in the vicinity of Willowmore. The present authors have located
two further nesting areas. One in the eastern Nama Karoo on the neighbouring farms of Clifton
(27.1 IS, 32.24E) and Thursford, WNW of Grahamstown, and the other in the southern Great Karoo
at the interface between the Succulent Karoo and the Nama Karoo on the farm Tierberg (33. OSS,
22. 1 6E), to the east of Prince Albert. The sites of nesting aggregations in these areas are bare patches
of horizontal to gently sloping clayey ground in karroid scrub with a strong succulent element. In all
instances, although there was an abundance of similar bare areas and although the bare areas selected
were extensive, even in small aggregations, the nests were crowded together, new nests being added
around the fringes of the aggregations.

Flight period

Collecting dates (collected records from Eardley 1994 and additional records from Albany
Museum) suggest that A. (Z.) piinctifrons is in flight in the southern and eastern Cape from early
summer, October, through to late summer, March. To the north in the northern Transvaal and
Zimbabwe it is possible that it flies through the winter as it has been collected from June to August.

Flower visiting

On the farm Clifton females and males of A. (Z.) piinctifrons regularly forage together with
A. ( M. ) atrocincta on the white flowers of Blepharis ct7/?£’/!5/.v ( Acanthaceae) in January and Eebruary.
In addition in October early flying males were taken on the same flowers at Clifton and on yellowish-
white flowers of a “mesem” (Mesembryanthemaceae) on Verdun (33.10S, 25.50E) in the
Kommadagga District to the north west of Clifton. Voucher specimens are listed under Grahamstown
and Kommadagga in Appendix 1 .

No forage plants of A. (Z.) piinctifrons were located on Tierberg.

Provision

Provision from the cells of nests excavated on Tierberg was a very wet mixture of pollen and
nectar. Some of the pollen was examined microscopially and was found to be of two types, one
relatively large and oval and the other relatively small and spherical, both smooth walled as seen with
the light microscope. Though comparisons with pollen from plants in flower were made it was not
possible to identify the pollen with any certainty.

353

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 7 29 APRIL 1996

Sheltering and sleeping

Sleeping by A. (Z. j punctifrons has not been observed by the present authors but has been recorded
at Willowmore by Brauns (1913). He found sleeping clusters of males and females mixed, up to 30,
on low bushes, the bees being firmly clamped onto the stems by means of their mandibles. More
rarely, males were found sheltering together with males of Amegilla niveata (Friese) (as Anthophora
niveata) in hollow stems, lying on the ground, and in dry open seed capsules of Datura. Actively
nesting females appeared to retire to their nests at night.

Description of the nest

Access to all nests was direct. In no instance was there any form of nest entrance turret. Three
nests of a nesting aggregation on Tierberg were excavated. The shaft, 6- 6.5 mm in diameter, in all
three nests initially sloped gently downwards before continuing subvertically at a constant diameter
to a depth of 95-105 mm below which it ended in one to three subvertical to sloping cells in linear
series (Fig. 3). In two of the nests there were secondary shafts in addition to the main shaft, in one a
single secondary shaft and in the other two and the initial of a third. The secondary shafts in turn each

Fig. 3. Vertical plans of the nests of Amegilla (Zehnmiegilla) punctifrons (Walker) investigated on the farm Tierberg (33. 088,
22.I6E). Scale bar = 10 cm.

354

GESS AND GESS: NESTING AND FLOWER VISITING OF SOME SOUTHERN AFRICAN ANTHOPHORINI

ended in a linear series of cells. Between the last completed cell in each series and the opening to the
next excavated shaft the shaft had been filled with earth and sealed off from the main shaft with a
mud-plate. Twelve cells in all were obtained. Each was 10 mm long and 7-7.5 mm in diameter at
mid-length. The walls were cemented and smoothed, and coated with wax.

Method of construction

Water is not used in nest excavation. A burrow is initiated at a shallow angle to the ground
surface. Soil is extracted using the mandibles, the bee continuously turning on its long axis. The spoils
of excavation are raked out of the nest and accumulate to one side of the nest. In densely populated
nesting aggregations there are no distinct tumuli, the entire surface of the compacted clayey ground
becoming covered with destructured soil. An entrance to a well established nest is concealed with
loose earth through which the bee digs when entering.

Associated insects

During the nesting season female Thyreus have repeatedly been observed in attendance at and
digging through closures of nests of A. (Z.) punctifrons on the farms Thursford and Clifton strongly
suggesting a parasitic association. A sample of eight was constituted of seven Thyreus alfkeni (Brauns),
three from Thursford and four from Clifton, and one Thyreus delumbutus (Vachal) from Clifton.
Brauns (1913) recorded that the latter (as Crocisa hraunsiana Friese) was present in large numbers
in association with the nesting aggregations of A. (Z.) punctifrons (as Anthophora circulata) at
Willowmore and, though he gives no further evidence, refers to it as a regular parasite.

It is of interest that T. delumbatus was collected in company with its putative host visiting Bowers
of B. copensis on Clifton.

A single female mutillid was observed digging through the loose earth closure of a nest on
Thursford suggesting a parasitic association.

Anthophora (Heliophila) praecox Friese

Geographic distribution

Anthophora ( Heliophila ) praecox Friese has been recorded from the western, southern and eastern
Cape Province, from the eastern Orange Free State and Lesotho (Eardley and Brooks 1989, Fig. 108).

Nesting areas, nest sites and density

Two nesting areas of A.(H.) praecox have been located, Hilton Farm (27.I5S, 32.2 IE),
Grahamstown and Tierberg Farm (33.08S, 22.I6E), Prince Albert. Nesting sites in both areas were
water cut vertical banks, in the former a river bank and in the latter the bank of an erosion gully. On
Tierberg the bee was also found nesting in an artificial vertical bank, the mud wall of a ruined building.
Nesting in the walls of buildings was previously noted by Brauns (1913) who observed that A. (H.)
praecox (as A. wartmanni var. praecox Friese) in Willowmore was often found nesting in large
numbers close together in walls constructed of unbaked bricks. The nests on Hilton and Tierberg
were also similarly crowded together.

355

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19. PT. 7 29 APRIL 1996

Flower visiting and provision

Anthophora (H.) praeco.x has been collected on the yellow flowers of Berkheya species
( Asteraceae) from the Olifants River Valley in the west to Hilton Farm, Grahamstown in the east. In
addition it has been collected on flowers of Sphalmanthns sp. (Mesembryanthemaceae) at Klein
Doom River, 10 km E Lemoenshoek, and on the blue flowers of Anchiisa capensis Thunberg
(Boraginaceae) on Hilton Farm. Voucher specimens are listed in Appendix 1.

This bee was not collected on flowers on Tierberg farm but pollen from provision from nest cells
was compared microscopically with that of plants growing in the vicinity of the nests and was found
to match that of Pentzia incana (Asteraceae).

The cells from the Hilton nests had been provisioned with a very wet but not fluid orange pollen
and nectar mixture which filled the cells to within 5 mm of the seal. The surface of the provision was
smooth.

Description of the nest

The nest, which is sited in a vertical compacted earthen bank, is a single-celled to multi-celled
burrow, in some instances with at least two cells in linear series. Access to the burrow is by a simple
entrance hole without any form of entrance turret. The cells are smoothed on the inside, lined with a
“varnish” like substance, and closed with a mud seal coated on both surfaces with the same substance
as is used for lining the cells.

Method of construction of the nest and oviposition

Nest initiation has not been observed. It was not clear whether the burrows were self-excavated
or pre-existing. Those at Hilton were in reddish “sandstone” and the cells had been lined with mud
clearly introduced into the nest, suggesting that the burrows might have been pre-existing. No such
difference between the substrate and the cell lining was observed in the Tierberg burrows. The burrows
in the erosion gulley were crowded together and apparently interlinked sharing more than one entrance
and a single entrance being entered by more than one female, suggesting nest sharing.

A single egg was obtained from a horizontally positioned cell from a Hilton nest. It was white,
curved, 3.5 mm long and 0.75 mm wide at mid-length and had been laid onto the vertical surface of
the provision. It was only attached by the upper end, the rest of the egg hanging free from the provision.

Associated insects

No direct evidence for nest parasitism was obtained; however, Thyreus calceatus (Vachal) was
commonly in attendance at the nests in the river bank at Hilton.

Anthophora (Parantegilla) epichariformis Gribodo

Geographic distribution

Anthophora (Paraniegilla) epichariformis Gribodo has been recorded from the southwestern
Cape (Eardley and Brooks 1 989). A nesting female and a male collected on the farm Tierberg (33. 088,
22.16E) to the east of Prince Albert in the southern Great Karoo (26.xi.-5.xii. 1987, F.W., S.K. and

356

GESS AND GESS: NESTING AND FLOWER VISITING OF SOME SOUTHERN AFRICAN ANTHOPHORINI

R.W.Gess) extends the known distribution eastwards.

Nesting area and nest site

One nest ofA.(P-) epichariformis was located on the farm Tierberg. It was situated near the base
of a high, subvertical, south-facing bank of the Tierberg River and sited beneath an overhang. The
river bank is constituted of a grey poorly sorted unconsolidated modern alluvium.

Flower visiting and provision

There appear to be no flower visiting records; however, pollen taken from the provision of the
three cells obtained from the excavated nest was all of one kind and matched that of a Pteronia species
(Asteraceae) growing in the vicinity of the nest.

The provision was a moist pollen and nectar mass.

Description of the nest and egg

The single nest obtained consisted of a burrow excavated in the river bank. There was no entrance
turret. The shaft penetrated the bank horizontally for 2 cm and then turned at right angles to run
subvertically downwards more-or-less parallel to the surface of the bank to a depth of 10 cm
terminating in three cells in linear series each sealed with mud. The walls of the cells were cemented
and lined with a waxy coating.

The egg, which is laid onto the surface of the provision, is curved, yellow and 4.5 mm long from
tip to tip across the arc.

Method of construction of the nest

Water did not appear to have been used for the excavation of the burrow. The nature of the
bonding agent used for cementing the walls of the cell and sealing the cells was not identified.

DISCUSSION OF NESTING BY AMEGILLA AND TWO SUBGENERA 0¥ ANTHOPHORA,

HELIOPHILA AND PARAMEGILLA

The most detailed description of the nesting of an AmegiUa is that of Houston (1991) for the
Australian AmegiUa (Asaropoda) daw son i (Rayment). Eurther nesting accounts for Australian
AmegiUa species are for AmegiUa (Asaropoda) sp. 2 (Houston 1991 ), AmegiUa (Asaropoda) sp.
( ?bomhiformis, Cardale \ 96Sh), AmegiUa (Asaropoda) sp. (Maynard \992), AmegiUa (Zonamegilla)
pidchra (Smith) (Michener I960, as A. salteri (Rayment); Cardale 1968a as AmegiUa (AmegiUa)
pulchra (Smith)). Eragmentary notes on several Australian species are given in Rayment ( 1935, 1951)
and on a Palaearctic species in Ferton (1920).

Nesting in aggregations as recorded ior A.(Zebr.) pnnctifrons seems to be a common feature of
AmegiUa. Single widely scattered nests as recorded for A. (M.) atrocincta is less common. However,
the nests of A. (A.) daw son i though they do occur singly and widely scattered are also found in
aggregations of up to 10 000. This raises the question of whether there are areas where populations
of A. (M.) atrocincta are large and whether in such areas this species may also nest in aggregations.

Excavation by AmegiUa may either be with (for example A. (M.) atrocincta and A. (A.) dawsoni)

357

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 7 29 APRIL 1996

or without the use of a wetting agent (for example A. (Zehr.) punctifrons) . Whereas water collection
by A. (M.) (itrocincta has been regularly observed, Houston states that no Australian solitary bees are
known to visit water. He suggests that nectar may be the wetting agent used in excavation and
construction by A. (A.) dawsoni. Unfortunately he did not test his hypothesis by analysing the mud
used in the construction of the turrets.

T urrets are constructed by some but not all Australian Amegilla. The turrets of A. (M. j atrocincta,
A. (A.) dawsoni t/nd Maynard’s A. (A.) sp. at least, apart from the probable difference in the bonding
agent, are initially similarly constructed. However, whereas turrets of up to 80 mm appear to be the
norm for A. (M.) atrocincta, 15-20 mm is usual for A. (A.) dawsoni and 10 mm for Maynard’s A. (A.)
sp.. Associated with the turret of A. (A.) dawsoni is a pronounced annular tumulus whereas there were
no visible deposits of discarded soil in the vicinty of the nests of A. (M.) atrocincta. Deposits of
discarded soil are, however, characteristic of A. (Zebr.) punctifrons which does not construct a turret
but keeps the nest entrance closed with loose material excavated from the shaft.

Multicellular burrows seem to be characteristic for Amegilla but the arrangement of secondary
shafts and cells differs markedly between species. The single-celled burrows of A. (M.) atrocincta,
A. (A.) dawsoni and Houston’s species 2 are similar in that they all terminate in a vertical cell with
a double seal above which is a soil filled antechamber. However, burrows with more than one cell
are markedly distinct. If the single two-celled nest of A. (M.j atrocincta is taken as typical for the
species then the secondary shaft in nests of this species diverges from the main shaft immediately
within the entrance, extends laterally and then downwards ending in a cell at approximately the same
depth as the first cell. However, the main shafts of A. (A.) dawsoni and Houston’s species 2 remain
unbranched until immediately above the antechamber at which point there is a lateral and downward
extension leading to a second cell. As each successive cell of A. (A.) dawsoni is deeper than that
preceeding it the main shaft appears to continue downward in steps. However, succeeding cells of
species 2 form a group all at the same depth. Whereas cells in linear series have not been recorded
for A. (M.) atrocincta and species 2 they seem to be usual for A. (Zon.) pulchra and A. {Zebr.)
punctifrons and occasionally two cells in sequence are constructed by A. (A.) dawsoni.

Lining of the cells with wax which is presumed to be derived from the Dufour’s gland secretion
seems to be general in the Anthophorini. A. (M.) atrocincta and A. (Asaropoda) species waterproof
the cell seal with wax. The way in which this is achieved appears to differ but this may reflect a
difference in interpretation. Houston considers the mud seal of A. (A.) dawsoni to be constructed
spirally and to be waxed on the lower surface just before the central aperture is finally sealed with
mud. From the small sample of A. {M.) atrocincta seals, all completed, available to the present authors
it appears that this species applies the wax together with the mud as the wax on the lower surface
extends across the final central application of mud and is also present on the upper surface. Both
species construct a second unwaxed seal above the first seal, fill the antechamber with soil and then
construct a third seal.

A knowledge of the nesting of more species is required before it will be possible to distinguish
between subgeneric and specific differences.

The notes on the nesting of Anthopliora (Heliophila) praecox and Anthophora (Paramegilla)
epichariformis presented here though scanty are of interest when considered together with the other
published observations for the two subgenera to which they belong.

Nesting in turretless burrows in aggregations in vertical to subvertical earthen banks has, in
addition to Anthophora (H.) praecox, been recorded for Anthophora (Heliophila) braunsiana Friese
(Rozen 1969) and Anthophora (Heliophila) rufolanata Dours (Rozen 1969 as Anthophora krehsi
Friese) in the eastern Cape Province, South Africa, and Anthophora f Heliophila) flexipes Cresson

358

GESS AND GESS: NESTING AND FLOWER VISITING OF SOME SOUTHERN AFRICAN ANTHOPHORINI

(Torchio and Youssef 1968 as Anthophora (Micrcmthophora) flexipes Cresson) and Anthophora
(Heliophila) peritomae Cockerell (Torchio 197! as Anthophora ( Micranthophora) peritoniae
Cockerell) in Utah, U.S.A. A sixth species of Anthophora (Heliophila), Anthophora (H.) vestita
Smith, has been recorded nesting in vertical earthen surfaces, walls built of unbaked, sundried clay
bricks (Brauns 1913). Unfortunately no details of the nests are given.

Entrance sharing as observed for A. (H.) praecox, has been recorded by Torchio ( 1971 ) for A.
(H.) peritomae Cockerell, similarly nesting in a bank honeycombed with burrows.

Instances of at least two cells arranged in linear series have been recorded for all but A. (H.)
rnfoianata. As this practice is variable it is possible that instances may be found to occur for this
species in addition.

Little seems to have been recorded on the nesting of Anthophora (Parainegilla). Ferton ( 1902,
1920) gives notes on the nesting of A. (Parainegilla) halneorinn Lepeletier and A. (Parainegilla)
ferriiginea Lepeletier in France. Both, like A. (P.) epicharifonnis excavate turretless, multicellular
nests. A. (P.) ferriiginea unlike the other two species seemingly nests in horizontal ground rather than
in vertical banks. As only a single nest of A. (P.) epicharifonnis is known it could not be established
whether or not the linear arrangement of the cells is the rule. In Ferton’s nests of A. (P.) ferriiginea
only some of the cells were arranged in linear sequence. It would therefore appear that even within
single nests of A. (Parainegilla) as in A. (Heliophila) this is a variable character. Brooks (1988)
mentions that Marikovskaya ( 1970) has described nests of two Eurasian species of A. (Parainegilla)
but gives no details.

ANALYSIS AND DISCUSSION OF FLOWER VISITING BY SOUTHERN AFRICAN

ANTHOPHORINI

The Anthophorini, that i^Ainegilla and Anthophora, are regarded as polylectic [collecting pollen
from a wide range of families] as a general rule, narrow oligolecty [collection of pollen from flowers
of a single family or even genus] being considered unusual (Brooks 1988). Most collectors when
recording flowers visited do not distinguish between pollen and nectar collection. The data for southern
African Anthophorini available to the authors in the present discussion should therefore be seen as
encompassing both types of visit indiscriminately. They are assembled from label data in the Albany
Museum collection (F.W., S.K., D.W., H.W. and R.W.Gess - 339 records, and C.F.Jacot Guillarmod
- 54 records) and from label data of other collectors (a total of 109 records) listed by Eardley and
Brooks ( 1989) and Eardley ( 1994). Records are available for 16 of the 31 species of Aniegilla and
10 of the 3 1 species of Anthophora represented in southern Africa. Twenty four dicot families and
three monocot families in total have been recorded as visited. Tables 1 and 2 indicate the families
known to be visited by each of the species. Immediately apparent are some interesting differences
between Aniegilla and Anthophora, and between the two subgenera ( Heliophila and Pyganthophora)
of Anthophora for which records are available.

Taking the 16 species of Aniegilla together, Acanthaceae appears to be markedly favoured being
recorded for 12 (75.00%) of the species as compared with one (10.00%) of the ten species of
Anthophora. By contrast seven (70. 00%) of the ten species of Anthophora were recorded from
Boraginaceae whereas of the 16 species of Aniegilla only one (6.25%) was recorded from this plant
family. Furthermore five (50.00%) of the ten species of Anthophora were recorded from Asteraceae
and from Sterculiaceae whereas of the 16 species of Aniegilla only six (37.50%) and two ( 12.50%)
respectively were recorded from these plant families. However, Asteraceae together with the equally
ranked Lamiaceae does come second as being most favoured after Acanthaceae by Aniegilla as a whole.

359

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 7 29 APRIL 1996

Table 1 . Flower families recorded as visited by 16 AmegiUa spp. together with the number and percentage of these species
visiting each flower family (the species here represented by numbers correspond to the species listed by number in
Appendix I ).

Flower families

Amegilla species

no.

%

of spp.

1

2

3

4

5

6

7

8

9

10

1 1

12

13

14

15

16

Acanthaceae

+

+

+

+

+

+

+

-

-

+

+

-

-

+

+

+

12

75.00

Asteraceae

-

-

+

-

-

+

+

-

-

+

+

-

-

-

-

+

6

37.50

Boraginaceae

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

1

6,25

Campanulaceae

-

-

-

-

-

-

-

-

-

-

+

-

-

-

-

2

12.50

Capparaceae

-

-

-

-

-

-

+

+

-

-

-

-

-

-

-

3

18.75

Convolvulaceae

-

-

+

-

-

-

-

-

-

-

-

-

-

-

-

-

1

6.25

Elatinaceae

-

-

-

-

-

-

+

-

-

-

-

-

-

-

-

-

1

6.25

Geraniaceae

-

-

-

-

-

-

+

+

-

-

-

-

-

-

-

+

3

18.75

Hydrophyllaceae

-

-

-

-

-

-

+

-

-

-

-

-

-

-

-

-

1

6.25

Lamiaceae

-

-

+

+

-

-

+

-

-

-

-i-

-

+

-

-

+

6

37.50

Malvaceae

-

-

-

-

-

-

+

-

-

-

-

-

-

-

-

-

1

6.25

Melastomataceae

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

1

6.25

Mesembryanthemaceae

-

-

-

-

-

-

+

-

-

-

-

-

-

+

+

4

25.00

Mimosaceae

-

-

-

-

-

-

-

-

-

-

-

-

-

-

+

2

12.50

Moringaceae

-

-

-

-

-

-

+

-

-

-

-

-

-

-

-

-

1

6,25

Papilionaceae

-

-

-

-

-

-

-

-

-

-

-

-

+

-

-

+

2

12.50

Pedaliaceae

-

-f

-

-

-

-

+

-

-

-

-

-

-

-

-

+

3

18.75

Plumbaginaceae

-

-

-

-

+

-

-

-

-

-

+

-

-

-

-

+

3

18.75

Proteaceae

+

1

6,25

Rosaceae

-

-

-

-

-

-

+

-

-

-

-

-

-

-

-

-

1

6.25

Scrophulariaceae

-

-

-

-

-

-

+

-

-

-

-

-

-

-

-

-

1

6.25

Selaginaceae

-

-

+

+

2

12.50

Solanaceae

+

1

6.25

StercLiliaceae

-

-

-

-

-

-

+

-

-

-

-

-

+

-

-

-

2

12.50

Zygophyllaceae

-

-

-

-

-

-

+

-

-

-

-

+

-

-

-

3

18.75

Liliaceae

-

-

-

-

-

-

-

-

-

-

-

-

+

-

-

+

2

12.50

no. of flower fams

for each bee sp.

1

2

6

0

2

2

15

3

1

2

5

1

4

1

2

14

The records for the subgenera Helioplula and Pyganthopbora suggest differences in forage flower
choice. For example, sampling of flower visitors to Asteraceae has yielded five species of Helioplula
but no Pyganthopbora. On the other hand sampling of Zygophyllaceae has yielded four species of
Pyganthopbora but no Heliopbila. The association with Asteraceae exhibited by these five Heliopbila
species of the Old World Vestita Species Group taken together with the comment of Brooks ( 1988),
reviewing the accounts of Torchio and Youssef ( 1968) and Torchio (1971), that there is a definite
preference by the New World Estebana Species Group for Asteraceae, strengthens an association
between the subgenus Heliopbila and Asteraceae.

360

GESS AND GESS: NESTING AND FLOWER VISITING OF SOME SOUTHERN AFRICAN ANTHOPHORINI

Table 2. Flower families recorded as visited by 1 0 Antliophora spp., 5 each of the subgenera Heliophila and Pygimthophora.

together with the number and percentage of these species visiting each flower family (the species here represented
by numbers correspond to the species listed by number in Appendix I ).

Flower families

Antliophora species

no. of

spp.

% of spp.

Heliophila

Pyganthophora

H

P

H+P

17

18

19

20

21

22 23

24 25

26

Acanthaceae

-

-

-

+

-

-

-

-

1

0

1

10.00

Asteraceae

+

+

+

+

+

-

-

-

5

0

5

50.00

Boraginaceae

-

-E

+

+

+

-

+ +

+

4

3

7

70.00

Campanulaceae

-

-

-

-

+

-

-

-

1

0

1

10.00

Lamiaceae

-E

-

-

+

-

-

-

-

2

0

2

20.00

Mesembryanthemaceae

-

-

-

-

-

+

+ +

-

0

3

3

30.00

Sterculiaceae

-

-

-

-

+

+ +

+

1

4

5

50.00

Solanaceae

-

-

-

-

-

-

-

0

1

1

10.00

Zygophyllaceae

-

-

-

-

-

-E +

+

0

4

4

40.00

Haemodoraceae

-

_

_

-

4-

-

-

-

1

0

1

10.00

Iridaceae

-

-

-

-

+

-

-

-

1

0

1

10.00

no. of flower fams
for each bee sp.

2

2

2

4

6

2 2

4 4

3

Clearly there are too few records available to draw any firm conclusions. However, the differences
in flower choice between genera and also between subgenera (despite the lack of distinction between
pollen and nectar collection) indicate that the southern African Anthophorini do not as a rule practise
indiscriminate broad polylecty.

ACKNOWLEDGEMENTS

The authors wish to thank: Messrs D.W., H.W. and R.W.Gess for assistance in the field; Mr
T.C. White of the farm Hilton, MrE.Norton of the farm Clifton and Mr J.Willets of the farm Thursford,
all Grahamstown district, and Mr A.C.Truter of the farm Verdun, Kommadagga district for allowing
them access to their land; Mr K. van Zyl of the Cape Department of Nature and Environmental
Conservation for permission to work in the Goegap Nature Reserve, Springbok; Mr P. Gordon of the
National Parks Board for permission to work in the Richtersveld National Park; and Dr S. Milton and
Dr R.Dean of the Karoo Biome Research Station at Tierberg, near Prince Albert for their hospitality.

Appreciative thanks are given to Dr R.W. Brooks of Snow Entomological Museum, University
of Kansas, Lawrence and Dr J.G.Rozen of the American Museum of Natural History, New York for
reading and commenting on a earlier draft of the manuscript.

Gratitude is expressed to the Foundation for Research Development for a running expenses grant
to F.W.Gess.

REFERENCES

Brauns, H. 1913. Biologie sudafrikanischer Apiden. Zeitschiift fiir wissenschaftUche hisektenhiologie 9(4): I 16-120 and
(6/7): 190-193.

361

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 7 29 APRIL 1996

Brooks. R.W. 1988. Systematics and phylogeny of anthophorine bees (Hymenoptera: Anthophoridae: Anthophorini). The
University of Kansas Science Bulletin 53(9): 436-575.

Cardale, J. 1968a. Nests and nesting behaviour of Amegilla (Ainegllla) pulchra (Smith) (Hymenoptera: Apoidea:
Anthophorinae). Australian Journal of Zoology 16: 689-707.

Cardale. J. 1968b. Observations on nests and nesting behaviour of Amegilla (Asarapoda) sp. (Hymenoptera: Apoidea:
Anthophorinae). Australian Journal of Zoology 16: 709-7 1 3.

Cardale. J. 1968c. Parasites and other organisms associated with nests of Amegilla Friese (Hymenoptera: Anthophorinae).
Journal of the Australian Entomological Society!: 29-34.

Eardley. C.D. and Brooks. R.W. 1989. The genus Anthophora Latreille in southern Africa (Hymenoptera:
Anthophoridae l.CntD/nD/Dgy Memoir. Department of Agricultural Development. Republic of South Africa No. 76:
1-55.

Eardley. C.D. 1994. The genus Amegilla Friese (Hymenoptera: Anthophoridae) in southern Africa. Entomology Memoir.
Department of Agricultural Development. Republic of South Africa No. 91 : 1-67.

Ferton. C. 1902. Notes detachees sur i'instinct des Hymenopteres melliferes et ravisseurs (2me Serie) avec la description
d'une nouvelle espece. Annales de la Societe entomologitiue de France 71: 499-53 1 .

Ferton. C. 1920. Notes detachees sur I'instinct des Hymenopteres melliferes et ravisseurs (9e Serie) avec la description de
deux espece nouvelle espece. Annales de hi Societe entomologU/ue de France 89: 329-375.

Houston. T.F. 1991. Ecology and behaviour of the bee Amegilla lAsaropoda) dawsoni (Rayment) with notes on a related
species (Hymenoptera: Anthophoridae). Records of the Western Australian Museum 15(3): 591-609.

Marikovskaya. T.P. 1970. Contribution to the study of the social bees of the Alma-Alta Preserve. Trudy Alma-Atin.skovo
Cosndarstveanovo Zapovednika 9: 21 I- 216 (in Russian).

Maynard. G. 1992. Notes on nests of Amegilla (Asaropoda) sp. (Hymenoptera: Anthophoridae). Australian Entomological
Magazine 19( I ): 33-35.

MiOiENER.C.D. 1 960. Observations on the behaviour of a burrowing bee (Amegilla) near Brisbane. Queensland (Hymenoptera.
Anthophorinae). Queensland Naturalist 16: 63-68.

Rayment. T. 1935. A Cluster of Bees. Sydney: Endeavour Press.

Payment. T. 195 1 . A critical revision of species in the genus Asaropoda by new characters. Memoirs of the National Museum
of Victoria No. 17: 65-80.

Rozen. J.G. 1969. The biology and description of a new species of African Thyreus. with life history notes on two species of
Anthophora ( Hymenoptera: Anthophoridae). Journal of the New York Entomological Society 77( I ): 51 -60.

