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J  ouirnsLi  or 

Hymenoptera 
esearch 


August  1995 


ISSN  #1070-9428 


CONTENTS 


BANKS,  D.  Male  nest  defense  in  the  digger  wasp  Cerceris  binodis  (Hymenoptera:  Spheci- 
dae)    


77 


CARVER,  M.  Euryischomyia  Girault  (Hymenoptera:  Chalcidoidea:  Aphelinidae:  Eriaporinae: 

EuryischiLni)    64 

FINNAMORE,  A.  T.  Revision  of  the  world  genera  of  the  subtribe  Stigmina  (Hymenoptera: 

Apoidea:  Sphecidae:  Pemphredoninae),  part  1    204 

GESS,  F.  W.  Descriptions  of  the  male  of  Riekia  nocatunga  Richards,  the  male  and  two  strik- 
ingly distinct  sympatric  colour  forms  of  Riekia  confluens  (Snelling)  and  the  male  of 
Rolandia  angulata  (Richards)  (Hymenoptera:  Vespidae:  Masarinae)  from  Aus- 
tralia           33 

GESS,  F.  W.,  S.  K.  GESS  and  R.  W.  GESS.  An  Australian  masarine,  Rolandia  angulata  (Rich- 
ards) (Hymenoptera:  Vespidae):  nesting  and  evaluation  of  association  with  Good- 
enia  (Goodeniaceae)    25 

GOKHMAN,  V.  E.  and  D.  L.  J.  QUICKE.  The  last  twenty  years  of  parasitic  Hymenoptera 

karyology:  an  update  and  phylogenetic  implications 41 

HANSSON,  C.  Revised  key  to  the  Nearctic  species  of  Chrysocharis  Forster  (Hymenoptera: 

Eulophidae),  including  three  new  species 80 

HEYDON,  S.  L.  A  review  of  the  North  American  species  of  Thinodytes  Graham  and  Mauleus 

Graham  (Hymenoptera:  Pteromalidae)    1 

LATTKE,  J.  E.  Revision  of  the  ant  genus  Gnamptogenys  in  the  New  World  (Hymenoptera: 

Formicidae) 137 

PULAWSKI,  W.  J.  The  wasp  genus  Tachytella  Brauns,  1906  (Hymenoptera:  Sphecidae)    .  .  .     121 


(Continued  on  back  cover) 


INTERNATIONAL  SOCIETY  OF  HYMENOPTERISTS 

Organized  1982;  Incorporated  1991 


OFFICERS  FOR  1995 

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Subject  Editors 
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ural History,  Central  Park  West  at  79th  Street,  New  York,  New  York  10024;  Secretary,  Systematic 
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Statement  of  Ownership 

Title  of  Publication:  Journal  of  Hymenoptera  Research. 

Frequency  of  Issue:  Once  a  year  (currently). 

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This  issue  was  mailed  25  September  1995 


J.  HYM.  RES. 
Vol.  4,  1995,  pp.  1-24 

A  Review  of  the  North  American  Species  of  Thinodytes  Graham  and 
Mauleus  Graham  (Hymenoptera:  Pteromalidae) 

Steven  L.  Heydon 

Bohart  Museum,  Department  of  Entomology,  University  of  California, 

Davis,  CA  95616-8584  USA 


Abstract. — The  Halticoptera-group  is  defined  as  containing  those  pteromaline  pteromalid  genera 
with  a  reticulate  body,  acarinate  pronotal  collar,  weakly  developed  notauli,  weakly  delimited 
frenum,  propodeum  with  the  median  carina  and  plicae  connected  posteriorly  by  a  W-shaped 
carina,  petiole  with  a  basal  flange,  and  the  hind  margin  of  the  first  gastral  tergite  sinuous  laterally 
and  usually  emarginate  medially.  Genera  included  in  this  group  are  Halticoptera  Spinola,  Halti- 
copterina  Erdos,  Andersena  Boucek,  Thinodytes  Graham,  Syntomopiis  Walker,  Mauleus  Graham,  and 
Ploskana  Boucek.  Thinodytes  and  Mauleus  are  revised  for  the  Nearctic  region  and  keys  to  the 
world's  described  species  are  given.  New  species  include  T.  caroticus  n.  sp.,  T.  cyzicopsis  n.  sp., 
T.  petiolatus  n.  sp.,  M.  cultratus  n.  sp.,  M.  iligneus  n.  sp.,  and  M.  venetus  n.  sp.  Polycystus  nigritus 
Howard  is  transferred  to  Mauleus  as  M.  nigritus  n.  comb,  and  Gastrancistrus  cephalon  Walker  is 
transferred  to  Thinodytes  as  T.  cephalon  n.  comb.  Bubekia  fallax  Gahan  n.  syn.  is  synonymized  with 
T.  cephalon  Walker. 


INTRODUCTION  are  Notoglyptus  Masi,  Sphegigaster  Spino- 
la, and  Schimitschekia  Boucek. 

The    Halhcoptera-group    is    herein    de-  ^            ^^^  Thinodytes  and  Mauleus,  the 

fined    as    containing    those    pteromaline  North  American  species  of  all  the  genera 

pteromalid  genera  with:  body  reticulately  ^^  ^^e  Halticoptera-group  occurring  in  the 

sculptured,  pronotal  collar  acarmate  (Figs  Nearctic  region  were  recently  reviewed 

21,  22),  notauli  weakly  developed  (Figs  ^^  ^^e  currently  being  studied.  Andersena 

21,  22),  frenum  weakly  delimited  (Figs  21,  includes  only  one  species  (Boucek  1993). 

22),  propodeum  with  a  sharp  median  ca-  Anderson  (1990)  reviewed  Halticopterina 

rina  and  the  plicae  connected  posteriorly  f^^  North  America  and  is  currently  revis- 

by  a  W-shaped  carina  (Figs  17-20),  peti-  i^g  Halticoptera.  The  world  species  of  No- 

ole  braced  basally  by  an  anteriorly  di-  togh/ptus  and  the  Nearctic  species  of  Si/n- 

rected  lateral  and  ventral  flange  (Figs  17-  tomopus  and  Sphegigaster  were  reviewed 

20),  and  hind  margin  of  first  gastral  ter-  by  Heydon  (1988),' Heydon  (1993),  and 

gite  sinuous  laterally  and  usually  emar-  Heydon  and  LaBerge  (1988),  respectively, 

ginate  medially  (Figs  21,  22).  Genera  of  This  paper  reviews  the  Holarctic  species 

this  group   include  Halticopitera  Spinola,  of  Thinodytes  and  Mauleus.  [Ploskana  was 

Halticopterina   Erdos,   Andersena    Boucek,  reviewed  by  Boucek  (1976).]  Research  by 

Thinodytes  Graham,  Syntomopus  Walker,  the  author  has  revealed  the  presence  of 

Ploskana   Boucek,  and  Mauleus  Graham,  numerous  species  of  both  these  genera  in 

Three  other  genera  that  show  many  of  the  Central  and  South  America, 

characteristics  of  the  Halticoptera-group  Thinodytes    previously    contained    two 

and   which   probably   are   more   or   less  described  species,  the  Palearctic  species 

closely  related  to  the  Halticoptera-^roup  T.  cyzicus  (Walker)  1839,  and  the  Carib- 


Journal  of  Hymenoptera  Research 


Figs.  1-10.  1,  Thinodytes  caroticus  n.  sp.,  female  hind  leg;  2,  Thinodytes  cephalon  (Walker),  2,  female  hind  leg; 
3,  4,  Thinodytes  cyzicopsis  n.  sp.,  female  head  dorsal  view,  4,  male  antenna;  5,  6,  Thinodytes  petiolatus  n.  sp.,  5, 
female  head  dorsal  view,  6,  male  antenna;  7,  Thinodytes  cyzicus  n.  sp.,  female  head  dorsal  view;  8,  Mauleus 
cultratus  n.  sp.,  female  habitus;  9,  Mauleus  iligneus  n.  sp.,  female  antenna;  10,  Mauleus  venetus  n.  sp.,  female 
antenna. 


bean  species  T.  clypeatus  (Girault)  1918. 
To  these  I  add  the  New  World  species  T. 
cephalon  (Walker)  1843,  n.  comb.,  and  de- 
scribe four  new  Nearctic  species — T.  car- 
oticus n.  sp.,  T.  cyzicopsis  n.  sp.,  T.  petiol- 
atus n.  sp.,  and  T.  santerna  n.  sp.  Mauleus 


was  created  for  the  species  M.  maderensis 
Graham,  1981  from  Madeira.  However, 
this  species  is  associated  with  plants  na- 
tive to  Mexico  and  is  suspected  to  be  na- 
tive to  the  New  World  (Boucek  and  Ras- 
plus  1991).  The  New  World  origin  of  this 


Volume  4,  1995 


Figs.  11-16.  11,  Thinodytes  petiolatus  n.  sp.,  female  clypeus;  12,  Thinodyies  cephahn  (Walker),  female  clypeus; 
13,  Maiik'iis  iligueus  n.  sp.,  male  clypeus;  14,  Synlomopiis  amerkanus  Ashmead,  female  head  dorsal  view;  15, 
Maidens  iligneiis  n.  sp.,  male  head  posterior  aspect;  16,  Halticoptera  sp.,  male  head  posterior  aspect. 


species  is  given  more  credence  by  the  new  Nearctic  species — M.  cultratus  n.  sp., 

presence  of  a  described  Neotropical  Man-  M.  iligneus  n.  sp.,  and  M.  venetus  n.  sp. 

/('HS  species,  M. ///^^r/fj/s  (Howard),  1897  n.  These   three  new   species  are  described 

comb,  and  the  author's  discovery  of  three  herein. 


Journal  of  Hymenoptera  Research 


Figs.  17-22.  17,  Tliinodijtes  ceplmlon  (Walker),  female  propodeum  and  petiole;  18,  Tlunodi/tes  ci/zicopsis  n.  sp.,  male 
propodeum  and  petiole;  19,  Tlii)iodi/tes  petiolatiis  n.  sp.,  female  propodeum  and  petiole;  20,  21,  Maidens  digneus  n. 
sp.,  20,  male  propodeum  and  petiole,  21,  female  habitus;  22,  Syntomopus  arpedes  Heydon,  female  habitus. 


METHODS  ^\^  jg  used  instead  of  clava.  In  addition, 

Terminology  generally  follows  that  of  the  gastral  tergites  are  numbered  T1-T7  be- 

Graham  (1969),  except  that  genal  concav-  ginning  with  the  first  tergite  after  the  pet- 

ity  is  used  instead  of  genal  hollow  and  iole.  The  following  abbreviations  are  used: 


Volume  4,  1995 


the  median  ocellar  diameter  is  MOD,  the 
ocellar-ocular  distance  is  OOL,  the  poste- 
rior ocellar  distance  is  POL,  the  lateral 
ocellar  distance  is  LOL,  the  multiporous 
plate  sensilla  are  MPP  sensilla,  the  lower 
ocular  line  is  LOcL,  and  the  antennal  ha- 
nicular  segments  are  Fl  through  F6.  The 
measurements  given  in  the  descriptions 
can  be  converted  to  millimeters  by  multi- 
plying by  0.02.  The  acronyms  for  the  mu- 
seums from  which  material  was  borrowed 
are  listed  in  the  acknowledgments  section. 

TJiinodytes  Graham 

Dicydus  Thomson,  1876:221,  253.  Type  species: 
Miscogaster  cyzicus  Walker,  1839:200;  by 
monotypy  (examined).  New  name  needed 
because  of  Dicydus  Walker,  1833:371,  455. 

Thinodytes  Graham,  1956:261.  Type  species: 
Miscogaster  cyzicus  Walker,  1839:200;  by  orig- 
inal designation  (examined).  Peck,  Boucek, 
and  Hoffer,  1964:41.  Graham,  1969:150,  167. 
Hedqvist,  1975:167.  Dzhanokmen,  1978:82. 
Boucek  and  Rasplus,  1991:32. 

Description. — Body  color  varying  from 
almost  wholly  black  to  metallic  green; 
scape  metallic  or  nonmetallic.  Head  and 
mesosoma  mostly  alveolate  except  sculp- 
turing sometimes  smooth  on  frenum  (T. 
cyzicopsis,  T.  petiolatiis,  and  T.  santenm) 
and  median  panels  of  propodeum  (T.  san- 
terna);  petiole  alveolate  in  species  with 
quadrate  to  elongate  petiole  (Figs  18-20), 
smooth  in  species  with  transverse  petiole 
(Fig.  17);  gastral  tergites  nearly  smooth. 
Clypeus  variable,  with  either  three  asym- 
metrically arranged  denticles  (Fig.  11)  (T. 
cyzicopsis,  T.  cyzicus,  T.  petiolatiis),  three 
symmetrically  arranged  denticles  (T.  san- 
terna  and  T.  clypeatus)  or  with  single  broad 
asymmetrically  placed  denticle  (Fig.  12) 
(T.  cephalon  and  T.  caroticus).  Head  with 
short  genal  concavity  often  present;  anten- 
nal torulus  above  LOcL.  Antenna  with  for- 
mula 1:1:2:6:3;  scape  cylindrical,  slender; 
length  of  flagellum  plus  pedicel  of  female 
less  than  head  width  (subequal  in  length 
in  T.  cyzicopsis),  equal  to  or  slightly  greater 
than  head  width  in  male;  funicular  seg- 


ments cylindrical;  MPP  sensilla  usually  in 
single  row,  their  length  about  equal  to 
length  of  funicular  segment  (Figs  4,  6);  fe- 
male club  simple  apically  and  with  small 
patch  of  micropilosity  on  ventral  side  of 
terminal  segment,  except  T.  cephalon  with 
terminal  spine  and  large  patch  of  micro- 
pilosity. Male  maxilla  with  stipites  unen- 
larged;  palps  slender  (Fig.  15).  Mesosoma 
arched  dorsally;  pronotum  with  collar 
very  short  medially,  anterior  edge  round- 
ed; mesoscutum  with  notaulus  incomplete 
or  extending  to  hind  margin  of  mesoscu- 
tum as  impressed  line;  scutellum  as  long 
as  wide,  lacking  anterior  median  sulcus, 
with  two  or  three  pairs  of  lateral  setae  (ex- 
cept T.  santenm  with  many  pairs),  frenum 
not  set  off  by  sulcus;  dorsellum  a  short 
transverse  ridge;  propodeum  with  width 
of  median  panels  about  1.6X  their  length 
(Figs  17-19),  plicae  and  median  carina  dis- 
tinct and  connected  posteriorly  by  W- 
shaped  carina  (Figs  17-19),  basal  fovea 
sometimes  bordered  mesally  by  short 
straight  carina  (Fig.  17)  (T.  clypeatus,  T.  cy- 
zicopsis, and  T.  cephalon),  or  by  long  sinu- 
ous carina  (Figs  18,  19)  (T.  cyzicopsis,  T. 
cyzicus,  and  T.  petiolatiis);  spiracles  ovate. 
Fore  wing  with  postmarginal  vein  about 
as  long  as  marginal  vein  but  postmarginal 
vein  sometimes  distinctly  shorter  (T.  car- 
oticus and  T.  cephalon);  stigma  small,  but 
its  height  half  the  distance  between  stigma 
and  anterior  wing  margin  in  T.  clypeatus; 
costal  cell  with  one  complete  and  two  or 
three  partial  distal  setal  rows;  basal  cell 
bare  except  sometimes  a  few  setae  distally 
(T.  clypK^atus  and  T.  cyzicus);  basal  vein  se- 
tose except  in  T.  cephalon;  speculum  de- 
veloped and  open  posteriorly.  Petiole 
braced  basally  by  lateral  and  ventral  an- 
teriorly directed  lamellate  flange  (Figs  17- 
19);  petiole  variable,  sometimes  trans- 
verse, unsclerotized  ventrally,  and  with- 
out lateral  setae  (Fig.  17)  (T.  caroticus,  T. 
cephalon,  and  T.  santcrna),  or  quadrate  to 
elongate  and  sclerotized  ventrally  (7.  cly- 
peatus, T.  cyzicopsis,  T.  cyzicus,  and  T.  pe- 
tiolatiis); lateral  setae  sometimes  present 


Journal  of  Hymenoptera  Research 


(Fig.  19)  (T.  cyzicopsis  and  T.  petiolatus); 
weak  median  carina  present  in  T.  clypea- 
tiis.  Gaster  of  female  ovate,  acuminate  api- 
cally,  1.4-1.8X  as  long  as  wide;  hypopy- 
gium  extending  V2  to  %  gastral  length; 
hind  margin  of  Tl  sinuous  laterally  and 
emarginate  or  straight  medially. 

Discussion. — Giving  a  few  characters  to 
separate  Thinodytes  from  the  other  genera 
of  the  Halticoptera-group,  particularly  Hal- 
ticoptera,  Maiileus,  and  Si/ntomopiis,  is  im- 
possible because  Thinodytes  is  what  is  left 
when  the  more  distinct  genera  of  the  Hal- 
ticoptera-group  are  characterized.  Apomor- 
phic  characters  among  related  genera  com- 
mon to  all  Thinodytes  species  such  as  the 
reticulate  body,  weakly  developed  notauli, 
poorly  delimited  frenum,  propodeum  with 
sharp  median  carina  and  plicae  connected 
posteriorly  by  W-shaped  carina,  petiole 
with  a  basal  bracing  consisting  of  an  ante- 
riorly directed  lateral  and  ventral  flange, 
and  the  hind  margin  of  the  first  gastral  ter- 
gite  being  sinuous  laterally  and  usually 
emarginate  medially  are  the  same  charac- 
ters defining  the  Hnlticoptera-group  itself. 
So  presently,  Thinodytes  can  only  be  de- 
fined negatively.  There  are  many  South 
American  species,  some  described  and 
some  not,  that  fit  within  the  present  defi- 
nition of  Thinodytes.  Once  these  are  inves- 
tigated, it  may  be  possible  to  divide  Thi- 
nodytes into  monophyletic  generic  units. 

Halticoptera  is  distinguished  from  Thinod- 
ytes by  a  number  of  apomorphic  character 
states  including  a  bidentate  clypeus,  the 
antennal  torulus  located  at  or  below  the 
LOcL,  the  scape  usually  nonmetallic,  the 
male  maxilla  with  lamellately  expanded 
palps  and  usually  with  another  lobe  on  the 
stipites,  and  a  median  longitudinal  carina 
on  the  petiole.  Thinodytes  has  the  clypeus 


with  variable  numbers  and  arrangements 
of  clypeal  denticles,  but  no  known  Thinod- 
ytes species  has  a  bidentate  clypeus.  The 
antennal  torulus  in  Thinodytes  is  located 
distinctly  above  the  LOcL  (except  in  T.  pe- 
tiolatus), and  the  scape  usually  has  metallic 
coloration.  The  male  maxilla  of  Thinodytes 
lacks  any  expansion  of  the  palps  or  any 
lobes  on  the  stipites.  Thinodytes  clypeatus  is 
the  only  species  of  Thinodytes  that  has  a 
median  carina  on  the  petiole.  Halticoptera 
species  are  commonly  bright  metallic 
green;  those  of  Thinodytes  are  usually  dark. 

Mauleus  is  distinguished  from  Thinodytes 
by  the  bidentate  clypeus  (Fig.  13),  the  dor- 
sum of  the  mesosoma  as  high  as  the  vertex 
(Figs  8,  21),  the  median  panels  of  the  pro- 
podeum short  (2X  as  wide  as  long)  (Fig. 
20),  and  the  lateral  flanges  of  the  petiole 
enlarged  and  thickened  (Fig.  20).  In  Thi- 
nodytes, the  clypeus  has  various  numbers 
and  arrangements  of  clypeal  teeth,  but  is 
never  bidentate,  the  dorsum  of  the  meso- 
soma is  distinctly  lower  than  the  vertex,  the 
median  panels  of  the  propodeum  are  lon- 
ger (Figs.  17-19),  and  the  basal  flanges  of 
the  petiole  are  lamellate  (Figs.  17-19). 

Syntomopus  is  distinguished  from  Thi- 
nodytes by  having  an  elongate  pronotum 
(length  about  V3  its  width)  (Fig.  22),  three 
broad  symmetrically  arranged  clypeal 
denticles  (Fig.  14),  and  usually  a  flattened 
mesosoma  (Fig.  22).  The  pronotum  of  Thi- 
nodytes is  much  shorter,  those  Thinodytes 
species  having  symmetrically  arranged 
denticles  have  them  fingerlike  rather  than 
broad  (Fig.  11),  and  the  mesosoma  is  nev- 
er so  flattened  as  it  usually  is  in  Syntomo- 
pus species. 

Biology. — The  known  hosts  of  Thinodytes 
species  are  all  small  Diptera  living  in 
plants  as  leaf  or  stem  miners. 


KEY  TO  HOLARCTIC  SPECIES  OF  THINODYTES  GRAHAM 

1.  Petiole  transverse,  smooth  (Fig.  17)   2 

-   Petiole  as  long  as  wide  or  longer  than  wide,  reticulate  (Figs  18-20)   4 

2.  Scape  and  legs  beyond  coxae  pale,  nonmetallic.  Frenum  and  median  panels  of  propodeum 
smooth.  Plicae  rounded  and  smoothly  convergent  posteriorly santema  n.  sp. 


Volume  4,  1995  7 

-  Scape  mostly  or  completely  metallic.  Legs  beyond  coxae  with  metallic  coloration  or  dark 
bands  on  femora  and  tibiae.  Frenum  and  median  panels  of  propodeum  weakly  to  strongly 
alveolate.  Plicae  with  distinct  angle  between  parallel  basal  part  and  convergent  posterior 
part 3 

3.  Hind  margin  of  Tl  emarginate  medially.  Dark  bands  on  tibiae  with  sharp  borders  (Fig.  1). 
Female  club  with  apex  simple    caroticus  n.  sp. 

-  Hind  margin  of  Tl  entire  medially.  Dark  bands  on  tibiae  with  diffuse  borders  (Fig.  2). 
Female  club  with  terminal  spine  at  apex cephalon  (Walker) 

4.  Clypeal  denticles  symmetrically  arranged.  Petiole  with  weak  median  carina.  Ovipositor 
exserted  for  length  equal  to  that  of  T7 clypeattis  (Girault) 

-  Clypeal  denticles  asymmetrically  arranged,  the  median  denticle  displaced  to  the  left  (Fig. 
11).  Petiole  without  a  median  carina  (Figs  18-19).  Ovipositor  sheaths  hardly  exserted   ....     5 

5.  Costal  cell  with  three  rows  of  setae  distally.  Eye  length  <4X  as  long  as  the  temple  length 
(Fig.  5).  Body  more  or  less  all  dark   cyzicus  (Walker) 

-  Costal  cell  with  two  rows  of  setae  distally.  Eye  length  >4X  as  long  as  the  temple  (Figs  3, 

7).  Body  dark  with  diffuse  metallic  patches  on  head  and  mesosoma 6 

6.  Male  with  terminal  segment  of  funicle  appearing  as  wide  or  wider  than  long  (Fig.  4). 
Petiole  usually  less  than  1.7X  as  long  as  wide  (Fig.  18).  Female  with  petiole  usually  less 
than  1.5X  as  long  as  wide.  Both  sexes  with  hind  margins  of  Tl  and  T2  as  long  medially 

as  laterally,  (eastern  United  States  and  Canada) cyzicopsis  n.  sp. 

-  Males  with  terminal  segment  of  funicle  appearing  longer  than  wide  (Fig.  6).  Petiole  usually 
more  than  1.7x  as  long  as  wide.  Female  with  petiole  usually  more  than  1.5  x  as  long  as 
wide  (Fig.  19).  Both  sexes  with  hind  margins  of  Tl  and  T2  usually  longer  laterally  than 
medially,  (far  western  United  States  and  Canada)    petiolatus  n.  sp. 


Thinodytes  caroticus  Heydon,  new  species 

(Fig.  1) 

Holotype,  female. — Color:  Body  dark 
blue-green  except  flagellum,  fore  tarsus, 
wing  veins  brown;  pretarsi  black;  knees, 
apical  K;  of  tibiae,  middle  and  hind  tarsi 
white,  border  between  the  light  ends  and 
dark  median  band  of  tibiae  sharp  (Fig.  1). 

Sculpture:  Clypeus  smooth;  head  and 
mesoscutum  delicately  and  regularly  al- 
veolate; scutellum,  frenum  finely  alveo- 
late; median  panels  of  propodeum  ob- 
scurely alveolate. 