Torchio. P.F. 1971. The biology of Anthophora (Micranthophora) peritomae Cockerell (Hymenoptera: Apoidea.
Anthophoridae). Los Angeles County Museum Contributions in Science 206: 1-14.

Torc'IIIO. P.F. and You.SSEF. N.N. 1968. The biology of Anthophora (Micranthophora) fle.xipes und its cleptoparasite. Zuco.v/u/u
macuUita. including a description of the immature stages of the parasite (Hymenoptera: Apoidea. Anthophoridae).
Journal of the Kansas Entomological Society 41(3): 289-302.

Appendix 1

Catalogue of flower visiting records for the Anthophorini (all records Albany Museum collection
except those marked * which are taken from Eardley and Brooks 1989 and Eardley 1994).

abbreviations - colours: B = blue: BV = bluish violet; G = green; O = orange; Pi = pink; PiV = pinkish violet; Pu = purple;
PuV = purplish violet; V = violet; W = white; WY = cream; Y = yellow;
sex; F = female; M = male.

areas; CP = Cape Province; Nam. = Namibia; Nat. = Natal; Tvl = Transvaal; Richtersveld N Park =
Richtersveld National Park.

362

GESS AND GESS: NESTING AND FLOWER VISITING OF SOME SOUTHERN AFRICAN ANTHOPHORINI

Anthophorini
Ainegilla Friese

1. Ainegilla (Ackinonopsis) mimadvena (Cockerell)

Acanthaceae

Isoglossa Derst.

*/. sp

2. Ainegilla (Aframegilla) nubica (Lepeletier)

Acanthaceae

Monechma Hochst.

M. moUissimiim (Nees) P.G.Mey.

Pedaliaceae

Sesamiim L.

*S. sp.

3. Ainegilla (Ainegilla) capensis (Friese)

Acanthaceae

Asteraceae (Compositae)

Othonna L.

*0. sp.

Geigeria Griesselich
*G. sp.

Convolvulaceae

Convolvulus L.

*C. sp.

Lamiaceae

Plectrantlnis L’ Merit.

*P. sp.

Melastomataceae

Dissotis Benth.
sp.

Selaginaceae

Walafrida E.Mey.

*VF. sp.

4. Ainegilla (Ainegilla) natalensis (Friese)

Acanthaceae

Lamiaceae

5. Ainegilla (Megainegilla) acraensis (Fabricius)

Acanthaceae

Blepliaris Juss.

B. capensis (L. /. ) Pers.

B. capensis (L. /. ) Pers.

- - - ? Durban, Nat.; Cockerell, 1930

F 1 Richtersveld N Park, CP;

FW,SK&RWGess, 24.ix.95

- M 1 S Leonardoville, Nam.; VBWhitehead, 1 1 .ii.84

- - - Wolkberg, Tvl; RHWatmough, -.iv.74

; Cockerell, 1908

; Cockerell, 1908

- F 1 Angola; Cockerell, 1907

- F 2 Katberg Pass, CP; VBWhitehead, 25.xi.85

Wolkberg, Tvl; RHWatmough, -.iv.74

- F 2 Chimanimani Mts, Moz.; RHWatmough, 3. iv. 72

- F I NEdenburg,Tvl;MJohannsmeier,-.-.-

Wolkberg, Tvl; RHWatmough, -.iv.74
Wolkberg, Tvl; RHWatmough, -.iv.74

W F 1 Grahamstown, CP; DWGess, 3.ii.81
W M 3 Grahamstown. CP; DWGess, 3. ii. 81

363

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 7 29 APRIL 1996

Plumbaginaceae

*' Limoniwn Mill.

6. AmegiUa (Micramegilla) atrocincta (Lepeletier)
Acanthaceae

- M 1 Bredasdorp. CP; CDEardley. 15.xi.82

Blepharis Juss.

B. capensis (L. /.) Pers.

W

M

1

Grahamstown, CP; FWGess, 5.i.79

B. capensis (L./.) Pers.

W

F

6

Grahamstown, CP; FW&DWGess, 7.i.79

B. capensis (L. /i) Pers.

w

M

8

Grahamstown, CP; FW&DWGess, 7.i.79

B. capensis (L. /.) Pers.

w

F

2

Grahamstown, CP; FW&DWGess, I5.i,81

B. capensis (L./.) Pers.

w

F

4

Grahamstown, CP; FW&DWGess, 3.ii.8l

B. capensis (L. /'. ) Pers.

w

F

5

Grahamstown, CP; FW.DW&RWGess, 8.ii.81

B. capensis (L. /.) Pers.

w

F

2

Waterford, CP; RWGess, 25.xi.87

Asteraceae (Conipositae)

Berkheya Ehrh.

B. helerophylla {Th.) O.Hoifm. Y M 1 Grahamstown, CP; FW&SKGess, 20.xi.90
7. Aniegilla (Micraniegilla) iiiveata (Frie,se)

Acanthaceae

Blepharis Juss.

B. capensis iL.f.) Pens.

Monechma Hochst.

M. inollissiimini (Nees) P.G.Mey. Pi F 5

W F I Waterford, CP; RWGess, 25.xi.87

M. mollissinmm (Nees) P.G.Mey. Pi M I

*M. ,sp.

*M. sp.

Peri.stophe Nees
P. cernua Nees
Petalidium Nees
*P. sp.

“acanth"

F

M

Richtersveld N Park, CP;

FW,SK,&RWGess, 18-24.ix.95
Richtersveld N Park, CP; FW.SK.&RWGess,
l8-24.ix.95

Usakos, Nam.; RHWatmough. 5.vii.75
Usakos, Nam.; RHWatmough, 5.vii.75

V F 1 Grahamstown, CP; FWGess, 3. xii. 81

- F 2 Ou Lutzputz, CP; VBWhitehead, 3l.vii.84
W M 2 Richtersveld N Park, CP; FW,SK,&RWGess,
24.ix.95

Aizoaceae: Mesembryanthema (Mesembryanthemaceae)

MesenihrYanthennim L.

*M. cry.stalinum Hook.f.

*M. sp.

*M. sp.

*M. sp.

"mesem”

"mesem”

Asteraceae (Conipositae)

Berkheya Ehrh.

B. carHmfoUa (DC.) Roessler
*6. rigida (Thunb.) H.Bol. &
Wolley-Dod ex Adamson &
Salter

W F 3 SW Kakamas, CP; VBWhitehead, 6.ii.84
- F I Jakkalsputs, CP; VBWhitehead, 21. xi.75
W F I Klinghardtberge, Nam.; RHWatmough, 19.x. 74
O F 2 Aurusberg, Nam.; RHWatmough, 23.x. 74
W F 3 Montagu/Matroosberg, CP;SK&RWGess, 4.xii.86
W F 2 Matroosberg, CP; RWGess, 4. xii. 86

Y M 1 ENE Ceres. CP; HWGess, 29-30.xi.89

Y F 2 SE Hopefield, CP; JGRozen er«/. , I5.X.72

364

GESS AND GESS: NESTING AND FLOWER VISITING OF SOME SOUTHERN AFRICAN ANTHOPHORINI

Senecio L.

S. rosmarinifolius L. /
Boraginaceae

Anchusa L.

A. capensis Thunb.

-4. capensis Thunb.
Lohostemon Lehm.

L. sp.

Y F 2

B F 1
B F 1

B F 4

Capparaceae

Cleome L.

C. paxii (Schinz) Gilg & Ben. Y F 1

Maeriia Forssk.

M. schinzii Pax W M 1

Elatinaceae

Bergia L.

B. glomerata L.f. G F 1

Geraniaceae

Geranium L.

*G. sp. -FI

*C. sp. - M 1

Hydrophyllaceae
Codon L.

C. royenii L. W F 1

Lamiaceae (Labiatae)

“labiate” V F 2

“labiate” B F I

^“labiate” B F I

Malvaceae

-FI

Mimosaceae

Acacia Mill.

A. caffra (Thunb.) Willd. WY M I

Moringaceae

Moringa Adans.

*M. ovalifoUa Dinter & Berger - F I

Pedaliaceae

Harpagophvtum DC. ex Meissn.

*H. procumhens mmch.)DC. - F 1

ex Meissn.

Sesanuim L.

triphrlluin Welw. ex Aschers, - F 1

Oudtshoorn, CP; FW&RWGess, 7-8.xii.86

Kamieskroon, CP; FW&SKGess, 9.x. 94
Kamieskroon, CP; FW.SK&RWGess, 28.ix.95

Clanwilliam/Graafwater, CP; FW&SKGess,
4.x. 90

Richtersveld N Park, CP;
FW,SK&RWGess, 16.ix.95

Vioolsdrif, CP; FW&SKGess, 3.X.85

Grahamstown, CP; FW&SKGess, 20.xi.90

Aussenkehr, Nam.; VBWhitehead. 16.ix.83
Aussenkehr, Nam.; VBWhitehead, I6.ix.83

Richtersveld N Park, CP; FW,SK&RWGess,
I9.ix.95

O LI berg Pass, Montagu, CP; FWGess, 3.xii.86
Nieuwoudtville, CP; FW&SKGess, 3-8.X.89
Aurusberg, Nam.; RHWatmough, 23.x. 74

Klinghardtberge, Nam.; RHWatmough,
I9.X.74

Oudtshoorn, CP; RWGess, 9-l2.xii.86

Gobabeb, Nam.; MLPenrith&SLouw, 18.ii.-
20.iii,83

E Kimberley, CP; VBWhitehead, I9.i.84

Derm, Nam.; VBWhitehead, I0.ii.84

365

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 7 29 APRIL 1996

*5. sp. -FI

*S. sp. -FI

Rosaceae

Neitnulopsis Brem. & Oberm.

*/V. sp. - F 2

Scrophulariaceae

Aptosimum Burch.

A. indivisum Burch, ex Benth. BV F I

A. spinescens (Thunb.) Weber PuV F I

Peliostonmm Benth.

P. leiicorrhizuni E.Mey. V F I

ex Benth.

W Boshoff, CP; VBWhitehead, I9.i.84
Harmonie, CP; VBWhitehead, 25.ii.80

Leonardville, Nam.; VBWhitehead, I5.iv.8l

Springbok, CP; FW&SKGess, 8.x. 94
Springbok. CP; SKGess, I5-2I.X.87

Richtersveld N Park, CP; FW.SK&RWGess,
I9.ix.95

Sterculiaceae

Hermannia L.

H. diseniiifolia Jacq.
H. diseniiifolia Jacq.
Zygophyllaceae

Tribulus L.

Y F I Nieuwoudtville, CP; FW&SKGess, 29.ix.94

Y F I Kamieskroon, CP; FW&SKGess, 1 0.X.94

M

*T. sp.

*T. sp.

*T. sp.

Zygophylhim L.

Z. sp.

Aniegilla (Micrimiegilla) veliilina (Friese)

Aizoaceae: Mesembryanthema (Mesembryantheniaceae)
Psilocinilon N.E.Br.

N Kakamas, CP; VBWhitehead, 7.ii.84
N Kakamas, CP; VBWhitehead, 7.ii.84
W Mariental, Nam.; RGOberprieler, 22.iii.8.J

Y F I Nieuwoudtville. CP; FW&SKGess, 2.X.89

*P. sp.

*P. sp.

'IStoeheria sp.

Capparaceae

Cleoine L.

C. paxii (Schinz) Gilg & Ben.

- F I Noordoewer, Nam.; VBWhitehead, 7.iii.80
-M3 Noordoewer, Nam.; VBWhitehead, 7. iii. 80
W F 4 Richtersveld N Park. CP; FW.SK&RWGess.
I6.ix.95

Y F 13 Richtersveld N Park, CP; FW.SK&RWGess,
I6.ix.95

Geraniaceae

Cenmiiim L.

*C. sp.

*G. sp.

9. Amegilla {Zehnmiegilla} hee/nianensis (Cockerell)
Capparaceae

Cleoine L.

*C. aiigiistifolia Forssk.
W. Amegilla (Zcbraiiiegilla) caleiis (Lepeletier)
Acanthaceae

Blepliai is Juss.

B. cupensis (L./. ) Pers.

- F I Aussenkehr. Nam.; VBWhitehead, 16. ix. 83

- M 2 Aussenkehr, Nam.; VBWhitehead. 16. ix. 83

- F 1 nr Vivo, Tvl; CDEardley. 3.ii.84

W F 1 Grahumstown, CP; FWGess, 7.i.79

366

GESS AND GESS: NESTING AND FLOWER VISITING OF SOME SOUTHERN AFRICAN ANTHOPHORINI

*B. diversispina (Nees) C.B.CI,
Justicia L.

*J.flava (Vahl.) Vahl,
Asteraceae (Compositae)

Zinnia L.

*Z. sp.

I \ . Amegilla (Zehramegilla) fallax (Smith)

Acanthaceae

“acanth”

*?_

Asteraceae (Compositae)

Vernonia Schreb.

V. fastigiata Oliv. & Hiern

Campanulaceae

Lobelia L.

L. sp.

Lamiaceae

Plectranthus L’ Merit.

P. sp.

H=7.

*7_

*7.

Plumbaginaceae

Limonimn Mill.

*L. sp.

\2. Amegilla (Zehramegilla) langi (Cockerell)

Zygophyllaceae

Tribidus L.

*T. sp.

1 3. Amegilla (Zehramegilla) ohsciiriceps (Friese)

Lamiaceae

"labiate”

Liliaceae

Trachyandra
*T. muricata (L/.) Kunth
Papilionaceae

Melolohliim Ecklon & Zeyher

M. sp.

Sterculiaceae

Hermannia L.

*H. sp.

\4. Amegilla (Zehramegilla) penicida Eardley
Acanthaceae

Blepharis Juss.

F - Langjan Nat. Res.,Tvl.; CDEardley, 2.ii.84

F - Langjan Nat. Res., Tvl.; CDEardley, 2. ii. 84

F I Lobito Bay, Angola; Cockerell, 1932

Pi F I Morgan Bay,CP; FW&SKGess, 30.iii.-4.iv.89

- - Wolkberg, Tvl; RHWatmough,

- M I Blyderivierspoort Dam, Tvl; CDEardley,

25-26.X.84

B M I Cape Peninsula, CP; FW,SK&DWGess,

7.iii.93

- F 1 Katberg, CP; VBWhitehead, 25.xi.85

B F 3 nr Loskop Dam, Tvl; RHWatmough,

B M I nr Loskop Dam, Tvl; RHWatmough,

- - Wolkberg, Tvl; RHWatmough,

- M I Bredasdorp, CP; CDEardley, 15. xi. 82

- F I KoopanSuid, CP; VBWhitehead, 1 3.ii.84

B F 3 Nieuwoudtville, CP; FW&SKGess, 3-8. X. 89

F I Clanwilliam, CP; VBWhitehead, 23.viii.84

WY M I Grahamstown. CP; FWGess, I2.X.77

F I Beaufort West, CP; VBWhitehead, 24.xi.85

367

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 7 29 APRIL 1996

B. capensis (L./ ) Pers.

W

F 8

Grahamstown, CP; FW&DWGess, 7.i.79

B. capensis (L./) Pers.

w

M 1

Grahamstown, CP; FW&DWGess, 7.i.79

B. capensis (L.f.) Pers.

w

F 1

Grahamstown, CP; FWGess, 8.ii.8l

B. capensis (L.f) Pers.

w

F 2

Grahamstown, CP; FWGess. I0.ii.86

\5. Ainegilla (Zebramegilla) punctifrons (Walker)
Acanthaceae

Blepharis Juss.

B. capensis (L./.) Pers.

W

M

2

Grahamstown. CP; FW&SKGess, 27.x. 72

B. capensis (L.f) Pers.

W

F

1

Grahamstown, CP; DWGess, 3.i.79

B. capensis (L./.) Pers.

W

F

1

Grahamstown, CP; FW&DWGess, 7.i.79

B. capensis (L.f.) Pers.

w

M

7

Grahamstown. CP; FW&DWGess, 7.i.79

B. capensis (L.f.) Pers.

w

F

6

Grahamstown, CP; DW&RWGess, I5.i.81

B. capensis (L.f.) Pers.

w

F

3

Grahamstown. CP; DWGess, 3.ii.81

B. capensis (L.f) Pers.

w

F

3

Grahamstown, CP; FW&SKGess, 8.ii.8l

Aizoaceae: Mesembryanthema (Mesembryanthemaceae)

''mesein” WYM I Kommadagga, CP; FW&SKGe.ss, 23.X.83

\6. Ainegilla {Zebramegilla} spUostoma (Cameron)

Acanthaceae

Blepluirls Ju.s.s,

B. capensis {L.f.) Peru. W M 2 Grahamstown, CP; FW&SKGe.ss, 27.x. 72

Aizoaceae; Mesembryanthema (Mesembryanthemaceae)

Mesemhryanthemwn L.

*M. cry.rtaliiuim Hook./i
Psilocaiilon N.E.Br.

P. acutisepahtm (Berger)

N.E.Br.

WF I Onrusrivier, CP; VBWhitehead, I7.i.76

WPi F 2 Heerenlogement. CP; FW.SK&RWGess, 8.X.93

P. cf. subnodosum (Berger)
N.E.Br.

“mesem"

Asteraceae (Compositae)

Senecio L.

S. linifoliits L.

S. UnifoUus L.

S. UnifoUiis L.

S. Ilnifolliis L.

S. UnifoUus L.
Campanulaceae

Cypliia Berg.

C. sp.

Geraniaceae

Pelargonium L' Herit
P. myrrhifoUum Ait.

P. myrrhifoUum Ait.
Lamiaceae (Labiatae)

Acrotome Benth.

A. inflata Benth.

A. inflata Benth.

Pi F 1 Graafwater, CP; FW.SK&RWGess, 6.X.95
W F I Montagu/Matroosberg, CP; FWGess, 4.xii.86

Y F 2 Grahamstown, CP; CFJacotGuillarmod, 27.i.75

Y M 2 Grahamstown, CP; CFJacotGuiliarmod, 27.i.75

Y M 3 Grahamstown, CP; CFJacotGuillarmod, 31. i. 73

Y F I Grahamstown, CP; CFJacotGuillarmod, 2. ii. 73

Y M 3 Grahamstown, CP; CFJacotGuillarmod, 2.ii.75

PiV M I Grahamstown, CP; FWGess, 21. iii. 78

- F I Oudtshoorn, CP; CFJacotGuillarmod, 10. X. 72

- M I Oudtshoorn. CP; CFJacotGuillarmod, 10.x. 72

BV M 5 Grahamstown, CP; SKGess, 17. iii. 78
BV M I Grahamstown, CP; FWGess, 21. iii. 78

368

GESS AND GESS: NESTING AND FLOWER VISITING OF SOME SOUTHERN AFRICAN ANTHOPHORINI

Salvia dentata Ait. B F 2

*S. sp. -Ml

Liliaceae

Aloe sp. Y F I

Miniosaceae

Acacia Mill.

A. cajfra (Thunb.) Willd. WY F I

A. karroo Hayne Y F I

Papilionaceae (Fabaceae)

Aspalathus L.

A. chortophila Ecklon & Zeyher Y F 1

Psoralea L.

P. pinnata L. B M 2

P. pinnata L. B F 2

P. pinnata L. B M 3

Pedaliaceae

Sesannnn L.

*5. sp. -FI

Plumbaginaceae

Limoni lint Mill.

Claiiwilliam, CP; FW&SKGess, 4.x. 90
Queenstown, CP; VBWhitehead, 29.xii.83

Grahamstown, CP; FWGess, 6.i.8l

Oudtshoorn, CP; RWGess, 9-l2.xii.86
Colesberg. CP; DWGess, I9.i.85

Riebeek East, CP; FW&SKGess, I4.xi.92

Grahamstown, CP; CFJacotGuillarmod, 2.ii.75
Grahamstown, CP; CFJacotGuillarmod. 9.ii.75
Grahamstown, CP; CFJacotGuillarmod, 9.ii.75

W Boshoff, CP; VBWhitehead, 19.iii.84

*L. sp.

Proteaceae

Paranonnis Salisb.

P. hracteokirus Salisb.

ex Knight

Solanaceae

Lyciiim L.

L. sp.

Zygophyllaceae

Zygophylhiin L.

Z. sp.

- F I Bredasdorp, CP; CDEardley, I5.xi.82

Pi F I Nieuwoudtville, CP; FW&SKGess, 30.ix.90

V M I Grahamstown, CP; FW&SKGess, 29.ix.77
- F I Queenstown, CP; VBWhitehead, -.i.64

Y M 3 Nieuwoudtville, CP; FW&SKGess, 28.ix.90

Anthophora Latreille

M . Anthophora (Heliophila) labrosa Friese

Asteraceae (Compositae)

Berkheya Ehrh.

B. heterophylla (Th.) O.Hoffm.
B. heterophylla (Th.) O.Hoffm.
B. heterophylla (Th.) O.Hoffm.
B. heterophylla (Th.) O.Hoffm.
B. heterophylla (Th.) O.Hoffm.
B. heterophylla (Th.) O.Hoffm.
B, heterophylla (Th.) O.Hoffm.
B. heterophylla (Th.) O.Hoffm.
B. sp.

Y F 2 Grahamstown, CP; FWGess, 1 2.x. 72

Y F 1 Grahamstown, CP; FW&DWGess, 16.x. 72

Y M 1 Grahamstown, CP; FW&DWGess, I6.X.72

Y F 1 Grahamstown, CP; FWGess, 25.x. 72

Y M 2 Grahamstown, CP; FW&SKGess, 15. xi. 77

Y F 2 Riebeek East, CP; FW&SKGess, 22.xi.82

Y M I Riebeek East, CP; FW&SKGess, 22.xi.82
YM I RiebeekEast, CP; FW&SKGess, I6.X.83

Y F I Clanwilliam, CP; FW&SKGess, 9.x. 90

369

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 7 29 APRIL 1996

Senecio L.

S. linifoliiis L.

S. linifoliiis L.

Lamiaceae (Labiatae)

"labiate"

\S. Anthopliora (Heliophilaj praeco.x Frie.se

Asteraceae (Compositae)

Berkheya Ehrh.

B. carUnifoHa (DC.) Roessler
B. heterophyllci (Th.) O.Hoffm.
B. heterophylla (Th.) O.Hoffm.
B. heterophylla (Th.) O.Hoffm.
B. sp.

Pentzia Thunb.

P. incana (Thunb.) Kuntze
Boraginaceae

Anchusa L.

A. capensis Thunb.

A. capensis Thunb.

A. capensis Thunb.

\9.Anthophora (Heliophila) nifolanala Dours
Asteraceae (Compositae)

Berkheya Ehrh.

B. heterophylla (Th.) O.Hoffm.
B. heterophylla (Th.) O.Hoffm.
B. heterophylla (Th.) O.Hoffm.
B. heterophylla (Th.) O.Hoffm.
B. heterophylla (Th.) O.Hoffm.
B. heterophylla (Th.) O.Hoffm.
Pteronia L.

P. incana (Burm.) DC
P. Incana (Burm.) DC
Senecio L.

S. linifoliiis L.

S. linifoliiis L.

S. linifoliiis L.

S. linifoliiis L.

S. linifoliiis L.

Boraginaceae

Anchusa L.

A. capensis Thunb.

A. capensis Thunb.

20. Anthophora (Heliophila) vestita Smith
Acanthaceae

Peristrophe Nees
P. cermia Nees
“acanth"

Y F I Grahamstown. CP; CFJacotGuillarmod, 25.i.75

Y F I Grahamstown, CP; CFJacotGuiilarmod. 2.ii.75

VMS Ouberg Pass. Montagu. CP; FWGess, 3. xii. 86

Y M I Theronsberg Pass.Ceres, CP; FWGess, 29-30.xi.89

Y F 1 Grahamstown, CP; FW&SKGess, 15. xi. 77

Y F 4 Riebeek East, CP; FW&SKGess, 22.xi.82
YM I RiebeekEast. CP; FW&SKGess, 16.X.83

Y F I Clanwilliam, CP; FW&SKGess, 9.X.90

Y F p Prince Albert, CP; SKGess, 26. xi. -3. xii. 87

B F I Grahamstown, CP; FWGess, 1 8.xi.77
B M 2 Grahamstown, CP; FWGess, 1 8. xi. 77
B M 1 Kamieskroon, CP; FW&SKGess, I l.x.94

Y F 2 Grahamstown, CP; FWGess, 1 2.x. 72

Y M 4 Grahamstown, CP; FWGess, 1 2.x. 72

Y F 5 Grahamstown, CP; FW&DWGess, 1 6.X.72

Y M 1 Grahamstown, CP; FWGess, 1 6.x. 72

Y M 3 Grahamstown, CP; FWGess, 25.X.72

Y F 1 RiebeekEast, CP; FW&SKGess, 1 6.X.83

Y F 2 Kamieskroon, CP; FW&SKGess, 1 7.ix.92

Y M I Kamieskroon, CP; FW&SKGess, 17.ix.92

Y F 10 Grahamstown, CP; CFJacotGuillarmod, 25.i.75

Y F 1 Grahamstown, CP; CFJacotGuillarmod, 27.i.75

Y M I Grahamstown, CP; CFJacotGuillarmod, 27.i.75

Y F 2 Grahamstown, CP; CFJacotGuillarmod, 31. i. 75

Y F 4 Grahamstown, CP; CFJacotGuillarmod, 2.ii.75

B M I Grahamstown, CP; FWGess, 18.xi.77
B M 1 Leliefontein. CP; FW.SK&RWGess, 2.X.95

V F 1 Grahamstown, CP; FWGess, 3. xii. 8 1

Pi F 6 Morgan Bay, CP; FW&SKGess, 30.iii.-4.iv.89

370

GESS AND GESS: NESTING AND FLOWER VISITING OF SOME SOUTHERN AFRICAN ANTHOPHORINI

Asteraceae (Compositae)

Berkheya Ehrh.

B. heterophylla (Th.) O.HolTm.
B. heterophylla (Th.) O.HolTm.
B. heterophylla (Th.) O.HolTm.
B. heterophylla (Th.) O.HolTm.
B. heterophylla (Th.) O.HolTm.
B. heterophylla (Th.) O.HolTm.
B. heterophylla (Th.) O.HolTm.
B. heterophylla (Th.) O.HolTm.
B. heterophylla (Th.) O.HolTm.

B. heterophylla (Th.) O.HolTm.
Cirsium Mill, emend. Scop.

C. vidi’are (Savi) Ten.

C. viilgare (Savi) Ten.

Senecio L.

S. linifolhis L.

S. liuifolhts L.

S. linifolhis L.

S. linifolins L.

S. linifolins L.

S. linifolins L.

S. linifolins L.

5. sp.

Boraginaceae

Anchnsa L.

A. capensis Thunb.

A. capensis Thunb.

Lamiaceae (Labiatae)

Acrotome Benth.

A. inflala Benth.

/I. inflata Benth.

21 . Anthophora ( Heliophila) wartmunni Friese
Asteraceae (Compositae)

Arctotis L.

A. /rter/.v Thunb.

Berkheya Ehrh.

6. heterophylla (Th.) O.Hoffm.

B. heterophylla (Th.) O.Hoffm.
Melalaslu R. Br.

M. mnricuta (L.) D.Don
M. ninricata (L.) D.Don
Pteronia L.

P. sp. B
P. sp. B
Senecio L.

Y F 8 Grahamstown. CPtFWGess, I2.X.72

Y M I Giahamstowii. CP; FWGess, I2.X.72

Y F 3 Grahamstown, CP;FW&DWGess, I6.X.72

Y F 3 Grahamstown, CP; FWGess, 23. X.72

Y F I Graham.stown, CP;FW&SKGess, I3.xi.77
YM I Grahamstown, CP; FW&SKGess, 1 3.xi.77

Y F I RiebeekEast,CP;FW,SK&DWGess,22.xi.82

Y M I RiebeekEast,CP;FW,SK&DWGess,22.xi.82

Y F 2 Grahamstown, CP; FW&SKGess, 20.ix.90

Y M 3 Grahamstown, CP; FW&SKGess, 20. ix. 90

Pu F I Grahamstown, CP; SKGess, 9.iii.78
Pu M 2 Grahamstown, CP; SKGess, 9. iii. 78

Y F 3 Grahamstown. CP; CFJacotGiiillarmod, 23. i. 73

Y M I Grahamstown, CP; CFJacotGiiillarmod, 23. i. 73

Y F I Grahamstown. CP; CFJucotGuillarmod, 27.1.73

Y F 2 Grahamstown, CP; CFJacotGmIlarmod, 3 1.1.73

Y M I Grahatnstown, CP; CFJacotGiiillarmod. 3 1 .1.73

Y F 2 Grahamstown, CP; CFJacolGiiillarmod,2.ii. 73

Y M I Grahamstown, CP; CFJacotGiiillarmod. 2.ii.73

Y F I Grahamstown, CP; FWGess, 28. xii. 86

B F 3 Grahamstown. CP; FWGess. 1 8. xi. 77
B M 7 Grahamstown, CP; FWGe.ss. 1 8.xi.77

BV M I Grahamstown. CP; FWGess, 3. iii. 78
BV M 2 Grahamstown, CP; SKGess, I 7. iii. 78

Y F I Clanwilliam, CP; FW&SKGess, 3.x. 90

YM I Grahamstown, CP; FWGess. 1 2.X.72

Y M 2 Grahamstown. CP; FWGess, 1 6. x.72

Pi F I Nieuwoudtville. CP; FW&SKGess, 29. ix. 90

Pi F I Nieuwoudtville, CP; FW&SKGess, 25. ix. 94

Y F I Nababeep, CP; FWGess, 12-1 3.X.89

Y M I Nababeep. CP; FWGess, 1 2- 1 3.x, 89

371

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 7 29 APRIL 1996

S. linifhlius L.

S. linifoUiis L.

S. linifolius L.

Boraginaceae

Anchusa L.