Structure:  Body  length  1.2  mm.  Head 
width  1.4X  height  (23:17),  2.3X  length  (23: 
10);  anterior  margin  of  clypeus  with  single 
broad  tooth;  malar  distance  5x  length  of 
genal  concavity;  eye  height  1.2X  length 
(10:8);  2.0X  malar  distance  (10:5),  eye 
length  4.0  X  temple  length  (8:2);  ratio  of 
MOD,  OOL,  POL,  LOL  as  2.0:3.0:6.5:3.0; 
vertex  rounding  regularly  into  occiput;  to- 
rulus  Vi  own  diameter  above  LOcL.  An- 
tenna with  length  of  pedicel  plus  flagel- 


lum  0.87X  head  width  (20:23);  relative 
lengths  of  scape,  pedicel,  annelli,  Fl-6, 
club  as  6.5:2.5:0.5:2.0:2.0:2.0:2.0:2.0:2.0:5.0; 
widths  of  Fl,  F6,  club  as  2:3:3;  apical  club 
segment  simple  apically,  with  micropilos- 
ity  ventrally.  Mesosoma  arched  dorsally, 
length  1.3X  width  (25:19);  dorsellum  short 
smooth  band;  propodeum  with  basal  fo- 
vea part  of  continuous  groove  across  an- 
terior margin  of  median  panels,  with  short 
weak  longitudinal  carina  crossing  groove 
halfway  between  plica  and  median  carina, 
with  spiracles  on  anterior  margin  of  pro- 
podeum, with  nucha  lunate  strip  and  car- 
inate  anteriorly.  Fore  wing  with  ratio  of 
lengths  of  submarginal,  marginal,  post- 
marginal,  stigmal  veins  as  19.0:11.5:9.0:5.0; 
stigma  small;  basal  cell  bare;  basal  vein 
with  row  of  4  setae.  Petiole  conical,  trans- 
verse, smooth,  with  median  carina.  Gaster 
fusiform,  length  1.8X  width  (30.0:16.5); 
hind  margin  of  Tl  emarginate  medially; 
ovipositor  sheaths  hardly  exserted;  hypo- 
pygium  extending  Vi  gastral  length. 


Journal  of  Hymenoptera  Research 


Allotype,  male. — Color  pattern  similar  to 
holotype  except  basic  body  color  dark 
blue;  fore  tarsi  pale  brown;  pale  portions 
of  legs  pale  yellow-brown  instead  of 
white.  Body  length  1.2  mm.  Antenna  with 
length  of  pedicel  plus  flagellum  0.98  X 
head  width  (21.0:21.5);  relative  lengths  of 
scape,  pedicel,  annelli,  Fl-6,  and  club  as 
6.0:2.5:0.5:2.0:2.0:2.0:2.0:2.0:2.0:6.0;  widths 
of  Fl,  F6,  club  as  2.5:3.0:3.0;  funicular  setae 
sparse,  reclinate.  Gaster  ovate,  length  1.5X 
width  (23:15). 

Variation. — The  body  length  of  females 
examined  varied  between  1.2  and  1.8  mm 
and  males  between  1.0  and  1.4  mm.  The 
color  of  the  dorsum  of  the  mesosoma  var- 
ies from  dark  blue-green  to  dark  blue.  The 
dark  bands  on  the  legs  are  always  distinct, 
but  the  intensity  of  their  metallic  colora- 
tion is  variable. 

Discussion. — Thinodi/tes  caroticus  most 
closely  resembles  T.  cephalon  because  both 
species  are  dark  in  color;  have  a  single, 
broad,  asymmetrically  placed  clypeal  den- 
ticle; and  have  smooth,  transverse  peti- 
oles. Thinodytes  caroticus  differs  from  T.  ce- 
phalon in  the  following:  1.  The  hind  mar- 
gin of  Tl  is  emarginate  in  T.  caroticus,  but 
straight  in  T.  cephalon.  2.  The  basal  vein  of 
the  fore  wing  is  setose  in  T.  caroticus,  but 
bare  in  T.  cephalon.  3.  The  median  panels 
of  the  propodeum  are  weakly  sculptured 
in  T.  caroticus,  but  distinctly  alveolate  in 
T.  cephalon.  4.  The  hypopygium  extends 
about  Vi  the  length  of  the  gaster  in  T.  car- 
oticus, but  about  %  the  gastral  length  in  T. 
cephalon.  5.  The  dark  bands  on  the  tibiae 
are  distinct  with  sharp  borders  in  T.  caro- 
ticus, but  are  less  distinct  and  have  diffuse 
borders  in  T.  cephalon.  These  distinctive 
tibial  color  bands  are  unique  to  T.  caroticus 
and  will  identify  the  species  at  a  glance 
(Fig.  1). 

Etymology. — The  species  name  comes 
from  the  Greek  karotikos,  meaning  stupe- 
fying or  soporific,  and  refers  to  the  general 
nondescript  appearance  of  this  species. 

Type  Material. — The  holotype,  allotype 
(both  UCDC)  and  one  male  paratype  were 


all  reared  by  the  author  from  the  leaf 
mines  of  Calycomyza  promissa  (Frick)  (Dip- 
tera:  Agromyzidae)  collected  30  June  1985 
on  the  South  Farms  of  the  University  of 
Illinois,  near  Champaign,  Illinois.  Fifty- 
nine  additional  paratypes  seen  were  col- 
lected as  follows  (CNCI,  INHS,  SEMC, 
UCDC,  USNM):  Bermuda.  DEVONSHIRE 
PARISH:  Devonshire  Marsh,  27.VI.1988,  1 
male.  FAGOT  PARISH:  Berry  Hill  Road, 
29.VI.1988,  2  females,  3  males;  Botanical 
Garden,  27. VI.  1988,  1  female;  Camdon 
Marsh,  29.VI.1988,  1  female.  SMITHS 
PARISH:  Spittal  Pond,  27.VI.1988,  1  fe- 
male. Canada.  ONTARIO:  Chatham,  1952 
(mass  reared  from  Hessian  fly  straw),  1 
male.  United  States.  CALIFORNIA:  Haw- 
thorne, IX.  1940  (ex  Aster  blotch),  3  fe- 
males, 1  male;  Jepson  Prairie  Preserve  (13 
km  s.  Dixon),  20.V.1983,  1  female;  Lake 
Hennessy  (11  km  ese.  St.  Helena), 
28.x.  1990,  1  female;  Los  Angeles  County 
(bred  from  dipterous  leaf  miner),  1  female; 
Sacramento,  16. IX.  1924  [ex  Agromyza  pus- 
ilia  {prob.= Liriomyza  pusilla)],  1  female;  So- 
quel,  26.VIII.1948  (ex  Agromyza  sp.),  2  fe- 
males, 1  male;  11  km  e.  St.  Helena  (Lake 
Hennessey),  28. X.  1990  (on  Baccharis),  1  fe- 
male, 7.IX.1991  (on  Heraculeum),  1  male;  6 
km  e.  Suisun  City,  11.VIII.1990,  2  males; 
1000  Palms,  29.III.1977,  1  female;  West- 
wood  Hills  (Los  Angeles  County), 
5.XI.1940  (ex  serpentine  leaf  miner  in  Zin- 
nia), 4  females,  1  male.  FLORIDA:  Braden- 
ton,  27.XI.1946  [ex  Liriomyza  pissilla 
{proh.= Liriomyza  pusilla)],  1  male;  Jackson- 
ville, 2  females.  GEORGIA:  Savannah, 
5. VI.  1943  (parasite  of  goldenrod  leafmin- 
er),  3  females.  ILLINOIS:  South  Farms  of 
the  University  of  Illinois,  19.V.1985,  1  fe- 
male; White  Heath,  24.IX.1939,  1  female. 
INDIANA:  4  miles  s.  New  Harmony  (Har- 
mony State  Park),  28. VI.  1983,  2  females;  2 
miles  s.  New  Lisbon,  14. VII. 1981,  1  male. 
IOWA:  Sioux  City  (reared  from  leaf  miner 
on  sunflower  leaf),  1  female,  5. VIII.  1921 
(reared  from  mine  of  leaf  on  sunflower),  1 
female.  KANSAS:  Lawrence,  14.V.1955,  1 
female.      MINNESOTA:      Albert     Lea, 


Volume  4,  1995 


26.VII.1960,  1  female.  NEW  MEXICO:  Ci- 
marron, 1909,  1  female;  Springer,  1909,  1 
female,  3  males.  TEXAS:  Clarendon, 
19. IX. 1905  (on  Grindelia  squarrosa  (Pursk) 
Dunal  (Compositae),  1  female;  Ennis, 
27.IX.1905  [ex  dipterous  leaf  miner  of  Ma- 
chaeranthera  annua  (Rydb.)  Shinner  (Com- 
positae)], 2  females;  Roma,  26. III. 1948  (ex 
pupa  of  dipterous  leaf  miner),  3  females, 
3  males. 

Biology. — This  species  has  been  reared 
from  leaf-mining  Agromyzidae,  mostly 
on  Compositae.  Known  hosts  include  Ca- 
lycomyza  promissa  (Prick)  and  Liriomyza 
pusilla  (Meigen).  Because  L.  pusilla  is  a  Pa- 
learctic  agromyzid  species  (Spencer  1976) 
and  is  not  known  from  the  Nearctic  region 
(Spencer  and  Steyskal  1986),  it  is  likely 
that  this  host  record  is  in  error.  There  is 
one  record  from  Chatham,  Ontario  from 
Hessian  fly  straw.  Thinodytes  caroticus  was 
also  reared  from  leaf  miners  on  Machaer- 
anthera  annua,  Zinnia,  goldenrod,  sunflow- 
er, and  "Aster".  It  has  been  reared  from 
both  linear-mining  and  blotch-making 
leaf  miners.  Other  plant  associations  of  a 
more  uncertain  nature  include  the  com- 
posites Baccharis  sp.,  Heracleum  sp.,  and 
Grindelia  squarrosa. 

Thinodytes  clypeatus  (Girault) 

Polycystus  clypeatus  Girault,  1918:128.  Holotype, 
female  (USNM);  Hym.  Type  No.  20682;  (ex- 
amined). 

Thinodytes  clypeatus  (Girault):  Heydon,  1989: 
193. 

Redescription. — Holotype,  female.  Color: 
Mesosoma,  petiole  black,  with  blue  tints 
on  propodeum  and  pleural  regions;  gaster 
dark  brown;  legs  with  basal  %  of  femora 
brown,  remainder  of  legs  white. 

Sculpture:  Dorsum  of  mesosoma  regu- 
larly and  delicately  reticulate;  median 
panels  of  propodeum  weakly  alveolate; 
petiole  alveolate. 

Structure:  Head  with  anterior  margin  of 
clypeus  with  three  symmetrically  ar- 
ranged fingerlike  denticles,  torulus  above 
LOcL.  Antenna  with  lengths  and  widths 


of  segments  as  follows  (in  iJim):  scape 
19.5X3.6:  pedicel  6.0x4.5:  annelli  2.6x3.6: 
Fl  5X6:  F2  4.6X6.0:  F3  5.2x6.2:  F4  4.6x6.5: 
P5  4.6X6.8:  F6  missing:  club  13.4X6.8;  club 
simple  apically.  Mesosoma  with  dorsum 
rather  flat,  length  1.4X  width  (33:24);  pro- 
notum  with  collar  width  11 X  length  (22: 
2),  sides  converging  posteriorly;  notauli 
shallow  posteriorly;  scutellum  with  two 
pairs  of  lateral  setae,  frenum  almost  indis- 
tinguishable from  remainder  of  scutellum; 
propodeum  with  length  of  median  panels 
0.45  X  width,  with  row  of  foveae  separated 
by  carinae  along  anterior  margin,  spiracle 
on  anterior  margin  of  propodeum,  nucha 
a  raised  smooth  crescent.  Fore  wing  with 
relative  lengths  of  submarginal,  marginal, 
postmarginal,  stigmal  veins  as  21:12:11:6; 
stigma  large,  height  Vi  distance  between 
stigma  and  anterior  margin  of  wing;  basal 
cell  with  one  seta;  basal  vein  with  three 
setae.  Petiole  length  1.2X  width  (6:5);  with 
weak  median  carina;  lacking  lateral  setae. 
Gaster  fusiform,  length  1.7X  width  (31.0: 
18.5);  Tl  emarginate  medially;  hypopy- 
gium  extending  %  length  of  gaster;  ovi- 
positor sheaths  exserted  for  length  equal 
to  that  of  T7. 

Discussion. — The  holotype  has  the  head 
removed  and  crushed  on  a  slide.  Thinod- 
ytes clypeatus  differs  from  all  other  de- 
scribed Thinodytes  species  because  it  has 
an  enlarged  stigma,  exserted  ovipositor 
sheaths,  and  a  median  carina  on  the  peti- 
ole. Thinodytes  clypeatus  has  three  sym- 
metrically arranged  denticles  like  Synto- 
mopus  species,  but  T.  clypeatus  differs  from 
Syntomopus  species  in  several  ways.  The 
denticles  of  T.  clypeatus  are  fingerlike  lobes 
like  those  of  the  other  Thinodytes  species 
with  three  denticles,  whereas  the  denticles 
of  Syntomopus  species  are  rather  broadly 
triangular  (Fig.  14).  The  pronotal  collar  of 
Thinodytes  clypeatus  is  short,  eleven  times 
as  wide  as  long;  the  pronotal  collar  in  Syn- 
tomopus is  only  about  three  times  as  wide 
as  long  (Fig.  22).  Most  species  of  Halticop- 
tera  have  a  median  carina  on  the  petiole, 
but  there  are  no  other  characters  to  indi- 


10 


Journal  of  Hymenoptera  Research 


cate  a  particularly  close  relationship  be- 
tween T.  clypeatus  and  Halticoptera. 

Distribution  and  Biology. — The  type  spec- 
imen was  reared  from  a  leaf  miner  on  corn 
on  2  May  1916,  on  St.  Vincent,  British 
West  Indies  by  F.  Watts.  De  Santis  (1979) 
reports  this  species  from  Barbados  also, 
and  gives  the  name  of  its  host  as  Agromyza 
parvicornis  Loew  (Diptera:  Agromyzidae). 

Thinodytes  cephnlon  (Walker), 

new  combination 

Figs  1,  17 

?Pteromnlus  Rhseo  Walker,  Walker  1839b:88-89. 
Lectotype,  female  (BMNH);  Hym.  Type  No. 
5.772  (examined). 

Gastmncistrus  cephalon  Walker  1843:30.  Lecto- 
type, male  (BMNH);  Hym.  Type  No.  5.661 
(examined). 

Bubekia  fallax  Gahan,  1933:114-116.  Holotype, 
female  (USNM);  Type  No.  44841  (examined). 
Allen  and  Painter,  1937:225.  Nikol'skaya, 
1937:25.  Peck,  1951:538.  Thompson,  1958:587. 
Peck,  1963:610-611.  Morrill  and  Kieckhefer, 
1971:1130.  Allen  and  Pienkowski,  1973:616, 
617.  Burks,  1979:789.  Hendrickson,  1979:300, 
302.  n.  syn. 

Discussion. — Thinodytes  cephalon  is  easily 
distinguished  by  its  single  broad  clypeal 
denticle,  apical  spine  on  the  female  club, 
notauli  traceable  to  the  hind  margin  of  the 
mesoscutum  as  impressed  lines,  bare  dor- 
sal vein,  smooth  and  short  petiole  (Fig. 
17),  and  first  gastral  tergite  having  the 
hind  margin  entire. 

A  possible  senior  synonym  of  T.  cephal- 
on is  Pteromalus  rhaeo  described  from  Chil- 
oe  Island,  Chile.  The  lectotype  female 
(designated  herein)  of  P.  rhaeo  is  unfortu- 
nately missing  the  head.  It  appears  similar 
to  specimens  of  T.  cephalon  examined,  of 
which  many  specimens  in  my  collection 
and  the  CNC  collection  were  from  Chiloe 
Island,  but  it  is  significantly  larger  than 
any  other  specimen  examined.  The  medi- 
an panels  of  the  propodeum  in  P.  rhaeo  are 
entirely  rugose  and  the  plicae  are  poste- 
riorly divergent.  In  other  T.  cephalon  ex- 
amined, the  median  panels  have  few  ru- 


gae which  are  located  mostly  posteriorly, 
and  the  plicae  are  parallel.  The  hind  tibia 
in  P.  rhaeo  is  almost  uniform  in  color;  in 
T.  cephalon,  the  hind  tibia  are  distinctly 
dark  over  the  middle  half  or  more.  Pter- 
omalus rhaeo  may  be  within  the  range  of 
variation  of  what  is  recognized  as  T.  ce- 
phalon, but  I  am  reluctant  to  formally  syn- 
onymize  the  name  until  more  is  known 
about  the  range  of  variation  of  T.  cephalon 
in  South  America. 

Distribution. — Thinodytes  cephalon  is  one 
of  the  most  commonly  collected  New 
World  pteromalids.  Its  distribution  ex- 
tends throughout  most  of  North  and 
South  America  (from  Chile  to  Canada  and 
also  Bermuda). 

Biology. — Thinodytes  cephalon  is  a  para- 
sitoid  of  dipterous  larvae  that  mine  leaves 
or  grass  stems.  Thinodytes  cephalon  was 
originally  described  from  puparia  of  the 
Hessian  fly,  Phytophaga  destructor  (Say) 
(Diptera:  Cecidomyiidae)(Gahan  1933).  It 
has  also  been  reared  from  the  wheat  stem 
maggot,  Meromyza  americana  Fitch  (Dip- 
tera: Chloropidae),  in  a  number  of  studies 
(Gahan  1933;  Allen  and  Painter  1937;  and 
Morrill  and  Kieckhefer  1971).  Allan  and 
Painter  report  that  T.  cephalon  probably 
oviposits  into  the  larval  stage  of  this  host. 
One  additional  host  added  herein  is  Lir- 
iomyza  trifoliearum  Spencer  (Diptera:  Agro- 
myzidae) on  alfalfa. 

Thinodytes  cyzicopsis  Heydon, 
new  species 
Figs.  3,  4,  18 

Holotype,  female. — Color:  Head,  mesoso- 
ma  black  with  metallic  green  patches  as 
follows:  entire  dorsellum,  pairs  of  spots 
alongside  median  ocellus,  lateral  hind  cor- 
ners of  pronotum  and  upper  epimeron; 
metallic  blue  patches  as  follows:  on  inner 
orbits,  anterior  part  of  lateral  lobe  of  scu- 
tum and  propodeum.  Antenna  with  scape, 
pedicel  dark  blue;  flagellum  black.  Petiole 
black.  Caster  brownish  black  with  metallic 
blue  reflections  dorsally,  green  ventrally. 
Legs  with  coxae  black  with  weak  blue  re- 


Volume  4, 1995 


11 


flections;  trochanters,  femora  brownish 
black  with  weak  green  reflections,  knees 
yellow,  tibiae  with  diffuse-edged  dark 
brown  band  extending  %  their  length;  fore 
tarsi  brown;  middle  and  hind  tarsi  with 
pretarsi  black,  remainder  yellow. 

Sculpture:  Head,  mesosoma,  regularly 
and  delicately  alveolate;  frenum,  propo- 
deum  shallowly  and  indistinctly  alveolate; 
petiole  finely  alveolate. 

Structure:  Body  length  1.6  mm.  Head 
width  1.2X  height  (27:22),  2.2  X  length  (27: 
12);  anterior  margin  of  clypeus  with  three 
minute  asymmetrically  arranged  apical 
denticles;  genal  concavity  absent;  eye 
height  1.4X  length  (13:9),  1.9X  malar  dis- 
tance (13:7),  length  4.5 X  temple  length  (9: 
2)  (Fig.  3);  ratio  of  MOD,  OOL,  POL,  LOL 
as  2.5:4.0:7.0:3.0;  vertex  rounding  smooth- 
ly into  occiput;  torulus  V2  own  diameter 
above  LOcL.  Antenna  with  length  of  ped- 
icel plus  flagellum  l.Ox  head  width  (27: 
27);  relative  lengths  of  scape,  pedicel,  an- 
nelli,  Fl-6,  club  as  11.0:3.0:1.0:2.5:3.0:2.75: 
2.75:2.5:2.5:7.0;  widths  of  Fl,  F6,  club  as 
3.0:3.5:3.5,  Fl^  appearing  quadrate,  F5-6 
slightly  transverse;  club  simple  apically, 
with  minute  patch  of  micropilosity  on  ter- 
minal segment.  Mesosoma  arched  dorsal- 
ly,  length  1.6X  width  (36:22);  pronotum 
with  humeral  angles  squared;  mesoscu- 
tum  with  notauli  shallow;  dorsellum 
bandlike,  short,  weakly  alveolate;  propo- 
deum  with  basal  fovea  bordered  mesally 
by  sinuous  carina;  nucha  a  raised  band, 
weakly  carinate  anteriorly;  spiracle  < 
0.5  X  own  diameter  from  anterior  margin 
of  propodeum.  Fore  wing  with  ratio  of 
lengths  of  submarginal,  marginal,  post- 
marginal,  stigmal  veins  as  25:13:14:8;  stig- 
ma small,  maximum  width  only  slightly 
more  than  stigmal  vein  width;  basal  cell 
bare;  basal  vein  setose.  Petiole  length  1.2X 
width  (6:5);  without  median  carina;  with 
one  pair  of  lateral  setae.  Gaster  length 
1.4X  width  (31:22);  hypopygium  extend- 
ing to  about  %  length  of  gaster;  ovipositor 
sheaths  hardly  exserted. 

Allotype,  male. — Color  similar  to  holo- 


type  except  frons  blue,  mesoscutum  with 
a  pair  of  large  diffuse  green  spots;  axilla 
green.  Body  length  1.3  mm.  Antenna  (Fig. 
4)  with  length  of  pedicel  plus  flagellum 
l.Ox  head  width  (24:23);  relative  lengths 
of  scape,  pedicel,  annelli,  Fl-6,  club  as  7.0: 
3.0:1.0:2.0:2.5:2.5:2.5:2.5:2:6.5;  widths  of  Fl, 
F6,  club  as  2.0:2.5:2.5;  setae  fine,  reclinate. 
Petiole  length  1.2X  width  (5:4)(Fig.  18). 
Gaster  ovate,  length  1.2X  width  (20.0: 
16.5). 

Variation. — The  body  color  varies  from 
mostly  dark,  like  the  holotype,  to  the  pat- 
tern shown  by  the  allotype  male.  The  in- 
tensity, size,  number,  and  hue  of  the  me- 
tallic spots  on  the  body  are  highly  vari- 
able. The  color  of  the  metallic  patches  var- 
ies from  coppery  green  to  green  to  blue- 
green.  The  body  length  ranges  between 
1.4  and  1.8  mm  in  females  and  1.2  and  1.7 
mm  in  males.  The  length  of  the  marginal 
vein  varies  from  0.9  to  1.2  times  the  length 
of  the  postmarginal  vein  but  averages 
about  equal  its  length  [x  =  1.02± 
(S.E.  =  )0.02  (n  =  12)].  The  petiole  averages 
1.39±0.045  (n=7,  range  1.2-1.5)  times  as 
long  as  wide  in  the  females  and 
1.57 ±0.061  (n  =  7,  range  1.2-1.9)  times  in 
males  (Fig.  18).  The  pair  of  lateral  setae  on 
the  petiole  are  sometimes  difficult  to  see 
because  they  tend  to  lie  near  the  anterior 
flanges  of  the  petiole. 

Discussion. -Thinodytes  cyzicopsis  resem- 
bles the  Palearctic  species  T.  cyzicus 
(Thomson),  but  differs  from  that  species  in 
the  following:  1.  Thinodytes  cyzicopsis  has 
a  series  of  metallic  patches  on  the  body, 
which  are  lacking  in  T.  cyzicus.  2.  The  ratio 
of  the  eye  length  to  temple  length  aver- 
ages 4.25±0.12  (n=6,  range  3.8^.5)  in  fe- 
male T.  cyzicofisis  (Fig.  3)  but  measured 
only  2.3  and  3.6  in  two  female  T.  cyzicus 
specimens  examined  (Fig.  5).  3.  The  costal 
cell  of  the  fore  wing  of  T.  cyzicopsis  has 
two  rows  of  setae  distally,  whereas  there 
are  three  rows  distally  in  the  costal  cell  of 
T.  cyzicus.  4.  The  petiole  has  a  pair  of  lat- 
eral setae  in  T.  cyzicopsis,  which  are  lack- 
ing from  T.  cyzicus. 


12 


Journal  of  Hymenoptera  Research 


Thinodytes  cyzicopsis  and  T.  petiolatus  are 
very  similar  species  and  specimens  cannot 
always  be  confidently  separated.  These 
two  species  are  distinct  from  other  Thinod- 
ytes in  their  common  possession  of  a  sim- 
ilar pattern  of  metallic  patches  on  the  head 
and  mesosoma.  Individuals  of  T.  cyzicopsis 
differ  from  those  of  T.  petiolatus  in  the  fol- 
lowing: 1.  The  ratio  of  eye  height  to  malar 
distance  in  T.  cyzicopsis  averages 
1.74±0.035  (n=8,  range  1.5-1.8)  in  females 
and  2.04  ±0.032  (n=9,  range  1.9  to  2.0)  in 
males;  in  T.  petiolatus  the  ratio  is  1.48  ±0.23 
(n=10,  range  1.4  to  1.6)  in  females  and 
1.77±0.038  (n=10,  range  1.6  to  2.0)  in 
males.  2.  All  funicular  segments  of  the 
male  antenna  are  transverse  to  quadrate  in 
T.  cyzicopsis;  all  funicular  segments  are 
usually  longer  than  wide  in  male  T.  pe- 
tiolatus. 3.  The  ratio  of  petiole  length  to 
width  in  T.  cyzicopsis  averages  1.39  ±0.045 
(n=7,  range  1.2  to  1.5)  in  females  and 
1.57±0.061  (n  =  7,  range  1.2  to  1.9)  in 
males;  in  T.  petiolatus  it  averages 
1.93±0.062  (n=10,  range  1.5  to  2.0)  in  fe- 
males (Fig.  19)  and  1.95±0.071  (n=10, 
range  1.6  to  2.4)  in  males.  4.  The  first  gas- 
tral  tergite  in  T.  cyzicopsis  is  about  as  long 
medially  as  laterally,  whereas  Tl  is  often 
shorter  medially  than  laterally  in  T.  petiol- 
atus. 5.  The  hind  margin  of  T2  is  straight 
to  just  noticeably  concave  in  T.  cyzicopsis, 
whereas  it  is  usually  distinctly  concave  in 
T.  petiolatus.  6.  The  terminal  segment  of 
the  male  funicle  appears  quadrate  to 
slightly  transverse  in  T.  cyzicopsis,  whereas 
it  appears  elongate  in  T.  petiolatus.  In  ad- 
dition to  these  structural  characters,  T.  cy- 
zicopsis is  found  east  of  the  Rocky  Moun- 
tains, and  T.  petiolatus  occurs  from  the 
Rockies  west. 