A. capensis Thunb.

A. capensis Tluinb.
Campainilaceae

Wahleuhergia Schrad. ex Roth
W. sp.

Haemodoi'aceae

Wacheiulorfia Bunn.

W. sp.

Iridaceae

HoinerUi Vent.

Y M I Grahanistown.CPiCFJacotGuillarmod, 25.i.75

Y F I Grahamstown.CP;CFJacotGiiillamiod, 3l.i.75

Y F I Grahamstown, CP; CFiacotGuillarmod, 2.H.75

B M 1 Kamieskroon,CP;FW,SK&RWGess,28.ix.95

B M I Leliefontein. CP; FW.SK&RWGess, 2.X.95

V F I Nieuwoudtville, CP; FW&SKGess, .30.ix.90

Y F 2 NieuwoLidtville, CP; FW&SKGess, 29.ix.90

7/. sp. Y F I Nieuwoudtville, CP; FW&SKGess, 28.ix.90

Scrophulariaceae

Oftia Adans.

O. fl/nanifl (L.) Bocq. W F 1 Graafwater, CP; FW&SKGess, 2 1 .ix.92

22. Anthophoni ( Pyganihophom) ahrochia Eardley & Brooks

Aizoaceae; Mesembryanthema (Mesembryanthemaceae)

Herrea Schwant.

H. sp.

H. sp.

H. sp.

Sterculiaceae

Hennannia L.

H. disermifoUa Jacq.
H. disermifoUa Jacq.
H. disermifoUa Jacq.
H. disermifoUa Jacq.
H. disermifoUa Jacq.
22. Anthophora {Pyganthophora) hraunsiana Friese
Solanaceae

Lyciiim L.

Y F 2 Leliefontein, CP; FW&SKGess, 13.ix.92

Y F 1 Springbok. CP; FW&SKGess, 8.X.94

Y F 3 Springbok, CP; FW.SK&RWGess, 27.ix.95

Y F 2 Springbok, CP; FW&SKGess, 8-1 0.ix.92

Y F 1 Springbok. CP; FW&SKGess, 3.X.94

Y F 3 Kamieskroon, CP; FW&SKGess, 10-11. x.94

Y F 2 Kamieskroon. CP; FW.SK&RWGess. 28.ix.95

Y F 3 Kamieskroon, CP; FW.SK&RWGess, 3.X.95

V M 2 Grahamstown, CP; FWGess, 9.iii.78

Leliefontein, CP; FW&SKGess, 13. ix.92

L. sp.

Zygophyllaceae

Zygophylhim L.

Z. divaricatwn Ecklon & Zeyher Y F I
24. Anthophora (Pyganthophora) diversipes Friese

Aizoaceae: Mesembryanthema (Mesembryanthemaceae)
Herrea Schwant.

W, sp. Y F 4 Leliefontein, CP; FW&SKGess, 1 3. ix.92

Boraginaceae

Anchusa L.

A. capensis Thunb. B M 1 Leliefontein, CP; FW,SK&RWGess, 2.x. 95

Loboslemott Lehm.

372

GESS AND GESS: NESTING AND FLOWER VISITING OF SOME SOUTHERN AFRICAN ANTHOPHORINI

L. trichotomus DC.

B

F 1

E Pakhuis Pass, CP; DWGess, 3.X.9I

L trichotomus DC.

B

M 2

E Pakhuis Pass, CP; DWGess, 3.x. 91

L. sp.

B

F 1

Clanwilliam, CP; FW&SKGess, 3.X.90

Sterculiaceae

Herniannia L.

H. disennifolia Jacq.

Y

F 1

Kamieskroon, CP; FW&SKGess, 1 l.x.94

H. disennifolia Jacq.

Y

F 1

Kamieskroon. CP; FW.SK&RWGess, 28. ix.95

H. disennifolia Jacq.

Y

F 1

Kamieskroon, CP; FW.SK&RWGess. 3.X.95

H. sp.

Y

F 2

Graafwater, CP; FW.SK&RWGess, 8.X.95

Zygophyllaceae

Zygophylliim L.

Z. divaricatiim Ecklon & Zeyher

Y

F 1

Leliefontein, CP; FW&SKGess. I3.ix.92

25. Anthophora (Pyganthophora) krugeri Eardley & Brooks

Aizoaceae: Mesembryanthema {Mesembryanthemaceae)
Herrea Schwant.

H. sp.

H. sp.

H. sp.

Boraginaceae

Anchusa L.

A. capensis Thunb.

Sterciiliaceae

Henmumia L.

H. disennifolia Jacq.

H. diseriiufolia Jacq.

H. vestita Thunb.

H. sp.

Zygophyllaceae

Zygopliylliiin L.

Z. divaricatiim Ecklon & Zeyher
26. Anthophora (Pygcinthophora) scludtzei Friese
Boraginaceae

ILohostemon Lehin.

Y F 2 Leliefontein, CP; FW&SKGess, 1 3.ix.92

Y F 4 Springbok, CP; FW&SKGess, 4-8.X.94

Y F 4 Springbok, CP; FW,SK&RWGess. 27.ix.95

B F 3 Leliefontein. CP; FW.SK&RWGess.2.x.95

Y F 4 Springbok, CP; FW&SKGess, 4-8.X.94

Y F 2 Springbok, CP; FW.SK&RWGess, 27. ix.95

Y F I Ladismith. CP; FW&SKGess, 21. viii.95

Y F 1 Graafwater, CP; FW.SK&RWGess, 7.X.95

Y F 1 Leliefontein. CP; FW&SKGess, 13. ix. 92

?L. sp.

Sterciiliaceae

Henminnia L.

H. cf. ameifoUu Jacq.
H. disennifolia Jacq.
Zygophyllaceae

Zygophylliim L.

Z. cf morgsana L.

Z. cf morgsana L.

WV F I Nieuwoudtville. CP; FW&SKGess, 25.ix.94

Y F I Nieuwoudtville, CP; FW&SKGess, 30. ix. 94

Y F I Nieuwoudtville. CP; FW&SKGess, 25.ix.94

Y F I Nieuwoudtville, CP; FW&SKGe.ss, 24.ix.94

Y F I Nieuwoudtville, CP; FW&SKGess, 26. ix. 94

Anthophora ( Panmiegilla) leonis Cockerell
Boraginaceae

Anchusa L.

A. capensis Thunb.

B M I Kamieskroon, CP; FW&SKG. IO.x.94

373

INSTRUCTIONS TO AUTHORS

LAYOUT of recent issues should be examined for details of acceptable style and format.

MANUSCRIPTS should be submitted on floppy disk accompanied by three print-outs
(spacing VA or 2). In exceptional cases typed manuscripts will be accepted. These should be
submitted in triplicate, typed on A4 size paper, double-spaced, on one side of the paper only
and with wide margins all round.

Only generic and specific names should be underlined to indicate italics. All other marking
up should be left to the editor.

Footnotes should be avoided as far as possible.

All measurements should be metric.

Systematic papers should follow the rules of the International code of zoological
nomenclature or the International code of botanical nomenclature, as applicable.

ILLUSTRATIONS should bear the author’s name, title of paper and number in soft pencil on
the reverse side. Diagrams, line drawings and photographs are all regarded as figures, should
be termed as such and should be numbered in a single series.

In addition to the original illustrations a photocopy of each should be submitted.

Diagrams and line drawings should be reducible to a size not exceeding 14 x 16 cm (17,5 cm
including captions). Photographs for half-tone blocks should be final size, of high contrast
and printed on glossy paper. Colour illustrations are only printed under special circumstances
and their financing must be arranged by the author.

A scale should appear with all illustrations or the magnification should be given in the
caption.

A list of captions should be submitted on a separate sheet.

TABLES should be submitted on separate sheets. Foldouts and tables printed sideways should
be avoided.

REFERENCES should be listed alphabetically at the end of the paper. Abbreviations of
periodical titles should conform with those in the World list of periodicals (4th ed.). Layouts
must be as follows:

Alexander, J. 1969. The indirect evidence for domestication. In: Ucko, P. J. and G. W.
Dimbleby eds. The domestication and exploitation of plants and animals. London:
Duckworth, pp. 123-129.

De Winter, B. 1969. Plant taxonomy today. S.Afr.J.Sci. 65 (8): 229-242.

JuBB, R. A. 1967. Freshwater fishes of southern Africa. Cape Town: Balkema.

GALLEY PROOFS are sent (when practicable) to authors for checking printer’s errors.
Adjustments of language, style and punctuation and any other amendments of the original
manuscript, if made at this stage, may be charged to authors.

AUTHOR’S COPIES (50) are issued free of charge. When more than one author is involved
the fifty free copies will be sent to the senior author. Additional copies ordered before printing
are supplied at a reduced cost.

CADAR, P.E.-K887

'■‘jiy-'-x'slV

T

m-

>-•

la^

;^5

.5*

(51-1

: /\^13 ISSN 0570-1880

Annals of the

Cape Provincial Museums

Natural History

Ann. Cape Prov. Mus. (nat. Hist.)

\Volume 19 Part 8 20th August 1997

Two small Orthocladiinae (Chironomidae, Diptera)
from the Western Cape Province, South Africa

I

A.D. HARRISON

Published jointly by the Cape Provincial Museums
at the Albany Museum, Grahamstown, South Africa

ANNALS OF THE CAPE PROVINCIAL MUSEUMS

These Annals are published in two series, Natural History and Human Sciences, the latter series
covering cultural history, ethnology, anthropology and archaeology. They are issued in parts at
irregular intervals as material becomes available.

The primary objective of these Annals is to disseminate the results of research work undertaken by
staff members of the Cape Provincial Museums. However, a manuscript by any author is eligible
for publication provided it is based wholly or partly on the collection/s of one or more of these
museums or if the material on which the work is based is wholly or partly deposited in one of
these museums.

Parts are obtainable from the Librarian of the Albany Museum. Correspondence in connection
with these Annals should be addressed to the Editor, Albany Museum, Grahamstown 6140.

Editor

Dr F. W. GESS

Assistant Editor
Dr S. K. GESS

Two Small Orthocladinae (Chironomidae, Diptera)
from the Western Cape Province, South Africa

by

A. D. HARRISON

(Freshwater Research Unit, Zoology Department, University of Cape Town,
Rondebosch, 7700 South Africa.)

ABSTRACT

All life stages of two small Orthocladiinae from mountain streams and rivers in the Western
Cape Province are described. Notocladius capicola, new genus and species, is very abundant and is
able to feed on very small particles scraped off the substratum and must play an important part in the
food chain; Parakiefferiella biloba Freeman, here described in detail for the first time, is far less
abundant and its gut is packed with diatom frustules.

INTRODUCTION

The Chironomidae form an important part of freshwater communities, with many of their larvae
feeding on algae and making available the energy from primary production to larger invertebrates
and vertebrates. Workers in the field of freshwater research also find chironomid communities useful
as indicators of ecological changes taking place in the environment; this makes their correct
identification, specially of their larvae and pupae very important. This paper deals with all life stages
of two small members of the sub-family Orthocladiinae, one is very common in mountain streams
and rivers and its larva feeds on very small particles scraped from the rocks. Nevertheless, the adult
has not been seen before and shows such peculiar features that a new genus and species has had to
be created for it.

METHODS

Adults were caught by sweeping vegetation near streams and rivers; larvae and pupae were caught
in nets with fine mesh (< 0.5 mm). All specimens were preserved in 70% or 80% alcohol and mounted
in Canada Balsam dissolved in cellosolve. Measurements were made with an eyepiece micrometer
and all drawings with a drawing tube on the microscope.

Sources of generic definitions for Parakiefferiella are given in the text. Morphological
terminology is from Sasther (1980).

Abbreviations used in this paper are;

AR = antennal ratio. Ratio of length of apical flagellomere to combined length of basal
flagellomeres.

LR = leg ratio. Ratio of length of tarsomere 1 to length of tibia.

SV = ‘Schenkel-Schiene-Verhaltnis’. Ratio of femur plus tibiae to tarsomere 1.

BV = ‘Beinverhaltnisse’. Combined length of femur, tibia and tarsomere 1 divided by length of
tarsomeres 2 to 5.

375

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 8 20 AUGUST 1997

DESCRIPTIONS
Notocladius Gen, nov.

Definition

Adult male. Small midge with wing length of about 1.00 nun. Head: antenna with 10 flagellomeres,
groove beginning at antennomere 3, apex without large subapical seta. No frontal tubercles. Eye
hairy, without dorsomesal extension; one outer vertical seta. Tentorium widest at basal half; clypeus
normal, palp segment 3 with at least one subterminal sensillum.Thorax: antepronotal lobes separated
by a shallow notch, no scutal tubercle; acrostichals absent; dorsocentrals erect, uniserial; 3 prealars;
2 scutellars. Wing: membrane without setae, very transparent, punctuation very fine; no anal lobe;
R1 and R2+3, later joined by R4+5, fusing with costa to form clavus which terminates before half the
wing length; costa not extended; weak false vein from RM to wing tip; Cu straight; wing veins
without setae except costa; squama bare. Legs: spurs and comb normal, no pseudospurs; no sensilla
chaetica on tarsomere 1 of mid and hind leg; tarsomere 4 short and slightly cordiform; no pulvillae.
Hypopygium: no anal point, virga absent; no superior or inferior volsellae; gonostylus simple
without crista dorsalis.

Adult female. Wing length about 1 . 1 mm. Head: antenna with 5 flagellomeres, apex without large
subapical seta. Eye similar to male, 1 outer vertical seta, no frontal tubercles. Palp similar to male
except that segments 2 and 3 are fused. Thorax similar to male. Wings similar to male but clavus
comparably longer extending more than halfway to wing tip. Legs similar to male but sensilla
chaetica present on tarsomere 1 of mid and hind leg. Genitalia: gonopophysis VIII simple, not
divided into ventro-lateral and dorso-medial lobes; gonocoxapodeme of VIII and coxostemapodeme
of IX prominent; gonocoxite IX closely applied to body with one seta; postgenital plate of X
triangular. Seminal capsules with short necks, ducts convoluted with common opening. Cerci small.

Pupa. Small (1.4 - 1.6 mm). Cephalothorax; setae - frontal setae absent, 2 antepronotals, 3 small
precomeals, 2 widely spaced dorsocentrals. No thoracic horn. Abdomen: posterior single row of
small hooks on tergites III - V and on stemites V - VII. Shagreen of small posteriorly pointing spines,
none on tergite I, sparse on tergites II - IV, denser on V - VIII forming broad anterior bands;
segmental setae small, anal lobe with 3 short macrosetae and no fringe, small lateral denticles
present.

Larva. Small, up to 1.5 mm. long. Antenna: 4 segmented, blade extending to apex of segment 2,
Lauterbom organs small, style small. Labmm: seta SI bifid, remaining setae simple; no labial
lamella; pecten epipharyngis of 3 subequal scales; some chaetulae lateralis enlarged and filmy with
ciliary fringe; premandible with 2 teeth, one thin and pointed, the other broad and blunt; no bmsh.
Mandible: apical tooth with 4 inner teeth, seta subdentalis small, seta interna small but with many
branches. Mentum: 1 median and 6 pairs of lateral teeth, ventromental plate distinct but small, no
beard. Maxilla: palpiger normal, chaetulae small, no pecten galearis, lacinal chaetae large, antaxial
seta present. Body: parapods normal; procercus about as high as wide with strong setae.

376

HARRISON: TWO SMALL ORTHOCLADIINAE (CHIRONOMIDAE, DIPTERA) FROM SOUTH AFRICA

Notocladius capicola spec. nov.

This species is based on a large number of males and females, numerous pupae, some with pharate
males, and numerous larvae caught with the pupae and assumed to belong to this species.

Adult male (N=5 mounted and many unmounted)

As per generic definition.

Body length, up to 1.2 mm.

Wing length. 1.0 mm.

Colour. Head and antennae brown; thorax: background creamy yellow, scutal stripes separate
and dark brown, setal pits large and light, preepistemum dark brown, legs brown, wings very
transparent and colourless; abdomen: tergites I-V dark brown, VI and VII with a narrow brown
anterior stripe and the rest yellowish, VIII and hypopygium brown.

Head (Fig. I). AR0.25, lOflagellomeres, apical flagellomere somewhat swollen (Fig. 2), terminal
setae small and colourless, mostly sensilla chaetica; eyes hairy; clypeus not swollen; palp segments
16, 12, 28, 62, 102, pm; 1 subapical sensilla on segment 3.

Thorax. Setation: antepronotals 0-1, dorsocentrals 9-11, prealars 3, 1 scutellar per side.

Wings (Fig 3). Anal lobe absent, squama bare, length of clavus/winglength (including
brachiolum) 0.3. A false vein from clavus almost to wing tip and a second between this and
M]+2- Setation: brachiolum 1, no setae on R or M veins.

Legs. Tarsomere 4 short and slightly cordiform (Fig 4). LR fore 0.85, mid 0.63, hind 0.68; SV
fore 2.5 , BV fore 4.0 . No sensilla chaetica on tarsomeres.

Hypopygium (Figs 5, 6 and 7). Anal point absent but 6 anal setae present (Fig. 5). Inferior volsella
absent but with row of strong setae in the position where it normally appears in other Orthocladinae
(Fig. 6). Gonostylus comparatively small but with large megaseta. Fig. 7 shows the apodemes.

Adult female (N=6 mounted, many unmounted)

As per generic definition.

Body length. 1.0 mm.

Wing length. 1.1 mm.

Colour. Similar to male but abdominal tergites unifonnly brown.

Head. AR 0.4, 5 flagellomeres. Head setae: 1 outer vertical per side. Palps: segments 2 and 3
are fused but can still be distinguished, 3 has a small terminal projection, lengths 28, 25, 59, 53,
93 pm. 1 subapical sensilla on 3.

Thorax. No scutal tubercle. Setation: lateral antepronotals nil, dorsocentrals 10, posterior prealars
3, scutellars 2 per side.

Wings. Similar to male but clavus longer, length of clavus/wing length (including brachiolum)
0.48, extending more than half-way to wing tip.

Legs. General structure similar to male including tarsomeres 4. LR fore 0.7, mid 0.6, hind 0.7.
Sensilla chaetica on tarsomere 1, midleg 12, hindleg 7.

Genitalia (Figs 8-10). Gonocoxite not divided into ventrolateral and dorsomedial lobes, but
apodeme lobe appears to be present (Fig. 8A); gonocoxapodeme (8B) is dark and slightly curved;
coxostemapodeme of IX (8C) dark, prominent and strongly bent; gonocoxite IX closely applied to
body with one seta (Figs 8 and 9); coxae small; tergite X with 2 low posterior protruberances each
with a long seta (Fig. 9). Seminal capsules oval, light brown, with short necks, ducts convoluted with
common opening (Fig. 10).

377

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 8 20 AUGUST 1997

Pupa (numerous specimens mounted some with pharate males or females)

As per generic description.

Colour. Greenish in life but occurs in small tent-like light grey tube or cocoon. Exuviae are very
light and transparent.

Cephalothorax. Dorsal surface covered with small tubercles; no thoracic horn. Setation: no
frontal setae, 2 antepronotals, 3 small subequal precomeals, two widely spaced dorsocentrals probably
representing one member each of the Dl,2 and D3,4 pairs.

Abdomen (Figs 1 1 and 12). Fine shagreen on tergites II-IV and denser, broad anterior bands on
V-VIII, no shagreen in conjunctives. Setation as in Fig. 11. Posterior hook rows on tergites III to V
and stemites V to VIII. Anal lobe with 3 short macrosetae subterminal and 2 terminal setae, no fringe
(Fig. 12).

Larva (numerous larvae mounted and unmounted)

Similar to generic definition

Colour. Dark green in life with head capsule and posterior claws appearing black.

Body length. Up to 1.5 mm, but difficult to measure as preserved larvae are invariably curved.
Head capsule. 300-312 pm.

Antenna (Fig. 13). AR 0.6, ring organ in basal third of segment 1, blade reaching nearly to base
of segment 3, Lauterbom organs and style small.

Labruni (Fig. 14). SI bifid but branches usually closely applied making this difficult to see, pecten
epipharynx with 3 subequal teeth, first pair of chaetulae lateralis scalelike, but others large and filmy,
fringed with ciliary projections forming part of a filtering apparatus with the modified
prementohypopharyngeal complex. Premandible (Fig. 15) with one thin pointed tooth and one broad
blunt tooth, without bmsh.

Mandible (Fig. 16).Teeth dark brown, seta subdentalis pointed, seta interna small with many fine
branches.

Mentum (Fig. 17). Somewhat elongate, teeth brown, median tooth slightly shorter than first pair
of lateral teeth in unworn specimen, ventromental plate distinct but narrow.

Maxilla (Fig 18). As in generic definition.

Body. Claws of anterior parapod serrated towards tip, postparapod claws simple, small and dark
brown, setae of procercus dark brown, only one pair of small setae in anal region. Body setae appear
to be absent.

Specimens examined. Adults: numerous males and females were netted in the evening or caught in
light traps beside the Elands River (33°44'S, 19°06'S), 27. iv. 1996, and netted drowned in the Elands
River and in the mountain zone of the Riviersonderend (34°03"S, 19°04'E) during ecological work
in 1994-95. Numerous pupae and larvae were available from R.Tharme’s ecological work in the
Elandsrivers, the Dutoitsldoof (33°56'S, 19°10'E), the Molenaars (33°43'S, 19°10'E), in the
Dutoitskloof Mountains and the mountain zone of the Riviersonderend during 1994-95, all in the
Western Cape Province.

Holotype and paratype males, paratype female and pupal and larval material, all mounted on
slides, deposited in the Albany Museum, Grahamstown, Eastern Cape Province.
Comments. This genus resembles species in the Corynoneura-gxoup by the presence of a clavus in
the wings of both sexes. According to Saether and Kristoffersen (1996) there is a distinct sexual
dimorphism in the formation of the clavus in the Corynoneura-growp. In males R4+5 plays no part in

378

HARRISON: TWO SMALL ORTHOCLADUNAE (CHIRONOMIDAE, DIPTERA) FROM SOUTH AERICA

Figs 1- 8. Notocladius capicola. Adult male: 1. head; 2. antenna, apical flagellomere; 3. wing; 4. tarsomeres 3, 4, 5;
5. anal segment; 6. hypopygium; 7. apodemes. Adult female. 8. genitalia, ventral, A apodeme lobe,

B gonocoxapodeme, C coxostemapodeme.

379

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 8 20 AUGUST 1997

Figs 9 - 18. Notocladius capicola. Adult female; 9. genitalia, lateral; 10. seminal capsules and ducts. Pupa: 11. abdomen;
12. anal lobe. Larva: 13. antenna; 14. labrum; 15. premandible; 16. mandible; 17. mentum; 18. maxilla.

380

HARRISON: TWO SMALL ORTHOCLADIINAE (CHIRONOMIDAE, DIPTERA) FROM SOUTH AFRICA

its formation but in certain females the base of this combined vein is thickened and in some it is fused
to the clavus, as in both sexes of Notocladius. These authors consider that the “false vein” arising
from the thickened base of R4+5 is in reality the extension of this vein and that the additional “false
vein” between R4^5 and M may represent a vestige of Mj. Their interpretation can be applied to the
wing of Notocladius but it is not suggested that this genus falls into the Corynoneura-grou^. This
type of wing structure appears to be associated with small body size.

Neglecting wing structure, Notocladius keys to somewhere near Lopescladius in the key of
Cranston et al. (1989). The pupae do not fit in anywhere in the key in Coffman et al. (1986) but the
larvae key into the Orthocladius-Cricotopus group in Cranston etal. (1983) because of the bifid seta
S I and the structure of the mentum. Of course, for practical reasons, keys are based on apomorphic
characters which have been developed as a result of ecological specialisation.

Ecology. The tiny larvae and pupae often occur in large numbers on the surface of stones and rocks
in the fast current in second or third order mountain streams and rivers in the Western Cape Province.
The dark green bodies and almost black head capsules of the free-living larvae make their comma-
like appearance most distinctive. They appear to be scrapers but their gut contains very few diatoms,
mostly amorphous matter; the filter-like mechanism of their mouth parts should help to gather very
small particles scraped off the surface, making them an important part of the food chain. On pupation
the larvae make tent-like cases mostly of silk-like fibres incorporating very little detritus, one end is
broader than the other and open and presumably faces downstream; no larval remains were found in
any of the large number of pupal cases examined.

Adult males swarm at dusk which proved to be a good time to net both males and females; both
were also caught in light traps.

So far N. capicola has only been found in the soft acid waters of the Table Mountain Sandstone
system of the Western Cape Province.

Parakiefferiella biloba Freeman

Eukiejferiella (Parakiefferiella) biloba. Freeman 1953
Nanocladius biloba. Freeman 1956
Parakiefferiella biloba. Freeman and Cranston 1980

The material studied consists of males, females, pupae, one with mature pharate female, and
larvae. Most specimens came from a small mountain stream on the Cape Peninsula which was reduced
to a trickle during the dry summer season. The fauna of the stream was very depauperate with very
few orthoclad species (including only one species of Parakiefferiella, P. biloba) being present. It was
therefore considered safe to assume that the only larvae of this genus present in a large numbers of
samples must be of this species.

Freeman (1953, 1956) describes the male but a more detailed description is given here.

Adult male (N = 5 mounted, numerous species unmounted)

Differs somewhat from the generic definition in Cranston et al. (1989).

Body length. 2.00 mm.

Wing length. 1.00 mm.

Colour. Unmounted specimens head, antennae and palps light brown; thorax; scutal stripes and
pre-epistemum light brown, background and legs yellowish, wings very light brown; abdomen and

381

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 8 20 AUGUST 1997

hypopygium fairly uniform light brown.

Head. AR 0.17,12 flagellomeres, 1-3 wider than the rest except for 1 1 and 12 which form a club,
which appears bilobed because of its irregular shape and the prominent groove (Fig. 19), numerous
sensilla chaetica on apical flagellomere; no frontal tubercles; eyes bare, small with no dorsal extension,
tentorium (Fig. 20) wider ventrally. Head setation: 3 postorbital setae; palp segments 16, 22, 34, 47,
59 pm, 2 subapical sensillae on segment 3, no segment with apical extension.

Thorax. Scutal tuft of microtrichia, diagnostic for the genus, is weak and lies in a small pale
patch, but also includes two very small acrostichal setae. Setation: lateral antepronotals absent,
dorsocentrals 9, posterior pre-alars 3, scutellars 2 per side.

Wings (Fig. 21). Similar to Freeman’s figures but he does not show that the retracted costa and
R veins are thickened; the costa is produced beyond R4+5; R2+3 runs close to R4+5 and does not end
clearly in the costa. Setation: R 1, Ri nil, R4^j 1 at tip. There is no anal lobe and one squamal setae
(Freeman’s specimens had 0-2 squamal setae).

Legs. All tarsomeres cylindrical. LR fore 0.9, mid 0.4, hind 0.5. SV fore 2.6, BV fore 2.8. No
sensilla chaetica on tarsomeres 1 of midleg and hindleg.

Hypopygium (Figs 22, 23 and 24). Similar to Freeman’s figures but he has not shown the virga
that is also found in most species of this genus; the small anal point has a large seta on either side
(Fig. 22), the inferior volsella is digitiform; the gonostylus (Fig. 23) is not as curved as that illustrated
by Freeman but its shape is largely a matter of aspect. The apodemes are shown in Fig. 24.

Adult female (N = 5 mounted, many unmounted)

Similar to generic definition in Saether (1977), except for structure of gonopophysis VIII.

Body length. 1.3 mm.

Wing length. 1 mm.

Colour. Similar to male.

Head. AR 0.5, 5 flagellomeres, last somewhat swollen; no frontal tubercles; eyes similar to male;
setation 2 outer verticals, 2 postorbitals. Palp segments 16, 22, 34, 47, 62 pm; 2 subapical sensillae
on segment 3.

Thorax. Scutal patch smaller than that of the male but also with 2 small acrostichal bristles;
setation: lateral antepronotals nil, dorsocentrals 8, posterior pre-alars 3, scutellars 1 or 2 per side.

Wings. Similar to male; setation: brachiolum 1, R 5, Rj 1, R4+g 2, Squama 1 or 2.

Legs. LR fore 0.7, mid 0.4, hind 0.5. No sensilla chaetica on any tarsomeres.