Type  Material. — The  holotype  (CNCI)  is 
from  Kouchibouquac  National  Park,  New 
Brunswick,  and  was  collected  9  August 
1977  by  S.  J.  Miller.  The  allotype  (USNM) 
is  from  Ithaca,  New  York,  and  was  col- 
lected on  vernal  alfalfa  on  22  June  1968  by 
A.  G.  Wheeler.  Sixteen  paratypes  were  col- 
lected as  follows  (CNCI,  UCDC,  USNM): 


Canada.  ALBERTA:  Elkwater  Lake, 
21.VII.1956,  1  female;  Lethbridge,  1  fe- 
male. NEW  BRUNSWICK:  Kouchibou- 
quac National  Park,  20.IX.1977,  1  female. 
NOVA  SCOTIA:  Alton,  IX.  1964  [ex  Phy- 
tobia  {Calycomyza)  solidaginis  on  Solidago],  1 
male;  Crosby,  31  .VII.  1952  (on  apple),  1 
male.  ONTARIO:  Ottawa,  22.VI.1972 
(swept  from  Salix  hlanda),  1  male, 
29.VI.1972  (swept  from  Salix  blanda),  3 
males,  24. VII. 1972  (swept  from  Salix  hlan- 
da), 1  male.  QUEBEC:  Lac  Brule, 
21.VII.1947,  1  female,  25.VII.1947  (swept 
from  Rosa  rugosa),  1  female,  9.VIII.1945,  1 
female.  United  States.  ILLINOIS:  2  miles 
e.  Shumway,  7.VII.1980,  1  female;  South 
Farms  of  the  University  of  Illinois,  nr. 
Champaign,  23.VI.1981,  1  male.  MASSA- 
CHUSETTS: Hopkinton,  9.VIII.1951  (ex 
Ilex  leaf  miner),  1  female.  NEW  MEXICO: 
Mesilla,  4. V.  1909,  1  female.  WEST  VIR- 
GINIA: Winchester,  16.VI.1964,  1  male, 
17.VI.1964,  1  male. 

Etymology. — The  species  name  is  de- 
rived from  the  species  name  of  Thinodytes 
cyzicus  and  the  Greek  suffix  -opsis,  mean- 
ing like  or  similar  in  appearance,  and  re- 
fers to  the  morphological  similarity  be- 
tween T.  cyzicopsis  and  T.  cyzicus. 

Biology. — Known  hosts  of  T.  cyzicopsis 
include  Phytobia  {Calycomyza)  solidaginis 
(Agromyzidae)[on  Solidago  sp.  (Composi- 
tae)]  and  an  Ilex  leafminer.  The  species  has 
also  been  taken  on  Salix  blanda  (Salica- 
ceae),  Rosa  rugosa  (Rosaceae),  and  alfalfa. 
Salix  and  Rosa  are  probably  primarily  nec- 
tar sources  because  neither  has  many  leaf- 
mining  agromyzids,  but  these  plants  are 
important  sources  of  nectar  and /or  hon- 
eydew  for  many  parasitic  Hymenoptera. 

Thinodytes  cyzicus  (Walker) 

Miscogaster  cyzicus  Walker,   1839a:200.   Lecto- 

type,  female  (BMNH);  Hym.  Type  No.  5.2570 

(examined). 
Syntomopus  cyzicus  (Walker):  Walker,  1846:28. 

Schmiedeknecht,  1909:376. 
Dicyclus  circulus  Thomson,  1876:253.  Lectotype, 

female  (LUND),  not  seen. 


Volume  4,  1995 


13 


Thinodytes  cyzicus  (Walker):  Graham,  1956:261. 
Graham,  1969:167.  Askew,  1970:380.  Hedqv- 
ist,  1975:180.  Boucek,  1977:56.  Kamijo,  1978: 
457.  Takada  and  Kamijo,  1979:21,  22,  23,  25. 
Hedqvist,  1983:167.  Boucek  and  Rasplus, 
1991:32. 


Discussion. — The  synonymy  of  Dicyclus 
circiilus  with  Thinodytes  cyzicus  is  accepted 
on  the  authority  of  Graham  (1969).  Thi- 
nodytes cyzicus  resembles  T.  cyzicopsis  and 
T.  petiolatus,  because  all  three  species  have 
three  small,  sharp,  asymmetrically  ar- 
ranged clypeal  denticles  (Fig.  11)  and  a 
long,  reticulate  petiole.  Besides  the  geo- 
graphic separation  of  their  ranges,  T.  cy- 
zicus can  be  distinguished  from  the  two 
Nearctic  species  by  its  body  color.  Thino- 
dytes cyzicus  is  uniformly  dark  and  lacks 
the  distinctive  diffuse  metallic  patches  on 
the  head  and  mesosoma  present  in  T.  cy- 
zicopsis and  T.  petiolatus.  Thinodytes  cyzicus 
also  has  three  rows  of  setae  distally  in  the 
costal  cell,  whereas  T.  cyzicopsis  and  T.  pe- 
tiolatus have  only  two  rows.  The  eye 
length  in  T.  cyzicus  is  4X  or  less  the  length 
of  the  temple  (Fig.  5),  whereas  T.  cyzicopsis 
and  T.  petiolatus  have  the  eye  length  more 
than  4X  the  length  of  the  temple  (Figs  3, 

7). 

Distribution. — Thinodytes  cyzicus  occurs 
throughout  the  Palearctic  region,  from 
Britain  to  Japan  (Graham  1969;  Boucek 
1970;  Kamijo  1978). 

Biology. — The  hosts  of  T.  cyzicus  are  all 
Agromyzidae.  Askew  (1970)  recorded  this 
species  from  an  agromyzid,  probably  Phy- 
tomyza  atricornis  Meigen,  on  Senecio  jaco- 
baea  Linnaeus.  Kamijo  (1978)  recorded  T. 
cyzicus  from  Chromatomyia  horticola  (Gou- 
reau)(Diptera:  Agromyzidae)  on  pea  and 
from  an  agromyzid  on  Lathy rus  maritimus. 
Takada  and  Kamijo  (1979)  recorded  T.  cy- 
zicus as  emerging  from  the  puparium  of 
Chromatomyia  horticola  and  speculated  that 
T.  cyzicus  may  parasitize  the  larval  stage 
of  its  host. 


Thinodytes  petiolatus  Heydon, 

new  species 

Figs  6,  7,  11,  19 

Holotype,  female. — Color:  Body  black  but 
frenum,  metanotum  coppery;  frons,  ver- 
tex, gena,  lateral  region  of  pronotum,  lat- 
eral lobe  of  mesoscutum,  middle  lobe  of 
mesoscutum  with  pair  of  large  diffuse  me- 
tallic patches,  anterior  lateral  corner  of  ax- 
illa dark  metallic  green;  gaster  with  green- 
ish reflections.  Antenna  with  scape  dark 
green;  remainder  brown,  pedicel  with 
weak  green  reflections.  Legs  with  coxae 
black  with  greenish  reflections;  femora 
brown  with  greenish  reflections;  remain- 
der of  legs  mostly  yellow-brown,  tibiae 
with  broad  diffuse  brown  band  mesally, 
apical  two  tarsi  brown.  Wing  veins  pale 
reddish  brown. 

Sculpture:  Clypeus  weakly  alveolate; 
face  alveolate,  cells  elongate  in  radiating 
fashion  from  clypeus;  remainder  of  head 
alveolate;  mesoscutum,  scutellum  deli- 
cately and  regularly  alveolate  except  fre- 
num smooth;  median  panels  of  propo- 
deum  alveolate;  petiole  finely  alveolate. 

Structure:  Body  length  1.8  mm.  Head 
width  1.4X  height  (28:21),  2.6 X  length 
(28.5:11.0);  clypeus  with  three  small  asym- 
metrically arranged  clypeal  denticles  (Fig. 
11);  weak  genal  concavity  extending  Vs 
malar  distance;  eye  height  1.3X  length 
(12.0:9.5),  1.5X  malar  distance  (12:8), 
length  4.8X  temple  length  (9.5:2.0)(Fig.  7) 
ratio  of  MOD,  OOL,  POL,  LOL  as  2:4:7:3 
vertex  rounding  regularly  into  occiput 
antennal  torulus  just  above  LOcL.  Anten- 
na with  length  of  pedicel  plus  flagellum 
0.91  X  head  width  (26:28.5);  ratio  of 
lengths  of  scape,  pedicel,  annelli,  Fl-6, 
club  as  11.5:3.5:1.0:2.5:2.5:2.5:2.5:2.5:2.5:6.0; 
widths  of  Fl,  F6,  club  as  2:3:3;  club  simple 
apically,  with  small  patch  of  micropilosity 
ventrally  on  apical  segment.  Mesosoma 
arched  dorsally,  length  1.7X  width  (37:22); 
notauli  shallow  posteriorly;  propodeum 
(Fig.  19)  with  basal  fovea  margined  mesal- 
ly by  long  sinuous  carina;  nucha  a  weakly 


14  Journal  of  Hymenoptera  Research 

sculptured  band,  carinate  anteriorly;  spi-  arate  these  two  species  are  given  in  the 

racle  <0.5X  own  diameter  from  anterior  discussion  section  for  T.   cyzicopsis.   The 

margin  of  propodeum.  Fore  wing  with  rel-  hind  margin  of  T2  in  many  specimens  of 

ative   lengths  of  submarginal,   marginal,  T.  petiolatus  is  distinctly  concave,  but  the 

postmarginal,  stigmal  veins  as  27.0:14.5:  visibility  of  this  character  depends  on  how 

15.0:9.0;  stigma  small;  basal  vein  with  row  the  specimen  has  dried.  This  character  is 

of  setae   along   length.   Petiole   (Fig.    19)  more  distinct  in  the  males  than  in  the  fe- 

length  2.0  X  width  (8:4);  without  median  males. 

carina;  with  two  pairs  of  lateral  setae.  Gas-  Type  Material: — The  holotype  (USNM) 

ter  ovate-acuminate,  length  1.4X   width  was  collected  at  Roseworth,  Idaho,  on  19 

(33:24);  hind  margin  of  Tl  strongly  sinu-  June  1931,  on  Salsola  pestifer  by  D.  E.  Fox 

ous  laterally,  emarginate  mesally;  ovipos-  and  the  allotype  (USNM)  was  collected  at 

itor  sheaths  hardly  exserted;  hypopygium  Twin  Falls,  Idaho,  on  5  August  1920  by  R. 

extending  %  gastral  length.  H.  Smith.  Sixty  paratypes  were  collected 

Allotype,  male.— Body  color  similar  to  as  follows  (CNCI,  INHS,  UCDC,  USNM): 
holotype  except  metallic  spots  more  dif-  Canada.  ALBERTA:  Banff  (Sunshine 
fuse  and  extensive;  frenum,  metanotum  Lodge,  7500'),  24. VII. 1962,  1  female;  Elk- 
blue-green;  dark  bands  on  tibiae  very  water,  9.VI.1956,  1  male;  Elkwater  Lake, 
weak.  Body  length  1.3  mm.  Antenna  (Fig.  21.VII.1956,  1  male;  Lethbridge,  7.VI.1956 
6)  with  length  of  pedicel  plus  flagellum  (swept  from  barley),  1  female,  5.VIII.1956 
1.1  X  head  width  (28:25);  ratio  of  lengths  (swept  from  barley),  1  male;  nr.  Leth- 
of  scape,  pedicel,  annelli,  Fl-6,  club  as  9.0:  bridge,  1924.1925,  1  female.  BRITISH  CO- 
3.5:1.0:2.5:3.0:3.0:3.0:3.0:3.0:8.0;  widths  of  LUMBIA:  Bowser,  28.V.1955,  1  female; 
Fl,  F6,  club  as  2:2:3;  setae  of  flagellomeres  McQueen  Lake,  10  miles  n.  Kamloops, 
fine,  reclinate.  Petiole  length  2.0  X  width  18.VI.1973,  1  male.  United  States.  ARI- 
(8:4).  Gaster  length  1.2X  width  (20:16);  ZONA:  Mesa,  3.VI.1958  (swept  from  alfal- 
truncate  apically.  fa),    1     male.    CALIFORNIA:    Albany, 

Variation. — The  body  color  varies  from  29. IV. 1958,  1  male;  Alpine  Lake,  VI. 1971, 

like  that  of  the  holotype  to  very  dark,  with  1  female;  Apple  Valley,  8.V.1955,  2  males; 

the  green  areas  on  the  head  and  the  me-  15  miles  w.  Baker,  6.V.1977,  2  females,  1 

sosoma  reduced  and  obscure,  the  frenum  male;  Boca,  22. VII. 1970,  1  female;  Bolinas, 

and  metanotum  green,  and  the  tibiae  dark  5.X.1975,  2  males;  Camino,  21.VII.1948  (ex 

metallic  green.  The  body  length  of  females  Phytomyza  aqiiilegiana),  10  females;  Cerro 

examined  varied  between  1.3  and  1.8  mm  Noroeste    (sw.    corner    of    Kern    Co.), 

and  males  varied  between  1.0  and  1.6  mm.  15.VII.1965,  1  female,  Cuyler  Harbor  (San 

The   sinuous   carina   on   the  propodeum  Miguel  Island),  11. VII. 1970,  1  male;  Dar- 

sometimes  extends  only  Va  the  length  of  win     Falls     (nr.     Panamint     Springs), 

the  propodeum.  The  row  of  setae  on  the  29. III. 1984  (on  Encelia),  1  male;  Emeryville, 

basal   vein   sometimes   curls   proximally,  28. V. 1958,   1    male;   Eureka   Dunes   (Inyo 

running  a  short  distance  along  the  cubital  Co.),    15.V.1979,    1    male;    Lake    Tenaya, 

vein.  23.VII.1949  (host  Recurraria  milleri),  1  fe- 

Disciission. — Thinodytes  petiolatus  and  T.  male;  Lake  Tahoe,  29. VI.  1927,  1   female; 

cyzicopsis  are  very  similar  species  and  are  Lily  Pond  (alpine  lake),  VI. 1971,  1  female; 

distinct  from  other  Thinodytes  species  in  Los  Angeles  {Agromyza  playptera  Thom.),  1 

the  similar  pattern  of  metallic  patches  on  female;  Los  Angeles  Co.,  1  male;  McClure 

an  otherwise  dark  body.  Individuals  of  T.  Beach  (Marin  Co.),  18.VII.1970,  1  male;  Mt. 

petiolatus  and  T.  cyzicopsis,  especially  fe-  Ingalls  (Placer  Co.),  11. VII.  1964,  1  male; 

males,    cannot   always   be    distinguished  near  Nicasio,  15.11.1991  (sweeping  Salix),  2 

with  absolute  certainty.  Characters  to  sep-  males;    22    miles    w.    Panamint    Springs, 


Volume  4,  1995  15 

7.V.1961,  2  males;  Placer  Co.,  Vlll,  1  male;  (5),  1  male;  Moscow,  6.VIII.1926,  1  male; 
Sagehen  Creek  (near  Hobart  Mills),  24.  Murtaugh,  29.V.1930  (3  &  5),  1  male;  Oak- 
VI.1970,  1  female,  21-25.VI.1982  (sweep-  ley,  7.VIII.1929  (1,  3  &  5),  1  female;  Rupert, 
ing  Primus),  1  male,  12.VII.1972,  1  male,  29.V.1930,  1  female;  Tuttle,  22.V.1931  (4), 
11-15.VII.1982,  1  female,  23.VII.1968,  1  fe-  1  female;  Twin  Falls,  7.VI.1930,  1  male; 
male,  1.VIII.1970,  1  female;  San  Bernardi-  Wendell,  22.V.1931  (5),  3  males.  MON- 
no  Co.,  San  Gorgonio  Wilderness,  TANA:  Big  Butte,  8.VIII.195?,  1  male.  OR- 
19. VII. 1982,  1  male;  Santa  Cruz,  EGON:  Corvallis,  15.VI.1981,  1  male, 
22.V11I.1948  (ex  PJnjtonn/za  sp.  B),  1  female;  26.VI.1985,  1  female,  3  males;  Near  Cor- 
Santa  Rosa,  25.V.1990,  2  males;  Sheppard  vallis  (St.  Mary's  Peak),  15.VIII.1984  (road- 
Pass  Trail  (Inyo  Co.),  16. VII. 1985  (on  side  vegetation),  1  female;  Eugene,  6- 
Sphenosciadium  capitellatum),!  male;  Shive-  12.VI.1984  (Malaise  trap),  1  female,  11 
ly,  19. VI. 1959,  1  female;  Sonora  Pass  males;  McKinzie  Pass  (Mt.  Washington 
(Mono  Co.,  9624'),  20.V1II.1960,  1  female;  Wilderness),  17.VIII.1984,  2  females, 
Soquel,  26.VIII.1960  (ex  Agromijza  sp.),  2  20.VIII.1984,  5  females,  2  males;  1  mile  w. 
females;  Titus  Canyon  (Death  Valley  Na-  McKinzie  Pass,  1  female,  1  male.  UTAH: 
tional  Monument),  24.III.1984  (on  Cownia  Logan  Canyon,  21. VII.  1976,  1  female; 
mexicana),  1  male;  Tomales  Bay  State  Park  Monte  Cristo,  6.VII.1976,  1  male;  My  ton,  3 
(Marin  Co.),  14.IV.1961,  1  male;  Trinidad,  males;  Wellsville,  13.V.1964,  1  male;  1.5 
24.IX.1977,  1  female;  Truckee,  20.VII.1970,  miles  w.  Wild  Horse  Butte  (Wild  Horse 
1  female;  White  Water,  Snow  Creek  Creek),  31.VII.1982,  1  female.  WASHING- 
(1500'),  29.III.1955,  1  male;  6  miles  w.  Bas-  TON:  San  Juan  Island  (Barney's  Place), 
setts  (Yuba  Pass),  9.VII.1970,  2  females,  2  23.VII.1944,  1  male.  Vancouver,  15.VI. 
males.  COLORADO:  Chambers  Lake  (Lar-  1911,  1  female,  1  male,  16.VI.1911,  1  male, 
imer  Co.),  16.VIII.1966,  1  male;  16  km  n.  20.VI.1911,  1  female.  WYOMING:  Kem- 
Colorado  Springs,  25. VII. 1991,  1  male;  Ft.  merer,  12. VII. 1985  {Eriogomim),  2  females, 
Collins,  20. VIII. 1895  (on  boxelder  foliage),  13.VII.1985  [Artemesia  spinosa  {?=spines- 
1  female,  1  male;  Glacier  Basin,  Rocky  cens  D.C.  Eaton],  1  male;  Snowy  Range, 
Mountain  National  Park,  24.VII.1977,  1  23.VIII.1951,  2  females,  1  male, 
male;  Echo  Lake  (Mt.  Evans,  10,500'),  Etymologxj. — The  species  name  refers  to 
4. VIII. 1961,  1  female.  IDAHO:  Boise,  the  long  petiole,  characteristic  of  this  spe- 
28.V.1984  (Malaise  trap),  2  males;  Buhl,  cies. 

27.V.1929    (1    &    50,    2    males;    Burley,  B/o/ogi/.— The  host(s)  of  T.  petiolatus  is 

14.VI.1930  (3),  1  female,  9. VII. 1931,  1  fe-  unknown  but  it  has  been  taken  in  associ- 

male;    Eden,    11. VIII. 1930    (2),    1    female;  ation  with  a  number  of  plants,  such  as  Ar- 

Hobbs  Butte,  22.V.1931,  1  male,  6. VI. 1931,  temesia  sp.,  Descurainia  sophia,  Eriogomim, 

1  male;  Hollister,  16.V.1931   (3),  1   male,  Salsola  pestifer,  and  Sisymbrium  altissimum 

20.V.1931,  2  males,  2.VI.1931,  1  female,  1  in  Idaho  and  Encelia  in  California.  It  has 

male,  5. VI. 1931  (3),  1  female,  7. VI. 1931  (3  also   been   collected    in   association   with 

&  5),  1  female,  13.VI.1931, 1  female;  Hubbs  crop  plants,  such  as  beets  in  Idaho  and 

Butte,  22.V.1931,  1  male,  6.VI.1931,  1  male;  barley  in  Alberta. 
Jerome,  11.VII1.1930  (2),  1  male;  Kimberly, 

27.VI.1931,  2   females;  Milner,  29.V.1930  Thinodytes  santerna  Heydon,  new  species 

Holotype,  female. — Color:  Body  black 
with  blue-green  reflections  on  vertex,  side 

'  The  host  plants  for  the  specimens  from  Idaho  are  j^^^^g    ^f   mesoscutum,    SCUtellum,    gaster; 

numbered  as  follows:  1=  A.?  rosae.  2=  Beta  viiharis  n        ■                  <-                       i    i          i 

L.  3=  Soplua  sophia  {=Descuraima  sophia  (L.)  V.B.  coppery  reflections  on  frenum  and  dorsel- 

Webb).  4=  Salsola  pestifer  A.  Nelson.  5=  N.(ortn  )  al-  lum;  yellow-green  reflechons  on  middle 

tissiiniim  or  S.  altissiina  {  =  Sisymbri\nu  altissimum  L.).  lobe  of  SCUtellum  and  propodeum.  Anten- 


16 


Journal  of  Hymenoptera  Research 


na  with  scape  yellow,  flagellum  brown 
dorsally,  brownish  white  ventrally.  Legs 
yellow  beyond  coxae,  tarsi  white,  pretarsi 
brown.  Fore  wing  veins  pale  brown. 

Sculpture:  Clypeus  alveolate;  frenum, 
dorsellum,  propodeum  smooth;  gaster 
with  T5-7  coriaceous,  remainder  smooth. 

Structure:  Body  length  1.6  mm.  Head 
width  1.4X  height  (28:20),  2.3  X  length  (28: 
12);  clypeus  with  three  small  symmetri- 
cally arranged  clypeal  denticles;  weak 
genal  concavity  extending  Va  malar  dis- 
tance; eye  height  1.3  X  length  (11:10),  2.2  X 
malar  distance  (11:6),  length  5.0  X  temple 
length  (10:2);  ratio  of  MOD,  OOL,  POL, 
LOL  as  2:3:6:3;  vertex  rounding  regularly 
into  occiput;  antennal  torulus  just  above 
LOcL.  Antenna  with  length  of  pedicel  plus 
flagellum  0.86  X  head  width  (24:28);  ratio 
of  lengths  of  scape,  pedicel,  annelli,  Fl-6, 
club  as  10:3:1:2:2:2:2:2:2:6;  widths  of  Fl, 
F6,  club  as  2:2:2;  club  simple  apically,  with 
small  patch  of  micropilosity  ventrally  on 
apical  segment.  Mesosoma  arched  dorsal- 
ly,  length  1.5  X  width  (32:22);  notauli  shal- 
low posteriorly;  propodeum  with  basal  fo- 
vea obscure,  nucha  lunate  strip  and  cari- 
nate  anteriorly,  spiracles  on  anterior  mar- 
gin of  propodeum.  Fore  wing  with 
relative  lengths  of  submarginal,  marginal, 
postmarginal,  stigmal  veins  as  24:12:12:6; 
stigma  small;  basal  vein  with  row  of  setae 
along  length.  Petiole  transverse,  smooth. 
Gaster  ovate  acuminate,  length  1.6X 
width  (36:23);  hind  margin  of  Tl  nearly 
straight,  slightly  convex  mesally;  hypo- 
pygium  extending  Vi  gastral  length. 

Discussion. — This  species  is  distinct 
from  all  other  Thinodytes  species  by  the 
characters  listed  in  the  key:  scape  and  legs 
beyond  coxae  pale,  nonmetallic;  frenum 
and  median  panels  of  propodeum  smooth; 
plicae  rounded  and  smoothly  convergent 
posteriorly.  In  these  characters,  T.  santerna 
is  phenetically  similar  to  species  of  Noto- 
glyptiis. 

Type  Material— The  holotype  (USNM) 
was  collected  3  April  1984,  at  the  Eagle 
Borax  Works   in   Death  Valley  National 


Monument,  Inyo  County,  California 
(USA)  by  E.  E.  Grissell  on  Distichlis  in  a 
brackish  marsh. 

Etymology. — The  specific  epithet  of  this 
species  is  from  the  Latin  noun  santerna, 
meaning  borax,  and  refers  to  the  locality 
where  the  type  specimen  was  collected. 

Biology. — Nothing  is  know  of  the  host(s) 
of  T.  santerna. 

Mauleus  Graham,  1981 

Type  Species:  Mauleus  maderensis  Gra- 
ham, 1981  (examined);  original  designa- 
tion. 

Description. — Body  very  dark  green  or 
blue;  scape  brownish  yellow,  nonmetallic. 
Head,  pronotum,  mesoscutum,  scutellum 
(including  frenum),  dorsellum,  median 
panels  of  propodeum,  petiole  alveolate; 
gastral  tergites  nearly  smooth.  Head  with 
clypeus  bidentate  (left  tooth  compound) 
(Fig.  13),  lateral  part  of  mouth  margin 
with  short  shallow  genal  concavity;  anten- 
nal torulus  IX  own  diameter  above  LOcL. 
Antenna  with  scape  cylindrical,  >6X  as 
long  as  wide;  flagellum  length  less  than 
head  width  in  females,  about  equal  to 
head  width  in  males;  funicular  segments 
cylindrical;  MPP  sensilla  in  single  row;  fe- 
male club  simple  apically  and  with  small 
patch  of  micropilosity  on  apical  segment. 
Male  maxilla  with  palps  slender,  stipites 
unenlarged.  Mesosoma  (Figs  8,  21)  arched 
dorsally;  pronotum  with  collar  short  (Figs 
23-25),  nearly  level  with  vertex  dorsally, 
anterior  edge  rounded;  mesoscutum  with 
notauli  shallow,  impressed  lines  at  most; 
scutellum  as  long  as  wide,  lacking  anterior 
median  groove,  frenum  indistinguishable 
from  remainder  of  scutellum;  dorsellum 
short,  length  about  equal  to  length  of 
ridge  across  anterior  margin  of  propo- 
deum; propodeum  (Fig.  20)  with  median 
panels  short  (width  about  2X  median 
length),  plicae  and  median  carina  well  de- 
veloped and  connected  posteriorly  by  W- 
shaped  carina;  spiracles  strongly  ovate  al- 
most linear.  Fore  wing  with  relative 
lengths  of  veins  as  follows:  submarginal  > 


Volume  4,  1995 


17 


Figs.  23-25.     23,  Maidens  maderensis  Graham,  female  pronotum  and  mesonotum;  24,  Maidens  iligneus  n.  sp., 
female  pronotum  and  mesonotum;  25,  Maidens  venetiis  n.  sp.,  female  pronotum  and  mesonotum. 


marginal  >  postmarginal  >  stigmal;  stig- 
ma small,  width  about  2X  width  of  stig- 
mal vein;  costal  cell  with  complete  row  of 
setae  and  sometimes  a  partial  second  row; 
basal  cell  bare;  basal  vein  setose;  specu- 
lum present,  open  posteriorly.  Petiole  (Fig. 
20)  longer  than  wide,  with  basal  flange 
thickened  laterally,  without  median  cari- 
na, without  lateral  setae.  Gaster  of  females 
lanceolate,  length  1.6  or  more  times  width; 
hypopygium  extending  V3  length  of  gaster 
or  more;  in  both  sexes,  hind  margin  of  Tl 
sinuous  laterally,  emarginate  medially 
(Figs  8,  21). 

Discussion. — This  genus  is  placed  in  the 
Halticoptera-group  as  defined  in  this  paper 
by:  the  rounded  pronotum,  shallow  no- 
tauli,  undifferentiated  frenum,  propo- 
deum  with  a  median  carina  and  plicae 
connected  by  a  W-shaped  carina,  reticu- 
late petiole  with  a  complete  basal  flange, 
and  Tl  that  is  sinuous  laterally  and  emar- 
ginate medially.  Maidens  and  Halticopitera 
exhibit  considerable  phenetic  similarity. 
The  clypeus  of  Mauleus  is  secondarily  bi- 
dentate  (Fig.  13);  the  left  hand  clypeal  den- 
ticle is  divided  by  a  sulcus  formed  as  a 
result  of  the  near  fusion  of  the  two  ap- 
proximated left  hand  denticles.  These  den- 
ticles are  shown  in  their  plesiomorphic 
separated  state  in  Thinodytes  (Fig.  11).  The 
same  compound  condition  of  the  left  den- 
ticle is  found  in  Halticoptera  Spinola  (Hey- 
don,  unpublished  data).   [In  contrast,  in 


the  unrelated  genus  Sphegigaster  Spinola, 
the  left  tooth  in  the  bidentate  clypeus  is  a 
single  unit  (Heydon,  unpublished  data)]. 
The  compound  clypeal  denticular  struc- 
ture and  the  nonmetallic  scape  are  possi- 
ble evidence  of  a  close  phylogenetic  rela- 
tionship between  Halticoptera  and  Mauleus. 
Halticoptera  is  well-defined  cladistically 
relative  to  Mauleus  by  the  very  low  inser- 
tion of  the  antennae  (at  or  below  the 
LOcL)  and  by  the  male  maxilla,  which  has 
the  terminal  two  segments  of  the  palps 
flattened,  expanded,  and  nearly  always 
yellow  and  often  the  stipites  also  expand- 
ed. Two  apomorphic  characters  readily 
define  Mauleus  relative  to  Halticoptera  and 
related  genera:  1.  The  propodeum  is  short- 
ened in  Mauleus  (median  panels  over  2X 
as  wide  as  long  in  Mauleus  species  exam- 
ined). 2.  The  basal  flanges  of  the  petiole 
are  exceptionally  large  and  thick,  giving 
the  petiole  a  connate  appearance.  The  bas- 
al flanges  in  related  genera  are  generally 
free-standing  lamella  and  the  petiole  is 
more  or  less  cylindrical. 

Biology. — Mauleus  iligneus  has  been 
reared  from  pupae  of  the  native  holly  leaf 
miner,  Phytouiyza  ilicicola  Loew  (Diptera: 
Agromyzidae).  Like  many  other  genera  of 
the  Miscogastrinae,  it  is  likely  that  Mau- 
leus species  are  parasitoids  of  the  pupal 
stages  of  leaf-mining  or  stem-mining  Dip- 
tera. 


18  Journal  of  Hymenoptera  Research 


KEY  TO  DESCRIBED  SPECIES  OF  MAULEUS  GRAHAM 

1.  Pronotal  collar  with  sides  parallel  in  dorsal  view  (Fig.  23) 2 

-  Pronotal  collar  with  sides  widest  near  anterior  margin  and  converging  posteriorly  in  dorsal 
view  (Figs  24,  25) 3 

2.  Vertex  and  mesoscutum  with  conspicuous  pale  setae.  Propodeum  with  reticulations  much 
less  coarse  than  on  scutellum,  median  panel  each  with  broad  shallow  groove  along  anterior 
margin nigritus  (Howard) 

-  Vertex  and  mesoscutum  with  indistinct  dark  setae.  Propodeum  with  reticulations  as  coarse 
as  on  scutellum,  median  panel  each  with  a  pair  of  elongate  sublateral  depressions  along 
anterior  margin maderensis  Graham 

3.  Propodeum  with  anterior  depression  between  basal  foveae;  median  panels  with  extensive 
area  of  weak,  almost  smooth  sculpture.  Pronotum  with  humeral  angles  acute,  coming  to 
blunt  points  in  dorsal  view  (Fig.  25).  MPP  sensilla  in  two  or  more  rows  on  funicular 
segments    venetus  Heydon 

-  Propodeum  without  distinct  anterior  depression  between  basal  foveae;  median  panels  al- 
most entirely  alveolate,  with  at  most  a  small  central  patch  of  weak  sculpturing.  Pronotum 
with  humeral  angles  either  squared  or  slightly  and  smoothly  convergent  posteriorly  (Fig. 
24).  MPP  sensilla  in  single  row  on  funicular  segments  (male  of  M.  cultratus  unknown)    ...     4 

4.  Eye  height  2.5-2.7 X  genal  distance.  Female  with  combined  length  of  head  and  mesosoma 
longer  than  gaster;  gaster  less  than  twice  as  long  as  wide;  hypopygium  extending  around 

%  gastral  length  (United  States) iligneus  Heydon 

-  Eye  height  2. 1-2.4  X  genal  distance.  Female  with  combined  length  of  head  and  mesosoma 
less  than  or  equal  to  length  of  gaster  (Fig.  8);  gaster  more  than  twice  as  long  as  wide; 
hypopygium  extending  to  near  tip  of  gaster  (southern  Mexico  to  Argentina) 

cultratus  Heydon 


Maidens  cultratus  Heydon,  new  species  Structure:  Body  length  (excluding  ovi- 

Fig.  8  positor  sheaths)  1.6  mm.  Head  width  1.2X 

height  (26:21),  2.2  X  length  (26:12);  genal 

Holotype.  female. -Color:  Head,  pleural  concavity  extending  Vs  malar  distance;  eye 

regions,    propodeum    dark   blue;    pehole  j^^-g^t  1.4X  length  (13.5:9.5),  2.2X  malar 

bluish  black;  collar,  dorsum  of  mesosoma  distance    (13.5:6.0),    length    4.8X    temple 

greenish   black;   gaster   dark   brown,   Tl  length;  ratio  of  MOD,  POL,  OOL,  LOL  as 

with  dark  blue  reflections.  Antenna  with  2:4:6:3;  torulus   IX  own  diameter  above 

scape  brownish  yellow  with  weak  metallic  lqcL.  Antenna  with  length  of  pedicel  plus 

reflections;  pedicel,  flagellum  brown.  Legs  flagellum  0.81  X  head  width  (21:26);  ratio 

with  coxae,  trochanters,  femora  dark  blue;  of  lengths  of  scape,  pedicel,  annelli,  Fl-6, 

tibiae  brown  except  basal  and  apical  tips  dub  as  9.0:3.5:1.0:1.5:2.0:2.0:2.0:2.0:2.0:5.0; 

brownish  yellow;  fore  tarsi  brown,  middle  widths  of  Fl,  F6,  club  as  2:3:3;  MPP  sparse, 

and  hind  tarsi  yellow-brown  with  pretarsi  only  one  or  two  visible  per  segment  from 

brown.  Wing  veins  pale  brown.  single  view.  Mesosoma  length  1.4X  width 

Sculpture:  Clypeus,  median  portion  of  (33:23);  pronotum  with  sides  converging 

face  finely  alveolate;  remainder  of  head,  posteriorly;    notauli    incomplete;    propo- 

mesonotum,   scutellum,   frenum,   median  deum  with  width  of  median  panels  2.2  X 

panels   of  propodeum   alveolate;   petiole  length  (11:5);  basal  fovea  a  shallow,  tri- 

finely  alveolate;  gaster  smooth  except  T5-  angular  depression;  plicae  fading  out  in 

7  weakly  coriaceous.  anterior  Vz;  spiracles  0.5  X  own  diameter 


Volume  4,  1995 


19 


from  anterior  margin  of  propodeum. 
Wing  with  relative  lengths  of  submargin- 
al,  marginal,  postmarginal,  stigmal  veins 
as  27:16:14:7;  costal  cell  with  single  com- 
plete row  of  setae;  basal  vein  setose  with 
one  seta  posteriorly  on  cubital  vein.  Peti- 
ole length  1.5X  width  (9:6);  sides  narrow- 
ing posteriorly.  Gaster  lanceolate  (Fig.  8), 
length  2.6 X  width  (42:16),  length  l.OX  that 
of  head  and  gaster  (42:42);  hypopygium 
reaching  to  apex  of  T7;  ovipositor  sheaths 
exserted  for  distance  equal  to  half  length 
of  hind  tibia. 

Variation. — The  color  of  the  head,  pleu- 
ral regions,  and  petiole  varies  from  dark 
blue,  as  in  the  holotype,  to  dark  green  and 
nearly  concolorous  with  the  dorsum  of  the 
mesosoma.  The  body  length  of  specimens 
from  the  type  locality  is  about  1.9  mm, 
whereas  the  length  of  the  specimen  from 
Ixtapan  is  2.3  mm,  the  one  from  Morelia 
is  2.4  mm,  and  the  female  from  Panama  is 
3.1  mm  in  length.  The  ovipositor  sheaths 
are  exserted  for  a  distance  equal  to  ¥i  to  Vi 
the  length  of  the  hind  tibia. 

Discussion. — Maidens  cultratus  can  be 
distinguished  from  M.  iligneus  by  the  char- 
acters given  in  the  discussion  section  for 
that  species. 

Eti/mologx/. — The  species  name  comes 
from  the  Latin  word  cultratus,  meaning 
knife-shaped,  and  refers  to  the  shape  to 
the  female  gaster. 

Type  Material. — The  holotype  female 
(CNCI)  and  one  paratype  female  were  col- 
lected from  San  Cristobal  de  las  Casas, 
Mexico,  on  29  June  1969.  Eight  other  para- 
type females  were  collected  as  follows 
(CASC,  CDAE,  CNCI,  SEMC,  USNM):  Ar- 
gentina. SALTA:  Rosario  de  Lerma,  4- 
8.XI.1983  (Malaise  trap).  Mexico.  CHIA- 
PAS: San  Cristobal  de  las  Casas,  1- 
12.V.1969;  MICHOACAN:  Morelia,  6.IX.- 
1938;  MEXICO:  Ixtapan,  9.VII.1954;  TA- 
MAULIPAS:  6  miles  n.  Ciudad  Victoria. 
Panama.  Chiriqui,  XII. 1946. 

Biology. — The  host(s)  of  this  species  are 
unknown. 


Mauleus  iligneus  Heydon,  new  species 
Figs.  13,  20,  21,  24 

Holotype,  female. — Color:  Like  that  of  M. 
cultrata  except  pedicel  not  metallic,  and 
middle  and  hind  tarsi  white. 

Sculpture.  Clypeus  and  immediate  vi- 
cinity finely  alveolate  (Fig.  13);  remainder 
of  head,  mesoscutum,  scutellum,  frenum, 
median  panels  of  propodeum  alveolate; 
petiole  finely  alveolate;  gaster  smooth  ex- 
cept T6  and  T7  coriaceous. 

Structure. — Body  length  1.8  mm.  Head 
width  1.3X  height  (29:23),  2.2X  length  (29: 
13);  eye  height  1.4X  length  (15.0:10.5), 
2.5 X  malar  distance  (15:6),  length  5.2 X 
temple  length  (10.5:2.0);  ratio  of  MOD, 
OOL,  POL,  LOL  as  2.5:4.0:7.0:3.0;  torulus 
located  IX  own  diameter  above  LOcL.  An- 
tenna with  length  of  pedicel  plus  flagel- 
lum  0.88 X  head  width  (25.5:29.0);  ratio  of 
lengths  of  scape,  pedicel,  annelli,  Fl-6, 
club  as  11.0:3.0:1.0:2.5:2.5:2.5:2.5:2.5:2.5:7.0; 
widths  of  Fl,  F6,  club  as  2.0:3.0:3.5.  Me- 
sosoma length  1.5X  width  (36.0:24.5);  pro- 
notum  with  humeral  angles  squared  but 
sides  convergent  posteriorly  (Fig.  24);  no- 
tauli  extending  to  hind  margin  of  mesos- 
cutum as  impressed  lines;  propodeum 
(Fig.  20)  with  plicae  fading  out  in  anterior 
half,  basal  fovea  extending  halfway  down 
median  panels  and  bordered  mesally  by 
carina,  nuchal  area  raised  but  acarinate 
anteriorly,  spiracles  almost  on  anterior 
margin  of  propodeum.  Fore  wing  with  rel- 
ative lengths  of  submarginal,  marginal, 
postmarginal,  stigmal  veins  as  25:15:11:6; 
costal  cell  with  one  complete  setal  row 
and  one  partial  row  distally;  basal  vein 
with  row  of  three  setae.  Petiole  (Fig.  20) 
length  1.3X  width  (8:6);  narrowing  poste- 
riorly; basal  flanges  large.  Gaster  fusiform 
(Fig.  21),  length  0.87X  length  of  head  and 
mesosoma  (39:45);  length  1.8X  width  (39: 
22);  hypopygium  extending  %  length  of 
gaster;  ovipc^sitor  sheath  exserted  for  a 
distance  equal  to/ length  of  hind  tibia. 

Allotype. — Male.  Color  similar  to  holo- 
type except  frons,  callus  green;  flagellum 


20  Journal  of  Hymenoptera  Research 

paler  ventrally;  tibiae  brownish  yellow,  allotype  (USNM),  and  an  additional  nine 
Body  length  1.8  mm.  Head  with  ocelli  rel-  female  and  seventeen  male  paratypes 
atively  larger,  ratio  of  MOD,  OOL,  POL,  (UCDC,  USNM)  were  reared  from  Phijto- 
LOL  as  3.0:3.0:7.5:3.5.  Antenna  with  mi/za  ilicicola  on  Ilex  opaca  Solander  at  Lex- 
lengths  of  pedicel  plus  flagellum  l.Ox  ington,  Kentucky  by  D.  A.  Potter  collected 
head  width  (31:31);  relative  lengths  of  16  May  1984.  An  additional  43  paratypes 
scape,  pedicel,  annelli,  Fl-6,  club  as  10.0:  were  collected  as  follows  (CNCI,  UCDC): 
3.5:1.0:3.0:3.5:3.0:3.0:3.0:3.0:8.0;  widths  of  ILLINOIS:  Cave-in-Rock  State  Park,  near 
Fl,  F6,  club  as  2.5:3.0:3.0;  setae  reclinate.  Cave-in-Rock,  4.VI.1981,  1  female.  KAN- 
Gaster  ovate,  length  1.8X  width  (42:  SAS:  Oswego,  17.V.1976,  1  female.  MARY- 
23)(Specimen  critical-point  dried  so  gaster  LAND:  College  Park,  V.1954  (ex  holly  leaf 
in  air-dried  specimen  will  be  shorter).  miner)  1  male,  VII-VIII.1937  (ex  Phytomyza 

Variation. — Length  of  female  specimens  ilicis),    4    females,    8     males;    Laurel, 

varies  between  1.5  and  2.1  mm  and  males  14.V.1965,  1  female.  NEW  JERSEY:  New 

between  1.1  and  1.8  mm.  Other  than  the  Brunswick,  26.V.1947  (ex  Phytomyza  ilici- 

rather  large  variation  in  size,  this  is  a  mor-  cola),  1  female.  TEXAS:  Houston,  8.XII.- 

phologically  uniform  species.  1929  (ex  leaf  miner  on  Ilex  vomitoria  Solan- 

Discussion. — Mauleus  iligneus   differs  der),  1  female,  1  male;  Sanderson,  9.V.- 

from  M.  cultratus  in  the  following:  1.  The  1912,  1   female.  VIRGINIA:  Norfolk,  V.- 

ratio  of  the  eye  height  to  the  genal  dis-  1929   (ex  Phytomyza   ilicis),  2  females,  6 

tance     is     more     in    M.     iligneus     [x=  males;  Richmond,  V.1938  (ex  P.  ilicis),  10 

2.56±(S.E.)0.022  (n  =  12);  range  2.5-2.7]  females,  4  males. 

than  in  M.  cultratus  [x  =  2.24  ±0.038  (n  =  7);  Biology. — This  species  has  been  reared 
range  2.1-2.4].  2.  Mauleus  iligneus  has  a  numerous  times  from  the  complex  of  Phy- 
partial  second  row  of  setae  in  the  costal  tomyza  mining  the  leaves  of  holly  [Potter 
cell  of  the  fore  wing,  which  is  lacking  from  and  Gordon  1985  (as  Sphegigaster  sp.),  Un- 
M.  cultratus.  3.  The  ratio  of  the  length  of  derhill  1943  (as  Sphegigastrinae,  new  ge- 
the  head  and  mesosoma  to  the  length  of  nus,  new  species),  and  Langford  and  Cory 
the  gaster  is  relatively  greater  in  M.  ilig-  1936  (as  Sphegigaster  sp.)].  It  may  also  be 
neus  [x  =  1.23  ±0.03  (n  =  10);  range  1.2-  the  species  called  Halticoptera  sp.  by  Kulp 
1.4] (Fig.  21)  than  in  M.  cultratus  (1968),  but  voucher  specimens  from  Kulp 
[x=0.918±0.035  (n=6);  range  0.8-1.0](Fig.  have  not  been  seen.  Host  records  prior  to 
8).  4.  The  ratio  of  the  gastral  length  divid-  1968  are  questionable  because  the  Phy to- 
ed by  its  width  is  less  for  M.  iligneus  myza  complex  on  holly  was  not  studied  in 
[x=1.76±0.04  (n=10);  range  1.7-1.9]  than  detail  until  then  (Kulp  1968). 
for  M.  cultratus  [x=2.56±0.11  (n=6);  range  Potter  and  Gordon  (1985)  reported  that 
2.2-3.1)].  5.  The  hypopygium  extends  to  Mauleus  iligneus  was  a  primary  parasite 
about  %  the  length  of  the  gaster  in  M.  //-  when  its  agromyzid  host,  the  native  holly 
/^news  (Fig.  21),  but  is  nearly  even  with  the  leafminer,  Phytomyza  ilicicola,  was  unpar- 
tip  of  T7  in  M.  cultratus  (Fig.  8).  6.  The  asitized,  but  was  a  facultative  hyperpar- 
ovipositor  sheaths  are  exserted  for  a  asite  on  Opius  striativentris  Gahan  (Hy- 
length  equal  to  Va  the  length  of  the  hind  menoptera:  Braconidae)  when  that  species 
tibia  in  M.  iligneus  (Fig.  21),  but  for  about  had  already  parasitized  the  agromyzid 
V3  to  Vi  the  length  of  the  hind  tibia  in  M.  maggot.  Potter  and  Gordon  reared  no 
cultratus  (Fig.  8).  Mauleus   iligneus   from   puparia   collected 

Etymology. — The  specific  epithet  for  just  a  few  days  prior  to  the  emergence  of 

Mauleus  iligneus  is  an  adjective  based  on  Phytomyza  ilicicola  and  speculate  that  Mau- 

the  Latin  word  ilex,  meaning  holly.  leus  iligneus  parasitizes  the  fly  late  in  its 

Type  Material. — The  holotype  (USNM),  pupal  stage. 


Volume  4,  1995 


21 


Mauleus  maderensis  Graham 

Fig.  23 

Mauleus  maderensis  Graham,  1981:8. 

Holotype,  female  (BMNH)  Hym.  Type 

No.  5.3454  (examined).  Boucek  and 

Rasplus,  1991:41. 

Diagnosis. — Mauleus  maderensis  differs 
from  the  three  newly  described  species  in 
that  it  has  the  sides  of  the  pronotum  in 
dorsal  view  as  for  most  other  Pteromali- 
dae — more  or  less  parallel,  but  weakly 
convex,  with  the  broadest  point  being 
about  halfway  between  its  anterior  and 
posterior  edge  (Fig.  23).  The  three  new 
species  of  Mauleus  described  in  this  paper 
have  the  pronotum  in  dorsal  view  broad- 
est near  its  anterior  margin  and  distinctly 
convergent  posteriorly.  Mauleus  luaderensis 
is  similar  in  propodeal  structure  to  the  Ca- 
ribbean species  M.  niritus.  However,  these 
two  species  are  easily  distinguished  by  the 
characters  given  in  the  key. 

Distribution. — Madeira  (Pico  das  Arru- 
das,  near  Sao  Martinho),  and  possibly 
Mexico  (Boucek  and  Rasplus,  1991)  be- 
cause the  species  is  associated  with  plants 
of  Mexican  origin  (Boucek,  pers.  comm.). 

Biology. — The  insect  host(s)  of  M.  mad- 
erensis remain  unknown. 

Mauleus  nigritus  (Howard), 
new  combination 

Polycystus  nigritus  Howard,  1897:142. 
Holotype,  female  (BMNH);  Hym.  Type 
No.  5.876  (examined).  Heydon,  1989:193. 

Diagnosis. — The  type  of  Mauleus  nigritus 
is  fragmented  with  most  of  the  antennae 
gone  and  the  gaster  and  petiole  mounted 
on  the  card  separately  from  the  remainder 
of  the  body.  Enough  is  remaining  how- 
ever, to  confirm  that  this  species  belongs 
in  Mauleus.  Mauleus  nigritus  differs  from 
the  three  newly  described  species  in  that 
it  has  the  sides  of  the  pronotum  in  dorsal 
view  more  or  less  parallel,  with  the  broad- 
est point  being  about  halfway  between  its 
anterior  and  posterior  edge.  The  three  new 
species  of  Mauleus  described  in  this  paper 


have  the  pronotum  in  dorsal  view  broad- 
est near  its  anterior  margin  and  distinctly 
convergent  posteriorly.  Mauleus  nigritus  is 
similar  in  pronotal  structure  to  M.  mader- 
ensis; however,  these  two  species  are  eas- 
ily distinguished  by  the  characters  given 
in  the  key. 

Distribution. — Mauleus  nigitus  is  known 
to  me  only  from  the  holotype  collected  at 
Baltazar,  on  the  windward  side  of  Grena- 
da (West  Indies)  by  H.  H.  Smith. 