Genitalia (Figs 25, 26 and 27). Gonopophysis VIII divided into a small dorsomesal lobe (Fig.
25 A) and a much larger ventrolateral lobe (Fig. 25 B); the small structure below the dorsomesal lobe
in the figure may be an apodeme lobe; gonocoxapodemes light in colour and joined (Fig. 25 C),
coxostemapodemes light but prominent (Fig. 25 D); tergite IX with two small posterior protrusions
each with one seta; gonocoxite IX welldeveloped with two long and two short setae (Figs 25 and 26),
segment X normal, postgenital plate small and rounded, cerci large. Seminal capsules ovoid, brown
but with a clearly demarcated light area at junction with spermathecal ducts which are long and
convoluted and open close to each other (Fig. 27).

Pupa (N = 3 mounted)

Does not follow generic definition of Coffman et al. (1986) in all respects.

Colour. Exuviae are fairly uniform light yellow, shagreen on abdomen darker.

382

HARRISON: TWO SMALL ORTHOCLADIINAE (CHIRONOMIDAE, DIPTERA) FROM SOUTH AFRICA

Cephalothorax. Dorsal surface finely pebbled with sculpturing becoming progressively finer
laterally. Frontal setae long, not on tubercles (Fig. 28), 2 median antepronotals, 3 small, short subequal
precorneal, 3 dorsocentrals, 1 anterior, and pair posterior; thoracic horn with a few terminal and
subterminal spines (Fig. 29).

Abdomen (Fig. 30). Shagreen on tergites I-VII of small posteriorly pointing spines with larger
spines forming anterior band and much larger spines forming narrow posterior band, fine, anteriorly
pointing spines in conjunctives 11-111 to VI-VIl (Fig. 31) ; no hook row on II; some short spines on
stemites IV-VII. No pedes spurii A or B. Abdominal setation as in Fig. 30, no fringe of taeniate setae
on any segment. Anal lobe truncated at apex, not tapering as in other species of this genus; 3 anal
macrosetae.

Larva (N = 5 mounted, many unmounted)

Similar to generic definition in Cranston et al. (1983) but differs in some respects

Colour. Head capsule light brown, claws very light, body yellowish when preserved.

Body length. \. 5 2.0 mm.

Head capsule length. 169 195 pm.

Antenna (Fig 32). AR 0.8; 6-segmented, segment 6 hair-like; blade reaching to base of segment
3 which is very short ( 1 .9 pm) and completely surrounded by large Lauterbom organs, style could
not be detected.

Labruni (Fig. 33). SI with 3 branches, remaining S setae simple; no labral lamella; pecten
epipharynx of 3 subequal scales; premandible with 2 apical teeth (Fig. 34).

Mandible (Fig. 35). Teeth brown, short apical tooth and 4 inner teeth; seta subdentalis and seta
interna both present.

Mentiim (Fig. 36, unworn specimen). Teeth brown, 2 median teeth and 5 lateral teeth, there is a
sign of an additional lateral tooth adpressed to each median tooth; ventromental plate very small.

Maxilla (Fig 37). Palpiger normal with numerous small lamellae at base, mostly behind the
palpiger in the figure; a pecten galearis could not be detected; the base of an antaxial seta present but
no seta; seta maxillaris simple.

Body. Parapods normal; procercus 2x as long as wide with 8 long setae; some simple body setae
up to 62 pm long; anal tubules shorter than parapod and somewhat conical in shape.

Specimens examined. Adults: numerous males and females from small waterfall, tributary of
Silvermine River, Cape Peninsula, 10 and ll.xi.l995, 24. ii. 1995, l.xii.l995, 9.1.1996; 1 female,
Kalk Bay waterfall. Cape Peninsula, 20.xii.l994; numerous males and females. Elands River, Du
Toil’s Kloof (33°44"S, 19°06'E), 27. iv. 1996; 1 male upper Riviersonderend (34°03'S, 19°04'E),
8.iv.l995. Pupae: small waterfall, tributary of Silvermine River 2 exuviae 24. xi. 1995, 1 pupa with
pharate female l.xii.l995. Larvae: numerous from the tributary of Silveraiine River at dates above.

Comments. The adult male, pupa and larva all differ from the generic definitions to some extent; the
male in that it has small acrostichal setae in the thoracic tuft of microsetae, the squama of the wing
is not bare; the pupa in that the anal lobes are truncated at the apex and not extended into tails; the
larvae in that the Lauterbom organs of the antennae are large and not small and the antennal style is
absent or much reduced and not strong. There are a few other minor differences, but none seem to
warrant a new genus being erected. It should be noted that neither the male nor the female adult has
the apical projection on palp segments as seen in the African species discussed by Ferrington and
Saether (1995).

383

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 8 20 AUGUST 1997

Figs 19- 28. Parakiefferiella biloba. Adult male: 19. flagellomeres 1 1 and 12; 20. tentorium; 21. wing; 22. anal point;

23 hypopygium; 24 apodemes. Adult female. 25. genitalia, ventral, A dorsomesal lobe, B ventrolateral lobe.
C gonocoxapodeme, D coxostemapodeme; 26 genitalia, lateral; 27. seminal capsules and ducts. Pupa:
28.frontal setae.

384

HARRISON: TWO SMALL ORTHOCLADIINAE (CHIRONOMIDAE, DIPTERA) FROM SOUTH AFRICA

Figs 29 - 31 .Parakiefferiella biloba. Pupa: 29 thoracic horn; 30. abdomen; 31. conjuctive II-III. Larva. 32. antenna;
33. labrum; 34.premandible; 35. mandible; 36. mentum; 37. maxilla.

385

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 8 20 AUGUST 1997

Ecology. P. biloba is found in first or second order mountain streams and small rivers. The larvae
from the small waterfall were inhabiting the moss and Scirpus digitatus in the water flow.

Distribution. Cape Peninsula mountains. Western Cape Province Fold Belt Mountains and source
streams of the Tugela River, Drakensberg Mountains, KwazuluNatal.

ACKNOWLEDGEMENTS

The author wishes to thank Miss Rebecca Tharme, Water Research Unit, University of Cape
Town, for help in collecting specimens.

REFERENCES

Coffman, W.P., Cranston, P.S., Oliver, D.R. and S,ether, O. A. 1986. The pupae of Orthocladiinae (Diptera: Chironomidae) of
the Holarctic region keys and diagnoses. In: Wiederholm T. ed. Chironomidae of the Holarctic region. Part 2. Pupae.
Ent. scand. Suppl. 28 pp. 147-296.

Cranston, P.S.. Oliver, D.R. and S.€THER, O.A. 1983. The larvae of Orthocladiinae (Diptera: Chironomidae) of the Holarctic
region keys and diagnoses. In: Wiederholm T. ed. Chironomidae of the Holarctic region, Part 1. Larvae. Ent. scand.
Suppl. 19 pp. 149-291

Cranston, P.S., Oliver, D.R. and SyCTHER, O. A. 1989. The adult males of Orthocladiinae (Diptera: Chironomidae) of the
Holarctic region keys and diagnoses. In. Wiederholm, T. ed. Chironomidae of the Holarctic region. Part 3. Adult
males. Ent. scand. Suppl. 34 pp. 165-352.

Ferrington, L.C. and S.«ther, O.A. 1995. Afrotropical species of Parakiefferiella Thienemann, with a review of species with
palpal projections. In: Cranston, P. ed. Chironomids: from genes to ecosystems CSIRO Australia. Pp 369-379.

Freeman. P. 1953. Chironomidae from Western Cape Province II. Proc. r. Entomol. Soc. bond. (B) 22: 201-213.

Freeman, P. 1956. A study of the Chironomidae (Diptera) of Africa south of the Sahara, Part 2. Bull. Brit. Mus. (Nat. Hist.)
Ent. 4: 287-368.

Freeman, P. and Cranston, P.S. 1980.Family Chironomidae. In: Crosskey, R.W. ed.. Catalogue of the Diptera of the Afrotropical
Region. London: British Museum (Natural History), pp. 175-202.

S.ETHER, O. A. 1977. Female genitalia in Chironomidae and other Nematocera. Bull Fish. Res. Bd Canada 197: 1-209.

S/Ether. O. a. 1980. Glossary of chironomid morphology terminology (Diptera: Chironomidae). Ent. Scan. Suppl. 14: 1-51.
S/ETUER, O. A. and Kristofferson , L. 1996. Chironomids with “M-fork”. A re-evaluation of the wing venation of the
Corynoneura-group (Insecta, Diptera, Chironomidae). Spixiana 19: 229-232.

386

INSTRUCTIONS TO AUTHORS

LAY OUT of recent issues should be examined for details of acceptable style and format.

MANUSCRIPTS should be submitted on floppy disk accompanied by three print-outs
(spacing VA or 2). In exceptional cases typed manuscripts will be accepted. These should be
submitted in triplicate, typed on A4 size paper, double-spaced, on one side of the paper only
and with wide margins all round.

Only generic and specific names should be underlined to indicate italics. All other marking
up should be left to the editor.

Footnotes should be avoided as far as possible.

All measurements should be metric.

Systematic papers should follow the rules of the International code of zoological
nomenclature or the International code of botanical nomenclature, as applicable.

ILLUSTRATIONS should bear the author’s name, title of paper and number in soft pencil on
the reverse side. Diagrams, line drawings and photographs are all regarded as figures, should
be termed as such and should be numbered in a single series.

In addition to the original illustrations a photocopy of each should be submitted.

Diagrams and line drawings should be reducible to a size not exceeding 14 x 16 cm (17,5 cm
including captions). Photographs for half-tone blocks should be final size, of high contrast
and printed on glossy paper. Colour illustrations are only printed under special circumstances
and their financing must be arranged by the author.

A scale should appear with all illustrations or the magnification should be given in the
caption.

A list of captions should be submitted on a separate sheet.

TABLES should be submitted on separate sheets. Holdouts and tables printed sideways should
be avoided.

REFERENCES should be listed alphabetically at the end of the paper. Abbreviations of
periodical titles should conform with those in the World list of periodicals (4th ed.). Layouts
must be as follows:

Alexander, J. 1969. The indirect evidence for domestication. In: Ucko, P. J. and G. W.
Dimbleby eds, The domestication and exploitation of plants and animals. London:
Duckworth, pp. 123-129.

De Winter, B. 1969. Plant taxonomy today. S.AfrJ.Sci. 65 (8): 229-242.

JuBB, R. A. 1967. Freshwater fishes of southern Africa. Cape Town: Balkema.

GALLEY PROOFS are sent (when practicable) to authors for checking printer’s errors.
Adjustments of language, style and punctuation and any other amendments of the original
manuscript, if made at this stage, may be charged to authors.

AUTHOR’S COPIES (50) are issued free of charge. When more than one author is involved
the fifty free copies will be sent to the senior author. Additional copies ordered before printing
are supplied at a reduced cost.

CADAR, P.E.-K887

SMITHSONIAN INSTITUTION LIBRARIES

3 9088 01429 5703

ISSN 0570-1880

I

I I

!

Annals of the

I

Cape Provincial Museums

Natural History

'Ann. Cape Prov. Mus. (nat. Hist.)

Volume 19 Part 9 20th September 1997

Phylogenetic Systematics of the Family Teloganodidae
(Ephemeroptera: Pannota)

W. P. MCCAFFERTY and T. Q. WANG

Published jointly by the Cape Provincial Museums
at the Albany Museum, Grahamstown, South Africa

ANNALS OF THE CAPE PROVINCIAL MUSEUMS

These Annals are published in two series. Natural History and Human Sciences, the latter series
covering cultural history, ethnology, anthropology and archaeology. They are issued in parts at
irregular intervals as material becomes available.

The primary objective of these Annals is to disseminate the results of research work undertaken by
staff members of the Cape Provincial Museums. However, a manuscript by any author is eligible
for publication provided it is based wholly or partly on the collection/s of one or more of these
museums or if the material on which the work is based is wholly or partly deposited in one of
these museums.

Parts are obtainable from the Librarian of the Albany Museum. Correspondence in connection
with these Annals should be addressed to the Editor, Albany Museum, Grahamstown 6140.

Editor

Dr F. W. GESS

Assistant Editor
Dr S. K. GESS

Phylogenetic Systematics of the Family Teloganodidae
(Ephemeroptera: Pannota)

by

W. P. MCCAFFERTY and T. Q. WANG
(Department of Entomology, Purdue University,

West Lafayette, IN 47907 USA)

ABSTRACT

Teloganodidae stat. nov. is recognized at the family rank and shown to be a sister group to
Ephemerellidae. Cladistic analysis of monophyletic species groups was performed, and the most
parsimonious cladogram is presented and discussed in terms of evolutionary trends and instances of
homoplasy. There are two major clades; these are recognized as the subfamilies Austremerellinae
subfam. nov. and Teloganodinae. Genera are defined cladistically by autapomorphies, and
sequencing conventions within the context of a strictly phylogenetic classification allow their
recognition at this rank. Two genera are recognized in Austremerellinae: Austremerella from
Australia and Vietnamella from southeastern Asia. Six genera are recognized in the Teloganodinae:
Ephemerellina, Nadinella gen. nov., Lithogloea, and Lestagella from southern Africa, and
Macafertiella and Teloganodes from southern Asia. Previous interpretations of Austremerella and
Lithogloea as subgenera of Ephemerellina were based on symplesiomorphies and are untenable. Each
genus is distinctive in both larval and adult stages, and a stage-correlated key is provided. Descriptive
and geographic accounts for each higher taxon are given, and known species listed. The new genus
Nadinella contains two nominal species: N. crassi (Allen and Edmunds) comb. nov. and N. brincki
(Demoulin) comb. nov. Australian and South African genera are shown to be archaic phylogenetic
relicts with origins in Gondwanaland prior to the initial breakup of that landmass. It is hypothesized
that the presence of Oriental lineages has resulted from Australasian interchange in the case of
Austremerellinae, and Africa to Asia transport via the Indian subcontinent in the case of
Teloganodinae.

INTRODUCTION

The family Teloganodidae stat. nov. is a group of pannote mayflies (McCafferty and Edmunds
1979) known from South Africa, the Orient, and Austrdia. These mayflies have not been generally
well known in the past, and incomplete and sometimes erroneous information has resulted in a history
of divergent interpretations. There are only 15 species currently described in the Teloganodidae.
This may in part be attributed to the relict nature of the older Gondwanan element of the family;
however, we anticipate that several more will be described from the more recent Oriental fauna within
the family. For example, we know of at least five undescribed species from southeast Asia at this
time.

The ecology of Teloganodidae is poorly known. Available collecting data, however, indicate
that larvae of all genera are restricted to mountain streams and rivers where they occur on stones or
vegetation in mostly swift currents. Our collecting records from South Africa, Sri Lanka, and Borneo
indicate that teloganodid larvae are commonly found on the vertical substrates of waterfalls. Thus,
an appropriate vernacular name for the family would be “the waterfall mayflies.” Harrison and Agnew
(1962) found that certain of the South African teloganodids were restricted to acid waters and gave

387

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

some additional ecological data. The only gut sample study (Tshemova 1972) indicates that the
larvae of the Oriental Vietnamella Tshemova are periphyton feeders. Differences we have found in
development of filtering setae on mouthparts of different genera, however, suggest that teloganodid
larvae may also be fine detritus collectors to various degrees. McShaffrey and McCafferty (1990a,
1990b) found a considerable range of feeding behavior in certain members of the related family
Ephemerellidae. The fact that Barnard (1932) has also reported teloganodid larvae from under stones
suggests detritus deposit feeding. According to Barnard (1932), some teloganodid larvae crawl out
of the water prior to emerging to the subimago, although it is not known if this is facultative or
obligatory behavior, or if it is widespread among species. Edmunds and McCafferty (1988) noted
that such behavior is predominant in primitive extant mayflies.

We will show that the Teloganodidae is a monophyletic lineage, and is a sister group to the larger,
generally more well known, and more apotypic Holarctic and Oriental family Ephemerellidae, within
which it was previously considered a subfamily (Allen 1965). Our conclusions are based on the
phylogenetic data we present herein. Furthermore, our linear higher classification of Teloganodidae
into subfamilies and genera is completely congruent with our cladistics data, in that it is strictly
phylogenetic and permits reproduction of the deduced cladogram of the taxa, as was advocated by
McCafferty (1991a).

The original concept of the family Ephemerellidae dates to Eaton (1883-88) with his Section 6
of Ephemerella, which included the Holarctic and Oriental genus Ephemerella Walsh and Oriental
genus Teloganodes Eaton, as they were known at that time. Klapalek (1909) first used the family
rank designation, but included only Ephemerella. Ulmer (1920) added the South American genus
Melanemerella Ulmer to the family, and Lestage ( 1 924) added the South African genus EphemerelUna
Lestage. Various other genera were described in the family up to the time that Edmunds and Traver
(1954), in their outline of higher reclassification, included the following world genera in the
Ephemerellidae: Ephemerella, EphemerelUna, Lithogloea Barnard (South Africa), Melanemerella,
Teloganella Ulmer (Oriental), Teloganodes, and Teloganopsis Ulmer (Oriental).

Demoulin (1955) removed Melanemerella from the family Ephemerellidae and placed it in a
new subfamily Melanemerellinae of Tricorythidae, and Edmunds et al. (1963) recognized this
subfamily but returned it to Ephemerellidae. Wang and McCafferty (1996a) recently removed
Melanemerella from Ephemerellidae by demonstrating that it is not a member of the Pannota but
probably an aberrant member of Leptophlebiidae.

Allen (1965) erected the subfamily Teloganodinae within the Ephemerellidae and included in it
the genera Teloganodes and EphemerelUna. He considered Austremerella Riek, from Australia, to
be synonymous with EphemerelUna', he did not mention Lithogloea. Demoulin (1970) described the
South African genus Lestagella Demoulin in the Ephemerellidae, but did not place it any subfamily.
He also placed Lithogloea as a subgenus of EphemerelUna. Tshemova (1972) described the Oriental
genus Vietnamella in the family Ephemerellidae, and although Allen (1980, 1984) placed this genus
in the Ephemerellinae, Edmunds and Murvosh (1995) correctly recognized that it belonged to the
Teloganodinae. Furthermore, Wang and McCafferty (1995) showed that those Oriental species placed
in EphemerelUna (Allen and Edmunds 1963a, Tshemova 1972) are in actuality species of Vietnamella.
Allen (1973) described the genus Manohyphella Allen from Madagascar and added it to the
Teloganodinae along with Teloganella. Although never stated by Allen, Lestagella was also, by
default, considered in the subfamily Teloganodinae, because he did not include it in Ephemerellinae
(Allen 1980, 1984).

388

McCAFFERTY and WANG; PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

In 1990, Hubbard listed the following genera in Teloganodinae: Ephemerellina, Lestagella,
Manohyphella, Teloganella, and Teloganodes. Since then, Wang et al. (1995) have removed
Teloganella from the Teloganodinae and Ephemerellidae, showing that it is instead a member of the
family Tricorythidae. Its classification previously had been argued by Edmunds and Polhemus ( 1 990)
and Peters and Peters (1993), but neither paper offered any supporting data. McCafferty and Wang
(1995) have also removed Manohyphella from Ephemerellidae and Teloganodinae, showing that it
also is a member of the Tricorythidae. McCafferty and de Moor (1995) re-established the genus
Lithogloea, thus adding it to the Teloganodinae. As mentioned above, Vietnamella was shown to be
a member of Teloganodinae (Edmunds and Murvosh 1995). Einally, the Sri Lankan genus
Macafertiella Wang was recently described in Teloganodinae by Wang and McCafferty (1996b).

Taking into account the complex history of the group, and the results of our cladistic analysis,
we recognize herein the following genera in the family Teloganodiddif. Aiistremerella, Ephemerellina,
Lestagella, Macafertiella, Nadinella gen. nov. from South Africa, Teloganodes, and Vietnamella.
The entire higher classification is shown in Table 1. We will show that the previous synonymies of
Austremerella with Ephemerellina, and Lithogloea with Ephemerellina were essentially based on
symplesiomorphies.

Table 1 . Higher classification and general distribution of the Teloganodidae

Family Teloganodidae

Subfamily Austremerellinae subfam. nov.

Genus Austremerella (Australian)

Genus Vietnamella (Oriental)

Subfamily Teloganodinae

Genus Ephemerellina (Southern Afrotropical)
Genus Nadinella gen. nov. (Southern Afrotropical)
Genus Lithogloea (Southern Afrotropical)

Genus Lestagella (Southern Afrotropical)

Genus Macafertiella (Oriental)

Genus Teloganodes (Oriental)

The extensive material that we have been able to bring together for comparative analysis has
been of vital importance in unravelling the systematics of this group. In particular, this has included
Oriental material donated from the George E. Edmunds, Jr. collection (including a recent gift from
T. Soldan from Vietnam), previously held at the University of Utah and now held in the Purdue
Entomological Research Collection (PERC herein). West Lafayette, Indiana; Australian material
loaned and donated by the Australian CSIRO (Australian National Collection, or ANC herein); South
African material loaned by the Albany Museum (AM herein); and the extensive material collected
in South Africa by the first author and Nadine McCafferty in 1990. The latter and donated material
resides in PERC.

Resolving the generic classification and assignment of species and stages in southern Africa
(where we recognize four endemic genera) was especially challenging. This was due to numerous
factors. Many species were originally incompletely described or characteristics were not described
accurately. Some species were subsequently misidentified in the literature, and larval and adult stages
were not always associated correctly. Some geographic records attributed to African teloganodines
proved to be misidentifications of tricorythid mayflies. We were able to recognize erroneous stage
correlations by employing a method discovered and used by McCafferty and Wang ( 1994) wherein
it was shown that the position and relative development of larval structures, such as tubercles, other

389

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

cuticular processes, and gills, can be represented in the adult stage by recognizable vestiges or
associated vestiges (e.g., gill socket vestiges). As an example, we demonstrate that adults assigned
to Ephemerellina bamardi Lestage by Barnard (1940) were misidentified and do not match adults
correctly assigned by him to this species earlier (Barnard 1932). Moreover, the misidentified adults
can be associated with larvae originally described as Ephemerellina crassi Allen and Edmunds
(1963a), which is the type of our new genus Nadinella. True larvae of E. bamardi were part of the
original series of larvae that had been incorrectly associated by Barnard (1932) with Lithogloea
harrisoni Barnard.

Although our higher classification of the Teloganodidae (Table 1) is based on results of our
phylogenetic research, which is essentially represented by a cladogram of species groups, we present
the cladistic methods, analysis, and discussion following presentation of the taxonomic keys and
systematic accounts of the taxa. The Key to Genera we present is a stage-associated key, meaning
that larvae and adults of each genus ideally are keyed together at the same place and in the same
sequence in the key. For each genus in the Accounts of Taxa, we give a generic synonymy and type
species, descriptions of larval and adult stages, diagnostic information, species included with specific
synonymies, distribution, material examined, and remarks regarding systematics and ecology. In
addition, 90 morphological figures, including a dorsal whole larval habitus of each genus, are included.

ACCOUNTS OF TAXA
Family Teloganodidae stat. nov.

Description of Mature Larvae

Eyes dorsal or dorsolateral (Figs 2-10), those of males (Figs 4, 10) divided into two distinct parts.
Mouthparts more or less prognathous, with branched, hairlike setae present, with variously sized
setules (e.g., Figs 47-64); labium relatively small. Forewingpads (Figs 2-10) fused with mesonotum
for most of their length, not juxtaposed posteriorly; posterior margin of mesonotum between apices
of wingpads with well-developed submedial lobes and medial V-shaped emargination. Hindwingpads
present. Simple, filamentous gills absent or present laterally on abdominal segment 1 ; lamellate gills
(Figs 2-10) dorsal and present on abdominal segments 2-5, 2-6, or 2-7; gills on abdominal segment
2 operculate (Figs 7-10), semi-operculate (Figs 4-6), or not operculate (Figs 2, 3); when gills on
abdominal segment 2 operculate or semi-operculate, then such gills more or less elongate-rounded
and always well separated from each other. Median caudal filament developed (three tailed) (Figs 2-
7) or reduced and rudimentary (two tailed) (Figs 8-10).

Description of adult

Eyes of male each divided into two distinct parts. Forewings (Figs 80-87) usually with many
short, detached, marginal intercalaries (Figs 80, 83-87) or sometimes with most short intercalaiies
attached (often to crossveins) (Figs 81, 82); one to four main CuA intercalaries present. Hindwings
present. Abdominal segment 2 with gill socket vestiges (Fig. 89). Male genitalia (Figs 90, 91) with
three segmented forceps; forceps segment 1 relatively long, more than twice as long as wide. Median
caudal filament developed (three tailed) or reduced (two tailed).

390

McCAFFERTY and WANG: PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

Diagnosis of the family

The Teloganodidae is distinct from other families of pannote mayflies. Larvae are distinguished
from the Ephemerellidae by the presence of gills on abdominal segment 2. They differ from other
families of Pannota with gills on abdominal segment 2 (e.g., Leptohyphidae, Tricorythidae, Caenidae)
by the posterior aspect of the mesonotum that has submedian lobes and a V-shaped median notch,
and by the subdivided eyes of the mature (pharate) males. Adults of Teloganodidae share generalized,
ancestral wing venation and derived divided male eyes with the Ephemerellidae, but differ from them
with respect to their distinctly more elongate basal segment of the male genital forceps. All adults
of Teloganodidae can be told from all adults of Ephemerellidae by possession of gill socket vestiges
on abdominal segment 2. The adults can be told from other pannote mayflies by their general cubital
venation, and, with the exception of Ephemerythus (Tricorythidae), by the presence of short marginal
intercalaries along the outer margin of the forewings.

Key to Genera

(Couplets 2-4 pertain to genera known from Australia and the Orient; couplets 5-7 pertain to genera
known form southern Africa)

1.

2.

3.

4.

Larva: Gills present on abdominal segment 7; gills on abdominal segment 2 not operculate or
semi-operculate (Figs 2, 3).

Adult: Forewings (Figs 80-82) with stigmatic costal interspace divided by secondary
longitudinal vein into upper and lower series of many cellules.

Subfamily Austremerellinae, 2

Larva: Gills absent on abdominal segment 7; gills on abdominal segment 2 operculate or
semi-operculate (Figs 4-10).

Adult: Forewings (Figs 83-87) with stigmatic costal interspace not divided into upper and
lower series of many cellules.

Subfamily Teloganodinae, 3

Larva: Head (Fig. 3) with pair of prominent, elongate, anteriorly directed cephalic horns
originating anterior to eyes; forefemora (Fig. 3) very broad and denticulate anteriorly.

Adult: Mesothorax without pair of long membranous processes posteriorly; head with
cephalic horn vestiges; forewings (Figs 81, 82) with most marginal intercalaries basally
attached.

Genus Vietnamella

Larva: Head (Fig. 2) without pair of cephalic horns; forefemora (Fig. 2) not as above.

Adult: Mesothorax (Fig. 88) ending posteriorly with pair of narrow-elongate, membranous
processes; head without cephalic horn vestiges; forewings (Fig. 80) with most marginal
intercalaries not attached basally.

Genus Austremerella

Larva: Two tailed, with median caudal filament reduced (Figs 8-10).

Adult: Two tailed, with median caudal filament reduced.

4

Larva: Three tailed, with well-developed median caudal filament (Figs 4-7).

Adult: Three tailed, with well-developed median caudal filament.

5

Larva: Gills present on abdominal segment 6; median row of dorsal abdominal tubercles well
developed (Fig. 8), with tubercle on tergum 10 longer than that of tergum 3.

Adult: Unknown; distribution may be limited to Sri Lanka.

Genus Macafertiella

391

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

— Larva: Gills absent on abdominal segment 6; median row of tubercles absent (Fig. 10) or only
poorly developed (Fig. 9), with tubercle on tergum 10, when present, shorter (Figs 9, 10) or
longer than that of tergum 3.

Adult: Not comparable because adult of Macafertiella unknown; generally distributed in
Southeast Asia from India to Philippines.

Genus Teloganodes

5. Larva: Lamellate gills present on abdominal segments 2-5; claws (Fig. 71) with two rows of
denticles; abdomen with pairs of dorsal tubercles (Figs 5, 74, 76) or broad, straight-margined
or slightly bifurcated posteromedial protuberances (Figs 75, 76) on at least some terga.

Adult: Abdominal terga 2-5 with gill socket vestiges; terga 3-5 each with small, broad
thickening medially, usually with two minute tubercles, or without apparent tubercle vestiges;
if tubercle vestiges absent, then IMP of forewings longer than MPj and both directly attached
to each other (Fig. 84).

Genus Nadinella gen. nov.

— Larva: Lamellate gills on abdominal segments 2-4 or 2-6; claws (Figs 70, 72, 73) with one
row of denticles; abdomen (Figs 4, 6, 7) without paired or broad, straight-margined or
bi-lobular, posteromedial dorsal tubercles or protuberances.