Biology. — The  insect  hosts  of  M.  nigritus 
remain  unknown. 

Mauleus  venetus  Heydon,  new  species 
Fig.  25 

Holotype,  female. — Color:  Body  black  ex- 
cept anterior  aspect  of  head,  collar,  me- 
soscutum  steel  blue  and  lateral  portions  of 
metanotum,  propodeum,  Tl  blue.  Anten- 
na with  scape,  ventral  side  of  pedicel  and 
Fl  brownish  yellow,  remainder  of  pedicel 
and  flagellum  brown.  Legs  with  coxae 
dark  blue;  femora  brown  with  metallic 
blue  reflections,  except  basal  and  apical 
tips  brownish  yellow;  tibiae  brownish  yel- 
low with  weak  brown  bands  mesally;  tarsi 
white  with  pretarsi  brown.  Fore  wing 
clear  with  veins  brownish  yellow,  paras- 
tigma  darker. 

Sculpture. — Clypeus  and  immediate  vi- 
cinity finely  alveolate;  remainder  of  head, 
mesoscutum,  scutellum,  frenum,  median 
panels  of  propodeum  alveolate;  petiole 
finely  alveolate;  gaster  smooth  except  T6 
and  T7  coriaceous. 

Structure. — Body  length  2.2  mm.  Head 
width  1.4X  height  (42:30),  2.5  X  length 
(42.0:16.5);  eye  height  1.5X  length  (20:13), 
2.0X  malar  distance  (20:10),  length  7.5X 
temple  length  (13:2);  torulus  2X  own  di- 
ameter above  LOcL;  ratio  of  MOD,  OOL, 
POL,  LOL  as  3.0:6.5:9.0:4.0.  Antenna  with 
length  of  pedicel  plus  flagellum  0.90  X 
head  width  (38:42);  relative  lengths  of 
scape,  pedicel,  annelli,  Fl-6,  club  as  15.0: 
4.0:1.0:4.5:4.5:4.5:4.0:4.0:3.5:8.0;  relative 
widths  of  Fl,  F6,  club  as  3.5:4.0:4.0;  MPP 
sensilla  in  two  rows  on  each  flagellar  seg- 


22 


Journal  of  Hymenoptera  Research 


merit;  terminal  club  segment  with  small 
ventral  patch  of  micropilosity.  Mesosoma 
length  1.5  X  width  (26.5:18.0);  pronotum 
with  humeral  angles  acute  (Fig.  25);  dor- 
sellum  short,  anterior  and  posterior  edges 
parallel;  propodeum  with  region  between 
basal  foveae  depressed  (this  depression 
shorter  mesally  than  laterally  and  bound- 
ed posteriorly  by  weak  carina),  plicae  fad- 
ing out  before  reaching  anterior  margin  of 
propodeum,  nuchal  region  raised,  mar- 
gined anteriorly  by  carina,  spiracles  al- 
most on  anterior  margin  of  propodeum. 
Fore  wing  with  relative  lengths  of  sub- 
marginal,  marginal,  postmarginal,  stigmal 
veins  as  37:23:15:8;  costal  cell  with  1  com- 
plete and  1  distal  partial  row  of  setae;  bas- 
al cell  bare;  basal  vein  with  row  of  setae; 
speculum  open  posteriorly.  Petiole  length 
1.1  X  width  (9:8).  Gaster  length  1.4X  width 
(43:31),  0.66  X  combined  length  of  head 
and  mesosoma  (43:65);  hypopygium  ex- 
tending about  Vi  gastral  length;  ovipositor 
sheaths  hardly  extending  beyond  hind 
margin  of  T7. 

Allotype,  male. — Similar  to  female  ex- 
cept: body  length  1.8  mm.  Head  with  ratio 
of  MOD,  OOL,  POL,  LOL  as  3.5:4.0:7.0:3.0. 
Antenna  with  length  of  pedicel  plus  fla- 
gellum  1.1  X  head  width  (37:34);  relative 
lengths  of  scape,  pedicel,  annelli,  Fl-6, 
club  as  11.0:3.0:1.0:4.5:4.5:4.0:4.0:4.0:4.0:9.0; 
relative  widths  of  Fl,  F6,  club  as  3.0:3.5: 
3.0;  MPP  sensilla  numerous  and  distrib- 
uted over  funicular  segments  in  many 
rows;  setae  reclinate,  nearly  absent.  Gaster 
length  1.1  X  width  (29:26). 

Diagnosis. — Mauleus  venetus  is  distin- 
guished from  the  other  two  Nearctic  Mau- 
leus species  by  the  acute  humeral  angles 
of  the  pronotum,  the  broad  depression 
along  the  anterior  margin  of  the  propo- 
deum, and  the  lack  of  sculpture  over  most 
of  the  median  panels  of  the  propodeum. 
Mauleus  venetus  most  closely  resembles  M. 
iligneus — the  coloration  of  the  two  species 
is  almost  identical  except  the  propodeum 
is  brighter  blue  in  M.  venetus.  In  addition, 
the  MPP  sensilla  of  M.   venetus   are  ar- 


ranged in  two  or  more  rows  on  each  fu- 
nicular segment  in  both  sexes,  whereas 
they  are  arranged  in  only  one  row  in  M. 
iligneus. 

Etymology. — The  species  name  comes 
from  the  Latin  word  venetus,  meaning 
blue  or  sea-blue,  and  refers  to  the  distinc- 
tive blue  propodeum  in  this  species. 

Type  Material. — The  holotype  (IRCW)  is 
a  female,  collected  in  Grant  Co.  (T6N, 
R6W,  SI  7),  Wisconsin  (USA),  in  a  malaise 
trap  exposed  7-14.VL1976.  The  allotype 
(UCDC)  was  collected  by  the  author  on 
the  South  Farms  of  the  University  of  Illi- 
nois, near  Champaign,  Illinois  (USA)  on 
26.V.1985.  Six  paratypes  were  collected  as 
follows  (CNCI,  IRCW,  UCDC,  USNM): 
Canada.  QUEBEC:  La  Trappe,  12.VII.1942, 
1  female.  United  States.  ILLINOIS:  Uni- 
versity of  Illinois  South  Farms,  near 
Champaign,  26.V.1985,  1  male.  MICHI- 
GAN: Midland  Co.,  2.VII.1943,  1  male. 
VIRGINIA:  Winchester,  16.VI.1964,  1 
male.  WISCONSIN:  Grant,  T6N,  R6W, 
S17,  3-8.VI.1976  (gypsy  moth  Malaise 
trap),  1  female,  14-21. VI.1976  (gypsy  moth 
Malaise  trap),  1  female. 

Biology. — The  host(s)  of  M.  venetus  are 
unknown. 

ACKNOWLEDGMENTS 

I  thank  Melissa  Bennett  and  two  anonymous  re- 
views for  their  careful  reading  of  this  manuscript.  I 
also  thank  the  following  people  for  the  loan  of  ma- 
terial used  in  this  study:  Dr.  J.  S.  Noyes,  The  Natural 
History  Museum,  London,  ENGLAND  (BMNH);  Dr. 
W.  J.  Pulawski,  California  Academy  of  Sciences,  San 
Francisco,  CA  (CASC);  Dr.  F.  G.  Andrews,  California 
State  Collection  of  Arthropods,  Sacramento,  CA 
(CDAE);  Dr.  G.  A.  P.  Gibson,  Canadian  National  Col- 
lection, Ottawa,  ON  (CNCI);  Dr.  W.  E.  LaBerge,  Illi- 
nois Natural  History  Survey,  Champaign,  IL  (INHS); 
S.  Krauth,  University  of  Wisconsin,  Madison,  WI 
(IRCW);  Dr.  R.  W.  Brooks,  Snow  Entomological 
Museum,  University  of  Kansas,  Lawrence,  KS 
(SEMC);  Dr.  E.  E.  Grissell,  United  States  National  Mu- 
seum, Washington,  D.C.  (USNM).  The  acronym  for 
the  collection  of  the  Bohart  Museum  at  the  University 
of  California,  Davis,  CA  is  UCDC. 


Volume  4,  1995 


23 


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(Hymenoptera:  Pteromalidae).  Journal  of  Hyme- 
noptera Research  2:107-116. 

Heydon,  S.  L.  and  W.  E.  LaBerge  1988.  A  review  of 
North  American  species  of  Sphegigaster  north  of 
Mexico  and  the  biology  of  their  hosts  (Hyme- 
noptera: Pteromalidae).  Journal  of  the  Kansas  En- 
tomological Society  61:  258-277. 

Howard,  L.O.  1897.  On  the  Chalcididae  of  the  island 
of  Grenada,  B.W.I.  Journal  of  the  Linnean  Society 
of  London.  Zoology  26:129-178. 

Kamijo,  K.  1978.  Chalcidoid  parasites  (Hymenop- 
tera) of  Agromyzidae  in  Japan,  with  description 
of  a  new  species.  Kontyu  46:455—469. 

Kulp,  L.  A.  1968.  The  taxonomic  status  of  dipterous 
holly  leaf  miners  (Diptera:  Agromyzidae).  Uni- 
versity of  Maryland  Agricultural  Experiment  Station 
Bulletin  A-155:l-42. 

Langford,  G.  S.  and  E.  N.  Cory.  1936.  The  holly  leaf- 
miner  and  its  control.  Proceedings  of  the  National 
Shade  Tree  Conference  13:109-112. 

Morrill,  W.  L.  and  R.  W.  Kieckhefer.  1971.  Parasitism 
of  the  wheat  stem  maggot  in  South  Dakota.  Jour- 
nal of  Economic  Entomology  64:1129-1131. 

Nikol'skaya,  M.  N.  1937.  The  chalcidoid  parasites 
(Hymenoptera)  of  some  injurious  flies  of  grain 
crops.  Entomological  Review  27:1-27. 

Peck,  O.  1951.  Superfamily  Chalcidoidea.  pp.  410- 
594.  In,  C.  F.  W.  Muesbeck,  K.  V.  Krombein,  and 
H.  K.  Townes.  Hymenoptera  of  America  North  of 


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Journal  of  Hymenoptera  Research 


Mexico,  Synoptic  catalog.  U.S.  Department  of  Ag- 
riculture, Agriculture  Monograph  2:1-1420. 

Peck,  O.  1963.  A  catalogue  of  the  Nearctic  Chalci- 
doidea  (Insecta:  Hymenoptera).  Canadian  Ento- 
mologist Supplement  30:1-1092. 

Peck,  O.,  Z.  Boucek,  and  A.  Hoffer.  1964.  Keys  to  the 
Chalcidoidea  of  Czechoslovakia.  Memoirs  of  the 
Entomological  Society  of  Canada  34:1-120. 

Potter,  D.  A.  and  F.  C.  Gordon.  1985.  Parasites  as- 
sociated with  the  native  holly  leafminer,  Phyto- 
myza  ilicicola  Loew  (Diptera:  Agromyzidae),  on 
American  holly  in  Kentucky.  Journal  of  the  Kansas 
Entomological  Society  58:727-730. 

Schmiedeknecht,  O.  1909.  Hymenoptera  family 
Chalcididae.  Genera  Insectorum  97:1-550. 

Spencer,  K.  A.  1976.  The  Agromyzidae  (Diptera)  of 
Fennoscandia  and  Denmark.  Fauna  Entomologica 
Scandinavica  5:1-606. 

Spencer,  K.  A.  and  G.  C.  Steyskal.  1986.  Manual  of 
the  Agromyzidae  (Diptera)  of  the  United  States. 
U.S.  Department  of  Agriculture,  Agriculture  Hand- 
book 638:1^78. 

Takada,  H.  and  K.  Kamijo.  1979.  Parasite  complex  of 
the  garden  pea  leaf-miner,  Phytomyza  horticola 
Goureau  in  Japan.  Kontyu  47:18-37. 


Thomson,  C.  G.  1876.  Hymenoptera  Scandinaviae.  4. 
Pteromalus  (Svederus).  Lund. 

Thomson,  W.  R.  1958.  Catalogue  of  parasites  and  pred- 
ators of  Insect  pests.  Section  2.  Host  parasite  cata- 
logue. Part  5.  Hosts  of  the  Hymenoptera  (Mis- 
cogasteridae  to  Trigonalidae),  Lepidoptera  and 
Strepsiptera.  p.  562-698.  Commonwealth  Insti- 
tute of  Biological  Control.  Ottawa,  Ontario. 

Underhill,  G.  W.  1943.  Some  insect  pests  of  orna- 
mental plants.  Virginia  Agricultural  Experiment 
Station  Bulletin  349:27-38. 

Walker,  F.  1833.  Monographia  Chalciditum.  Ento- 
mologist's Magazine  1:367-384,  455^66. 

Walker,  F.  1839a.  Monographia  Chalciditum,  1.  Lon- 
don. 

Walker,  F.  1839b.  Monographia  Chalciditum,  2.  Lon- 
don. 

Walker,  F.  1843.  Descriptions  of  Chalcidites  discov- 
ered near  Conception,  in  South  America,  by  C. 
Darwin,  Esq.  Annals  and  Magazine  of  Natural  His- 
tory 12:30-32. 

Walker,  F.  1846.  List  of  the  specimens  of  Hymenopterous 
insects  in  the  collection  of  the  British  Museum.  Part 
I-Chalcidites.  London. 


J.  HYM.  RES. 

Vol.  4,  1995,  pp.  25-32 


An  Australian  Masarine,  Rolandia  angulata  (Richards) 

(Hymenoptera:  Vespidae):  Nesting  and  Evaluation  of 

Association  with  Goodenia  (Goodeniaceae) 

Friedrich  W.  Gess,  Sarah  K.  Gess  and  Robert  W.  Gess 
Albany  Museum,  Grahamstown,  South  Africa 


Abstract. — An  account  is  given  of  some  aspects  of  the  nesting  of  Rolandia  angulata  (Richards) 
(Masarinae).  This  wasp  nests  in  multicellular  burrows  in  compacted  sandy  soil.  The  sloping  en- 
trance to  the  burrow,  not  surmounted  by  a  turret,  is  concealed  beneath  a  pebble,  a  plantlet  or  a 
suitable  item  of  debris.  The  main  thrust  of  the  shaft  is  vertically  downwards,  however,  at  intervals 
it  curves  outwards  to  end  in  a  horizontal  cell  so  that  each  cell,  except  the  last  excavated  and 
therefore  deepest  one,  appears  to  be  accessed  by  a  lateral  shaft.  The  cells  are  unlined.  The  archi- 
tecture of  the  nest  and  the  method  of  its  construction  are  discussed.  The  association  between 
Rolandia  angulata  and  Goodenia  (Goodeniaceae)  flowers,  the  source  of  nectar  and  pollen,  is  evalu- 
ated. It  is  shown  that  the  association  with  Goodenia  pinnatifida  Schldl.,  at  least,  is  mutually  bene- 
ficial. Indeed  it  is  suggested  that  in  some  areas,  at  some  times,  R.  angulata  may  be  the  most 
important  potential  pollinator  of  this  plant. 


The  genus  Rolandia   Richards,   1962   is  (28.03S,    148.30E),    and    85    km    E    of   St 

listed  by  van  der  Vecht  and  Carpenter  George    (28.03S,    148.30E)    (27-29.X.1993, 

(1990)  as  a  junior  synonym  of  Metaparagia  F.W.,  S.K.  and  R.W.  Gess). 

Meade-Waldo,  1911.  As  the  analysis  on  There  is  only  one  published  observation 

which  this  is  based  has  not  yet  been  pub-  on  the  nesting  of  Rolandia — that  of  R.  ma- 

lished  the  generic  name  Rolandia  is  used  culata    entering    a    turretless,    simple, 

in  the  present  paper.  Rolandia  is  restricted  oblique,  blindly  ending  burrow  in  sandy 

to  Australia.  Four  species  are  known,  two,  ground  (Houston,  1984).  However,  Hous- 

R.  maculata  (Meade-Waldo)  and  R.  hous-  ^^^  is  currently  investigating  further  the 

toni  Snelling,  from  Western  Australia,  a  riesting  of  this  species  and  has  made  some 

third,  R.  borreriae  Snelling,  from  Northern  ^^^^i^^  available  for  comparison. 

Territory,    and    the    fourth,    R.    angulata  ^^  investigations  concerning  R.  angii- 

(Richards),  the  subject  of  this  paper,  from  ^"^"  published  m  the  present  paper  were 

Queensland  and  New  South  Wales.  undertaken   by    the   authors    during    the 

r,  .^    J-              1  I      u        u                    J   J  course  of  a  fieldtrip  to  Australia  in  Octo- 

Rolandia    angulata    has    been    recorded  ,        ^^^^    w       , 

£                   .u         ..           r-i            1      J          J  ber  1993.  Voucher  specimens  have  been 

from    southwestern    Queensland    and  ,         .      ,  .      ,       »„ 

,,         ^          - ,         o      .1     T»r  ,         r  deposited  in  the  Albany  Museum  (R.  an- 

northwestern    New    South    Wales,    from  [,.,<-              ^      .           i  .i     i 

n                u      no  o/ic     1/ir  /.niri    /o-  u     j  gulata,  the  forage  plants,  and  the  bee  visi- 

Cunnamulla    [28.04S,    145.40E     (Richards  ;        «.     i.u     r               i     x.  x    .^u      a      ..    i- 

.^^„        T^.  ,  .            ,             ,  ^     ,.                .  tors  to  the  forage  plants),  the  Australian 

1968  as  Riekia  auQulata  and  SneWine,  1986)  xt  .•        it         ..  V-  n     ..•         /-     u          /o 

,             ,           '^                               t>          /  National  Insect  Collection,  Canberra   [R. 

m  the  north  to  90  km  W  of  Cobar,  Barnato  ,,,^,,,,^,,1    and    the    Australian    National 

Tanks    [31.38S,    144.59E]    (about   400   km  Herbarium,  Canberra  (the  forage  plants). 

south  of  Cunnamulla)  (Richards  1968)  in 

the  south,  and  from  three  sites  to  the  east  NESTING 

of  Cunnamulla,  80  km  E  of  Cunnamulla  Description  of  the  Nesting  Areas. — Two 

(28.04S,  145.40E),  27  km  W  of  St  George  nesting  areas  of  Rolandia  angulata  were  lo- 


26 


Journal  of  Hymenoptera  Research 


Figs.  1-4.  1,  Eucalyptus  woodland  between  St  George  and  Cunnamulla,  southwestern  Queensland;  2,  Acacia 
scrub  80  km  east  of  Cunnamulla,  southwestern  Queensland;  3,  a  nesting  site  of  Rolandia  ai^gulata  27  km  west 
of  St  George,  southwestern  Queensland;  4,  a  nest  entrance  of  Rolandia  angulata  beneath  a  piece  of  plant  debris 
(approximately  X  3.6). 


cated  between  St  George  and  Cunnamulla 
in  southern  Queensland,  one  27  km  west 
of  St  George  on  28  October  and  the  other 
80  km  east  of  Cunnamulla  on  29  October, 
the  former  in  dry  open  woodland  domi- 
nated by  Eucalyptus  (Fig.  1)  and  the  latter 
in  dry  open  scrub  dominated  by  Acacia 
(Fig.  2).  Both  areas  had  recently  received 
rain  in  the  form  of  localized  thundershow- 
ers  which  had  resulted  in  a  growth  of  an- 
nuals which  were  in  flower  on  the  road 
verges  and  in  the  low  lying  areas.  The  soil 
in  both  areas  was  sandy,  compact  but  fri- 


able. It  was  increasingly  moist  at  least  to 
the  depth  of  the  deepest  nest  cells,  38  cm. 
Flozvers  Visited. — The  only  previous  rec- 
ord of  flower  visiting  by  R.  angulata  is  that 
of  Richards  (1968)  for  females  collected  on 
Goodenia  cycloptera  R.Br,  in  C.Sturt  (Good- 
eniaceae)  at  Barnato  Tanks.  In  the  present 
study  females  and  males  were  collected 
and  observed  visiting  the  yellow  flowers 
of  a  prostrate  herb  also  identified  as  G.  cy- 
cloptera (Fig.  11)  and  an  erect  herb  Good- 
enia pinnatifida  Schldl.  (Fig.  6)  in  the  two 
areas  where  nests  were  discovered  and 


Volume  4,  1995 


27 


also  in  the  area  85  km  east  of  St  George. 
At  the  most  easterly  site,  that  is  85  km  east 
of  St  George,  both  species  of  Goodenia 
were  common.  Travelling  westwards  to 
Cunnamulla  it  was  observed  that  G.  cy- 
cloptera  rapidly  became  uncommon 
whereas  G.  puinntifida  became  increasingly 
common  and  at  the  site  80  km  east  of  Cun- 
namulla was  the  most  common  roadside 
herb.  The  abundance  of  R.  angiilata  fol- 
lowed a  similar  pattern  to  that  of  G.  pin- 
natifida. 

Visits  to  the  flowers  by  female  R.  angii- 
lata were  abundant  by  lOhOO  and  contin- 
ued through  the  heat  of  the  day  and  the 
afternoon.  Visits  became  fewer  in  the  late 
afternoon  and  ceased  after  IThOO.  Males 
were  observed  to  patrol  the  flowers  and 
only  later  in  the  day  to  visit  them  for  nec- 
tar. 

All  plants  flowering  together  with  the 
two  Goodenia  species,  most  notably  several 
Asteraceae  and  a  Wahlenbergia  species 
(Campanulaceae),  were  sampled  for  flow- 
er visitors.  None  was  being  visited  by  R. 
angulata. 

Provision. — Provision  in  the  form  of  a 
firm,  white  pollen  loaf  was  obtained  from 
each  of  three  cells.  Pollen  from  the  loaves 
was  examined  microscopically  and  found 
to  match  that  obtained  from  the  Goodenia 
flowers. 

Mate  location. — During  the  morning 
males  patrolled  the  flowers  where  they 
sought  the  females.  They  did  not  alight  on 
the  flowers  but  rested  on  a  neighbouring 
plant,  for  example  on  a  grass  stem,  or  on 
the  ground.  Numerous  attempted  copu- 
lations at  flowers  were  observed  and  a  sin- 
gle attempt  to  copulate  with  a  nest-exca- 
vating female  was  noted.  Several  instances 
of  a  male  and  female  grappling  on  the 
ground  next  to  plants  and  of  "hot  pur- 
suit" were  noted. 

Description  of  the  Nest. — The  nest  (n  =  8) 
consists  of  a  subterranean  burrow  (Fig.  5) 
excavated  in  horizontal  sandy  soil  in  a 
clearing  (Fig.  3).  The  entrance  is  concealed 
beneath  a  pebble,  a  plantlet  or  a  suitable 


item  of  debris  (Fig.  4).  It  is  a  simple  hole 
3-4  mm  in  diameter,  not  surmounted  by 
a  superstructure.  For  approximately  the 
first  10  mm  the  shaft  slopes  gently  down- 
wards. Thereafter  its  main  thrust  is  verti- 
cally downwards,  however,  at  intervals  it 
curves  outwards  to  end  in  a  horizontal  cell 
so  that  each  cell,  except  the  last  excavated 
and  therefore  deepest  one,  appears  to  be 
accessed  by  a  lateral  shaft.  These  "lateral 
shafts"  radiate  out  through  360°  each 
deeper  than  that  preceding  it. 

The  diameter  of  the  shaft  narrows  at  a 
depth  of  approximately  80  mm  and  then 
continues  constant.  The  first  cell  in  the 
nests  investigated  was  at  a  depth  of  be- 
tween 180  and  370  mm,  the  "lateral 
shafts"  were  30  mm  in  length,  and  the 
cells  13  mm  long  and  4  mm  in  diameter 
at  the  widest  point.  A  lateral  shaft  is 
packed  with  sand  after  the  cell  which  ter- 
minates it  has  been  provisioned.  Up  to 
seven  cells  per  nest  were  recorded.  The 
cells  are  unlined. 

Method  of  Construction  of  the  Nest  and 
Provisioning. — Water  is  not  required  for 
the  excavation  of  the  nest  as  the  sandy 
soil,  though  compact,  is  friable.  The  sand 
extracted  from  the  burrow  is  carried  out 
of  the  shaft  held  between  the  head  and  the 
prosternum,  the  genae  being  fringed  with 
ammochaetae.  Whilst  excavation  is  in 
progress,  the  female,  when  leaving  the 
nest,  backs  out.  During  the  initial  stages  of 
burrow  excavation  the  extracted  sand  is 
dropped  in  flight  in  a  more  or  less  con- 
stant area  to  one  side  of  the  entrance  and 
about  120  mm  from  it.  As  the  burrow  be- 
comes deeper  the  extracted  sand  is 
dropped  further  from  the  nest  in  a  con- 
stant arc  about  250  mm  from  the  entrance. 
The  sand  being  scattered,  there  is  no  de- 
tectable accumulation  and  so  no  tumulus 
develops. 

The  presence  of  a  recently  hatched  larva 
in  a  sealed,  fully  provisioned  cell  indicat- 
ed that  mass  provisioning  is  practised. 
The  sand  used  for  packing  a  shaft  leading 
to  a  cell  is  almost  certainly  obtained  with- 


28 


Journal  of  Hymenoptera  Research 


in  the  nest  during  the  excavation  of  the 
shaft  leading  to  the  next  cell. 