Adult: Abdominal terga 2-4 or 2-6 with gill socket vestiges; terga 3-5 (or more) with no
dorsal tubercle vestiges or with only single median tubercle vestige; if tubercle vestiges
absent, then IMP of forewings shorter than MP^ and basally detached from MP2 (Fig. 86).

' 6

6. Larva: Head (Fig. 7) margined with long setae anteriorly; gills on abdominal segment 2
(Fig. 7) operculate.

Adult: Abdominal segment 5 without gill socket vestiges; IMP of forewings shorter than
MP2 (Fig. 86).

Genus Lestagella

— Larva: Head (Figs 4, 6) not margined with long setae; gills on abdominal segment 2 (Figs 4,
6) semi-operculate, with part of following gill pair exposed.

Adult: Abdominal segment 5 with gill socket vestiges; IMP of forewings shorter (Fig. 85) or
longer (Fig. 83) than MP2.

7

7. Larva: Abdomen (Fig. 4) with median dorsal row of sharp tubercles; Gills absent on
abdominal segment 1 .

Adult: Abdominal terga 3-6 (Fig. 89) each with small, median tubercle vestige; IMP of
forewings longer than MP2, and MP, attached directly to IMP (Fig. 83).

Genus Ephemerellina

— Larva: Abdomen (Fig. 6) with single median row of dorsal tubercles represented by broad-
based and attenuated extensions of the posterior tergal margins; Gills present on abdominal
segment 1.

Adult: Abdominal terga 3-6 without tubercles; IMP of forewings shorter than MPj and
detached from it (Fig. 85).

Genus Lithogloea

Subfamily Austremerellinae subfam. nov.

Diagnosis

Larvae of the Austremerellinae may be told from those of the Teloganodinae by the presence of
gills on abdominal segment 7 (Figs 2, 3). Gills on abdominal segment 2 are not operculate or semi-

392

McCAFFERTY and WANG; PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

operculate in Austremerellinae (Figs 2, 3). The inner margins of the cerci have elongate, relatively
dense, and sometimes forked, hairlike setae along the medial margin (Figs 77, 78). Larvae of
Austremerellinae also possess a double row of small, sharp tubercles on abdominal terga (Figs 2, 3),
whereas only in the South African genus Nadinella gen. nov. can a double row of tubercles be found
in the Teloganodinae, and these tend to be blunt. Adults of Austremerellinae possess forewings with
a complex stigmatic area, being longitudinally subdivided by a secondary vein into extensive upper
and lower rows of cellules between the Costa and Subcosta (Figs 80-82). Forewings of
Austremerellinae also have more elongate intercalary veins in that area between IMP and CuA, and
have three or more CuA intercalary attachments to CuA. Adults also possess gill socket vestiges on
abdominal segment 7, although they may be difficult to discern.

Genus Austremerella Riek

(Figs 2, 1 1, 20, 29, 38, 47, 56, 77, 80, 88)

Austremerella Riek, 1963:50. (Larva and adult) Type: Austremerella picta Riek, by original
designation.

EphemerelUna {sxxhgems Austremerella), Allen, 1965:264.

Description of mature larva

Head (Fig. 2) without cephalic horns. Labrum (Fig. 1 1 ) subquadrate, approximately twice as
broad as long, with short, scattered setae over entire dorsal surface. Mandibles (Figs 20, 29) robust;
incisors oriented distally; medioapical setal patch of setae developed on left mandible (Fig. 29).
Maxillae (Fig. 38) with palpi absent. Superlinguae of hypopharynx (Fig. 47) moderately developed,
slightly concave laterally and not extending beyond lingua distally; apical margin of lingua convex.
Labium (Fig. 56) with well-divided and apically narrowed glossae and paraglossae; palpal segment
3 longer than width of segment 2. Pronotum (Fig. 2) short, more than twice as broad as long, not
produced anterolaterally. Forefemora (Fig. 2) moderately broad. Tarsal claws with one row of
denticles. Simple filamentous gills absent on abdominal segment 1; lamellate gills present on
abdominal segments 2-7; gills on abdominal segment 2 (Fig. 2) not operculate or semi-operculate.
Paired dorsal abdominal tubercles present (Fig. 2). Caudal filaments not banded; median caudal
filament well developed (three tailed).

Description of adult

Head without pair of cephalic horn vestiges. Mesothorax ending posteriorly with pair of elongate
membranous processes (Fig. 88). Forewings (Fig. 80) with IMP longer than MP2; MP2 attached
directly to IMP; most marginal intercalaries free, not attached. Abdominal segments 2-7 with gill
socket vestiges. Median caudal filament well developed (three tailed).

Diagnosis

The larvae of Austremerella can be distinguished from those of Vietnamella by their lack of
cephalic horns, maxillary palpi, and gills on abdominal segment 1. Furthermore, segment 3 of the
labial palpi are much longer; the mandibles are not modified apically as in Vietnamella (Figs 21, 30);
and the labrum of Austremerella has short setae scattered over the entire dorsal surface, whereas in
Vietnamella, the setae are confined to the distal half of the dorsal surface of the labrum. The adults

393

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

of Austremerella have a unique pair of membranous filaments as described above, and most marginal
intercalary veins in the forewings are typical of the family and also Ephemerellidae, in that they are
unattached. Most short marginal intercalaries of the forewings of Vietnamella are attached to other
veins.

Species included

Austremerella picta Riek

Austremerella picta Riek, 1963:50.

Ephemerellina picta (Riek), Allen, 1965:264.

Distribution

Australia: Queensland.

Material examined

Austremerella picta: AUSTRALIA: Larvae, Queensland, Flaggy Cr., Mistake Mtns. via Laidley,
S. E. Old, 11-11-1973, S. R. Monteith, ANC, PERC. Female adult paratype, Queensland, Lamington
Natl. Park, VIII- 11-1942, E. F. Riek, PERC.

Remarks

This monotypic genus is defined by the following autapomorphies (see also Phylogenetics,
below): the elongation of labial palpal segment 3, and the presence of unusual mesothoracic processes
in the adults. Within the subfamily Austremerellinae, they are further defined by loss of the gills on
abdominal segment 1 and the loss of maxillary palpi. Otherwise, the genus appears to be the most
plesiotypic genus within the family (with the possible exception of Ephemerellina of the subfamily
Teloganodinae), retaining a preponderance of plesiomorphic character states. Riek (1963) did not
notice the pair of gills on abdominal segment 7 of the larvae, and this led to some historical confusion
about the integrity of the genus. Allen (1965) incorrectly considered it a subgenus of Ephemerellina.
The male adults of A. picta remain unknown. Above, we provide new collecting data associated with
the larvae. Unfortunately, the only ecology known of Austremerella is that larvae have been found
in mountain streams.

Genus Vietnamella Tshernova
(Figs 3, 12, 21, 30, 39, 48, 57, 69, 78, 81, 82)

Vietnamella Tshernova, 1972:366. (Larva) Type: Vietnamella thani Tshernova, by original
designation.

Vietnamella, You and Su, 1987:176. (Adult).

Vietnamella, Wang and McCafferty, 1995:193. (Revision).

Description of mature larva

Head (Fig. 3) with prominent, anteriorly directed cephalic horns anterior to eyes. Labrum (Fig.
12) with lateral margins convergent distally, approximately twice as broad as long, with scattered

394

McCAFFERTY and WANG; PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

moderately long setae in distal half dorsally. Mandibles (Figs 21, 30) narrow basally, broad apically;
incisors fused and oriented more or less laterally; medioapical patch of setae developed on left
mandible. Maxillae (Fig. 39) with three-segmented palpi. Superlinguae of hypopharynx (Fig. 48)
moderately developed, rounded laterally, and slightly shorter than lingua; apical margin of lingua
straight. Labium (Fig. 57) with short and broad, apically truncate glossae and paraglossae; palpal
segment 3 very short, with length less than width of segment 2. Pronotum (Fig. 3) about 1.5 times
as broad as long, anterolateral comers produced into processes. Tarsal claws (Fig. 69) with one
denticle. Simple, filamentous gills present on abdominal segment 1; lamellate gills present on
abdominal segments 2-7; gills on abdominal segment 2 (Fig. 3) not operculate or semi-operculate.
Paired dorsal abdominal tubercles present (Fig. 3). Caudal filaments not banded; median caudal
filament well developed (three tailed).

Description of adult

Head with pair of cephalic horn vestiges. Mesothorax without pair of elongate membranous
processes. Forewings (Figs 81, 82) with IMP nearly as long as MP2; MP2 attached to IMP near base;
most short marginal intercalaries are attached to crossveins or longitudinal veins. Abdominal
segments 2-7 with gill socket vestiges. Median caudal filament well developed (three tailed).

Diagnosis

The larvae of Vietnamella can be distinguished from those of Austremerella by the presence of
cephalic horns, restricted setae on the labmm, fused and laterally oriented incisors on the mandibles,
maxillary palpi, the shorter terminal labial palpal segments, the truncate glossae and paraglossae, the
longer prothorax, the single denticle of the tarsal claws, and gills on abdominal segment 1 . The adults
of Vietnamella have vestiges of the larval cephalic horns on the head, and they do not have the
specialized mesothoracic filaments present in Austremerella.

Species included

Vietnamella dabieshanensis You and Su

Vietnamella dabieshanensis You and Su, 1987:176.

Vietnamella guadunensis Zhou and Su, 1995:48.

Vietnamella omata (Tshemova)

Ephemerellina omata Tshemova, 1972:368.

Vietnamella omata (Tshemova), Wang and McCafferty, 1995.

Vietnamella qingyuanensis Zhou and Su, 1995:47.

Vietnamella sinensis (Hsu)

Ephemerella sinensis Hsu, 1936:325.

Ephemerellina sinensis (Hsu), Allen and Edmunds, 1963a: 15.

Vietnamella sinensis (Hsu), Wang and McCafferty, 1995:193.

Vietnamella thani Tshemova

Vietnamella thani Tshemova, 1972:367.

Distribution

The genus is known from Vietnam and southern and southeastern China (Yunnan, Anhui, Fujian,
Jiang Xi, and Zhejiang Provinces).

395

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

Material examined

Vietnamella thani: VIETNAM: Larvae, VinhPhuProv., R. Dan, Tran Dao, X- 17-1 984, T. Soldan,
PERC.

Remarks

This genus is defined by the following autapomorphies (see also Phylogenetics, below): the
presence of cephalic horns in the larvae (including their vestiges in the adults), the laterally oriented,
fused mandibular incisors, the apically truncate paraglossae, the single claw denticle, and the fact
that most short marginal intercalaries in the forewings are attached either to crossveins or longitudinal
veins. The relatively large number of autapomorphies of Vietnamella shows that it is a relatively
apotypic genus within an old subfamily. It is phonetically so different from other teloganodids that
we consider it aberrant. It is not surprising that all workers previous to Edmunds and Murvosh ( 1995)
considered it in Ephemerellidae (Ephemerellinae) . Wang and McCafferty (1995) showed that all
Oriental species that were previously known as adults and that were called Ephemerellina were in
fact Vietnamella. The taxonomy of species is somewhat dubious at the present because species have
not been adequately compared with each other. Eor example, the known alate forms of V. omata
(unknown as larvae) may prove to be associated with the larvae of V. thani (unknown in alate stages).
In addition, V. sinensis and V. dabieshanensis have been taken from the same general area in the
Dabie Mountains of southeastern China, and they might prove to be synonymous, and Zhou (pers.
comm.) has indicated that he believes that V. guadunensis and V. qingyuanensis are actually immature
V. dabieshanensis.

Little is known of the habitat or habits of this genus, although Tshemova (1972) indicated that
the larvae of V. thani were periphyton feeders. Adaptive convergences in mouthpart structure found
in Vietnamella and the genera Lestagella and Teloganodes of the subfamily Teloganodinae (reduced
mandibles, and well-developed filtering setae marginally on the labrum, on short superlinguae, and
on paraglossae and somewhat fused glossae) suggest that there are close feeding similarities, including
filtering ability in the three genera.

Subfamily Teloganodinae

Diagnosis

Larvae of the Teloganodinae may be distinguished from those of the Austremerellinae by the
absence of gills on abdominal segment 7, by the presence of either operculate (Figs 7-10) or semi-
operculate (Figs 4-6) gills on abdominal segment 2, and their shorter and more sparsely setaceous
cerci (Fig. 79). All larvae lack maxillary palpi (Figs 40-46) and tubercles or horns on the head (Figs
4-10). Also, larvae either possess a double row of blunt tubercles (Figs 5, 74-76), a single row of
sharp or blunt tubercles (Figs 4, 6, 8, 9), or no dorsal tubercles (Figs 7, 10) on the abdominal terga.
Adults of Teloganodinae do not have a forewing stigmatic area that is divided by a secondary
longitudinal vein into two rows of several cellules, and there are fewer intercalary veins between IMP
and CuP (Figs 83-87). They also lack gill socket vestiges on abdominal segment 7 (Fig. 89).

396

McCAFFERTY and WANG; PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

Genus Ephemerellina Lestage

(Figs 4, 13, 22, 31, 40, 49, 58, 65, 70, 79, 83, 89)

Ephemerellina Lestage, 1924:346. (Adult). Type: Ephemerellina barnardi Lestage, by original
designation.

Ephemerellina, Barnard, 1932:251. (Larva).

Ephemerellina, Allen, 1965:293. (Incl. Austremerella).

Ephemerellina, Demoulin, 1970:123. (Incl. Lithogloea).

Ephemerellina, McCafferty and de Moor, 1995:472. (Excl. Lithogloea).

Ephemerellina, Wang and McCafferty, 1995:193. (Revision).

Description of Mature Larva

Head (Fig. 4) without well-developed marginal fringe of setae. Labrum (Fig. 13) broadly
subquadrate, apical width approximately three times length, with scattered short setae over entire
dorsal surface; apical margin broadly and gradually emarginate. Mandibles (Figs 22, 31) broadly
robust; inner and outer incisors divergent; long mediolateral seta absent; outer incisor with at least
one sharp denticle and apicomedial patch of setae developed on left mandible (Fig. 31). Maxillae as
in Fig. 40. Superlinguae of hypopharynx extended distally beyond apices of lingua [Fig. 49, Fig. 12g
of Demoulin (1970)], lateral margins nearly straight; lingua convex apically. Labium (Fig. 58) with
well-demarcated glossae and paraglossae; paraglossae somewhat bluntly pointed medioapically.
Prostemum (Fig. 65) with bi-lobular, spinous process medially. Forefemora (Fig. 4) relatively narrow.
Tarsal claws (Fig. 70) with single row of denticles. Simple, filamentous gills absent on abdominal
segment 1 ; lamellate gills present on abdominal segments 2-6; gills on abdominal segment 2 (Fig. 4)
semi-operculate. Abdomen (Fig. 4) with single row of relatively narrow-elongate and sharp tubercles
dorsally; posterolateral processes only poorly developed and approximate to base of following
segment. Caudal filaments banded; median caudal filament well developed (three tailed).

Description of adult

Forewings (Fig. 83) with IMP longer than MPj and both veins directly attached to each other in
basal half of wing. Abdominal segments 2-6 (Fig. 89) with gill socket vestiges; abdominal terga
3-5 (sometimes others also) each with small, conical, medial tubercle vestige (Fig. 89). Median
caudal filament well developed (three tailed).

Diagnosis

The larvae of Ephemerellina are distinguished from those of other genera of Teloganodinae by
their possession of a unique, spinous and bi-lobular, prostemal process, and by their possession of a
labrum (Fig. 13) that is subquadrate, relatively broad, covered by short setae over nearly its entire
dorsal surface, and somewhat emarginate along the apical margin. Larvae may be further distinguished
from other African genera of Teloganodinae by their single row of relatively narrow-elongate dorsal
abdominal tubercles. A combination of characteristics must be used to diagnose the adults of
Ephemerellina from those of other Teloganodinae (see Key to Genera, above). They differ from
other African genera of Teloganodinae by the presence of a single median row of conical tubercle
vestiges on the abdominal terga.

397

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

Species Included

Ephemerellina bamardi Lestage, 1924:348.

Ephemerellina sp. Allen and Edmunds, 1963a: 15.

Distribution

South Africa: Western Cape Province.

Material examined

Ephemerellina barnardi: SOUTH AFRICA: Male and female adults, Gt. Winterhoek Mts., 4500
ft., XI- 19 16, AM. Larva, Gt. Winterhoek Mts., IX, XI- 1932, K. H. Barnard, AM. Larva, Cedarbergen,
1-1930, K. H. Barnard, AM. Larvae, West Cape, Jonkershoek Mts., waterfall of 2nd trib. Eerste R.,
IX-28-1990, W. P. andN. McCafferty, PERC. Larva, Great Berg R., Lower forest reserve, Driefontein
Bridge, VII- 19- 1950, AM. Larva, Viljoen’s Pass, Elgin, S side, X-29-1931, K. H. Barnard, AM.

Remarks

This monotypic genus is defined by the autapomorphic spinous, bi-lobular, prostemal process
in the larvae (see also Phylogenetics, below). It can be viewed as the most plesiotypic genus of the
Teloganodinae, in that it retains the greatest number of plesiomorphic character states. It also rivals
Austremerella of the Austremerellinae as the most plesiotypic genus of the entire family. There have
been various attempts to place representatives of other lineages in this genus. Allen (1965) considered
Austremerella a subgenus of Ephemerellina. Demoulin (1970) considered Lithogloea a subgenus of
Ephemerellina, and also placed species, either as subgenus Ephemerellina or subgenus unknown in
Ephemerellina that actually belong to Nadinella gen. nov. Prom our cladistic analysis, it is now clear
that symplesiomorphy was the basis of all of the above interpretations. Allen and Edmunds (1963a)
and Tshemova (1972) placed certain species, now known to belong to Vietnamella, in the genus
Ephemerellina. This latter situation was resolved by Wang and McCafferty (1995), and the historically
confused taxonomy of African Teloganodidae is resolved herein. In Lestage’s (1924) original
description of the adults, he incorrectly reported two terminal segments of the male genital forceps.
This was corrected by Barnard (1932), who at the same time, provided description of the larval stage.

Ephemerellina is known from temperate mountain streams in the extreme southwest of the
African continent. We consider it a temperate Gondwanaland relict (see Biogeography discussion
under Phylogenetics, below). The first author and N. McCafferty collected larvae in abundance from
moss on vertical rock faces of waterfalls of small mountain tributaries of the Eerste River near
Stellenbosch. Based on data from small tributaries and waterfalls, it appears that the habitat is similar
to that of Nadinella gen. nov. (see below). King (1981) reported some spatial and temporal data for
E. bamardi in the Eerste River. She found larvae throughout the year and most commonly from
stones in the upper reaches of the river. Barber-James and de Moor (pers. comm.) indicated that
larvae have been found on the straplike leaves (ca. 1 mm width) of Isolepis (Cyperaceae), where this
plant occurs in mats in swift current of acidic streams of the Western Cape.

398

McCAFFERTY and WANG: PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

Genus Nadinella gen. nov.

(Figs 5, 14, 23, 32, 41, 50, 59, 66, 71, 74-76, 84, 90)

Type species

Ephemerellina crassi Allen and Edmunds.

Description of mature larva

Head (Fig. 5) without well-developed marginal fringe of setae. Labrum (Fig. 14) subquadrate
and relatively narrow, with apical width only approximately 1.5 times length; setae in transverse
medial third of dorsum; apical margin deeply notched medially. Mandibles (Figs 23, 32) somewhat
narrow in basal half and gradually broadening apically; incisors separated but not divergent; left
mandible with outer incisor with three semi-blunt poorly demarcated denticles and without
medioapical setal patch (Fig. 32). Maxillae as in Fig. 41. Superlinguae of hypopharynx (Fig. 50)
moderately developed, rounded laterally, and subequal in length to lingua; lingua with apical margin
nearly straight. Labium (Fig. 59) with well-divided glossae and paraglossae; glossae rounded apically;
paraglossae bluntly pointed apically. Prostemum (Fig. 66) without medial spinous process.
Forefemora (Fig. 5) relatively narrow. Tarsal claws (Fig. 71) with two rows of denticles. Simple,
filamentous gills present or absent on abdominal segment 1; lamellate gills present on 2-5; gills on
abdominal segment 2 (Fig. 5) semi-operculate. Abdomen (Fig. 5) with two rows of blunt dorsal
tubercles, sometimes coalescing on some segments into short and broad, apically straight-margined
or rounded processes (Figs 74-76); posterolateral processes poorly developed and approximate to
base of following segment. Caudal filaments banded; median caudal filament well developed
(three tailed).

Description of adult

Forewings (Fig. 84) with IMP longer than MPj, and both veins directly attached to each other
in basal half of wing. Abdominal segment 2-5 with gill socket vestiges; abdominal terga 3-5
(sometimes others also) with small transverse thickening medially, each bearing two small, conical
tubercle vestiges [see Figs 8c, 8d of Barnard (1940)]; tubercle vestiges may not be evident in some
specimens. Male genitalia as shown in Fig. 89. Median caudal filament well developed (three tailed).

Diagnosis

The larvae of Nadinella can be distinguished from larvae of all other genera of Teloganodinae
by the presence of two rows of denticles on the claws, the relatively narrow and deeply notched
labrum, and the double row of dorsal abdominal tubercles. Among the African genera of
Teloganodidae, it can further be differentiated by the presence of lamellate gills on abdominal
segments 2-5. Ephemerellina and Lithogloea larvae have lamellate gills on abdominal segments 2-
6, and Lestagella larvae have lamellate gills on abdominal segments 2-4. Some Nadinella larvae
have the double tubercles coalesced into a single, broad, posteromedian protuberance. Lithogloea
larvae have single, somewhat broad, medial tubercles that may be blunt or sharp apically. In those
Nadinella larvae in which the paired tubercles have become coalesced into a broad protuberance.

399

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

some bifurcation will usually still be evident on at least some of the segments. If no bifurcation is
evident, then often some, if not most, of the protuberances will be very broad and straight along their
apical margin [Figs 75, 76, Fig. 131 in Demoulin (1970)]. In any case, it would be prudent not to rely
solely on the presence of the double row of tubercles for identifying Nadinella.

The adults of most Nadinella are distinct among the Teloganodinae, in that they have short, broad
medial processes, with small paired tubercle vestiges evident on at least abdominal terga 3-5. They
may be differentiated among the African genera of Teloganodidae by the presence of gill socket
vestiges on abdominal segments 2-5 . The former characteristic may not be evident on adults associated
with those few larvae that have coalesced tubercles (see discussion, above), and the latter character
can be difficult to see in some specimens. Because Nadinella has a relatively elongate IMP that is
directly attached with MP2, it cannot be confused with Lithogloea or Lestagella, even though the
dorsal abdominal tubercle vestiges may not be evident in any of these genera. The male genitalia of
N. crassi and L. harrisoni are also distinctive, particularly with regard to the shape of the penes
(Figs 90, 91).

Etymology

Nadinella is named for Nadine McCafferty, who was instrumental in carrying out the field work
in South Africa that facilitated the study of Teloganodidae.

Species included

Nadinella brincki (Demoulin) comb. nov.

Ephemerellina brincki Demoulin, 1970:125.

Nadinella crassi (Allen and Edmunds) comb. nov.

Ephemerellina crassi Allen and Edmunds, 1963a: 12.

Distribution

South Africa: Western Cape Province.

Material examined

Nadinella crassi: SOUTH AFRICA: Holotype larva, paratype larvae and male subimago. Great
Berg R., French Hoek Reserve, XI-20, 21-1950, A. D. Harrison, PERC. Larva, Gt. Winterhoek,
IX,XI-1932, K. H. Barnard, PERC. Larvae, West Cape, Jonkershoek Mts., waterfall of 2nd trib.
Eerste R., IX-28-1990, W. P. and N. McCafferty, PERC. Larvae, West Cape, Jonkershoek Mts.,
waterfall of 1st trib. Eerste R., IX-28-1990, W. P. and N. McCafferty, PERC. Larvae, West Cape,
Jonkershoek St. Forest, rivulet trib. Eerste R. IX-28-1990, W. P. and N. McCafferty, PERC. Larvae,
West Cape, Jonkershoek Mt. Reserve, Eerste R. near bridge at end of dirt road, IX-28-1990, W. P.
and N. McCafferty, PERC. Subimago with larval exuviae. Great Berg R., railway bridge above Groot
Drakenstein, XI-1 1-1953, AM.

Nadinella brincki: SOUTH AFRICA: Larva, Eerste Stellenbosch, X-25-1930, AM (incorrectly
labeled by K. H. Barnard as Lithogloea harrisoni).

Nadinella sp.: SOUTH AFRICA: Kruig R., ni-8-1960, AM.

400

McCAFFERTY and WANG: PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

Remarks

This genus is defined by the autapomorphic double row of denticles on the tarsal claws (see also
Phylogenetics, below). The deeply notched condition of the labrum also appears to be autapomorphic
within the Teloganodidae, but we do not know how consistent it will prove to be on a generic level.
Demoulin (1970) originally considered N. crassi as “Ephemerellina (subg.?).” He also described at
that time what he termed as the “simplex” form of N. brincki and placed it in Ephemerellina (subgenus
Ephemerellina). This latter species, however, clearly belongs to Nadinella, having the double row
of claw denticles, lamellate gills on abdominal segments 2-5, etc. It is atypical of most of the
individuals we have seen of Nadinella, in that the usual double row of dorsal abdominal tubercles
has coalesced into a single broad and straight-margined tubercle, or protuberance. We do not know
if this represents a variation of N. brincki or perhaps another species. Presumably, it lacks the simple
gills on abdominal segment 1, although this is not clear from Demoulin’ s account. We have seen
variations of N. crassi (a species with simple gills on abdominal segment 1) with similar, and even
more extreme, tubercle variation (Figs 74-76). Barnard (1940) included an adult of Nadinella as part
of a series of Ephemerellina barnardi Lestage. The presence of a double row of tubercle vestiges on
the terga of that specimen [Figs 8c, 8d of Barnard (1940)] clearly excludes it from Ephemerellina.

Nadinella is restricted to temperate mountain streams in the extreme southwest of the African
continent. We consider it a temperate Gondwanaland relict (see Biogeography discussion under
Phylogenetics, below). The first author collected larvae in abundance from moss growing on the
rock faces of waterfalls of tributaries of the Eerste River near Stellenbosch, as well as from habitats
with less gradient. We have also seen collecting labels for Nadinella larvae that specifically noted
that they had been taken in Wardia moss on rocks.

Genus Lithogloea Barnard

(Figs 6, 15, 24, 33, 42, 51, 60, 67, 72, 85, 91)

Lithogloea Barnard, 1932:252. (Larva and adult). Type: Lithogloea harrisoni Barnard, by

monotypy.

Ephemerellina (subgenus Lithogloea), Demoulin, 1970:128.

Lithogloea, McCafferty and de Moor, 1995:472.

Description of mature larva

Head (Fig. 6) without well-developed marginal fringe of setae. Labrum (Fig. 15) with apical
margin nearly straight and lateral margins slightly convergent apically; dorsal surface with irregular
transverse rows of setae at midlength and apical fourth; basal width approximately twice length.
Mandibles (Figs 24, 33) narrowed, slightly broader apically; inner and outer incisors divergent; long
mediolateral seta absent; left mandible with outer incisor with at least one sharp denticle and without
apicomedial patch of setae (Fig. 33). Maxillae as in Fig. 42. Superlinguae of hypopharynx (Fig. 51)
with apical margins approximately subequal with that of lingua, lateral margins rounded; lingua
convex apically. Labium (Fig. 60) with well-divided glossae and paraglossae; paraglossae somewhat
bluntly pointed medioapically. Prostemum (Fig. 67) without bi-lobular, spinous process. Forefemora
(Fig. 6) relatively narrow. Tarsal claws with single row of denticles. Simple filamentous gills present
on abdominal segment 1; lamellate gills on abdominal segments 2-6; gills on abdominal segment 2

401

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

(Fig. 6) semi-operculate. Abdomen (Fig. 6) with single row of poorly developed, short, usually broad-
based tubercles dorsally; posterolateral processes well developed, elongate-subtriangular, sharply
attenuated, and well separated from base of following segment; middle segments with short lateral
setae (Fig. 6). Caudal filaments banded; median caudal filament well developed (three tailed).

Description of adult

Forewings (Fig. 85) with IMP shorter than MPj and detached from it. Abdominal segments 2-
6 with gill socket vestiges; abdominal terga without tubercle vestiges. Male genitalia as shown in
Fig. 91. Median caudal filament well developed (three tailed).

Diagnosis

The larvae of Lithogloea are distinguished from those of other genera of the Teloganodinae by
their well-developed, elongate, and remote abdominal posterolateral processes, which are widely
separated from the base of the following segment. At least one species of the Oriental genus
Teloganodes also has posterolateral processes nearly as developed as those of the Lithogloea, and
Lestagella has moderately developed posterolateral processes. In the latter genera, however, gills
are absent on abdominal segment 6, there is a well-developed setal fringe on the head, and lateral
abdominal setae are much longer. With respect to dorsal abdominal tubercles, there are some aberrant
forms of Nadinella that could be confused with Lithogloea. This situation is discussed in the Diagnosis
of Nadinella, above. Adults of Lithogloea are distinguished from those of other genera of
Teloganodinae by the combination of an IMP in the forewings that is much shorter than MP2 and
detached from it, and the presence of gill socket vestiges on abdominal segments 2-6. The nearly
apically truncate penes (Fig. 91) of Lithogloea should also be of some aid in differentiating the adult
in Africa.