The  fully  fed  larva  spins  a  white  cocoon 
which  completely  fills  the  cell.  Like  the 
sealed  cell  it  is  therefore  rounded  at  the 
inner  end  and  truncate  at  the  outer  end. 

Discussion  of  Nesting. — The  nest  of  R.  an- 
gulata  is  essentially  similar  to  that  of  R. 
maculata,  which  is  described  by  Houston 
(pers.  comm.)  as  a  vertical  burrow  in 
sandy  soil,  about  300  mm  deep  and  un- 
lined,  with  unlined  cells  at  the  lower  end. 
Thus  the  basic  nest  type  for  these  two  spe- 
cies of  Rolnndia  can  be  defined  as  a  mul- 
ticellular sub-vertical  burrow  in  horizontal 
ground  excavated  by  the  nester,  without 
an  entrance  turret  and  with  excavated 
cells  not  containing  constructed  cells. 
When  compared  with  the  seven  basic  nest 
types  recognized  for  the  Masarinae  as  a 
whole  by  Gess  and  Gess  (1992)  this  ap- 
pears to  fit  Nest  type  1  except  for  the  lack 
of  an  entrance  turret.  When  the  method  of 
construction  is  compared  with  that  of  the 
species  listed  for  this  nest  type,  a  basic  dif- 
ference is  apparent.  Water  is  used  in  the 
excavation  and  construction  of  Nests  of 
type  1,  like  nests  of  types  2  and  3,  whereas 
the  nests  of  the  two  Rolandia  species  are 
excavated  without  the  use  of  water.  This 
is  possible  due  to  the  friable  nature  of  the 
soil  in  which  they  are  sited.  As  noted  in 
Gess  and  Gess  (1992)  nesting  in  friable  soil 
in  the  Vespidae  is  probably  derived  rather 
than  primitive  as  in  the  Pompilidae  and 
Sphecidae.  The  nest  type  of  the  two  Rolan- 
dia species  is  therefore  seen  as  a  sub-type 
which  can  be  derived  from  Nest  type  1. 

Both  species  of  Rolandia  carry  sand,  ex- 
tracted from  the  shaft,  out  of  the  burrow 
held  between  the  head  and  the  proster- 
num,  the  genae  being  fringed  with  am- 


Fig.  5.  Plan  of  vertical  section  of  a  nest  of  Rola7tdia 
attgulata.  Cells  2 — 5  followed  cell  I,  radiating  out  in 
sequence  through  360°  and  at  successively  greater 
depths. 


50mm 


29 


Figs.  6-10.  6,  Goodenia  pinnatifida,  an  erect  herb;  7,  a  flower  of  Goodenia  piiiiiatifida,  indusium  concealed  by 
the  bases  of  the  adaxial  petals;  8,  Rolandia  angulata  entering  a  flower  of  Goodenia  pinnatifida,  indusium  of  the 
flower  exposed  (X  2.8);  9,  Rolandia  angulata  in  nectar  drinking  position  in  a  flower  of  Goodenia  pinnatifida, 
indusium  of  the  flower  fitting  snugly  over  the  wasp's  mesosoma  (X  2.8);  10,  Rolandia  angulata  withdrawing 
from  a  flower  of  Goodenia  pinnatifida,  showing  dusting  of  pollen  on  the  head,  dorsum  of  the  prothorax,  and 
anterior  part  of  the  mesoscutum  (X  2.8). 


mochaetae.  The  possesion  of  ammochae- 
tae  fringing  the  genae  is  a  generic  char- 
acter (Snelling  1986)  and  it  is  therefore  ex- 
pected that  all  members  of  the  genus 
excavate  their  nests  in  a  similar  manner. 
The  only  other  vespid  genus  recorded  as 
having  a  psammophore  for  the  removal  of 
the  spoils  of  excavation  is  Pterochihis  (Eu- 
meninae)  (Bohart  1940)  for  which  nesting 
in  vertical  burrows  in  friable  soil  by  two 
species  has  been  described  (Isely  1914  and 
Evans  1956). 

EVALUATION  OF  ASSOCIATION 
WITH  FLOWERS  VISITED 

Both  R.  angulata  males  and  females  ob- 
tain nectar  and  possibly  pollen  for  their 


own  nourishment  and  females  collect  nec- 
tar and  pollen  for  provisioning  their  nest- 
cells  apparently  solely  from  Goodenia  flow- 
ers. They  are  therefore  probably  depend- 
ant on  Goodenia  flowers.  To  determine 
whether  or  not  the  association  between 
the  wasp  and  the  flowers  is  mutually  ben- 
eficial or  not,  that  is  whether  or  not  the 
wasp  in  addition  pollinates  the  flowers, 
requires  a  consideration  of  the  functional 
morphology  of  the  flowers,  the  behaviour 
of  the  wasps  in  the  flowers,  and  wasp/ 
flower  fit. 

Goodenia  flowers  are  distinctly  two 
lipped.  The  lower  wings  of  the  two  ad- 
axial petals  are  differentiated  in  their  low- 
er parts  into  auricles  which  envelop  the 


30 


Journal  of  Hymenoptera  Research 


Figs.  11-13.  11,  Goodenia  cycloptera,  a  prostrate  herb;  12,  three  flowers  of  Goodenia  cycloptera,  indusium  con- 
cealed by  the  bases  of  the  adaxial  petals;  13,  Rolandia  angulata  in  nectar  drinking  position  in  a  flower  of  Goodenia 
cycloptera,  indusium  of  the  flower  pressed  down  on  the  wasp's  folded  wings  (X  3.3). 


indusium  in  the  flower  (Figs  7  and  12)  so 
that  it  is  only  exposed  when  an  insect  vis- 
itor pushes  the  corolla  lobes  apart  as  it 
seeks  the  nectar  at  the  base  of  the  flower 
(Figs  8  and  13).  The  indusium  is  a  cup  at 
the  top  of  the  style  and  the  surrounding 
stigmatic  initial  that  collects  and  retains 
pollen  from  the  stamens,  which  dehisce  in 
the  bud,  and  presents  it  thus  to  the  pollen 
vector  (Carolin,  Rajput  and  Morrison 
1992).  Later  the  stigmatic  initials  mature 
and  grow  out  of  the  indusium  and  collect 
pollen  from  pollen  vectors. 


R.  angulata  when  visiting  the  flowers  for 
nectar  always  alights  on  the  lower  lip  and 
then  pushes  its  way  in  beneath  the  indu- 
sium so  that,  if  the  flower  is  in  the  pollen 
presenting  phase,  the  wasp  receives  pollen 
on  its  head.  When  it  is  in  the  nectar  drink- 
ing position  in  a  flower  of  G.  pinnatifida, 
the  mesosoma  (Fig.  9)  is  snugly  capped  by 
the  indusium  but,  when  it  is  in  a  flower 
of  G.  cycloptera,  which  is  deeper,  it  is  the 
folded  wings  which  are  pressed  beneath 
the  indusium  (Fig.  13).  On  emerging  from 
a  flower  of  G.  pinnatifida  it  can  be  clearly 


Volume  4,  1995 


31 


seen  to  have  been  well  dusted  with  pollen 
not  only  on  the  head  but  also  on  the  dor- 
sum of  the  prothorax  and  the  anterior  part 
of  the  mesonotum  (Fig.  10).  When  coming 
thus  laden  with  pollen  to  a  flower  with 
receptive  stigmas  the  wasp  would  be  ide- 
ally suited  to  pollinate  it.  It  is  not  clear, 
however,  whether  R.  angulata  would  effec- 
tively pollinate  the  deeper  flowers  of  G. 
cycloptera.  It  is  possible  that  pollen  may  be 
transferred  from  its  head  to  a  receptive 
stigma  but  the  pollen  received  on  the 
wasp's  wings  is  unlikely  to  be  successfully 
transferred. 

It  is  concluded  that,  within  its  limited 
distribution,  R.  angulata  is  a  potential  pol- 
linator, at  least,  of  the  widespread  species 
G.  pinnatifida  with  which  it  therefore  has  a 
mutually  beneficial  association.  However, 
at  none  of  the  sites  was  R.  angulata  the  sole 
visitor  to  the  Goodenia  flowers. 

At  the  most  easterly  site  a  second  ma- 
sarine,  Riekia  nocatunga  Richards,  which 
was  also  recorded  from  Goodenia  fascicu- 
laris  F.Muell.  &  Tate  at  Kondar  to  the 
southeast,  was  a  relatively  abundant  vis- 
itor to  G.  pinnatifida.  Furthermore  the 
flowers  of  both  species  of  Goodenia,  par- 
ticularly those  of  G.  cycloptera,  were  vis- 
ited abundantly  by  bees.  The  most  com- 
mon species  was  Leioproctus  {Chri/socolle- 
tes)  moretonianus  (Cockerell)  (Colletidae) 
which  was  also  recorded  from  Goodeni- 
aceae  at  several  sites  in  Queensland  by 
Michener  (1965).  Less  common  visitors 
were  a  second,  but  slightly  smaller,  spe- 
cies of  Leioproctus  {Chrysocolletes)  and  a 
species  of  Megachile  (Megachilidae).  Rel- 
atively uncommon  visitors  were  six  fur- 
ther species  of  Megachile,  an  additional 
coUetid,  a  halictid,  a  few  anthophorids 
and  the  honey-bee.  Travelling  westwards 
the  bees  were  uncommon  at  the  site  27 
km  west  of  St  George  and  at  the  site  80 
km  east  of  Cunnamulla  all  but  the  second 
species  of  Leioproctus  (Chrysocolletes)  were 
absent.  At  the  latter  site  a  third  much 
smaller  species  of  Leioproctus  was  also  re- 
corded.   However,   R.   angulata   was    the 


only  abundant  visitor  suggesting  that  in 
some  areas,  at  some  times,  it  may  be  the 
most  important  potential  pollinator  of  G. 

pinnatifida. 

ACKNOWLEDGMENTS 

The  following  are  thanked  with  appreciation:  Terry 
Houston  of  the  Western  Australian  Museum  for  per- 
mission to  quote  his  unpublished  observations  con- 
cerning the  nesting  of  Rolandia  maciilata;  Jo  Cardale 
of  the  Australian  National  Insect  Collection,  Canberra 
for  facilitating  the  identification  of  forage  plants;  Kir- 
sten  Cowley,  Australian  National  Herbarium,  Can- 
berra for  the  identifications  of  the  forage  plants;  Alan 
Weaving  of  the  Albany  Museum  for  producing  black 
and  white  negatives  from  the  authors'  colour  trans- 
parencies for  Figs  1-4  and  6-13;  and  the  South  Afri- 
can Foundation  for  Research  Development  for  a  roll- 
ing support  grant  to  F.W.Gess,  which  made  possible 
the  fieldwork  in  Australia.  The  manuscript  was  re- 
viewed by  Karl  V.  Krombein  and  an  anonymous  re- 
viewer, who  are  thanked  for  their  constructive  com- 
ments. 

LITERATURE  CITED 

Bohart,  R.M.  1940.  A  revision  of  the  North  American 
species  of  Pterocheilus  and  notes  on  related  gen- 
era (Hymenoptera,  Vespidae).  Annals  of  the  En- 
tomological Society  of  America  33(1):  162-208. 

Carolin,  R.C.,  M.T.M.  Rajput,  and  D.  Morrison.  1992. 
Goodeniaceae.  In:  George,  A.S.  ed.,  Flora  of  Aus- 
tralia, volume  35:  Brunoniaceae,  Goodeniaceae.  Can- 
berra: Australian  Government  Publishing  Ser- 
vice, pp.  4-300. 

Evans,  H.E.  1956.  Notes  on  the  biology  of  four  spe- 
cies of  ground-nesting  Vespidae  (Hymenoptera). 
Proceedings  of  the  Entomological  Society  of  Washing- 
ton 58(5):  265-270. 

Gess,  F.W.  and  S.K.  Gess.  1992.  Ethology  of  three 
southern  African  ground  nesting  Masarinae,  two 
Celonites  species  and  a  silk-spinning  Quartinia 
species,  with  a  discussion  of  nesting  by  the  sub- 
family as  a  whole  (Hymenoptera:  Vespidae). 
Journal  of  Hymenoptera  Research  1(1):  145-155. 

Houston,  T.F.  1984.  Bionomics  of  a  pollen-collecting 
wasp,  Paragia  tricolor  (Hymenoptera:  Vespidae: 
Masarinae),  in  Western  Australia.  Records  of  the 
Western  Australian  Museum  11(2):  141-151. 

Isely,  D.  1914.  The  biology  of  some  Kansas  Eumen- 
idae.  Kansas  University  Scietice  Bulletin  (2)8(7): 
233-309. 

Meade-Waldo.  G.  1911.  Notes  on  the  family  Masar- 
idae  (Hymenoptera),  with  descriptions  of  a  new 
genus  and  three  new  species.  Annals  and  Maga- 
zine of  Natural  History  (8)8:  747-750. 

Michener,  CD.  1965.  A  classification  of  the  bees  of 
the  Australian  and  South  I'acific  regions.  Bulletin 


32 


Journal  of  Hymenoptera  Research 


of  the  American  Museum  of  Natural  History  130:  1- 
362. 

Richards,  O.W.  1962.  A  revisional  study  of  the  Masarid 
wasps  {Hymenoptera,  Vespoidea).  London:  British 
Museum  (natural  History). 

Richards,  O.W.  1968.  New  records  and  new  species 
of  Australian  Masaridae  (Hymenoptera:  Vespo- 
idea). Journal  of  the  Australian  Entomological  Soci- 
ety 7:  101-104.' 


Snelling,  R.R.  1986.  The  taxonomy  and  nomenclature 
of  some  Australian  paragiine  wasps  (Hymenop- 
tera: Masaridae).  Contributions  in  Science  378:  1- 
19. 

van  der  Vecht,  J.  and  J.M.  Carpenter.  1990.  A  cata- 
logue of  the  genera  of  the  Vespidae  (Hymenop- 
tera). Zoologische  Verhandelingen  26:  1-62. 


J.  HYM.  RES. 

Vol.  4,  1995,  pp.  33-10 

Descriptions  of  the  Male  of  Riekia  nocatiinga  Richards,  the  Male  and 

Two  Strikingly  Distinct  Sympatric  Colour  Forms  of 

Riekia  confluens  (Snelling)  and  the  Male  of 

Rolandia  angiilata  (Richards)  (Hymenoptera: 

Vespidae:  Masarinae)  from  Australia 

Friedrich  W.  Gess 
Albany  Museum,  Grahamstown,  6140,  South  Africa 


Abstract. — The  male  of  Riekia  nocatunga  Richards  from  southern  Queensland,  the  male  and  two 
strikingly  distinct  sympatric  colour  forms  of  Riekia  confluens  (Snelling),  comb,  nov.,  from  Western 
Australia,  and  the  male  of  Rolandia  angnlata  (Richards)  from  southern  Queensland  are  described. 
The  characters  distinguishing  R.  nocatunga  and  R.  confluens  are  discussed. 


The  Australian  Masarinae  and  their  nat- 
ural history  are  relatively  poorly  known. 
During  a  recent  collecting  trip  to  Australia 
by  the  author,  S.K.Gess  and  R.W.Gess, 
material  collected  included  males  of  three 
species,  Riekia  nocatunga  Richards,  Riekia 
confluens  (Snelling),  comb,  nov.,  and  Rolan- 
dia angnlata  (Richards),  hitherto  described 
only  from  the  female  sex.  Two  strikingly 
distinct  sympatric  colour  forms  of  Riekia 
confluens  were  found  to  occur.  One  of 
these  was  previously  unknown  and  helps 
to  elucidate  the  identity  of  a  single  male 
Riekia  collected  by  T.F.Houston  and  dis- 
cussed by  Snelling  (1986)  who  was  unable 
to  place  it  in  a  species. 

Riekia  Richards,  1962  and  Rolandia  Rich- 
ards, 1962  have  been  sunk  as  junior  sub- 
jective synonyms  of  Metaparagia  Meade- 
Waldo,  1911  by  van  der  Vecht  and  Car- 
penter (1990)  on  the  basis  of  an  as  yet  un- 
published cladistic  analysis  by  Carpenter. 
Until  such  time  as  this  analysis  is  pub- 
lished and  may  be  studied,  the  present  au- 
thor considers  it  best  to  continue  to  accept 
Riekia  and  Rolandia  as  genera  in  their  own 
right. 

Institutions  in  which  the  material  stud- 
ied   is   deposited   are:    Albany   Museum, 


Grahamstown,  South  Africa  (AMG);  Aus- 
tralian National  Insect  Collection,  Canber- 
ra (ANIC);  Western  Australian  Museum, 
Perth  (WAM). 

Riekia  nocatunga  Richards 
Riekia  nocatunga  Richards,  1962:  55-57,  female. 

This  species  was  described  from  7  miles 
N  of  Nocatunga  [on  maps  as  Nockatunga] 
[27.40S,  142.40E],  Queensland  (not  New 
South  Wales  as  in  Richards  1962;  correct 
in  Richards  1968)  (holotype  female  and  4 
female  paratypes)  and  Bourke  [30.09S, 
145.59E],  New  South  Wales  (1  female 
paratype). 

Subsequently  an  additional  five  females 
from  90  km  W  of  Cobar,  Barnato  Tanks 
[31.38S,  144.59E],  New  South  Wales,  were 
attributed  to  this  species  by  Richards 
(1968).  He  noted  that  they  differed  from 
his  original  description  with  regard  to  the 
pale  markings  of  which  he  gave  some  de- 
tails. Two  of  these  females,  in  the  collec- 
tion of  the  Natural  History  Museum,  Lon- 
don, have  been  examined  by  the  present 
author.  They  are  confirmed  as  being  R.  no- 
catunga. They  are  less  melanistic  than  the 
specimens  originally  described.  The  dark 
areas  are  less  extensive  and  some,  in  par- 


34  Journal  of  Hymenoptera  Research 

ticular  those  on  the  pronotum  and  tergites  each  side  of  mesoscutum  where  its  fur- 

1  and  2,  are  brownish  rather  than  black,  rows  meet  pronotum  (usually  but  not  al- 

The  light  areas  are  more  extensive  and  ways  present  in  females)  absent  in  both 

their  colour  is  a  strong  yellow  rather  than  males  examined, 

pale  lemon-yellow.  Antennal   scape   (with   radicle)   very 

Snelling  (1986)  figured  and  briefly  dis-  slightly  longer  than  interantennal  dis- 
cussed a  male  Riekia  from  43  km  ENE  Lan-  tance;  first  flagellomere  less  than  half  (0.4) 
dor  Homestead  (25.08S,  116.54E)  in  West-  as  long  as  scape  (with  radicle),  one  and  a 
ern  Australia.  Whereas  it  agreed  generally  third  times  as  long  as  wide  at  its  distal 
with  the  original  description  of  R.  nocatun-  end,  and  one  and  one  fifth  as  long  as  the 
ga,  Snelling  stated  that  there  was  no  cer-  second  flagellomere.  Last  three  flagellom- 
tainty  that  it  was  the  opposite  sex  of  that  eres  flattened  and  slightly  concave  be- 
species  and  that  he  suspected  that  it  was  neath;  ultimate  flagellomere  narrowing 
not.  He  listed  some  discrepancies,  allow-  apically  and  distinctly  curved  to  form  a 
ance  being  made  for  sexual  differences,  hook. 

and  stated  that  more  material  would  have  Tergite  7  evenly  rounded  apically. 

to  be  available  before  the  specific  status  of  Genitalia  (Figs  3  and  4);  parameral  spine 

the  male  could  be  determined.  broad  and  thick,  evenly  curved  over  its 

Recently  collected  Riekia  material,  of  rel-  length  and  not  hook-like  apically;  ventral 

evance  to  the  clarification  of  the  above  tax-  process  of  each  paramere  relatively  broad 

onomic  question,  consists  of  associated  fe-  and  short,  subtriangular,  without  a  nar- 

males  and  males  from  both  Queensland  row  elongation;  digitus  short, 

and    Western    Australia.    The    specimens  Length  7.5-7.8  mm;  length  of  forewing 

from  Queensland  are  certainly  R.  nocatun-  5.8  mm,  hamuli  14. 

ga,  as  established  by  the  comparison  by  New    Material    Examined. — Queensland: 

the  author  of  several  of  the  females  with  Lundavra   State  School,  Kondar  (28. OSS, 

the  holotype  in  the  Australian  National  In-  149.51E),  25.X.1993  (F.W.,S.K.&  R.W.Gess) 

sect  Collection,  Canberra.  The  specimens  11  females,  1  male  (in  yellow  flowers  of 

from  Western  Australia  on  the  other  hand  Goodenia  fascicularis   F.Muell.    &    Tate, 

represent   two   strikingly   distinct   colour  Goodeniaceae);  Southwood  Road,  western 

forms  of  Riekia  confluens.  One  is  superfi-  boundary   of  Southwood   National   Park 

cially  similar  looking  to  Riekia  nocatunga  (27.56S,    149.30E),    26.x. 1993    (F.W.,S.K.& 

but  morphologically  distinct.  The  males  of  R.W.Gess)  2  females  (at  water);  85  km  E 

this  form  show  the  characters  noted  by  of  St  George  (28.03S,  148.30E),  27.X.1993 

Snelling  for  his  male.  (F.W.,S.K.&  R.W.Gess)  8  females,  1  male 

Male. — (Figs  3  and  4).  Colour  pattern  (in  yellow  flowers  of  Goodenia  pinnatifida 

very  similar  to  that  of  female  but  differing  Schldl.,  Goodeniaceae).  Two  females  and 

in  that  the  following  parts  are  pale  lemon-  1  male  in  ANIC;  2  females  in  WAM;  rest 

yellow:  small  streak  on  scapes  distally,  en-  of  material  in  AMG. 

tire  clypeus  other  than  for  narrow  lateral 

and  anterior  margins,  single  large  frontal  Rekia  confluens  (Snelling)  comb.  nov. 

spot  between  and  above  antennal  sockets, 

.1                .    .         1                  ,  ..  Paravia  confluens  Snelling,  1986:  14  and  Fie.  28, 

narrow    streak    marginmg    lower    orbits,  c       \                                                      &      ' 

elongate  spot  in  lower  half  of  each  ocular  p.  ,  •          ^     n-       mo^   ^   o      j  t-      mm 

^ ,        ^             .                       .            ^  ^  Riekia  sp.  Snelling,  1986:  6,  8  and  Figs  10,  19- 

sinus  (not  rising  above  upper  limit  of  fron-  21  male 

tal  spot  and  separated  from  it  by  about 

width  of  antennal  socket),  uninterrupted  Snelling  (1986)  erroneously  described 

but  posteromedially  narrowly  emarginate  confluens  as  a  species  of  Paragia  Shuck- 

pronotal  band  connecting  humeri.  Spot  on  ard,  1837  despite  the  fact  that,  in  his  own 


Volume  4,  1995 


35 


Figs.  1-4.  Riekia  nocatunga.  1,  dorsofrontal  view  of  vertex  and  dorsal  view  of  anterior  third  of  thorax  of 
female  (X17.6);  2,  dorsal  view  of  posterior  third  of  thorax  of  female  (X17.6);  3-4,  ventral  and  ventrolateral 
views  of  genitalia  of  male  (X57). 

Figs.  5-8.  Riekia  confluens.  5,  dorsofrontal  view  of  vertex  and  dorsal  view  of  anterior  third  of  thorax  of  female 
(X17.6);  6,  dorsal  view  of  posterior  third  of  thorax  of  female  (X17.6);  7-8,  ventral  and  ventrolateral  views  of 
genitalia  of  male  (X63);  [all  lemon-yellow  colour  form] 


generic  key,  the  species  with  its  uncon- 
stricted  gastral  tergum  2  runs  down  to 
Riekia  Richards,  1962.  Further,  with  the 
exception  of  the  number  of  spine-Hke  se- 
tae at  the  apex  of  the  mid-  and  hindtibiae 
(a  specific  character)  the  characters  given 
by  him  in  his  diagnosis  of  the  species  are 


all  common  to  both  coiiflnciii^  and  nocn- 
tungn  and  may  therefore  be  considered 
to  be  generic  characters  pertaining  to 
Riekia.  Similarly  the  confluence  of  the 
postocular  and  preoccipital  carinae 
(leading  to  the  name  confluens),  or  alter- 
natively the  absence  of  the  postocular  ca- 


36 


Journal  of  Hymenoptera  Research 


rina,  is  a  character  shared  with  nocatiin- 
ga.  Conversely,  the  same  characters  set 
confluens  (and  nocatiinga)  generically 
apart  from  Paragia. 

Material  collected  at  a  single  site  near 
Carnarvon,  Western  Australia,  at  distanc- 
es of  165  km  and  260  km  in  a  westerly  and 
west-south-westerly  direction  respectively 
from  the  type  localities  of  confluens  (16  km 
WSW  Lyons  River  Homestead,  24.38S, 
115.20E — paratype  female,  and  36  km  ESE 
Minnie  Creek  Homestead,  24.00S, 
115.42E — holotype  female)  consists  of  53 
females  and  4  males.  The  material  is  di- 
visible into  two  very  distinct  groups,  one 
black  and  reddish-brown  and  the  other 
black  and  lemon-yellow.  Whereas  intra- 
group  variability  is  negligible,  inter-group 
differences  with  respect  to  colour  gener- 
ally and  to  colour  pattern  on  the  metaso- 
ma  in  particular  are  striking. 