Species included

Lithogloea harrisoni Barnard, 1932:253.

Ephemerellina harrisoni (Barnard), Demoulin, 1970:129.

Lithogloea harrisoni Barnard, McCafferty and de Moor, 1995:472.

Distribution

South Africa: Western Cape. There are unconfirmed records of Lithogloea from the Eastern
Cape Prov. and Kwazulu-Natal Prov., South Africa (Crass 1947), Swaziland (Stander 1963), and the
Northern Transvaal Prov., South Africa and Malawi (Harrison and Agnew 1962). We have seen
material labeled Lithogloea spp. in the Albany Museum from the Crocodile River (Eastern Transvaal),
Schageni, Karino, the Itawa River (ZambiaX Usutu River (Swaziland), and Malawi all collected in
the 1950’s and 60’ s. These all represent misidentified material of Ephemerythus Gillies
(Tricorythidae). Ephemerythus was not described until 1960 (Gillies 1960), and up to that time had
been known as “ephemerellid genus ?” (e.g., Kimmins 1955). It remains poorly known by non-
specialists. A record of Lithogloea from Zaire by Marlier (1954) is most likely attributable to the
baetid genus Acanthiops Waltz and McCafferty (see Barber- James and McCafferty 1997). Whereas
it is possible that Lithogloea eventually may be confirmed from temperate and mountainous areas of
the Eastern Cape and Kwazulu-Natal provinces of South Africa, we have been unable to substantiate

402

McCAFFERTY and WANG: PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

such a distribution at this time, and a more northern range in Africa can neither be substantiated nor
is it predicted by the general biogeography of Teloganodidae (see Biogeography under Phylogenetics,
below). In fact, the only materials of actual teloganodines that we have examined from outside the
Western Cape are Lestagella from the Eastern Cape, misidentified and mislabeled as Lithogloea, and
all of what Crass referred to as Lithogloea may be attributable to Lestagella. This is understandable
because Lestagella was not known until 1970 (Demoulin 1970) and was not generally recognized
after that. From all of the above we must assume that Lithogloea is restricted to the Western Cape
of South Africa.

Material examined

Lithogloea harrisonv. SOUTH AFRICA: Male and female adults, Gt. Winterhoek, XI- 1932, K.
H. Barnard, AM. Subimagos, Gt. Drakenstein. Larva, West Cape, Jonkershoek Mts., Eerste R. nr
bridge at end of dirt rd, IX-28-1990, W. P. and N. McCafferty, PERC.

Remarks

This genus is defined by the autapomorphic well-developed, elongate abdominal posterolateral
processes (see also Phylogenetics, below). The bluntly fused and pointed maxillae that lack denticles
also appear to be unique within the family; the maxillae of Lestagella are also reduced, but differently.
Barnard (1932) originally described Lithogloea based on L. harrisoni. Later he (Barnard 1940)
recognized part of the material he had considered as L. harrisoni as another species L. penicillata,
which was later recognized as another genus, Lestagella, by Demoulin (1970).

Little is known of the ecology of this species. It was not taken from waterfalls in the Jonkershoek
Mountains by the first author as was Nadinella and Ephemerellina, but was found cohabiting with
them in other habitats with less gradient. King (1981) and King et al. (1988) provided some spatial
and temporal data with respect to L. harrisoni in the Jonkershoek mountain reaches of the Eerste and
Long rivers. Larvae have recently been taken on the 1 mm wide leaves of Isolepis (Cyperaceae),
where this plant occurs in mats in swift currents of certain streams in the Western Cape (Barber-
James and de Moor, pers. comm.). The range of Lithogloea is somewhat dubious at this time (see
Distribution, above), and records other than those from the Western Cape need to be authenticated
with further study of material. Such records are likely attributable to Lestagella, because that genus
was not recognized as a separate genus until after the questionable records appeared. Since workers
were presumably using Barnard ( 1 940) for identification, they would have easily confused Lithogloea
and Lestagella.

Genus Lestagella Demoulin

(Figs 7, 16, 25, 34, 43, 52, 61, 68, 72, 86)

Lestagella Demoulin, 1970:130. (Larva and adult). Type: Lithogloea penicillata Barnard, by
original designation.

Description of mature larva

Head (Fig. 7) with well-developed marginal fringe of setae, long anteriorly, shorter laterally.
Labrum (Fig. 16) with rounded lateral margins and slightly emarginate apical margin; dorsum with

403

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

medial transverse row of setae; basal width nearly three times length. Mandibles (Figs 25, 34) narrow
throughout and atrophied apically; mediolateral elongate seta present, not bristlelike, sometimes
broken or missing; left mandible without apicomedial setal patch (Fig. 34). Maxillae as in Fig. 43.
Superlinguae of hypopharynx (Fig. 52) shorter than lingua, with straight lateral margins; apical margin
of lingua with small notch medially. Labium (Fig. 61) with poorly divided glossae and paraglossae;
paraglossae rounded apically. Prostemum (Fig. 68) without bi-lobular, spinous process medially.
Forefemora (Fig. 7) broad. Tarsal claws (Fig. 73) with single row of denticles. Simple filamentous
gills present on abdominal segment 1; lamellate gills present on abdominal segments 2-4; gill on
abdominal segment 2 (Fig. 7) fully operculate. Abdomen (Fig. 7) without dorsal abdominal tubercles;
posterolateral processes subtriangular and moderately developed, not elongate and sharply attenuated,
and moderately separated from base of following segment; middle segments with long lateral setae
(Fig. 7). Caudal filaments banded; median caudal filament well developed (three tailed).

Description of adult

Forewings (Fig. 86) with IMP shorter than MP2 and detached from it. Abdominal segments 2-
4 with gill socket vestiges; abdominal terga without dorsal abdominal tubercle vestiges. Median
caudal filament well developed (three tailed).

Diagnosis

Lestagella larvae can be told from larvae of other Teloganodinae by the reduced and atrophied
mandibles, the modified apices of the maxillae, and the medial notch of the lingua of the hypopharynx.
Other than these mouthpart characteristics, which will require slide mounting to examine efficiently
and may possibly prove to be species specific, the combination of the three tails and the well-developed
marginal fringe of setae on the head will distinguish the larvae of Lestagella from all others. The
adults of Lestagella can be told from those of all other teloganodines by the combination of three
tails, the relatively short and detached IMP in the forewings, and the presence of gill socket vestiges
only on abdominal segments 2-4.

Species included

Lestagella penicillata (Barnard).

Lithogloea penicillata Barnard, 1940:637.

Distribution

South Africa: Western Cape and Eastern Cape. Unusual specimens from the Amatola Mountains
of the Eastern Cape were tentatively identified as Lithogloea harrisoni by Crass (1947). Although
we could not find that material, we maintain that the record is attributable to either Lestagella or
Ephemerythus (family Tricorythidae). Other materials we have seen labeled as Lithogloea are actually
Lestagella. The latter two genera have been commonly misidentified as Lithogloea (see Distribution,
under Lestagella).

404

McCAFFERTY and WANG: PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)
Material examined

Lestagella penicillata\ SOUTH AFRICA: Larvae, South Cape, Wit R., IX- 1 8- 1989, AM. Larvae,
West Cape, JonLershoek Mts., waterfall of 2nd trib. Eerste R., IX-28-1990. W. P. andN. McCafferty,
PERC. Larva, West Cape, Jonkershoek Mts., waterfall of 1st trib. Eerste R., IX-28-1990, W. P. and
N. McCafferty, PERC. Larvae, West Cape, Jonkershoek Mts. Reserve, Eerste R., nr bridge at end
of dirt rd, IX-28-1990, W. P. and N. McCafferty, PERC. Larvae, West Cape, upper Kirstenbosch
NatT. Bot. Card., Skeleton Gorge Stream, X-1-1990, W. P. and N. McCafferty, PERC. Larvae, West
Cape, Kirstenbosch NatT. Bot. Gard., Window Stream at rd, IX-29-1990, W. P. and N. McCafferty,
PERC. Larva, Eastern Cape, Madonna and Child, X-7-1989, AM. Subimago and larvae, Tweck’s
Pont, IX-1933, K. H. Barnard, AM. Larvae, Gt. Winterhoek Mts., IX-1 1-1932, K. H. Barnard, AM.

Remarks

This genus is defined by the autapomorphic atrophy of the apical mandibular armature (see also
Phylogenetics, below). The notched lingua of the hypopharynx also appears unique among the
Teloganodidae. Lestagella belongs to a clade that also includes the Oriental genera Macafertiella
and Teloganodes. As such, it represents a transition between the archaic African fauna and the Oriental
teloganodines (see Biogeography under Phylogenetics, below). Barnard (1940) originally considered
this genus as part of Lithogloea; however, Demoulin (1970) recognized its distinctiveness and
established the genus.

Although Lestagella is known from a number of mountain streams in the Western and Eastern
Cape, little is known of its ecology.

Genus Macafertiella Wang

(Figs 8, 17, 26, 35, 44, 53, 62)

Macafertiella Wang (in Wang and McCafferty), 1996:15. (Larva). Type: Macafertiella insignis
Wang and McCafferty, by original designation.

Description of mature larva

Head (Fig. 8) with well-developed marginal fringe of setae, setae longest posterior to antennae.
Labrum (Fig. 17) with distally convergent lateral margins and slightly emarginate apical margin;
dense transverse row of long filtering setae located at approximately 3/4 distance from base and
curved laterally, following marginal shape of labrum; basal width approximately three times length.
Mandibles (Figs 26, 35) narrow throughout, slightly broader apically; long mediolateral seta present,
bristlelike; left mandible with outer incisors blunt and not divergent, and with sparse medioapical
patch of setae (Fig. 35). Maxillae as Fig. 44. Superlinguae of hypopharynx (Fig. 53) extending
beyond lingua, with straight lateral margins; apical margin of lingua straight with slightly produced
area medially. Labium (Fig. 62) with poorly divided glossae and paraglossae; glossae nearly
completely fused medially; paraglossae rounded apically. Prostemum without bi-lobular, spinous
process medially. Forefemora (Fig. 8) relatively narrow. Tarsal claws with single row of denticles.
Simple filamentous gills absent on abdominal segment 1; lamellate gills present on abdominal
segments 2-6; gill on abdominal segment 2 (Fig. 8) fully operculate. Abdomen (Fig. 8) with single
row of sharp medial tubercles dorsally; posterolateral processes poorly developed and approximate

405

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

to base of following segment. Cerci not banded. Median caudal filament vestigial (two tailed).

Adult unknown

Diagnosis

Larvae of Macafertiella are distinguished from those of other Teloganodinae by the unique
labrum, but also by the combination of being two tailed and having gills on abdominal segment 6.
Although initially Wang and McCafferty (1996b) indicated there were size differences between the
abdominal tubercles of Macafertiella and Teloganodes, the range in size of Teloganodes tubercles
has since been found to include the size of those of Macafertiella. The adults are unknown, but
assuredly will have a reduced median caudal filament as well as larva-associated vestiges of the
median row of dorsal abdominal tubercles and gill socket vestiges on abdominal segment 2-6.
Together, these should allow recognition of the unknown adult and prevent confusion with the genus
Teloganodes, which may be taken sympatrically and is similarly two tailed, but which has no gill
socket vestiges on abdominal segment 6.

Species included

Macafertiella insignis Wang and McCafferty, 1996:16.

Distribution
Sri Lanka.

Material examined

Macafertiella insignis: SRI LANKA: Larval holotype and paratype, Belihuloya Region, Veli-
Oya, trib. of Walawe-Ganga, 700 m, XII-8-1970, F. Starmiihlner, PERC. Larva paratype, Kitilgala
Region, Rambukpoth-Oya, nr Pitawela, trib. Kelani-Ganga, 650 m, XII-27-1970, F. Starmiihlner,
PERC.

Remarks

This genus is defined by the apomorphic labrum and labral setation (see also Phylogenetics,
below). Although Macafertiella is clearly a sister group of Teloganodes and a member of a rather
distinctive clade consisting also of Lestagella, it is somewhat aberrant, particularly with respect to
mouthparts and legs. It has been found cohabiting with undescribed species of Teloganodes as well
as T. tristis in Sri Lanka. There remains the possibility that the larvae of M. insignis is associated
with an adult from Sri Lanka that was named Teloganodes major by Eaton (1884). If this proves to
be true, then T major would become the type of Macafertiella. Virtually nothing is known of the
ecology of Macafertiella.

Genus Teloganodes Eaton

(Figs 9, 10, 18, 19, 27, 28, 36, 37, 45, 46, 54, 55, 63, 64, 87)

Teloganodes Eaton, 1882:208. (Adult). Type: Cloe tristis Hagen, by original designation.
Teloganodes, Ulmer, 1939:627. (Larva).

406

McCAFFERTY and WANG: PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

Description of mature larva

Head (Figs 9, 10) with well-developed marginal fringe of setae, longer setae posterior to antennae
(Fig. 9) or over entire margin (Fig. 10). Labrum (Figs 18, 19) with tapering lateral margins, and
slightly emarginate apical margin; dorsum with transverse area of setae at midlength area or slightly
beyond midlength; basal width approximately 2.5 times length. Mandibles (Figs 27, 28, 36, 37)
narrow; long mediolateral seta present and bristlelike; left mandible with incisors separate and
juxtaposed with denticles of outer incisor blunt (Fig. 36), or with denticles of outer incisor fused and
blunt (Fig. 37). Maxillae (Figs 45, 46) often with small lateral nodules. Superlinguae of hypopharynx
(Figs 54, 55 ) shorter than lingua, with lateral margins slightly concave; apical margin of lingua convex.
Labium (Figs 63, 64) with glossae and paraglossae very poorly divided, glossae nearly completely
fused medially; paraglossae rounded apically. Prostemum without bi-lobular, spinous process
medially. Forefemora (Figs 9, 10) relatively broad. Tarsal claws with single row of denticles. Simple,
filamentous gills on abdominal segment 1 absent; lamellate gills present on abdominal segments 2-
4 or 2-5; gills on abdominal segment 2 (Figs 9, 10) fully operculate. Abdomen with single row of
poorly to well-developed dorsal tubercles (Fig. 9) or without dorsal tubercles (Fig. 10); posterolateral
processes poorly (Fig. 9) to moderately developed (Fig. 10), if moderately developed, then
posterolateral processes slightly upturned and middle segments with long lateral setae (Fig. 10). Cerci
not banded. Median caudal filament reduced (two tailed).

Description of adult

Forewings (Fig. 87) with IMP shorter than MP2 and detached from it. Abdominal segments 2-
4 or 2-5 with gill socket vestiges; abdominal terga with or without median row of tubercle vestiges.
Median caudal filament reduced.

Diagnosis

The larvae of Teloganodes are distinguished from those of other Teloganodinae by the
combination of being two tailed and having no gills on abdominal segment 6. The adults of
Teloganodinae can be told from those of other Teloganodinae by the combination of the two-tailed
condition and the absence of gill socket vestiges on abdominal segment 6.

Species included

Teloganodes dentata Navas, 1931:19.

Teloganodes lugens Navas, 1933:17.

Teloganodes major Eaton, 1884: 136.

Teloganodes tristis (Hagen).

Cloe tristis Hagen, 1858:476.

Teloganodes tristis (Hagen), Eaton, 1884:135.

Distribution

Southern Asia: China: Zhejiang Province (Navas 1933); Hong Kong (Dudgeon 1990); India:
Maharashtra Province (Navas 1931); Indonesia: Borneo, Celebes, Java (Ulmer 1939), Lombok,
Sumatra (Ulmer 1939); Malaysia: Sabah, West Malaysia; Philippines; Sri Lanka.

407

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

Material examined

(All material deposited in PERC; all material larvae, except where noted).

Teloganodes tristis: MALAYSIA (EAST): Sabah, Sungai Moyog, 3 mi E of Penampung, IX-
27-29-1978, G. F and C. Edmunds. Sabah, Sungai Moyog at trib. 8 mi E Penampung, X-1-1978, G.
F. and C. Edmunds. MALAYSIA (WEST): Larvae and male subimagos, Selangor, large trib. Sungai
Selangor, 6 mi NE Kota Kuba Baharu (Mile 44), XI- 1-1987, G. F. and C. Edmunds. SRI LANKA:
Male adults and larva, Ceylon, Newara Eliya Dist., Bakers Falls nr Farr’s Inn, Horton Plains, 7000',
VIII-28-1968.

Teloganodes spp: INDONESIA (BORNEO): Kalimantan Timur Prov., Waterfall and stream, 1 1
km NE of Samarinda, VIII-27-1985, J. T. and D. A. Polhemus. INDONESIA (CELEBES): Sulawesi
Selatan Prov., Pattunuang R., 7 km SW of Bantimurung, X-13-1985, 0-100 m, J. T. and D. A.
Polhemus. Sulawesi Tengah Prov., stream 10 km SE Kamarora, Lore Lindu Nat. Park, 830 m, X-8-
1985, J. T. and D. A. Polhemus. Sulawesi Utara Prov., Toraut R., Dumoga-Bone Nat. Park 0°34'N,
123°53-54'E, IX-3-5-1985, D. A. Polhemus. Sulawesi Utara Prov., upper Metelanga R., 10 km S.
of Doloduo, IX-4-7-1985, J. T. and D. A. Polhemus. Sulawesi Utara Prov., forest stream S. of Lake
Mala, 1200 m, IX-1 1-1985, D. A. Polhemus. INDONESIA (LOMBOK): Nusa Tenggara Barat Prov.,
Aik Jut R., 1 km N of Sesaot, 30 km NE Mataram, X-23-1985, 350 m, J. T. and D. A. Polhemus.
MALAYSIA (EAST): Sabah, Sungai Moyog at tributary 8 mi E of Penampung, X-1-1987, G. F. and
C. Edmunds. Sabah, Sungai Moyog, 3 mi E of Penampung, IX-27-29-1987, G. F. and C. Edmunds.
Sabah, Sungai Tuaran, Tamparuli, X-3-1987, G. F. and C. Edmunds. Sabah, Liwagu, Moyog, N of
Kundassan, 915 m, VIII-16-17-1972, G. F. and C. Edmunds. Sabah, Liwagu R. at bridge, Ranau,
335 m, VIII-1 1-16-1972, G. F. and C. Edmunds. Pahang, Fraser Hill, Jeriau Falls (20°C), VIII-30-
1978, G. F. and C. Edmunds. MALAYSIA (WEST): Perak, Sungai Jor: Cameron Highlands Road,
19 Prov., IX-22-23-1978, G. F. and C. H. Edmunds. Larvae, male and female subimagos, Selangor,
large trib. Sungai Selangor, 5-10 mi. NE Kota Kuba Baharu (Mile 42, 43, 44, 48), 31-VIII-78, IX-
1-13-1978, G. F. and C. Edmunds (24°C). Selangor, Sungai Tua, 2-6 mi. N. Batu Caves, IX- 14-
15-1978, G. F. and C. Edmunds. Selangor, trib. of Sungai Gombak, 16 mi., 27-VIII-78, G. F. and
C. Edmunds. Selangor, Sungai Kanching, Templer Park, VIII-28-1978, G. F. and C. Edmunds.
Trengganu, Kampong Sungai Tong, IX-9-1978, G. F. and C. Edmunds. PHILIPPINES: Leyte, Leyte
Prov., Lusig R. at Hilusig, VII-15-1985, J. T. and D. A. Polhemus. Leyte, Leyte Prov., Pangusungan
R., N. of Baybay, CL 1988, VII-17-1985, J. T. and D. A. Polhemus. Luzon, Benguet Prov., 7 km,
Asia Hot Springs Rd., VII-7-1985, J. T. and D. A. Polhemus. Luzon, Benguet Prov., stream below
Camp John, Hay hydro, nr. Tuba mines, VII-8-1985, 900 m, J. T. and D. A. Polhemus. Luzon,
Pangasinan Prov., Bayaling R., 100 m, 15 km E. of Bauang, VII-6-1985, J. T. and D. A. Polhemus.
Luzon, Quezon Prov. Nat. Botanic Gardens, Llavac, VII-1 1-1985, J. T. and D. A. Polhemus.
Mindanao, South Cotabato Prov., Lahit R., nr. Lake Sebu, VII-19-1985, J. T. and D. A. Polhemus.
Mindanao, Zamboanga del Sur Prov., Bituti R., 7 km NW of Zamboanga City, 100 m, VII-22-1985,
J. T. and D. A. Polhemus. Larva, female adult, Mindoro, Mindoro Oriental Prov., Apararai Cr., 22
km SW of Calapan, VII- 13- 1985, J. T. and D. A. Polhemus. SRI LANKA: Deniyaya Region, Meda-
dola trib. of Gin-Ganga R., Sinharaja Range 1000 m, XI-9-1971, F. Starmiihlner. Deniyaya Region,
Nagahaketa-Dola trib. of Nilwala-Ganga R. 500 m, XI-13-1970, F. Starmiihlner. Deniyaya Region,
Campden Hill Dola trib. of Gin-Gana R., 700 m, XI-9-1970, F. Starmiihlner. Deniyaya Region,
Nagahaketa-Dola trib. of Nilwala-Ganga R., 500 m, XI-13-1970, F. Starmiihlner. Deniyaya Region,
Campden Hill Dola trib. of Gin-gana R., 700 m, XI-1 1-1970, F. Starmiihlner. Kitulgala Region,

408

McCAFFERTY and WANG: PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

Ambukpoth-Oya, near Pitawela, trib. of Kelani-Ganga, 650 m, XII-26-27-1970, F. Starmiihlner.
Kitulgala Region, Kelani Ganga near resthouse by Kitulgala, XII-28- 1970, F. Starmiihlner. Kitulgala
Region, Hal-Oya near Ginigathhena, trib. of Kelani-Ganga 700 m, XII-27-1970, F. Starmiihlner.
Maskeliya Region, Gartmore Dola 2000 m, at waterfall XI-29-1970, F. Starmiihlner. Maskeliya
Region, backwaters of Gartmore Dola waterfall, 1800 m, Xl-30-1970, F. Starmiihlner. Maskeliya
Region, Mocha R., trib. of Maskeliya R., dam by Adam’s Peak estate, 1800 m, Xl-28-1970, F.
Starmiihlner, PERC. Maskeliya Region, Maskeliya R. at base of Adam’s Peak, about 1 km above
Maskeliya impoundment 1800 m, Xll-7-1970, F. Starmiihlner. Maskeliya Region, Hakgala-Dola
brook above Hakgala, near Nuwara Eliya 2000 m, Xll-2-1970, E. Starmiihlner. Ratnapura Region,
Ira-Handha-Pana-Ela, right bank fork trib. of Kalu-Ganga R., 100 m, Xl-23-1970, F. Starmiihlner.

Remarks

This genus is defined by the loss of gills on abdominal segment 6 within the Macafertiella +
Teloganodes lineage (see also Phylogenetics, below). It is a relatively diverse and highly apotypic
genus, showing some variability in characters that are stable in other genera of Teloganodidae. Eaton
( 1 882) named the genus for Sri Lankan adults that had previously been described under the genus
Cloe by Hagen (1858). Eaton (1884) added an additional species from Sri Lanka, T. major, also
known only from the adult, and figured a forewing of an unnamed species from West Malaysia. We
can assume that Eaton’s generic placement of these additional species was correct, since he was aware
of the fact that Teloganodes lacks a middle tail. Nonetheless, his T. major could eventually prove to
be the adult of Macafertiella, which is also known from Sri Lanka. Ulmer (1924) provided the first
male adult description. Navas (1931) described a species from India, T. dentata, and one from China,
T. lugens, both based only on adults. Ulmer (1940) described the larval stage of Teloganodes for the
first time as T. tristis, from Sumatra and Java. We have examined larvae from Sri Lanka that are
almost identical to those described by Ulmer, and therefore consider his species identification correct.
Hubbard and Pescador (1978) reported T. tristis from the Philippines.

Very little is known of the ecology of the genus Teloganodes. Hubbard and Peters (1984),
however, indicated that, in Sri Lanka, larvae were taken from 90-2100 m, always on stones in swift
currents in small to large streams (sometimes at waterfalls), with water temperatures ranging from
14°C to about 26°C. Our records also show that Teloganodes is common in waterfalls, also having
been taken from waterfalls in Borneo by J. T. and D. A. Polhemus and from waterfalls in Sri Lanka
by F. Starmiihlner (see Material examined, above). These data are in general agreement with data
available on the African Teloganodinae, resulting from collections by the first author and N.
McCafferty of Ephemerellina and Nadinella in abundance at waterfalls.

PHYLOGENETICS

Cladistics

Cladistic methodology for deducing phylogeny is after Ross (1937, 1974) and Hennig (1950,
1966). Operational taxonomic units (OTUs) analyzed consisted of eight distinctive species groups
(Fig. 1) that each appeared monophyletic, based on their possession of unique or specialized
characteristics (cladistic analysis substantiated monophyly for each OTU). The outgroup for
determining character state polarity consisted of all other pannote mayflies.

Thirty comparative characters with different character states distributed among the OTUs were
used in the analysis, and these are presented in terms of their apomorphic and plesiomorphic states
in Table 2. Among the cladistic characters, 20 have only one apomorphy expressed in the

409

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

Teloganodinae; three are phenoclinal within the Teloganodidae in that there are two stepwise
apomorphies, the first of which is transitional to the next (see apomorphies 12 and 24; 21 and 26; 22
and 27); and two are bi-directional, in that the two apomorphies cited are different and independent
(see apomorphies 6 and 23; 8 and 17).

Results of the cladistic analysis are given in the form of a most parsimonious cladogram
(Fig. 1) deduced from our data. Numbers in Figure 1 refer to all synapomorphies that define the
particular clade or lineage where they appear and correspond to those listed in Table 2. Some of
these character states are given in abbreviated form in the table, but details can be found in the text
and figures.

Character Evolution

The deduced cladogram (Fig. 1 ) represents the most parsimonious arrangement that could be
generated from our data; it is not, however, the only alternative. Thus, in accepting the parsimonious
cladogram, convergences with respect to some character states must also be accepted. Below, we
present an interpretation of character state evolution, particularly in regard to homoplasy, that must
be accounted for in light of the deduced phylogeny of the Teloganodidae.

Multistate phenoelinal characters are often highly indicative of phylogenetic relationships (Ross
1974). We interpret that this is also the case in the Teloganodidae. One phenocline involves the
relative operculate nature of the gills on abdominal segment 2 [see apomorphies 12 and 24 (Fig. 1,
Table 2)], in which the semi-operculate condition (12) is intermediate to the full operculate condition
(24). A parallel transition is seen in the phylogeny of the Timpanoga complex (Ephemerellidae)
(McCafferty and Wang 1994). The second phenocline involves the development of the long
mediolateral seta on the mandibles [see apomorphies 21 and 26 (Fig. 1, Table 2)], in which the
appearance of a long hairlike seta (21) is intermediate to that seta becoming bristlelike (26). Such a
bristlelike seta has been independently derived in the Prosopistomatidae . The third phenocline
involves the degree of fusion of the glossae [see apomorphies 22 and 27 (Fig. 1, Table 2)], in which
the poorly divided condition (22) is intermediate to the nearly completely fused condition (27).
Numerous other synapomorphies express the same branching sequence as demonstrated by these
phenoclines.

Lestagella, Nadinella, Ephemerellina, and Lithogloea all possess distinctive black bands on the
larval caudal filaments, and this characteristic might very likely represent an apomorphy. Nonetheless,
the most apotypic members (Macafertiella and Teloganodes) of the clade that also
includes these four (Fig. 1) do not have the black banding. Because of the distinctive synapomorphies
that define the entire large clade [apomorphies 1 1-12 (Fig. 1, Table 2), we hypothesize that the black
banding was secondarily lost in the ancestor of Macafertiella and Teloganodes, both of which also
lost the developed middle tail. The appearance of the black banding in various members of the family
Ephemerellidae and Neoephemeridae, for example, indicates that this is probably an adaptive
characteristic.

From the parsimonious cladogram (Fig. 1), it appears that the loss of certain gill pairs occurs
independently, and therefore must either not be used, or used with caution, for deducing phylogeny.
In the case of the Teloganodidae larvae, the loss of the already reduced gills on abdominal segment
1 is one example of not being usable. The loss of this gill pair evidently has occurred independently
in three different lineages; Austremerella, Ephemerellina, one species of Nadinella, and in the
Macafertiella + Teloganodes lineage. Although the absence of these simple filamentous gills may
help distinguish individual lineages, the character must be discounted for phylogenetic deduction.