The  distinctness  of  the  two  sympatric 
groups  and  the  notable  absence  of  any  in- 
termediate forms  initially  led  to  the  belief 
that  two  species  were  represented.  How- 
ever, no  morphological  characters  could 
be  found  that  supported  this  view  and  it 
is  concluded  that  the  apparent  difference 
in  overall  facies  is  a  product  of  the  differ- 
ences in  colour  and  colour  pattern.  The 
reddish-brown  colour  form,  consistent 
with  the  description  of  Snelling's  two  fe- 
males (particularly  with  that  of  the  para- 
type), is  represented  in  the  present  mate- 
rial by  females  only,  the  lemon-yellow  col- 
our form  by  both  sexes.  To  facilitate  intra- 
specific  comparison,  descriptions  limited 
to  colour  pattern  are  given  of  both  female 
forms  whereas  for  the  purpose  of  inter- 
specific comparison  with  nocatiinga  the  de- 
scription of  the  male  is  given  in  greater 
detail. 

In  passing  it  is  noted  that  in  Snelling's 
description  it  is  incorrectly  stated  that  the 
probasitarsus  is  slightly  more  than  twice 
wider  than  long.  This  is  clearly  an  inad- 
vertent error  as  it  is  in  fact  slightly  less 
than  half  as  wide  as  long. 


Reddish-brown  (RB)  Colour  Form 

Female. — Black.  The  following  reddish- 
brown:  mandibles  other  than  for  teeth  and 
extreme  base,  upper  half  of  clypeal  disc,  a 
variously  developed  transverse  band 
(seemingly  formed  of  a  pair  of  medially 
fused  spots)  between  and  above  antennal 
sockets  and  narrowly  separated  ventrally 
(where  sometimes  angularly  emarginate) 
from  clypeal  marking,  variously  sized 
elongate  spot  in  upper  half  of  each  ocular 
sinus  (rising  to  level  of  anterior  ocellus 
and  exceeding  upper  margin  of  frontal 
marking)  [median  frontal  transverse  band 
and  lateral  spots  sometimes  broadly  fused 
above],  a  large  oval  spot  behind  eyes  dor- 
sally,  entire  upper  surface  of  prothorax 
other  than  for  narrow  streak  bordering 
posterior  margin,  tegulae  except  for  clear 
testaceous  central  spot  and  narrow  mar- 
gin, scutellar  disc  other  than  for  its  ante- 
rior margin,  a  large  spot  on  raised  part  of 
axillae  and  a  minute  spot  between  it  and 
tegula,  a  diffuse  spot  on  metanotum  me- 
dially, a  large  spot  on  angles  of  propo- 
deum,  large  spot  on  mesopleuron  below 
tegula,  distal  end  of  mid-  and  hindcoxae, 
part  of  foretrochanter  and  whole  of  mid- 
and  hindtrochanters,  femur,  tibia  and  tar- 
someres  of  all  legs,  gastral  tergite  1  other 
than  for  anterior  declivity,  tergite  2  other 
than  for  a  variously  developed  anterior 
black  band  which  may  be  triangularly 
produced  in  the  middle  and  for  a  pair  of 
widely  separated  diffuse  dark  markings 
on  posterior  margin,  an  anteriorly  point- 
ing triangular  spot  on  posterior  margin  of 
tergite  3,  tergite  4  other  than  for  a  narrow 
anterior  black  band,  tergite  5,  diffuse 
transverse  posterior  band  on  gastral  ster- 
nites  2-5  (those  of  sternites  2  and  3  and 
sometimes  4  strongly  and  widely  anteri- 
orly produced  in  the  middle). 

Lemon-yellow  (LY)  Colour  Form 

Female. — (Figs  5  and  6).  Black.  The  fol- 
lowing reddish-brown:  tip  of  mandibles 
except  actual  teeth,  flagellomeres  1-10  be- 


Volume  4,  1995  37 

neath,  trochanters  of  mid-  and  hindlegs,  eral   and   anterior   margins,   single   large 

proximal  half  to  three  quarters  of  outer  as-  frontal  spot  between  and  above  antennal 

pect    of    femora    (where    moderately    to  sockets,  lower  aspects  of  mid-  and  hind- 

heavily   black   suffused)    and    inner   and  coxae,  entire  outer  aspect  of  midfemora, 

lower  aspect  of  same,  inner  and  lower  as-  entire  outer  aspect  of  all  tibiae  (including 

pect  and  distal  end  of  tibiae,  all  tarsomer-  distal  ends),  narrow  longitudinal  streak  on 

es,  posterolateral  portions  of  dark  mark-  all  basitarsi,  transverse  posterior  band  on 

ings  on  sternites  2-4  and  sternite  6  apical-  gastral  sternites  2-5  (all  with  median  lobe 

ly  (all  black-suffused).  The  following  lem-  strongly  and  widely  anteriorly  produced, 

on-yellow:  proximal  half  to  two-thirds  of  together  giving  the  effect  of  a  wide  me- 

mandibles,    subapical    spots    on    scapes,  dian  longitudinal  band), 

clypeus  other  than  for  narrow  lateral  and  Antennal   scape   (with   radicle)   very 

anterior  margins  and  in  some  specimens  slightly    longer    than    interantennal    dis- 

arcuate  black  lines  on  distal  half,  a  pair  of  tance;  first  flagellomere  slightly  more  than 

irregularly  shaped  frontal  spots  between  half  (0.53)  as  long  as  scape  (with  radicle), 

and  above  antennae,  a  narrow  streak  on  one  and  three  quarter  times  as  long  as 

lower    inner    orbits    flanking    clypeus,    a  wide  at  its  distal  end,  and  twice  as  long  as 

broad  mark  in  ocular  sinus  rising  to  level  the  second  flagellomere.  Last  three  flagel- 

of  anterior  ocellus,  an  oval  spot  behind  lomeres  flattened  and  slightly  concave  be- 

eyes  dorsally,  uninterrupted  but  postero-  neath;    ultimate    flagellomere    narrowing 

medially   narrowly   emarginate   pronotal  apically  and  distinctly  curved  to  form  a 

band  connecting  humeri,  posterior  corner  hook. 

of  pronotum,  tegulae  except  for  clear  tes-  Tergite  7  truncate  and  narrowly  trans- 

taceous  central  spot  and  narrow  margin,  a  verse  apically. 

large  transverse  suboval  shield-like  spot  Genitalia  (Figs  7  and  8);  similar  to  those 
on  scutellum  (not  quite  reaching  anterior  of  R.  nocatunga  but  differing  in  detail  par- 
margin),  a  small  spot  on  raised  part  of  ax-  ticularly  with  respect  to  the  form  of  the 
illae,  a  large  spot  on  angles  of  propodeum,  inwardly  directed  lobe  of  the  volsella  be- 
large  spot  on  mesopleuron  below  tegula,  low  the  digitus. 

a  small  spot  on  mesosternum  anterior  to  Length  7.8-8.3  mm;  length  of  forewing 

coxal  cavities,  a  small  lateral  spot  on  mid-  5.5-5.7  mm;  hamuli  12-13. 

and  hindcoxae,  distal  half  of  outer  aspect  Material  Examined. — Western  Australia: 

of  forefemora,  distal  spot  on  outer  aspect  8  km  NE  of  Carnarvon  (24.51S,  113.45E) 

of  mid-  and  hindfemora,  outer  aspect  of  on  road  to  Bibbawarra  Bore,  3-5. x. 1993 

all  tibiae  (except  distal  ends),  wide  ante-  (F.W.,S.K.&  R.W.Gess)  16  RB  females,  29 

riorly  trilobed  posterior  band  on  gastral  LY  females  and  4  LY  males  (in  yellow 

tergites  1-5,  tergite  6  (other  than  for  vari-  flowers  of  Lechenaiiltia  sp.,  Goodeniaceae), 

ously  developed  posteriorly  pointing  V-  4  RB  females,  1  LY  female  (at  water),  2  RB 

shaped  black  mark  leaving  lateral  and  me-  females,  1  LY  female  (without  biological 

dian  spots  or  almost  eliminating  these),  data).  Two  RB  females,  2  LY  females  and 

transverse  posterior  band  on  gastral  ster-  1  LY  male  in  both  ANIC  and  WAM;  rest 

nites  2-5  (those  of  sternites  2  and  3  tri-  of  material  in  AMG. 

lobed    with    median    lobe    strongly    and  Discussion. — R.  confliieiis  may  be  distin- 

widely  anteriorly  produced).  guished  from  R.  nocatunga  in  both  sexes 

Male. — (Figs  7  and  8).  Colour  pattern  by  the  colour  pattern  and  the  below  indi- 

very  similar  to  that  of  female  but  differing  cated  morphological  characters, 

in  that  the  following  parts  are  pale  lemon-  In  R.  confluens  the  ratio  of  POL  (distance 

yellow:  entire  anterior  aspect  of  scapes,  between    posterior   ocelli):OOL    (distance 

entire  clypeus  other  than  for  narrow  lat-  between  a  posterior  ocellus  and  nearest 


38  Journal  of  Hymenoptera  Research 

eye  margin)  is  1:1.3-1.4,  whereas  in  R.  no-  the  species  differ  in  the  male  sex  in  the 

catunga  it  is  1:1.0-1.1.  relative  proportions  of  the  first  two  flagel- 

In  R.  confliiens  the  pilosity  of  the  pro-  lomeres,  in  the  form  of  gastral  tergite  7 

notum  is  only  moderately  dense,  the  in-  and  in  slight  differences  in  the  genitalia, 

dividual   hairs   being    slender   and    only  Present  data  would   indicate  that  the 

slightly  curved  apically  whereas  in  R.  no-  distributions  of  the  species  are  widely  sep- 

catunga  the  pilosity  is  much  denser  and  arated,   R.    nocatungci   occurring   in   New 

the  individual  hairs  are  more  robust  and  South  Wales  and  Queensland  and  R.  con- 

markedly  and  evenly  curved.  fluens  in  Western  Australia. 

In  R.  confluens  the  scutellum  (Fig.  6)  has 

its  disc  posteriorly  widely  and  evenly  to  Rolandia  angulata  (Richards) 

bluntly  rounded  and  abruptly  declivitous  ^.^^^.^  ^,^^^^^^^^  j^.^j^^^^^^  ^^^^.  ^q^_^q2,  female, 

to  the  metanotum  which  it  slightly  over-  ^^,^„^^,.^  ^^^^^^^^^  (Richards),  Snelling,  1986:  8. 
hangs;  the  posterolateral  free  edge  of  each 

lateral  wing  is  almost  straight.  The  meta-  This  species  was  described  from  Cun- 

notum  in  its  median  third  is  not  depressed  namulla  [28.04S,  145. 40E],  Queensland 

middorsally  and  is  therefore  of  greatest  (holotype  female  and  1  female  paratype) 

depth  in  the  middle;  it  is  vertical,  not  vis-  and   from  90   km  W  of  Cobar,   Barnato 

ible  from  above.  In  R.  nocatunga  the  scu-  Tanks    [31.38S,    144.59E]    (about   400   km 

tellum  (Fig.  2)  has  its  disc  posteriorly  nar-  south  of  Cunnamulla),  New  South  Wales 

rowly  rounded  and  gradually  descendant  (12  female  paratypes).  Snelling  (1986)  re- 

to  metanotum  which  it  does  not  overhang;  corded  an  additional  8  females  from  Cun- 

the  posterolateral  free  edge  of  each  lateral  namulla.    Hitherto   the   male   appears   to 

wing  is  widely  and   shallowly  concave,  have  been  unknown.  One  of  the  present 

The   metanotum   in   its   median   third   is  females  was  compared  with  the  holotype 

somewhat  depressed  middorsally  and  is  in  the  Australian  National  Collection  of 

therefore  of  even  depth;  it  is  sloping,  vis-  Insects,  Canberra,  and  found  to  be  identi- 

ible  from  above.  cal. 

In  R.  confluens  gastral  tergite  1  is  two  Male. — (Figs  9  and  10).  Colour  pattern 

and  two-fifths  to  two  and  a  half  times  very  similar  to  that  of  female,  differing 

wider  than  long  whereas  in  R.  nocatunga  chiefly  in  that  the  yellow  markings  are 

it  is  two  and  a  quarter  times  wider  than  lighter  in  hue  and  brighter  and  contrast 

long.  more    strongly    with    the    light    reddish 

In  R.  confluens  the  number  of  spine-like  brown  areas.  The  bright  yellow  mandi- 

setae  on  the  apex  of  the  mesotibia  is  1  bles,  clypeus  and  transverse  lower  frontal 

whereas  in  R.   nocatunga  it  is  2-4,  most  marking,  and  the  distinct  median  and  lat- 

commonly  3;  the  number  of  such  setae  on  eral  spots  on  the  gaster  facilitate  recogni- 

the  apex  of  the  metatibia  is  1-2,  most  com-  tion  of  the  male  even  in  the  field, 

monly  1,  and  2-3  respectively.  Antennal  scape   (with  radicle)   shorter 

The  species  differ  in  the  female  sex  in  than  interantennal  distance;  first  flagello- 

the  form  of  the  pronotum  as  seen  from  mere  three  times  as  long  as  broad  at  its 

above.  In  R.  confluens  (Fig.  5)  the  anterior  distal  end,  about  two-thirds  as  long  as 

and  lateral  margins  describe  a  semicircle  scape  (with  radicle)  [three-quarters  as  long 

whereas    in    R.    nocatunga    (Fig.    1)    the  as  scape  without  radicle],  and  a  little  more 

"shoulders"  are  anterolaterally  produced  than  twice  as  long  as  second  flagellomere; 

so  that  the  anterior  and  lateral  margins  do  flagellomeres  3-9  subequal  in  length  and 

not  describe  a  semicircle  but  the  anterior  with  the  exception  of  3  wider  than  long; 

margin  appears  subtransverse.  3-7  increasing  in  width;  7  and  8  of  same 

As  may  be  seen  from  the  descriptions,  width;  9  as  wide  as  6;  10  slightly  longer 


Volume  4,  1995 


39 


Figs.  9-10.     Rolandia  angulata.  9,  posterior  view  of  right  fore  leg  of  male  (X44.5);  10,  ventral  view  of  genitalia 
of  male  (X44.5). 


than  preceding  flagellomeres,  narrowing 
and  apically  rounded;  distal  flagellomeres 
not  modifled  beneath. 

Foreleg  (Fig.  9);  process  of  foretrochan- 
ter  large,  compressed,  broadly  digitiform 
in  posterior  view  and  slightly  forwardly 
curved,  its  apex  attaining  level  of  angulate 
base  of  forefemur;  forefemur  in  posterior 
view  broad  and  ventrally  strongly  angu- 
late at  base  and  widely  but  shallowly  in- 
curved from  there  to  about  midlength, 
with  ventral  margin  angulate  along  its 
length  but  becoming  lamellate  at  basal  an- 
gle, with  ventral  surface  basally  (i.e.  ad- 
jacent to  lamellate  ventral  angle)  excavat- 
ed. 

Genitalia  (Fig.  10);  parameral  spines 
apically  sharply  pointed  and  ventrally  re- 
curved; ventral  process  of  each  paramere 
tapering  towards  apex  in  lateral  view  but 
in  ventral  view  more  or  less  of  even  width 
over  most  of  its  length  and  apically  nar- 
rowly rounded;  digitus  and  inwardly  di- 
rected lobe  of  volsella  as  figured. 

Length  8.7-9.2  mm;  length  of  forewing 
6.3-6.5  mm,  hamuli  16-18. 

Material  Examined. — Queensland:  85  km 
E  of  St  George  (28.03S,  148.30E),  27.X.1993 


(F.W.,S.K.&  R.W.Gess)  6  females  (in  yel- 
low flowers  of  Goodenia  cycloptera  R.Br.in 
C.Sturt,  Goodeniaceae),  7  females  (in  yel- 
low flowers  of  Goodenia  pinnatifida  Schldl., 
Goodeniaceae);  27  km  W  of  St  George 
(28.03S,  148.30E),  28.X.1993  (F.W.,S.K.& 
R.W.Gess)  31  females,  10  males  (in  yellow 
flowers  of  Goodenia  cycloptera  R.Br.in 
C.Sturt,  Goodeniaceae),  1  female  (in  yel- 
low flowers  of  Goodenia  pinnatifida  Schldl., 
Goodeniaceae),  2  females  (nesting  in 
sandy  soil),  2  females,  3  males  (without  bi- 
ological data);  80  km  E  of  Cunnamulla 
(28.04S,  145.40E),  29.X.1993  (F.W.,S.K.& 
R.W.Gess)  49  females,  31  males  (in  yellow 
flowers  of  Goodenia  pinnatifida  Schldl., 
Goodeniaceae),  4  females  (nesting  in 
sandy  soil),  9  females,  2  males  (without  bi- 
ological data).  Three  females  and  3  males 
in  ANIC;  3  females  and  3  males  in  WAM; 
rest  of  material  in  AMG. 

Discussion. — Snelling  (1986)  was  correct 
in  his  assumption  that  the  males  of  R.  an- 
gulata would  differ  from  those  of  the  very 
similar  R.  houstoni  in  the  same  characters 
as  do  the  females.  Of  particular  note  is  the 
scutellum  which  in  both  sexes  of  R.  an- 
gulata slopes  evenly  to  the  metanotum  and 


40 


Journal  of  Hymenoptera  Research 


lacks  a  definite  posterior  face.  The  males 
run  down  satisfactorily  in  Snelling's  key, 
allowance  being  made  for  the  longer  than 
broad  second  flagellomere. 

ACKNOWLEDGEMENTS 

The  author  wishes  to  thank  Ian  Naumann  and  Jo 
Cardale  of  the  Australian  National  Insect  Collection, 
Canberra  for  hospitality  in  their  department  and  for 
facilitating  the  comparison  of  some  of  the  present 
specimens  with  type  material  in  their  care,  Kirsten 
Cowley  of  the  Australian  National  Herbarium,  Can- 
berra for  identifying  the  forage  plants,  Suzanne  Lewis 
of  the  Natural  History  Museum,  London  for  lending 
material  under  her  care,  and  Robin  Cross  of  the  Elec- 
tron Microscopy  Unit,  Rhodes  University,  Grahams- 
town  for  producing  the  scanning  electron  micro- 
graphs reproduced  as  Figs  1-10.  An  earlier  version  of 
the  manuscript  was  reviewed  by  Karl  V.  Krombein 
of  the  Smithsonian  Institution,  Washington  and 
James  M.  Carpenter  of  the  American  Museum  of  Nat- 
ural History,  New  York.  The  latter,  in  particular,  is 
thanked  for  his  constructive  critique  and  for  drawing 
attention  to  a  potential  synonymy  arising  from  the 
fact  that  R.  confluens  had  been  described  by  Roy  Snell- 
ing  as  a  species  of  Paragia,  a  genus  to  which  the  wasp 


in  question  quite  clearly  does  not  belong.  Gratitude 
to  the  South  African  Foundation  for  Research  Devel- 
opment is  expressed  for  a  rolling  support  grant  which 
made  the  field  work  in  Australia  possible. 

LITERATURE  CITED 

Meade-Waldo,  G.  1911.  Description  of  new  species 
of  African  Diploptera  in  the  collection  of  the  Brit- 
ish Museum.  Annals  and  Magazine  of  Natural  His- 
tory (8)8:  448-457. 

Richards,  O.W.  1962.  A  revisional  study  of  the  Masarid 
ivasps  (Hymenoptera,  Vespoidea).  London:  British 
Museum  (natural  History). 

Richards,  O.W.  1968.  New  records  and  new  species 
of  Australian  Masaridae  (Hymenoptera:  Vespo- 
idea). Journal  of  the  Australian  Entomological  Soci- 
ety 7:  101-104." 

Shuckard,  W.E.  1837.  Descriptions  of  new  exotic  acu- 
leate Hymenoptera.  Transactions  of  the  Entomolog- 
ical Society  of  London  2:  68-82. 

Snelling,  R.R.  1986.  The  taxonomy  and  nomenclature 
of  some  Australian  paragiine  wasps  (Hymenop- 
tera: Masaridae).  Contributions  in  Science  378:  1- 
19. 

van  der  Vecht,  J.  and  Carpenter,  J.M.  1990.  A  cata- 
logue of  the  genera  of  the  Vespidae  (Hymenop- 
tera). Zoologische  Verhandelingen  26:  1-62. 


J.  HYM.  RES. 
Vol.  4,  1995,  pp.  41-63 

The  Last  Twenty  Years  of  Parasitic  Hymenoptera  Karyology: 
An  Update  and  Phylogenetic  Implications 

Vladimir  E.  Gokhman  and  Donald  L.  J.  Quicke 

(VEG)  Botanical  Garden,  Moscow  State  University,  Moscow  119899,  Russia; 
(DLJQ)  Department  of  Biology,  Imperial  College  at  Silwood  Park,  Ascot,  Berks  SL5  7PY,  U.K. 


Abstract. — A  survey  of  karyological  data  in  the  parasitic  Hymenoptera  shows  a  considerable 
uniformity  in  chromosome  numbers  within  higher  taxonomic  groupings,  with  most  chalcidoids 
having  n  =  5-6  and  most  cynipoids,  ichneumonoids  and  those  very  few  species  of  Diapriidae 
and  Scelionidae  examined  to  date  having  n  =  10-12.  The  Encyrtidae  and  Eurytomidae  differ  from 
other  chalcidoids  examined  to  date  in  having  higher  n  values  (8-11).  The  braconid  subfamilies 
Aphidiinae  {ii  -  4-7),  Charmontiinae  (n  =  5)  and  Exothecinae  (»  =  6)  are  shown  to  have  lower 
values  than  do  other  braconids.  New  data  are  presented  for  32  species,  including  the  first  records 
of  chromosome  numbers  for  the  Eupelmidae,  Bethylidae,  Diapriidae,  Ormyridae,  the  ichneumonid 
subfamilies  Banchinae,  Pimplinae,  and  Tryphoninae,  and  the  braconid  subfamilies  Aphidiinae, 
Charmontinae,  Doryctinae,  Exothecinae,  Macrocentrinae,  Meteorinae,  and  Miracinae.  Chromo- 
some number  data  are  discussed  in  relation  to  current  views  on  Hymenoptera  phylogeny. 


INTRODUCTION 

There  has  long  been  a  tendency  for  tax- 
onomists  to  ignore  many  potential  sources 
of  systematic  evidence  in  favour  of  more 
traditional  morphological,  and  nowadays 
molecular,  data.  Whilst  this  lack  has  not 
been  ubiquitous  amongst  taxonomists, 
where  it  has  occurred  it  may  have  been 
due  to  a  number  of  reasons  including  lack 
of  understanding  of  novel  characters,  lack 
of  facilities  for  investigating  them  and  lack 
of  availability  of  information  about  such 
characters.  In  this  respect,  it  is  not  sur- 
prising that  chromosome  number  and  oth- 
er karyological  features  have  played  little 
role  in  considerations  of  hymenopteran 
phylogeny,  and  this  is  especially  true  for 
the  parasitic  families.  Reasons  for  this  are 
manifold.  Hymenopteran  chromosomes 
have  traditionally  been  studied  in  imma- 
ture stages  in  which  dividing  cells  are 
common,  but  larvae  of  parasitic  wasps  are 
perhaps  especially  difficult  to  identify  and 
obtain  except  when  species  are  in  culture. 
Recent  developments  in  the  study  of  hy- 


menopteran   chromosomes    (Imai    et    al. 

1988,  Baldanza  et  al.  1991b),  have  opened 
the  possibility  of  greatly  expanding  our 
knowledge  of  parasitic  wasp  karyology. 
Further,  it  is  now  possible  to  obtain  good 
results  working  with  adult  tissues  such  as 
ovaries  (Gokhman  1985,  1990a),  and  such 
protocols  have  allowed,  for  the  first  time, 
an  examination  of  chromosome  number 
and  structure  in  natural  populations. 

Karyological  studies  have,  nevertheless, 
played  an  important  role  in  the  systemat- 
ics  and  species  level  taxonomy  of  various 
groups  of  the  non-parasitic  Hymenoptera 
including,  sawflies,  ants,  bees  and  social 
wasps  (principally  Polistinae)  and  these 
have  been  reviewed  in  detail  elsewhere 
(see  for  example,  Kerr  1972,  Kerr  and  da 
Silveira  1972,  Naito  1982,  Imai  et  al.  1977, 
Moritz  1986,  Pompolo  and  Takahashi 
1987,  1990,  Hoshiba,  Matsuura  and  Imai 

1989,  Costa  et  al.  1993).  Wide  variation  in 
n  is  well  known  among  the  ants  (Formic- 
idae)  and  ranges  from  1  to  47  (Imai  and 
Taylor  1989,  Imai  et  al.  1990),  but  is  modal 
at  11  (Imai  et  al.  1988).  Indeed,  the  greater 


42 


Journal  of  Hymenoptera  Research 


part  of  this  range  can  even  be  found  with- 
in the  single  ant  genus  Myrmecia.  How- 
ever, within  the  parasitic  Hymenoptera 
there  appears  to  be  be  rather  more  consis- 
tency, and  for  most  famiUes  the  observed 
range  of  variation  is  far  narrower. 