410

McCAFFERTY and WANG: PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

Table 2. Character states used to hypothesize phylogenetic relationships of the higher taxa of Teloganodidae [numbered
apomorphies are cited on the cladogram (Fig. 1 )]

Apomorphy

Plesiomorphy

1. Male forceps segment 1 elongate (Figs 90, 91).

2. Forewing stigmatic area subdivided into
rows of cellules (Figs 80-82).

3. Cerci medially with long and dense setae (Figs 77, 78).

4. Cephalic horns present (Fig. 3).

5. Mandibular incisors laterally oriented (Figs 21, 30).

6. Paraglossae apically truncate (Fig. 57). | [see 23]

7. Forewings with most short intercalaries
attached basally (Figs 81, 82).

8. Larval claws with single basal denticle.

(Fig. 69). t [see 17]

9. Labial palpal segment 3 elongated (Fig. 56).

10. Adult mesothorax with posterior processes (Fig. 88).

1 1 . Gills on abdominal segment 7 absent.

12. Gills on abdominal segment 2 covering large
portion of following gills (Figs 4-10).* [see 24]

13. Larval prostemum with spinous bi-lobular
process (Fig. 65).

14. Labrum with dorsal setae in medial
rows (Figs 14-19)

15. Mandibles narrowed (Figs 23-28).

16. Left mandibles with medioapical setal patch
reduced (Figs 32-36).

17. Larval claws with two rows of denticles
(Fig.71).t [see 8]

18. Forewings with IMP shorter than, and
detached from, MP2 (Figs 85-87).

19. Larval abdominal posterolateral processes
elongate-triangular and widely

separated from base of following segment (Fig. 6)

20. Larval cephalic setal fringe well developed,
at least anterior to eyes (Figs 7-10).

21. Mandibles with elongate mediolateral seta
(Figs 34-37).* [see 26]

22. Glossae and paraglossae poorly
divided (Figs 61-64).* [see 27]

23. Paraglossae rounded apically (Figs 61-64). t [see 6]

24. Gills on abdominal segment 2 fully operculate
(Figs 7-10).* [see 12]

25. Incisors and molae of mandibles atrophied (Figs 25, 34).

26. Mandible with long mediolateral seta well
developed and bristlelike (Figs 35-37).* [see 21]

27. Glossae nearly fused (Figs 62-64).* [see 22]

28. Median caudal filament reduced in both
adults and larvae (Figs 8-10).

29. Labrum with dense, curved row of long
filtering setae dorsally (Fig. 17).

30. Gills 6 absent.

* One of two phenoclinal apomorphies of the character,
t One of two bi-directional apomorphies of the character.

I Forceps segment 1 short.

2'. Stigmatic area not modified

as such (Figs 83-87).

3^ Cerci without such long and dense setae (Fig. 79).

a' . Cephalic horns absent.

5 . Incisors distally oriented (e.g.. Figs 20, 29).

6' . Paraglossae somewhat pointed (e.g.. Figs 56, 58).

1'. Most marginal intercalaries
free (e.g.. Figs 80, 83).

8 . Claws with single row of
denticles (Figs 70, 72, 73).

9'. Segment short (e.g.. Figs 77, 78).

10^. Mesothorax not modified.

I I Gills 7 present.

12^. Gills 2 not covering more

than half of following gill (Figs 2, 3)

13^. Prostemum without process
(Figs 66-68).

14^. Labrum with scattered setae
dorsally (Figs II, 13).

15^. Mandibles robust (Figs 20, 22).

16 . Left medioapical setal patch
developed (Figs 29-31).

17'. Claws with one row of
denticles (Figs 70, 72-73).

18 '. IMP long and directly

attached with MPj (Figs 80-84).

19'. Posterolateral processes
moderately developed
and approximate (Figs 2, 3, 4, 5, 8, 9).

20'. Setal fringe not developed
(Figs 2, 4-6).

21'. Mandibles without such
such seta (Figs 29-33).

22 '. Glossae and paraglossae deep
divided (Figs 56-60).

23'. Paraglossae somewhat pointed (Figs 56, 58-60).

24'. Gills 2 semi-operculate
(Fip4-6).

25 '. Incisors and molae not reduced.

26'. Seta not bristlelike (Fig. 34).

27'. Glossae poorly divided (Fig. 61).

28'. Three tailed (Figs 2-7).

29'. Setae not as developed
(Figs 14-16, 18, 19).

30'. Gills 6 present.

411

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

because only in the case of the common ancestry of Macafertiella and Teloganodes would it have
served as a supplementary defining synapomorphy. The loss of gills on abdominal segment 6 is an
example of an apomorphy that must be used with caution. It cannot be used to deduce common
ancestry of lineages, because, based on parsimony, it evidently occurred convergently in Nadinella,
Lestagella and Teloganodes. It does serve, however, as a defining autapomorphy for the Teloganodes
lineage, and as such is critical to defining Teloganodes as a monophyletic genus distinct from its
sister Macafertiella lineage.

There are certainly instances when the loss of gills on particular abdominal segments is of
importance for deducing common ancestry. The loss of gills on abdominal segment 2, for example,
is a synapomorphy that defines the entire family Ephemerellidae, and the loss of gills on abdominal
segment 7 in Teloganodidae is a synapomorphy that helps define one of two major, most basal clades
within the family [see apomorphy 1 1 (Fig. 1, Table 2)] that we consider a separate subfamily (see
below).

It might be argued that the reduction of the median caudal filament is so widespread in disparate
lineages in Ephemeroptera, and thus so highly subject to homoplasy, that it is of little or no use in
deducing phylogeny. It is, moreover, highly probable that in certain lineages the median caudal
filament becomes secondarily re-developed and lengthened from the reduced condition (see
McCafferty 1979). Nonetheless, loss of the middle tail in certain individual monophyletic groups of
mayflies recognized at the family level, such as among Teloganodidae or Heptageniidae larvae, is
an obvious independent evolutionary event that provides cladistic information. The common ancestry
of Macafertiella and Teloganodes was deduced in part by using the only instance of this loss in the
Teloganodidae. Furthermore, this common ancestry is supported by two other synapomorphies [see
apomorphies 26-28 (Fig.l and Table 2)]. On the other hand, if one were developing a cladogram of
the entire family Baetidae, the innumerable instances of reduction of the middle tail would obviously
constitute misleading information for deducing major lineages.

Dorsal abdominal tubercles are developed on the larval abdomens of many mayflies, but are
probably most common among the Ephemerellidae and Teloganodidae. Such tubercles have certainly
developed as independent apomorphies in such groups as the genus Acanthiops of the family Baetidae
(see Waltz and McCafferty 1987, Barber-James and McCafferty 1997), and the genus Anepeorus of
the family Heptageniidae (as Spinadis in Edmunds and Jensen 1974, Edmunds et al. 1976). As first
suggested by Demoulin (1967), the general habitus of the larvae of certain Acanthiops are amazingly
similar to some Teloganodidae, but this is especially true for the recently discovered Macafertiella
because of its single row of tubercles and two-tailed condition. Such tubercles are even found
independently developed in other aquatic insect orders, such as certain stoneflies (see e.g., lilies 1961,
Hynes 1970).

With respect to the current study, we hypothesize that dorsal abdominal tubercles were present
in the immediate common ancestor of the Teloganodidae and Ephemerellidae because of their relative
pervasiveness in these groups. We also infer that ancestrally they occurred in two rows on the
abdominal terga. This inference derives from the fact that the double row condition is found throughout
the Ephemerellidae and in three lineages of Teloganodidae. Based on other data, the teloganodids
lineages with double rows represent one of the two basally derived clades within the Teloganodidae
(Austremerella -t- Vietnamella), and the somewhat plesiotypic Nadinella of the latter’s sister group.
In other Teloganodidae, the double tubercle rows have either become a single median row, or tubercles
are secondarily lost. A glimpse of how this evolution has perhaps easily occurred is seen in the
variation of tubercles found in individual larvae of Nadinella (Figs 5, 74-76), where on various terga,
the tubercles of a pair can be broadened and juxtaposed medially, coalesced into a single broad

412

McCAFFERTY and WANG: PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

protuberance, and sometimes appear as a single median tubercle. In other teloganodids, there is either
a single row of dorsal abdominal tubercles (Ephemerellina, Lithogloea, Macafertiella, and plesiotypic
species of Teloganodes) or such tubercles have entirely disappeared (Lestagella and apotypic species
of Teloganodes). Considering the cladogram, the single row condition appears to have preceded the
loss of tubercles which has occurred independently in Lestagella and within Teloganodes.

A distinctive setal fringe is developed on the entire cephalic margin of the Lestagella +
Macafertiella + Teloganodes clade. It is independently developed, but only laterally in Vietnamella.
Other cases of homoplasy are found in other families of mayflies, e.g., Haplohyphes in the
Leptohyphidae (see Lugo-Ortiz and McCafferty 1995), Dicercomyzon in the Tricorythidae [present
but not figured by Demoulin (1954a)], and Epeorus in the family Heptageniidae (see e.g., Edmunds
et al. 1963).

Abdominal posterolateral processes are well developed in many different lineages of
Ephemerellidae [to the extreme in Timpanoga (see McCafferty and Wang 1994)] and some other
families of mayflies. They are also well developed independently in Lithogloea, somewhat in
Lestagella, and in at least one species of Teloganodes within the Teloganodidae. In Lithogloea, the
processes are sharply attenuated; in Lestagella and Teloganodes, they are not as attenuated.

Although it is somewhat difficult to determine the exact plesiomorphic state of the mandibular
incisors among teloganodids by using outgroup methodology, a generalized form of the armature of
the more variable left mandible appears to be represented by the mandibles of Nadinella (Fig. 32)
and most species of Teloganodes (Fig. 36), for which almost exact matches can be found among the
Ephemerellidae (see e.g.. Fig. 40 of Allen and Edmunds (1963b)]. In these mandibles, the incisors
are separate, only slightly divergent from each other, oriented distally, and the outer incisor has three
blunt denticles. Only slight modifications of this plan are found in Ephemerellina, Lithogloea, and
Macafertiella. However, in Lestagella (Figs 25, 34), the incisors are completely reduced; in
Vietnamella (Figs 21, 30), the incisors are fused and laterally oriented; and in at least one species of
Teloganodes (Figs 28, 37), the outer incisor is blunt and without denticles and the inner incisor is
highly reduced. Differences in the incisors were of no use cladistically, possibly owing to the adaptive
nature of such structures. Caution must always be used in assessing the condition of the incisors of
mayfly mandibles, even for diagnostic use, because they can become worn in long-lived instars.
There is also evidently some change in characteristics from young to older instar larvae, for example,
see comparative mouthpart figures of Lestagella provided by Demoulin (1970, Fig. 15).

Other characteristics of the mandibles did provide valuable cladistic information, including the
phenoclinal development of the long mediolateral seta as discussed above. In addition, there has
been a strong tendency for the body of the mandible to become narrowed in teloganodids [see
apomorphy 15 (Fig. 1) (Table 2)]. This may be phenoclinal, but we cannot be sure. In the clade
having the narrowed mandible, Nadinella is most apotypic and also has a mandible that does not
appear as narrowed as others in the clade. Outside of this clade, Vietnamella also shows some
narrowing of the mandibles, and while this may be viewed as generally convergent, it is different
from that seen in the others because the mandibles are abruptly narrowed in the basal two-thirds, with
the apices remaining broad. A small patch of setae occurs on the medial margin near the apices of
the mandibles of Ephemerellidae and Teloganodidae; however, it is lost in the left mandible of the
clade made up of Nadinella, Lithogloea, Lestagella, Macafertiella, and Teloganodes. The only
incongruent character state distribution is that it occurs weakly developed in Macafertiella. Since
so many other synapomorphies (Fig. 1) corroborate the phylogenetic placement of Macafertiella, we
can only assume that the setal patch was re-developed subsequently in Macafertiella.

413

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

Other characters of the mouthparts were of some value, but quite predictably were always subject
to some degree of homoplasy. Dorsal setal patterns on the labrum evolved from generally scattered
dorsal setae, similar to that found in Austremerella and Ephemerellina, which happen to be the most
plesiotypic lineages in both of the major clades. Setae become restricted to irregular, transverse rows
in the midregion of the labrum [see apomorphy 14 (Fig. 1, Table 2)]. The most ordered setal row is
found in Macafertiella amongst those with this condition. In Vietnamella, the generally scattered
setae have independently become absent from the basal half of the labrum, but remain relatively
scattered throughout the distal half.

As mentioned above the phenoclinal development of fusion in the glossae of the labium is
instructive, but in addition to this, the apical shape of the paraglossae is important. The plesiomorphic
condition is one in which the paraglossae are bluntly pointed apically, and sometimes this point is
oriented medioapically. In the clade of Lestagella + Macafertiella + Teloganodes, the apices of the
paraglossae have become narrowly rounded. A different apomorphy of this bi-directional character
is seen in Vietnamella, in which the apices of the paraglossae have become flat and truncate [see
apomorphies 6 and 23 (Fig. 1, Table 2)].

Maxillary palpi are absent in all but Vietnamella among the Teloganodidae, perhaps indicating
that the palpi were lost independently in Austremerella and the common ancestor of the other major
clade (Fig. 1 ). An alternative explanation for this would be that the palpi were absent in the common
ancestor of the family Teloganodidae, but re-appeared in the aberrant Vietnamella lineage. Generally,
structures are not expected to reappear (Ross 1974) but there are exceptions. Small nodule-like
rudiments of the maxillary palpi are present on the maxillae of many Teloganodes.

Finally, with respect to wing venation in the Teloganodidae, the appearance of a free, additional
longitudinal vein in the stigmatic area of the forewing, with numerous small cells, or cellules, on
either side of it (Figs 80-82) is an important synapomorphy for the Austremerella + Vietnamella
lineage [see apomorphy 2 (Fig. 1, Table 2)]. Somewhat similar modifications of the forewing have
evolved independently in some other mayflies. For example, in certain Heptageniidae genera, the
stigmatic area contains a secondary longitudinal vein, but cellules are not as developed, and in certain
genera of a number of families, stigmatic crossveins may be more or less anastomosed, sometimes
giving the impression of cellules (see Edmunds et al. 1976). Within the Teloganodidae, in fact, the
forewings of African genera of Teloganodinae show some slight reticulation or tendency for such in
the stigmatic area (Figs 83-86). Although this cannot be confused with the Austremerella +
Vietnamella synapomorphy, it does indicate that the common ancestor of the Teloganodidae probably
had some slight reticulation in the stigmatic area. The shortening of IMP compared to MPj, and its
detachment from MPt in the fore wings is a stable and important apomorphy for establishing Lithogloea
in the Lithogloea -i- Lestagella + Macafertiella + Teloganodes clade [see apomorphy 18 (Fig. 1,
Table 2)].

Phylogenetic Classification

Although cladistic methods have recently been employed to hypothesize phylogeny in
Ephemeroptera (e.g., McCafferty 1979, McCafferty and Edmunds 1979, Peters 1980, Savage and
Peters 1983, Dominguez 1995), it is only more recently that phylogenetic classifications have been
strictly derived from cladistic data, or cladograms (Bae and McCafferty 1991; McCafferty 1991a,
1991b; McCafferty and Wang 1994; Lugo-Ortiz and McCafferty 1996; Barber-James and McCafferty
1997). Based on our cladistic analysis of the Teloganodidae, we also present a strictly phylogenetic
classification as elaborated and endorsed in general by Wiley (1981) and in particular with respect

414

McCAFFERTY and WANG: PHYLOGENETICS OE TELOGANODIDAE (EPHEMEROPTERA)

to Ephemeroptera by McCafferty (1991a). We also incorporate sequencing conventions (Nelson
1972, 1973), which were thoroughly discussed by McCafferty and Wang (1994) with respect to their
application to the Timpanoga complex (Ephemerellidae).

The strictly phylogenetic hierarchical classification of the Teloganodidae is given in Table I. As
such, the cladogram (Fig. 1 ) is entirely reproducible from the linear classification. The most basal
bifurcate branching in the cladogram (Fig. 1) is reflected in the classification as the subfamilial
delineation.

Within the subfamily Teloganodinae (Fig. 1), sequential branches of species group OTUs are
recognized as separate genera by sequencing. All such genera are further defined by autapomorphies.
This is both a phylogenetic and practical classification because all the genera as such are easily
distinguishable in both the larval and adult stages (see the Key to Genera, above). It would be tempting
to recognize some highly specialized species in Teloganodes as a separate taxon, either genus or
subgenus; however, the remainder of Teloganodes would be left without an autapomorphy to define
them, and thus would essentially become a non-allowable paraphyletic group.

Biogeography

The Teloganodidae can be viewed as a group of Old World Gondwanan origin based on its
restricted distribution in the Orient, Australia and southern Africa. Its sister family Ephemerellidae
is essentially Holarctic and Faurasian. Edmunds (1972, 1975, 1981, 1982), Bae and McCafferty
(1991), and McCafferty et al. (1992) have argued that mayflies are ideal candidates for studying
historical biogeography because of the order’s antiquity, restriction to freshwater as larvae, short-
lived and fragile nature of alate stages, and conservative vagility. Such qualities may be expected to
be particularly amenable to reflecting ancient patterns affected primarily by vicariance (Croizat et al.
1974), as could be expected by Southern Hemisphere lineages old enough to have participated in
continental drift. By superimposing the distributions of the genera of Teloganodidae (Table 1) on
the cladogram of the genera (Fig. 1), it is possible to hypothesize certain aspects of the historical
biogeography of the various lineages (Ross 1974, Rosen 1975).

We hypothesize a Gondwanan origin for the Teloganodidae prior to the initial breakup of
Gondwanaland. The family’s restriction to the Old World Southern Hemisphere and tropical Asia
would suggest this. We hypothesize that the subfamily Austremerellinae originated prior to the
breakup of Gondwanaland, subsequently becoming isolated in Australia and more recently dispersing
through Wallacea to the tropical Orient [most probably circa 25 to 17 mya (million years ago) (Raven
and Axelrod 1974, Hamilton 1979)], where it is now represented by the relatively aberrant genus
Vietnamella. The subfamily is represented in Australia by the phylogenetic relict Austremerella, with
an ancestry that possibly dates to the Triassic. While dispersal from Australia to the Orient has been
relatively uncommon and relatively recent (Raven and Axelrod 1972, Hamilton 1979), models of
earth history [with Australia and the Orient at somewhat opposite ends of Pangaea (see e.g., Pielou
1979)], preclude other explanations at this time.

We hypothesize that the subfamily Teloganodinae arose in Gondwanaland and subsequently
became isolated in temperate South Africa. A considerable biota in temperate South Africa is regarded
as Gondwanan in origin and has been referred to as the palaeogenic [not to be confused with the
Paleogene period of Tertiary (see e.g., Stanley 1989)], or old, element in the Afrotropics (e.g., Brinck
1955a, 1955b; Stuckenberg 1962; Harrison 1965). Ephemerellina, Nadinella, and Lithogloea are
restricted to the mountainous southern temperate area of the Western Cape Province of South Africa.
Lestagella apparently has a somewhat more extensive range that includes mountainous areas of

415

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

extreme southeastern Africa as well. We have been able to substantiate Eastern Cape records of
Lestagella. Reports of Lithogloea outside of the Western Cape, however, are apparently erroneous
(see especially Distribution undtx Lithogloea and Lestagella in the Account of Taxa, above). In any
case, all African genera qualify as temperate Gondwanaland, phylogenetic relicts, the origin of which
can be traced to 135 to 200 mya (see, e.g., Pielou 1979) and possibly much earlier.

Harrison (1965) considered the leptophlebiid mayfly genera Aprionyx Barnard and
Castanophlebia Barnard as South African representatives of an “old-element, palaeo-endemic” fauna
because they were thought to have affinities with South American and Australasian Leptophlebiidae.
Peters and Edmunds (1964) suggested that the South African leptophlebiid genera Aprionyx and
Adenophlebia Eaton had Paleoantarctic affinities. Harrison (1965) included Ephemerellina and
Lithogloea (the only South African genera of Teloganodidae recognized at that time) as only
candidates for inclusion in his “old-element” category because their broader biogeographic affinities
were not known outside of South Africa. Our phylogenetic data on the Teloganodidae substantiate
the inclusion of African teloganodids in his biogeographic grouping.

The African genus Lestagella and two, more apotypic. Oriental genera Macafertiella and
Teloganodes make up the most derived clade in the subfamily Teloganodinae. We hypothesize that
the origin of this clade was temperate South Africa, and that it originated relatively early. Whereas
all other genera of African Teloganodidae are evidently restricted to the Western Cape, we have
examined specimens of Lestagella from the Eastern Cape and also believe that other specimens from
the Amatola Mountains, Eastern Cape, tentatively identified as Lithogloea harrisoni by Crass (1947)
are either Lestagella or Ephemerythus (Tricorythidae). Harrison and Agnew (1962) noted that
teloganodids had been seen from the Transvaal and as far north as Malawi, but those reports are
attributable to Ephemerythus based on our examination of material (see Distribution midtr Lithogloea
and Lestagella, in the Accounts of Taxa, above). Lestagella, therefore, appears to be both part of the
temperate Gondwanaland element but also may be typical of many such groups in Africa in having
become distributed eastward and sometimes northward along mountain chains (Harrison 1965). The
geography of Lestagella would appear to be compatible with the transitional nature of this lineage
depicted by its phylogenetic position (Fig. 1) and its hypothesized more recent participation as a
faunistic link between the African and Oriental members of the subfamily Teloganodinae [the trans-
Indian Ocean track (Craw 1988) as seen in the Teloganodidae].

The Macafertiella + Teloganodes lineage became isolated and radiated in tropical Asia. Given
our hypothesis of a temperate southern Africa origin for the Teloganodinae, the trans-Indian Ocean
track of this group could be explained by movement of the Indian subcontinent northward from
southern Africa-Madagascar landmass [beginning circa 150 mya and culminating circa 45 mya (e.g.,
see Kummel 1970)]. Dispersal during pluvial periods through Asia Minor [perhaps circa 17 mya
(Raven and Axelrod 1974)], as for example has been hypothesized for certain mayfly genera such as
Afrornera (McCafferty and Gillies 1979) and Povilla (Hubbard 1984), does not appear to be a viable
possiblity. This is because teloganodids are entirely absent from Asia Minor and east Africa, where
they would be expected if there had been such a dispersal. There are examples of a strong Afro-
Oriental affinity in many other lineages of mayflies, sometimes with genera distributed exclusively
in southern Africa and the Orient (e.g., see Gillies 1957, Peters et al. 1964, McCafferty and Edmunds
1973, Edmunds 1979, Waltz and McCafferty 1994, Provonsha and McCafferty 1995). The Afro-
Oriental relationship may be special among the teloganodids and some Leptophlebiidae because of
their general restriction to mountain streams. Today, the Oriental teloganodid fauna apparently
consists of two independently derived lineages of Teloganodidae, represented by the subfamilies
Austremerellinae and Teloganodinae.

416

McCAFFERTY and WANG: PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

Based on all of the above, one might expect Teloganodidae (esp. Teloganodinae) to be a
reasonable candidate for distribution in Madagascar. However, as pointed out by Edmunds (pers.
comm.), one should not always expect a uniform biota in all the divided parts of a former landmass,
as for example is the case with Tricorythus Eaton (Tricorythidae), because few genera and species
are widespread on the present landmasses. In the case of the Teloganodidae, Madagascar is evidently
not represented in the trans-Indian Ocean track. On the other hand, in the case of the Teloganella
complex of genera (Manohyphella, Provonshaka McCafferty and Wang, and Teloganella), which
are currently regarded in the Tricorythidae (McCafferty and Wang 1995, Wang et al. 1995), it is
Africa that is excluded from representation in their trans-Indian Ocean track.

Consideration of the relict genera of Teloganodidae in South Africa and Australia indicates that
they are typical of other known phylogenetic, or evolutionary, relicts among the Ephemeroptera in
several respects. Interesting comparisons can be made with the Amphinotic families such as
Ameletopsidae, Oniscigastridae, Nesameletidae-Rallidentidae, and Coloburiscidae that consist only
of genera that are phylogenetic relicts of Transantarctica and are now found only in Australia, New
Zealand, and southern temperate South America (see Edmunds 1975, McCafferty 1991a). The genera
are small, with only one or very few species each, and they are highly distinctive but at the same time
demonstrate some degree of convergence among relatives. These common characteristics of such
taxa are predictable by their age and the decimation of related species (after Gould 1989). Such
phylogenetic relicts should probably be referred to as “survivors” rather than relicts, as has been aptly
pointed out by Udvardy (1969). Such archaic forms figuratively have outlived all other members of
once possibly thriving taxa, and on a grand scale, they fit the final stage of the taxon cycle elaborated
by Ricklefs and Cox (1972), in that descendant species have dwindled, leaving only a few perhaps
fortuitous survivors.

None of the Transantarctic, Amphinotic families of mayflies mentioned above are found in Africa
(or Madagascar or the Indian subcontinent). On the other hand, no Teloganodidae show the
Transantarctic pattern. In generally accepted models of the breakup of Gondwanaland (e.g., Wegener
1929, Kummel 1970, Colbert 1973), Africa separated from Transantarctica prior to the breakup of
Transantarctica, respectively into West and East Gondwana. From this, one can conclude that the
teloganodid lineages represented in South Africa and Australia can be traced to southern Pangaea, at
least to 200 mya. This makes them as old or older than the strictly Amphinotic groups, which may
have originated subsequently in West Gondwana, showing the classical patterns that are apparent in
some more well known organisms, for example the flightless birds (see Cracraft 1973). This being
the case, some adjustment may be required among ephemeropterists with respect to any “conventional
wisdom” that extant pisciform mayflies are the oldest living mayflies.

Unfortunately, scant Ephemeroptera fossil data are available from the Southern Hemisphere to
shed additional light on these theories (see McCafferty 1990). Triassic mayfly fossils known from
South Africa are not clearly related to any extant mayflies (Riek 1976, Hubbard and Riek 1977), and
fossils from Australia are no older than the Lower Cretaceous (Jell and Duncan 1986). McCafferty
(1997), however, has recently found an African fossil from Lower Cretaceous Lebanese amber that
belongs to a clade of Atalophlebiine Leptophlebiidae containing both Amphinotic and Afrotropical
extant genera. That paleonological data undoubtedly substantiates the primitive position of the clade
hypothesized from phylogeny by Peters and Edmunds (1970). The only ancient fossil that has been
assigned to anything possibly remotely related to the Teloganodidae was a partial larva of Turfanerella
tingi (Ping), from the Jurassic in China, which was placed in the Ephemerellidae by Demoulin ( 1954b).
Edmunds (1972), however, noted that this fossil was probably a pisciform mayfly. Despite the dearth
of paleontological information available, cladistics of the phylogenetic relicts of Ephemeroptera have
provided rather compelling data for inferring historical biogeography of the Teloganodidae.

417

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

ACKNOWLEDGEMENTS

We sincerely thank Peter Cranston (Canberra, New South Wales), Ferdy de Moor (Grahamstown,
South Africa), and George Edmunds (Salt Lake City, Utah) for providing us with invaluable specimens
for study. We also thank Arwin Provonsha (West Lafayette, Indiana), Bob Waltz (Indianapolis,
Indiana), Michael Hubbard (Tallahassee, Florida) and George Edmunds for critically reading the
manuscript, and Helen Barber-James (Grahamstown, South Africa) for providing valuable
information. We especially thank Nadine McCafferty (West Lafayette, Indiana) for helping to collect
teloganodids in South Africa, and Ferdy de Moor for generally facilitating field work and research
in South Africa. Research was supported in part by the South African Foundation for Research
Development. This paper has been assigned Purdue Agricultural Research Program Journal Number
15025, and partial funding for the publication of this monogragh has been provided by a separate
grant from the South African Foundation for Research Development.

418

McCAFFERTY and WANG: PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

4-8 Vietnamella

2-3

9-10

Austremerella

13

Ephemerellina

11-12

17

Nadinella

14-16

19

Lithosloea

18

25

Lestagella

20-24

29 Macafertiella

26-28

30 Telosanodes

Fig. 1. Cladogram of monophyletic species groups of Teloganodidae.

419

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

Fig. 2. Austremerella picta, larval habitus.

420

McCAFFERTY and WANG: PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

Fig. 3. Vietnamella thani, larval habitus.

421

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

Fig. 4. Ephemerellina bamardi, larval habitus.

422

McCAFFERTY and WANG: PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

Fig. 5. Nadinella crassi, larval habitus.

423

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

Fig. 6. Lithogloea harrisoni, larval habitus.

424

McCAFFERTY and WANG: PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

Fig. 7. Leslagella penidllata, larval habitus.

425

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

Fig.8. Macafertiella insignis, larval habitus.

426

McCAFFERTY and WANG: PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

Fig. 9. Teloganodes tristis, larval habitus.

427

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

Fig. 10. Teloganodes sp., larval habitus.

428

w

McCAFFERTY and WANG: PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

Figs 11-28. Labrum and right mandible. 11-19. Labrum, dorsal. 11. Aiistremerella picta. 12. Vietnamella thani.

13. Ephemerellina barnardi. 14. Nadinella crassi. 15. Lithogloea harrisoni. 16. Lestagella penicillata.
17. Macafertiella insignis. 18. Teloganodes tristis. 19. Teloganodes sp. 20-28. Right mandible. 20. A. picta.
21. V. thani. 22. E. bamardi. 23. N. crassi. 24. L. harrisoni. 25. L. penicillata. 26. M. insignis. 27. T. tristis.
28. T. sp.

429

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

Figs 29-46. Left mandible and right maxilla. 29-37. Left mandible. 29. Austremerella picta. 30. Vietnamella thani.

31. Ephemerellina barnardi. 32. Nadinella crassi. 33. Lithogloea harrisoni. 34. Lestagella penicillata.
35. Macafertiella insignis. 36. Teloganodes tristis. 37. Teloganodes sp. 38-46. Right maxilla. 38. A. picta.
39. V. thani. 40. E. bamardi. 41. N. crassi. 42. L. harrisoni. 43. L. penicillata. 44. M. insignis. 45. T. tristis.
46. T. sp.

430

McCAFFERTY and WANG: PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

Figs 47-64. Hypopharynx and Labium. 47-55. Hypopharynx. 47. Austremerella picta. 48. Vietnamella thani.

49. Ephemerellina barnardi. 50. Nadinella crassi. 51. Litliogloea harrisoni. 52. Lestagella penicillata.
53. Macafertiella insignis. 54. Teloganodes tristis. 55. Teloganodes sp. 56-64. Labium. 56. A. picta.
57. V. thani. 58. E. bamardi. 59. L. harrisoni. 60. L. penicillata. 61. M. insignis. 62. T. tristis. 63. T. sp.

431

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

Figs 65-79. Larval structure. 65-68. Prosternum. 65. Ephemerellina barnardi. 66. Nadinella crassi.

67. Lithogloea harrisoni.6&. Lestagella penicillata. 69-73. Claw. 69. Vietnamella thani. 70. E. barnardi.
71. V crassi. 72. L. harrisoni. 73. L. penicillata. 74-76. N. crassi, dorsal abdominal tubercle variations.
77-79. cercus, middle segments (left: medial margin; right: outer margin). 77. Aiistremerella picta. 78. V. thani.
79. E. barnardi.

432

McCAFFERTY and WANG: PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

Figs 80-91. Adult structures. 80-87. Forewing. 80. Austremerella picta. 81. Vietnamella omata [redrawn from Tshemova
(1972)]. 82. V. dabieshanensis [modified from You and Su (1987)]. 83. Ephemerellina baniardi. 84. Nadinella
crassi. 85. Lithogloea harrisoni. 86. Lestagella peniciUata. 87. Tehganodes tristis. 88. Thoracic nota, A. picta
(arrow: process). 89. Abdomen, lateral, E. bamardi (top arrows: tubercle vestiges; bottom arrows: gill socket
vestiges). 90-91. Male genitalia. 90. N. crassi. 91. L. harrisoni.

433

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

REFERENCES

Allen, R. K. 1965. A review of the subfamilies of Ephemerellidae (Ephemeroptera). J. Kansas ent. Soc. 38: 262-266.

Allen, R. K. 1973. New Ephemerellidae from Madagascar and Afghanistan. Pan-Pacif. Ent. 49: 160-164.

Allen, R. K. 1980. Geographic distribution and reclassification of the sub-family Ephemerellinae (Ephemeroptera,
Ephemerellidae) In: Flannagan, J. F. and Marshall, K. E., eds., Advances in Ephemeroptera biology. New York:
Plenum, pp. 71-91.

Allen, R, K. 1984. A new classification of the subfamily Ephemerellinae and the description of a new genus. Pan-Pacif. Ent. 60:
245-247.

Allen, R. K. and Edmunds, G. F., Jr. 1963a. New and little known Ephemerellidae from southern Asia, Africa and
Madagascar. Pacif. Insects 5: 1 1-22.

Allen, R. K. and Edmunds, G. F., Jr. 1963b. A revision of the genus Ephemerella (Ephemeroptera: Ephemerellidae) VI. The
subgenus Serratella in North America. Ann. ent. Soc. Am. 56: 583-600.

Bae, Y. j. and McCafferty, W. P. 1991. Phylogenetic systematics of the Potamanthidae (Ephemeroptera). Trans. Am. ent. Soc.
117:1-143.

Barber-james, H. M. and McCafferty, W. P. 1997. Review and a new species of the African genus Acanthiops (Ephemeroptera:
Baetidae). Ann. Limnol. In press.

Barnard. K. H. 1932. South African may-flies (Ephemeroptera). Trans. R. Soc. S. Afr. 20: 201-259.

Barnard, K, H, 1940. Additional records and descriptions of new species of South African alder- flies (Megaloptera), may-flies
(Ephemeroptera), caddis-flies (Trichoptera), stone-flies (Perlaria) and dragon-flies (Odonata). Ann. S. Afr. Mus. 32:
609-661.

Brinck, P. 1955a. A monograph of the whirligig beetles of southern Africa. Gyrinidae. In: Hanstrdm, B. et al., eds.. South African
animal life Vol 1. Uppsala: Almquist and Wicksells, pp. 329-518.

Brinck, P. 1955b. Odonata. In: Hansrdm, B. et al„ eds.. South African animal life Vol. 2. Uppsala: Almquist and Wicksells, pp.
191-233.

Colbert, E. H. 1973. Wandering lands and animals. New York: Dutton.

Cracraft, j. 1973. Continental drift, paleoclimatology, and the evolution and biogeography of birds. J. Zool. Land. 169: 455-545.

Crass, R, S. 1947. The may-flies (Ephemeroptera) of Natal and the Eastern Cape. Ann. Natal Mus. 11: 37-110.

Craw, R. 1988. Panbiogeography: method and synthesis in biogeography. In: Myers, A. A. and Giller, P. S., eds., Analytical
biogeography. London: Chapman and Hall, pp. 405-435.

Croizat, L., Nelson, G. and Rosen, D. E. 1974. Centers of origin and related concepts. Syst. Zool. 23: 265-278.

Demoulin, G. 1954a. Descriptions preliminaire d’un type larvaire nouveau d’Ephemeropteres Tricorythidae du Congo Beige. Bull.
Inst. r. Sci. nat. Belg. 30 (6): 1-4.

Demoulin, G. 1954b. Les Ephemdropteres jurassiques du Sinkiang. Bull. Annls Soc. r. ent. Belg. 90: 322-326.

DEMOULIN, G. 1955. Melanemerella brasiliana Ulmer, Ephemerellidae ou Tricorythidae? (Ephemeroptera). Bull. Annls Soc. r. ent.
Belg. 91: 214-216.

Demoulin, G. 1967. Description de deux larves atypiques de Baetidae (Ins. Ephemeroptera). Bull. Annls Soc. r. ent. Belg. 103:
226-232.

Demoulin, G. 1970, Ephemeroptera des faunes ethiopienne et malgache. In: Hanstrdm, B et al., eds.. South African animal life
Vol. 14. Stockholm: Swedish National Science Research Council, pp. 24-170.

Dominguez, E. 1995. Cladistic analysis of the Ulmeritus-Ulmeritoides group (Ephemeroptera, Leptophlebiidae), with descriptions
of five new species of Ulmeritoides. J. N. Y. ent. Soc. 103: 15-38.

Dudgeon, D. 1990. Determinants of the distribution and abundance of larval Ephemeroptera (Insecta) in Hong Kong running
waters. In: Campbell, I. C. ed.. Mayflies and stoneflies life histories and biology. Dordrecht: Kluwer, pp. 221-232.

Eaton, A. E. 1882. An announcement of new genera of the Ephemeridae. Entomologist’s, mon. Mag. 18: 207-208.

Eaton, A. E. 1 883-88. A revisional monograph of recent Ephemeridae or mayflies. Trans. Linn. Soc. Land., 2nd Ser. Zool. 3:
1-346.

Edmunds, G, F., Jr. 1972. Biogeography and evolution of Ephemeroptera. A. Rev. Ent. 17: 21-42.

Edmunds, G. F., Jr. 1975. Phylogenetic biogeography of mayflies. Ann. Mo. bot. Gdn 62: 251-263.

434

McCAFFERTY and WANG: PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

Edmunds, G. F., Jr. 1979. Biogeographical relationships of the Oriental and Ethiopian mayflies. Itv. Pasternak, K. and Sowa, R.

eds., Proceedings of the second international conference on Ephemeroptera. Warsaw: Panstwowe Wydawnictwo
Naukowe, pp. 11-14.

Edmunds, G. F., Jr. 1981. Discussion. In: Nelson, G. and Rosen, D. E. eds., Vicariance biogeography. New York: Columbia Univ.
Press, pp. 287-297.

Edmunds, G. E., Jr. 1982. Historical and hfe history factors in the biogeography of mayflies. Am. Zoo/. 22: 371-374.

Edmunds, G. F., Jr. and Jensen, S. L. 1974. A new genus and subfamily of North American Heptageniidae (Ephemeroptera). Proc.
ent. Soc. Wash. 76: 495-497.

Edmunds, G. F., Jr. and Mccafferty, W. P. 1988. The mayfly subimago. A. Rev. Ent. 33: 509-529.

Edmunds, G. F., Jr. and Murvosh, C. M. 1995. Systematic changes in certain Ephemeroptera studied by R. K. Allen. Pan-Pacif
Ent. 71: 157-160.

Edmunds, G. F., Jr. and Polhemus, D. A. 1990. Zoogeographic patterns among mayflies (Ephemeroptera) in the Malay
Archipelago, with special reference to Celebes. In: Knight, W. J. and Holloway, J. D., eds.. Insects and the rain forests
of south east Asia (Wallacea). London: R. ent. Soc. Lond., pp. 49-56.

Edmunds, G. F., Jr. and Traver, J. A. 1954. An outline of reclassification of the Ephemeroptera. Proc. ent. Soc. Wash. 56:
236-140.

Edmunds, G. F., Jr., Allen, R. K. and Peters, W, L. 1963. An annotated key to the nymphs of the famihes and subfamilies of
mayflies (Ephemeroptera). Univ. Utah biol. Ser. 13: 1-49.

Edmunds, G. F., Jr., Jensen, S. L. and Berner, L. 1976. Mayflies of North and Central America. Minneapolis: Univ. Minnesota
Press.

Gillies, M. T. 1957. New records of species of Euthraulus Barnard (Ephemeroptera) from east Africa and Oriental region. Proc.
R. ent. Soc. Lond. 26: 43-48.

Gillies, M. T. 1960. A new genus of Tricorythidae (Ephemeroptera) from East Africa. Proc. R. ent. Soc. Lond. 29: 35-40.

Gould, S. J. 1989. Wonderful life: the Burgess Shale and the nature of history. New York: Norton.

Hagen, H. 1858. Synopsis der Neuroptera Ceylons. Verh. zool. bot. Ges. Wien 8: 471-488.

Hamilton, W. 1979. Tectonics of the Indonesia region. USGS FTof. Rep. 1078, Washington, D. C.: U. S. Government Printing
Office.

Harrison, A. D. 1965. Geographical distribution of riverine invertebrates in southern Africa. Arch. Hydrobiol. 61: 387-394.

Harrison, A. D. and Agnew, J. D. 1962. The distribution of invertebrates endemic to acid streams in the Western and Southern
Cape Provinces. Ann. Cape Prov. Mus. 2: 273-291.

Hennig, W. 1950. Grundzuge einer Theorie der phylogenetischen Systematik. Berlin: Deutscher Zentralverlag.

Hennig, W. 1966. Phylogenetic systematics. Urbana: Univ. Illinois Press.

Hsu, Y.-C. 1936. New Chinese mayflies from Kiangsi Province (Ephemeroptera). Peking nat. Hist. Bull. 10: 319-326.

Hubbard, M. D. 1984. A revision of the genus Povilla (Ephemeroptera: Polymitarcyidae). Aquat. Insects 6: 17-35.

Hubbard, M. D. 1990. Mayflies of the world: a catalog of the family and genus taxa (Insecta: Ephemeroptera). Gainesville:
Sandhill Crane Press.

Hubbard, M. D. and Pescador, M, L. 1978. A catalog of the Ephemeroptera of the Philippines. Pacif. Insects 19: 91-99.

Hubbard, M. L. and Peters, W. L. 1984. Ephemeroptera of Sri Lanka: an introduction to their ecology and biogeography.
In: Fernando, C. H., ed.. Ecology and biogeography in Sri Lanka. The Hague: W. Junk, pp. 257-274.

Hubbard, M. D. and Riek, E. F. 1977. New name for a Triassic mayfly from South Africa (Ephemeroptera). Psyche 82: 260-261.

Hynes, H. B. N. 1970. The ecology of running waters. Toronto: Univ. Toronto lYess.

Julies, J. 1961. Gebirgsbache in Europa und in Sudamerika — ein limnologischer Vergleich. Verh. int. Verein. theor angew. Limnol.
14:517-523.

Jell, P. A. and Duncan, P. M. 1986. Invertebrates, mainly insects from the freshwater. Lower Cretaceous, Koonwarra fossil bed
(Korumburra Group), South Gippsland, Victoria. Mem. Assoc. Australasian Paleontol. 3: 1 1 1-205.

Kimmins, D. E. 1955. Ephemeroptera from Nyasaland, with descriptions of three new species and some interesting nymphal forms.
Ann. Mag. nat. Hist. (12) 8: 859-880.

King, J. M. 1981. The distribution of invertebrate communities in a small South African river. Hydrobiologia 83: 43-65.

King, J. M., Day, J. A., Hurly, M.-P., Henshall, H. and Davies, B, R. 1988. Macroinvertebrates in a southern African mountain
stream. Can. J. Fish, aquat. Sci. 45: 2168-2181.

KlapAlek, F. 1909. Ephemerida, Eintagsfliegen. In: Brauer, A., ed., Die Siisswasserfauna Deutschlands Vol. 8. Jena: Gustav
Fischer, 1-32.

435

ANN. CAPE PROV. MUS. (NAT. HIST.) VOL. 19, PT. 9 20 SEPTEMBER 1997

Kummel, B. 1970. History of the earth. San Francisco: Freeman.

Lestage, J. a. 1924. Les Ephemeres de I’Afrique du Sud. Catalogue critique et systematique des especes connues et description
de trois genres nouveaux et de sept especes nouvelles. Revue zool. afr. 12: 316-352.

Lugo-ortiz, C. R. and Mccafferty, W. P. 1995. Contribution to the taxonomy of the Leptohyphidae (Insecta: Ephemeroptera) of
Central America. Stud. Neotrop. Fauna Environ. 30: 165-176.

Lugo-ortiz, C. R. and Mccafferty, W. P. 1996, Phylogeny and classification of the Baetodes complex (Ephemeroptera:
Baetidae). J. N. Am. benthol. Soc. 15: 367-380.

Marlier, G. 1954. Recherches hydrobiologiques dans les riviers du Congo Oriental. II. Etude ecologique. Hydrobiologia 6:
225-264.

McCafferty, W. P. 1979. Evolutionary trends among the families of Ephemeroidea. In: Pasternak, D. and Sowa, R. eds..
Proceedings of the second international conference on Ephemeroptera. Warsaw: Panstwowe Wydawnictwo Naukowe,
pp. 45-50.

McCafferty, W. P. 1990. Ephemeroptera. In: Grimaldi, D. A., ed.. Insects from the Santana Formation, Lower Cretaceous, of
Brazil. New York: Am. Mus. nat. Hist., Bull. 195. pp. 20-50.

McCafferty, W. P. 1991a. Toward a phylogenetic classification of the Ephemeroptera (Insecta): a commentary on systematics.
Ann. ent. Soc. Am. 84: 343-360.

McCafferty, W. P. 1991b. The cladistics, classification, and evolution of the Heptagenioidea (Ephemeroptera). In: Alba-Tercedor,
J. and Sanchez-Ortega, A. eds.. Oveniew and Strategies of Ephemeroptera and Plecoptera. Gainesville: Sandhill Crane
Press, pp, 87-102.

McCafferty, W. P. 1997. Discovery and analysis of the oldest mayflies (Insecta, Ephemeroptera) known from amber. Bull. Soc.
Hist, nat., Toulouse. In press.

McCafferty, W. P, and de Moor, F. C. 1995. South African Ephemeroptera: problems and priorities. In: Corkum, L. D, and
Ciborowski, J. J. H. eds.. Current directions in research on Ephemeroptera. Toronto: Canadian Scholars’ Press,
pp. 463-476.

McCafferty, W. P. and Edmunds. G. F., Jr. 1973. Subgeneric classification of Ephemera (Ephemeroptera: Ephemeridae).
Pan-Pac. Ent. 49: 300-307.

McCafferty, W, P. and Edmunds, G. E., Jr. 1979. The higher classification of the Ephemeroptera and its evolutionary basis. Ann.
ent. Soc. Am. 72: 5-12.

McCafferty. W. P and GILLIES, M. T. 1979. The African Ephemeridae (Ephemeroptera). Aquat. Insects 1: 169-178.

McCafferty, W. P and WANG, T.-Q. 1994. Phylogenetic classification of the Timpanoga complex (Ephemeroptera:
Ephemerellidae). J. N. .Am. benthol. Soc. 13: 569-579.

McCafferty, W. P. and WANG, T.-Q. 1995. A new genus and species of Tricorythidae (Ephemeroptera: Pannota) from
Madagascar. Ann. Limnol. 31: 179-183.

McCafferty, W. P. Elowers, R. W. and Waltz, R. D. 1992. The biogeography of Mesoamerican mayflies. In: Darwin, S. P.

and Weldon, A. L. eds.. Biogeography of Mesoamerica: proceedings of a symposium. New Orleans: Tulane Stud. Zool.
Bot., Suppl. Publ. 1. pp. 173-193.

McShaffrey, D. and McCafferty, W. P 1990a. Eeeding behavior and related functional morphology of the mayfly Ephemerella
needhami (Ephemeroptera: Ephemerellidae). J. Insect Behav. 3: 673-688.

McShaffrey, D. and McCafferty, W. P 1990b. Ecological association of the mayfly Ephemerella needhami (Ephemeroptera:
Ephemerellidae) and the green alga Cladophora (Chlorophyta: Cladophoraceae). J.freshwat. Ecol. 6: 383-394.

Navas, L. 1931. Comunicaciones entomologicas. 14, Insectos de la India. Revta Acad. Cien. exact, fi's.-quim. nat. Zaragosa 15:
12-41.

Navas, L. 1933. Neuropteres et insectes voisons. Chine et pays environnants. Mus. Heude Notes Ent. China 1: 1-23.

Nelson, G. J. 1972. Phylogenetic relationships and classification. Syst. Zool. 21: 227-231.

Nelson, G. J, 1973. Classification as an expression of phylogenetic relationships. Syst. Zool. 22: 344-359.

Peters, W. L. 1980. Phylogeny of the Leptophlebiidae: an introduction. In: Elannagan, J. F. and Marshall, K. E., eds., Advances in
Ephemeroptera biology. New York: Plenum, pp. 33-41.

Peters, W. L. and Edmunds, G. E,, Jr. 1964. A revision of the generic classification of the Ethiopian Leptophlebiidae
(Ephemeroptera), Trans. R. ent. Soc. Land. 116: 225-253.

Peters, W. L. and Edmunds, G. E., Jr. 1970. Revision of the generic classification of the Eastern Hemisphere Leptophlebiidae
(Ephemeroptera). Pacif. Insects 12: 157-240.

436

McCAFFERTY and WANG: PHYLOGENETICS OF TELOGANODIDAE (EPHEMEROPTERA)

Peters, W. L. and Peters, J. G. 1993. Status changes in Leptohyphidae and Tricorythidae (Ephemeroptera). Aquat. Insects 15:
45-49.

Peters, W. L,, Gillies, M. T. and Edmunds, G. E., Jr. 1964. Two new genera of mayflies from the Ethiopian and Oriental regions
(Ephemeroptera: Leptophlebiidae). Proc. R. ent. Soc. Land. 33: 117-124.

PlELOU. E. C. 1979. Biogeography. New York: John Wiley and Sons.

Provonsha, a. V. and McCafferty, W. P. 1995. New brushlegged caenid mayflies from South Africa (Ephemeroptera:
Caenidae). Agimt. Insects 17: 241-251.

Raven, P. H. and Axelrod, D. 1. 1972. Plate tectonics and Australasian paleobiogeography. Science 176: 1379.

Raven, P. H. and Axelrod, D, 1. 1974. Angiosperm biogeography and past continental movements. Ann. Mo. hot. Gdn 61:
539-673.

Ricklefs, R. E. and Cox, G. W. 1972. Taxon cycles in the West Indian avifauna. Am. Nat. 106: 195-219.

Riek, E. E. 1963. An Australian mayfly of the family Ephemerellidae (Ephemeroptera). J. ent. Soc. Qd 2: 48-50.

Riek, E. F. 1976. An unusual mayfly (Insecta: Ephemeroptera) from the Triassic of South Africa. Palaeontol. Afr. 19: 149-151.

Rosen, D. E. 1975. A vicariance model of Caribbean biogeography. Syst. Zool. 24: 431-464.

Ross, H. H. 1937. A generic classification of the Nearctic sawtJies (Hymenoptera, Symphyta). Illinois biol. Monogr. 15: 1-173.

Ross, H. H. 1974. Biological systematics. Reading, Massachusetts: Addison-Wesley.

Savage, H. M. and Peters, W. L. 1983. Systematics of Miroculus and related genera from northern South America
(Ephemeroptera: Leptophlebiidae). Trans. Am. ent. Soc. 108: 491-600.

Stander, G. j. 1963. Hydrobiology of polluted waters. C.S.I.R. Spec. Rep. No. Wat. 27: 3-16.

Stanley, S. M. 1989. Earth and life through time. San Francisco: Freeman.

Stuckenberg, B. R. 1962. The distribution of the montane palaeogenic element in the South African invertebrate fauna. Ann. Cape
Prov. Mus. 2: 190-204.

Tshernova, O, A. 1972. Some new Asiatic species of mayflies (Ephemeroptera, Heptageniidae, Ephemerellidae). Ent. Obozr. 51:
604-614.

Udvardy, M. D. F. 1969. Dynamic zoogeography. New York: Van Nostrand Reinhold.

Ulmer, G. 1920. Neue Ephemeropteren. Arch. Naturgesch. 85: 1-80.

Ulmer, g. 1924. Ephemeropteren von den Sunda-lnseln und den Philippinen. Treubia 6: 28-91.

Ulmer, G. 1939-40. Eintagsfliegen (Ephemeropteren) von den Sunda-lnseln. Arch. HydrobioL. Siippl. 16: 443-692.

Waltz, R. D. and McCafferty, W. P. 1987. New genera of Baetidae (Ephemeroptera) from Africa. Proc. ent. Soc. Wash. 89:
95-99.

Waltz, R. D. and McCafferty, W. P. 1994. Cloeodes (Ephemeroptera: Baetidae) in Africa. Aquat. Insects 16: 165-169.

Wang, T.-Q. and McCafferty, W. P. 1995. Specific assignments in Ephemerellina and Vietnamella (Ephemeroptera:
Ephemerellidae). Ent. News 106: 193-194.

Wang, T.-Q. and McCafferty, W. P. 1996a. Redescription and reclassification of the south American mayfly Melanemerella
brasiliana (Ephemeroptera: Leptophlebiidae). Ent. News 107: 99-103.

Wang, T.-Q. and McCafferty, W. P. 1996b. New genus of Teloganodinae (Ephemeroptera: Pannota: Ephemerellidae) from
Sri Lanka. Ann. Linmol. 32: 15-18.

Wang, T.-Q., McCafferty, W. P. and Edmunds, G. F., Jr. 1995. Larva and adult of Teloganella (Ephemeroptera: Pannota) and
assessment of familial classification. Ann. ent. Soc. Am. 88: 324-327.

Wegener, A. 1929. The origin of continents and oceans. New York: Dover.

Wiley, E. O. 1981. Phylogenetics: the theory and practice of phylogenetic systematics. New York: John Wiley & Sons.

You, D.-S. and Su, C.-R. 1987. A new species of Vietnamella from China (Ephemeroptera: Ephemerellidae). Acta Zootaxon.
Sinica 12: 176-179 [in Chinese],

Zhou, C. and Su, C. 1995. Two new species of Vietnamella from China (Ephemeroptera: Ephemerellidae). J. Nanjing Nortn. Univ.
18: 47-49.

437

n

r

' I'i '■ -I — ' ■ - ■ '■' .;, ,1'-'

■'"■ •1-^ T 5f< ■*,!<) . I 'H

■■' ' " ' V.;' *• '•:**'* ;.,

■'(l•(. 'IJV* <' ' '• it'’ ’'-'V ^ 'iWprt'^^;t;^wR i'[\V ; .'( ,1'

••■i- ' i> •’••> I ,i -■

* ’f.l- fif ’.Srv.'i.^ l.K.ilit'iW'i'

-• •i.w,,’* '. •v«'' •j/'.-.' '

I ••■■ " 3i ■ '1. ■ I 'v I ’f (^1. K .-{

-*'1^

Nri ' ' «a»' ' - ' ■ ,, u. ,'(u> II ''r'. ,t t

■■■'>■ <1 ' .■' I './ijVwOM

' ^ ‘ ,. V*- ' M *' *t I*)

• . ■, -• i '

" •. ■ .v:’ A a,} ‘ ■ ^

• ’ ■;• ( V* 'U l.,lAi .» '»■ M!tl ■•''.■. 'I t ti.v

, ., ■, ' ■■ yi;>v,,,fU-; 1;,,

I .•■<'

' <■ , II I 't

K f i

‘ 'ff-

aiol,

" I- ’VSt' . ;i: .

■ 1 •'Vj,.'*

•ViV

'r,*

M'Y

'1 lu.. It '# ^^wi¥l

■ " ■■<>.'.

■ ■ ' i ilia. V JM

. » •> I, /a^«*

, . ;■. ,r..'l .ill ijsaiir^!)^ ,

.; ■..ii'iv ' y? , -Ji*’' l,'’iM .Vk.it ,..i

:-.' - •• t I' ■ *!■

■| ,; I - 't r ■r’.lirl' f

' *; ';^>ri '',a

, -li "
< iV

* J i'

' 'r I . ,T

V* ti*

; l‘ • . 4

.7/

/•'.v

• • >?

ii'*i

» 1 N*f*- ’ -

. “ -1

i*.

INSTRUCTIONS TO AUTHORS

LAY OUT of recent issues should be examined for details of acceptable style and format.

MANUSCRIPTS should be submitted on floppy disk accompanied by three print-outs
(spacing VA or 2). In exceptional cases typed manuscripts will be accepted. These should be
submitted in triplicate, typed on A4 size paper, double-spaced, on one side of the paper only
and with wide margins all round.

Only generic and specific names should be underlined to indicate italics. All other marking
up should be left to the editor.

Footnotes should be avoided as far as possible.

All measurements should be metric.

Systematic papers should follow the rules of the International code of zoological
nomenclature or the International code of botanical nomenclature, as applicable.

ILLUSTRATIONS should bear the author’s name, title of paper and number in soft pencil on
the reverse side. Diagrams, line drawings and photographs are all regarded as figures, should
be termed as such and should be numbered in a single series.

In addition to the original illustrations a photocopy of each should be submitted.

Diagrams and line drawings should be reducible to a size not exceeding 14 x 16 cm (17,5 cm
including captions). Photographs for half-tone blocks should be final size, of high contrast
and printed on glossy paper. Colour illustrations are only printed under special circumstances
and their financing must be arranged by the author.

A scale should appear with all illustrations or the magnification should be given in the
caption.

A list of captions should be submitted on a separate sheet.

TABLES should be submitted on separate sheets. Foldouts and tables printed sideways should
be avoided.

REFERENCES should be listed alphabetically at the end of the paper. Abbreviations of
periodical titles should conform with those in the World list of periodicals (4th ed.). Layouts
must be as follows:

Alexander, J. 1969. The indirect evidence for domestication. In: Ucko, P. J. and G. W.
Dimbleby eds, The domestication and exploitation of plants and animals. London:
Duckworth, pp. 123-129.

De Winter, B. 1969. Plant taxonomy today. S.AfrJ.Sci. 65 (8): 229-242.

JuBB, R. A. 1967. Freshwater fishes of southern Africa. Cape Town: Balkema.

GALLEY PROOFS are sent (when practicable) to authors for checking printer’s errors.
Adjustments of language, style and punctuation and any other amendments of the original
manuscript, if made at this stage, may be charged to authors.

AUTHOR’S COPIES (50) are issued free of charge. When more than one author is involved
the fifty free copies will be sent to the senior author. Additional copies ordered before printing
are supplied at a reduced cost.

CADAR. P.E.-K887

SMITHSONIAN INSTITUTION LIBRARIES

3 9088 01429 5711

913 yrgsiiR

10/03/ 815750

i

I

\

i I

I

I

I

i

.,4

(

V',"