Outside  of  the  Hymenoptera,  both  chro- 
mosome numbers,  structure  and  size  have 
all  been  found  useful  in  phylogenetic  re- 
construction, usually  but  not  exclusively 
in  combination  with  other  morphochar- 
acters,  and  this  has  been  particularly  true 
of  plant  systematics.  For  example,  George 
&  Geethamma  (1992)  have  recently  pro- 
posed a  phylogeny  of  jasmines  based  on 
chromosome  numbers  and  assumed  poly- 
ploidy events.  Among  the  insects,  chro- 
mosome numbers  have  been  used  exten- 
sively in  the  systematic  treatment  and 
phylogeny  of  the  Hemiptera  (Blackman 
1980,  Emeljanov  &  Kirillova  1989,  1991). 
Kuznetsova  (1985),  for  example,  has  con- 
cluded that  the  homopteran  subfamily  Or- 
geriinae  is  monophyletic  based  on  an  au- 
tosomal fusion  giving  2m =27  (in  males) 
compared  with  the  plesiomorphic  value  of 
2n  =  29  (in  males),  and  similarly  that  the 
tribe  Almanini  of  the  Orgeriinae  is  mono- 
phyletic on  the  basis  of  an  autosome-sex 
chromosome  fusion  (giving  2n=26). 

In  this  paper  we  review  the  current  state 
of  knowledge  of  the  karyology  of  parasitic 
wasps  and,  in  addition,  provide  new  data 
for  approximately  32  taxa  including  the 
first  records  of  chromosome  numbers  for 
the  Eupelmidae,  Bethylidae,  Diapriidae, 
Ormyridae,  the  ichneumonid  subfamilies 
Banchinae,  Pimplinae,  and  Tryphoninae, 
and  the  braconid  subfamilies  Aphidiinae, 
Charmontinae,  Doryctinae,  Exothecinae, 
Macrocentrinae,  Meteorinae,  and  Miraci- 
nae. 

MATERIALS  AND  METHODS 

New  karyotypic  data  were  obtained  us- 
ing ovarian  tissues  of  adult  wasps  which 
were  either  collected  from  the  wild  at  Sil- 
wood  Park,  Berkshire,  U.K.,  during  early 
September  1995,  or  obtained  from  labora- 


tory cultures.  Chromosome  preparations 
were  obtained  according  to  the  schedule 
described  in  Appendix  I.  Voucher  speci- 
mens for  Ichneumoninae  are  deposited  in 
the  collection  of  Moscow  State  University, 
those  for  other  taxa  are  in  the  Natural  His- 
tory Museum,  London. 

RESULTS  AND  DISCUSSION 

The  last  review  of  chromosome  number 
in  the  parasitic  Hymenoptera  was  by  Cro- 
zier  (1975)  at  which  time  values  were 
known  for  only  about  twenty  species 
(with  data  published  before  1930  not  in- 
cluded for  the  reason  of  insufficient  reli- 
ability); his  data  are  summarized  in  Figure 
1.  Since  then,  chromosomes  of  nearly  140 
additional  species  have  been  studied 
(Gokhman  1994),  and  for  this  review  we 
have  investigated  32  more  in  order  to  help 
fill  in  a  number  of  gaps  and  to  confirm 
some  previous  findings.  These  data  are 
presented  in  Table  1  and  summarized  in 
Figure  2.  The  great  majority  of  published 
chromosome  numbers  come  from  studies 
on  the  ichneumonid  subfamily  Ichneu- 
moninae by  Gokhman  (1985,  1987,  1989, 
1990a,  1990b,  1991a,  1991b,  1993a),  though 
they  also  include  a  substantial  body  of  ev- 
idence especially  relating  to  various  fam- 
ilies of  Chalcidoidea.  Within  the  better 
studied  parasitoid  families  (i.e.  Ichneu- 
monidae,  Cynipidae,  Torymidae)  chro- 
mosome numbers  were  found  to  be  rela- 
tively stable,  and  were  generally  uniform 
at  the  generic  level.  This  contrasts  mark- 
edly with  observations  for  many  aculeate 
Hymenoptera  which  often  show  striking 
chromosomal  variability  even  within  gen- 
era (Imai  and  Taylor  1989,  Imai  et  al. 
1990). 

The  apparent  bimodality  in  haploid 
number  of  parasitic  Hymenoptera  appar- 
ent in  Crozier's  small  sample  (Fig.  1)  now 
appears  to  be  well  founded  (Fig.  2).  How- 
ever, the  exact  modal  numbers  are  slightly 
different  (n  =  5  and  11  compared  with  n 
=  5  and  10  of  Crozier),  probably  due  to 
the  strong  bias  in  favour  of  the  ichneumo- 


Volume  4,  1995 
14  -T- 

12  - 

10 


43 


6  - 


2  -- 


+ 


+ 


+ 


+ 


4- 


3    4    5    6    7     8    9    10    11    12    13    14    15    16    17 

Haploid  chromosome  number 

Fig.  1.     Histogram  of  haploid  chromosome  numbers  for  parasitic  Hymenoptera,  data  modified  after  Crozier 
(1975)  to  take  into  account  subsequent  taxonomic  changes. 


nid  subfamily  Ichneumoninae  in  the  pres- 
ent enlarged  data  set.  As  will  be  apparent 
from  the  figures,  the  peak  in  haploid  val- 
ues around  5  largely  comprises  members 
of  the  Chalcidoidea  and  there  is  only 
slight  overlap  between  these  and  the  other 
peak  centred  around  11.  The  same  is  true, 
though  less  pronounced,  when  the  data 
are  plotted  for  genera  rather  than  species 
(Figure  3:  each  n  value  in  a  genus  being 
represented  only  once  in  the  histogram). 
Members  of  three  of  the  chalcidoid  fami- 
lies investigated  to  date,  the  Eurytomidae 
(Eiiri/toma),  Encyrtidae  (6  species  in  three 
genera)  and  Aphelinidae  (one  species, 
Pteroptrix  (=  Archenomiis)  orientalis  Silves- 
tri,  out  of  5  investigated;  see  below)  differ 
from  other  chalcidoids  in  having  haploid 
numbers  ranging  between  8  and  11,  whilst 
members  of  the  braconid  subfamilies 
Aphidiinae  {Aphidius,  Diaretiella,  Ephcdriis, 
Praon),  Charmontinae  [Charmon)  and  Ex- 
othecinae  [Rhysipolis)  are  atypical  in  hav- 
ing haploid  numbers  of  4  to  7,  6  and  5 
respectively,  compared  with  other  ichneu- 


monoids  whose  haploid  n  values  range 
from  8  to  17. 

GENOMIC  SIZE  AND  ITS 
IMPLICATIONS 

Rasch  et  nl.  (1975,  1977)  calculated  the 
haploid  genomic  size  of  the  braconid 
wasps,  Habrobracon  juglandis  and  H.  seri- 
nopae  both  to  be  0.154).16  x  10  ^^g  dNA 
and  that  of  the  pteromalid,  Nnsonin  (as 
Mormoniclla)  vitripennis  to  be  0.33-0.34  X 
10  '-^g.  These  values  translate  (using  an 
average  molecular  weight  of  a  base  pair 
as  660)  to  base  pair  numbers  of  approxi- 
mately 1.4  X  10**  and  3.0  X  10^  base  pairs 
for  the  ichneumonoid  and  the  chalcidoid 
respectively.  More  recently.  Bigot  et  al. 
(1991)  using  DNA  reassociation  kinetics 
calculated  the  haploid  genomic  sizes  of 
the  ichneumonid  Diadroniiis  piilchclliis  and 
the  chalcidoid,  Eiipclmus  viiillcti,  as  1-2  X 
10'^  base  pairs  and  10  X  10"^  base  pairs  re- 
spectively. Unfortunately,  no  data  for  the 
numbers  of  chromosomes  are  available  for 
Eupelmus  vuiUeti  but  we  have  investigated 


44 


Journal  of  Hymenoptera  Research 


Table  1.    Chromosome  number  in  parasitic  wasps 


Taxon 


2/!* 


Reterence(h)t 


Diaprioidea 

Diapriidae 
Beli/ta  depressa  Thomson 
Cinetus  lanceolatus  Thomson 
Scelionoidea 

Scelionidae 
Telejtomus  fariai  Lima 
Chalcidoidea 

Aphelinidae 
Apheliniis  mali  Haldeman 
Aphytis  mytilaspidus  (Le  Baron) 
Archenonnis  orientalis  Silvestri 
Encarsia  berlesei  Howard 
Encarsia  pergandiella  Howard 

Chalcididae 
Bradiyineria  intermedia  Nees 
B.  lasus  Burks 

B.  ovata  Say 

Dirhiniis  himalayanus  Westwood 
Encyrtidae 
Ageniaspis  fiiscicoUis  Dalman 
Copidosoma  buyssoni  Mayr 

C.  gelechiae  Howard 
C.  gelechiae  Howard 

C.  truncatelliim  (Dalman) 

(=  Ifloridanum  Ashmead) 
C.  floridanum  Ashmead 

C.  floridanum  Ashmead 
Apoanagyrus  lopezi  (DeSantis) 

Eulophidae 
Cirrospilus  diallus  Walker 
Colpoclypeiis  florus  Walker 
Melittobia  chalybii  Ashmead 
Tetrastichus  gigas  Burks 
T.  megachilidis  Burks 

Eupelmidae 
Anastatus  catalonicus  Bolivar 
Macroneura  vesicularis  (Retzius) 

Eurytomidae 
Eudecatoma  bigidtata  (Swederus) 
Eurytoma  californica  Ashmead 

Leucospidae 
Leucospis  affinis  Say 

Ormyridae 
Ormynis  sp. 

Pteromalidae 
Anisoptcromalus  calandrae  (Howard) 
Coelopisthia  extenta  (Walker) 
Dibrachys  sp. 

Lariophagus  distinguendus  Foerster 
Muscidifurax  zaraptor  Legner 
Nasonia  vitripennis  (Walker) 


8 
10 


10 


6 
6 
5 
6 
6 

5 
5 

9 
10 


?6 

7 

5 
5 
5 

5 
5 


16  present  papertt 

20  present  paper 


20  Dreyfus  &  Breuer  1944 


5 

10 

Viggiani  1967 

5 

10 

Rossler  &  De  Bach  1973 

11 

22 

Baldanza  et  al.  1991a 

5 

10 

Baldanza  et  al.  1991b 

6 

12 

Hunter  et  al.  1993 

3 

6 

Hung  1986 

5 

10 

Hung  1986 

5 

10 

Hung  1986 

5 

10 

Amalin  et  al.  1988 

c.  10 

C.20 

Silvestri  1908;  Martin  1914 

12 

24 

Silvestri  1914 

11 

22 

Hegner  1915 

10 

20 

Patterson  1921 

10 

20 

Hunter  &  Bartlett  1975 

8 

16 

Leiby  1922;  Patterson  1917,  1921 
Patterson  &  Porter  1917 

11 

22 

Strand  &  Ode  1990 

10 

20 

Dijken  1991 

12  present  paper 

12  Dijkstra  1986 

10  Schmieder  1938 

12  Goodpasture  1974 

12  Goodpasture  1974 

10  present  paper 

10  present  paper 

18  present  paper 

20  Goodpashire  1974 

12  Goodpasture  1974 

?12  present  paper 

14  present  paper 

10  present  paper 

10  Goodpashjre  1974 

10  Gershenzon  1968 

10  Goodpashare  1974 

10  Gershenzon  1946,  1968;  Pennypacker 

1958;  Whiting  1960,  1968;  Wahr- 

man  &  Zhu  1993 


Volume  4,  1995 


45 


Table  1.    Continued 


Taxon 


Reference(s)t 


Nasonia  vitripennis  (Walker) 

5  +  0-1 

IB      10 

Nasonia  vitripiennis  (Walker) 

6 

12 

Ptewmalus  puparum  L. 

5 

10 

P.  venustus  Walker 

5 

10 

Torymidae 

Monodontoinerus  dementi  Grissell 

6 

12 

M.  moniivagus  Ashmead 

6 

J2 

M.  obscurus  (Westwood) 

4 

8 

M.  obscurus  (Westwood) 

6 

12 

M.  saltuosus  Grissel 

5 

10 

Tory  mils  baccharidis  Huber 

6 

12 

T.  californicus  (Ashmead) 

6 

12 

T.  capillaceus  Huber 

6 

12 

T.  koebelci  Huber 

5 

10 

T.  occidentalis  Huber 

6 

12 

T.  tubicola  Osten-Sacken 

6 

12 

T.  umbilicatus  Gahan 

5 

10 

T.  vesiculi  Moser 

6 

12 

T.  warreni  (Cockerell) 

6 

12 

Trichogrammatidae 

Trichogmmma  cliilonis  Ischii 

5 

10 

T.  dcicvi  Pinto  &  Oatman 

5 

10 

T.  dendrolimi  Matsumura 

5 

10 

T.  evanescens  Westwood 

5 

10 

T.  nubialale  Ertle  &  Davis 

5 

10 

T.  pretiosum  Riley 

5 

10 

T.  spp.  (7  strains) 

Cynipoidea 
Cynipidae 
Andricus  curvator  Hartig 
A.  feciindator  Hartig 
A.  kollari  Hartig 
A.  quercuscalicis  Burgsdorf 
Aulacidea  hiemcii  Bouche 
Biorrhiza  pallida  Olivier 
Callirhi/tis  palmiformis  Ashmead 
Cynips  divisa  Hartig 
Diastrophus  nebulosus  Osten-Sacken 
Diplolepis  elganteriae  Hartig 
D.  nervosum  Curtis 
D.  rosae  L. 

D.  rosae  L. 

D.  spinosissimae  Girault 

Dryocosmus  kuripliilus  Yasumatsu 

Dryophanta  erinacea  Mayr 

Neurotcrus  laeviusculus  Schenck 

N.  numismalis  Fourcroy 

N.  quercusbaccarum  L. 

Trigo)iaspis  megaptera  Panzer 
Xestopluvics  potcntillac  Retzius 


10 


Nur  et  al.  1988;  Werren  1991 

Goodpasture  1974 

Guhl  &  Dozortseva  1934,  Dozortseva 

1936 

McDonald  &  Krunic  1971 

Grissell  1973b;  Goodpasture  1975a 
Goodpasture  1975a 
Goodpasture  1975a 
McDonald  &  Krunic  1971 
Grissell  1973b;  Goodpasture  1975a 
Goodpasture  &  Grissell  1975 
Goodpasture  &  Grissell  1975 
Goodpasture  &  Grissell  1975 
Goodpasture  &  Grissell  1975 
Goodpasture  &  Grissell  1975 
Goodpasture  &  Grissell  1975 
Goodpasture  &  Grissell  1975 
Goodpasture  &  Grissell  1975 
Goodpasture  &  Grissell  1975 

Hung  1982 

Stouthamer  &  Kazmer  1994 

Liu  &  Xiong  1988 

Hung  1982 

Hung  1982 

Hung  1982;  Stouthamer  &  Kazmer 

1994 

Fukada  &  Takemura  1943  (cited  by 

Hung  1982) 


10 

20 

Dodds  1938;  Sanderson  1988 

10 

20 

Dodds  1938 

10 

20 

Hogben  1920;  Sanderson  1988 

10 

?20 

Sanderson  1988 

10 

20 

Dodds  1938 

10 

20 

Dodds  1938 

w 

20 

Goodpasture  1979b 

10 

20 

Sanderson  1988 

10 

20 

Hegner  1915' 

9 

27(3n) 

Sanderson  1988 

9 

18 

Sanderson  1988 

9 

18 

Henking  1892;  Stille  &  Davring  1980; 
Sanderson  1988 

?12 

?24 

Schleip  1909 

9 

18 

Sanderson  1988 

10 

20 

Abe  1994 

c.  12 

24 

Wieman  1915 

10 

20 

Sanderson  1988 

10 

20 

Dodds  1938 

10 

20 

Doncaster  1910,  1911,  1916;  Dodds 
1938 

10 

20 

Dodds  1938 

10 

20 

Dodds  1938 

46 


Journal  of  Hymenoptera  Research 


Table  1.    Continued 


Taxon 

;;' 

2n* 

Reference(s)t 

Eucoilidae 

Pseudeucoila  bochei  Weld 

10 

20 

Jungen  cited  in  Crozier  1975 

Ichneumonoidea 

Braconidae 

Aphidius  rhopalosiphi  De  Stefani  Perez 

7 

14 

present  paper 

Biosteres  carbonarius  (Nees) 

U 

28 

present  paper 

Charmon  cruentatiis  Haliday 

5 

10 

present  paper 

Cotesia  gloineratiis  (L.)  (as  Apanteles) 

12 

24 

Hegner  1915i 

Dacnusa  sp. 

17 

34 

present  paper 

Diaeretiella  rapae  Mcintosh 

6 

12 

present  paper 

Ephedrus  sp. 

17 

?14 

present  paper 

Habwbracon  hebetor  Say 

10 

20 

Torvik-Greb  1935 

H.  jiiglandis  Ashmead^ 

10 

c.  20 

Torvik-Greb     1935;    Speicher     1937; 
Rasch  et  al.  1977 

H.  jiiglandis  Ashmead^ 

11 

22 

Whiting  1927 

H.  pectinophorae  Watanabe 

10 

20 

Inaba  cited  in  Makino  1959 

H.  serinopae  Ramakrishna 

10 

20 

Rasch  et  al.  1977 

Heterospihis  prosopidis  Viereck 

17 

34 

present  paper 

Macrocentrus  thomcicus  (Nees) 

13 

26 

present  paper 

Meteorus  versicolor  Wesmael 

8 

16 

present  paper 

Meteorus  gyrator  Thunberg 

10 

20 

present  paper 

Meteorus  pallipes  Wesmael 

10 

20 

present  paper 

Mirax  sp. 

10 

20 

present  paper 

Phaenocarpa  persimilis  Papp 

17 

34 

Prince  &  Stace  (cited  by  Crozier  1977) 

Praon  abjectum  Haliday 

4 

8 

present  paper 

Rln/sipolis  decorator  (Haliday) 

6 

12 

present  paper 

Ichneumonidae 

Aethercerus  discolor  Wesmael 

11 

22 

Gokhman  1985 

Ae.  dispar  Wesmael 

12 

24 

Gokhman  1991 

Ae.  nitidus  Wesmael 

11 

22 

Gokhman  1990a 

Ae.  raniui  Gokhman 

11 

22 

Gokhman  1991 

Agrothereutes  extrematus  (Cresson) 

10 

20 

Koonz  1939 

Aophis  pulchricornis  (Gravenhorst) 

13 

26 

Gokhman  1990b 

Aptesis  puucticollis  (Thomson) 

8 

16 

Gokhman  1990a 

Baeosemus  dent  if er  Gokhman 

11 

22 

Gokhman  unpubl.  obs. 

Baranisobas  ridibundus  (Gravenhorst) 

11 

22 

Gokhman  1990b 

Chasmias  niotatorius  (Fabricius) 

17 

34 

Gokhman  1985 

Coelichneiimon  cyaniventris  (Gravenhorst) 

13 

26 

Gokhman  1990a 

C.  sugillatorius  (L.) 

13 

26 

Gokhman  1990a 

Colpognathus  celerator  (Gravenhorst) 

11 

22 

Gokhman  unpubl.  obs. 

Cratichneumon  viator  (Scopoli) 

14 

28 

Gokhman  1985 

Diadroinus  prosopius  Holmgren 

11 

22 

Gokhman  1990a 

D.  pulchellus  Wesmael 

11 

22 

Hedderwick  et  al.  1985 

D.  subtilicornis  (Gravenhorst) 

11 

22 

Gokhman  1990b 

D.  troglodytes  (Gravenhorst) 

11 

22 

Gokhman  1990a 

D.  varicolor  Wesmael 

11 

22 

Gokhman  pers.  obs. 

Dicaelotus  sp.  nr.  parvulus  (Gravenhorst) 

11 

22 

Gokhman  1990a 

D.  piimilis  (Gravenhorst) 

11 

22 

present  paper 

Diphyus  latebricola  (Wesmael) 

12 

24 

Gokhman  1990a 

D.  raptorius  (L.) 

12 

24 

Gokhman  1990a 

Dirophanes  callopus  (Wesmael) 

9 

18 

Gokhman  1987 

D.  fidvitarsis  (Wesmael) 

10 

20 

Gokhman  1990a,b 

D.  invisor  (Thunberg) 

10 

20 

Gokhman  1987 

Dyspetes  arrogator  Heinrich 

10 

20 

present  paper 

Volume  4,  1995 


47 


Table  1.    Continued 


Taxon 


Ret'erence(s)+ 


Ephialtes  manifestator  (L.) 

15 

30 

Eurylahiis  tonms  Wesmael 

10 

20 

Gelis  sp. 

13 

26 

Gli/pta  lapponica  Holmgren 

9 

18 

Heterischnus  nigricollis  (Wesmael) 

11 

22 

H.  tnincator  (Fabricius) 

11 

22 

Homothenis  locutor  (Thunberg) 

11 

22 

Ichneimion  albiger  Wesmael 

12 

24 

/.  auip}iibolus  Kriechbaumer 

12 

24 

/.  bucciilcntus  Wesmael 

12 

24 

/.  confiisor  Gravenhorst 

12 

24 

/.  crassifemiir  Thomson 

12 

24 

/.  croceipes  Wesmael 

12 

24 

/.  extensorius  L. 

12,13 

24,  26^ 

/.  cf.  extensorius  L. 

11 

22 

/.  formosus  Gravenhorst 

11 

22 

/.  gracilentus  Wesmael 

12 

24  (25) 

/.  gracilicomis  Wesmael 

11 

22 

/.  ingratiis  Hellen 

12 

24 

/.  inquinatus  Wesmael 

13 

26 

/.  iiisidiosus  Wesmael 

12 

24 

/.  liigens  Gravenhorst 

12 

24 

/.  melanotis  Holmgren 

12 

24 

/.  mmutorius  Desvignes 

12 

24 

/.  molitorius  L. 

12 

24 

/.  nereni  Thomson 

11 

22 

/.  sarcitorius  L. 

11 

22 

/.  stramentarius  Gravenhorst 

12 

24 

I.  submarginatus  Gravenhorst 

10 

20 

/.  subquadmtiis  Thomson 

12 

24 

/.  suspiciosus  Wesmael 

12 

24 (26) 

/.  validicornis  Holmgren 

22 

24 

Lissonota  sp. 

11 

22 

Mastnis  smithii  (Packard) 

13 

26 

Oroiwtus  binotatus  (Gravenhorst) 

11 

22 

Orthocentrus  sp. 

14 

28 

Orthopelma  mediator  (Thunberg) 

11 

22 

Patrodoides  chah/beatiis  (Gravenhorst) 

8 

16 

Phaeogencs  melaiiogonos  (Gmelin) 

11 

22 

Ph.  nigridens  Wesmael 

11 

22 

Ph.  semivulpimis  (Gravenhorst) 

9 

18 

Ph.  spiuiger  (Gravenhorst) 

11 

22 

Polysphmcta  tiiberosa  Gravenhorst 

9 

18 

Pseudoambhjteles  homocerus  (Wesmael) 

9 

18 

Stenichneumon  cidpator  (Schrank) 

14 

28 

Sycaonia  sicaria  (Gravenhorst) 

11 

22 

Syspasis  alboguttata  (Gravenhorst) 

11 

22 

S.  scutellator  (Gravenhorst) 

11 

22 

Tycherus  australogemimis  Gokhman 

11 

22 

T.  bellicontis  (Wesmael) 

10, U 

20(21) 

T.  dilleri  Ranin 

n 

22 

T.  fuscicornis  (Wesmael) 

n 

22 

T.  ischiomelinus  (Gravenhorst) 

9 

18 

present  paper 
Gokhman  1987 
present  paper 
present  paper 
Gokhman  1990b 
Gokhman  unpubl.  obs. 
Gokhman  1990b 
Gokhman  1990a 
Gokhman  1990b 
Gokhman  1993a 
Gokhman  1985 
Gokhman  1985 
Gokhman  1990b 
Gokhman  1993a 
Gokhman  1990a 
Gokhman  1990a 
Gokhman  1993a 
Gokhman  1990a 
Gokhman  1990b 
Gokhman  1993a 
Gokhman  1990a 
Gokhman  unpubl.  obs. 
Gokhman  1990a 
Gokhman  1987 
Gokhman  1990a 
Gokhman  1990a 
Gokhman  1990a 
Gokhman  1987 
Gokhman  1990a 
Gokhman  1990b 
Gokhman  1993a 
Gokhman  unpubl.  obs. 
present  paper 
Koonz  1936 
Gokhman  1987 
Gokhman  1990a 
Hogben  1920 
Gokhman  1993a 
Gokhman  1990b 
Gokhman  1990b 
Gokhman  unpubl.  obs. 
Gokhman  1990a 
present  paper 
Gokhman  unpubl.  obs. 
Gokham  1985 
Gokham  19990b 
Gokhman  1985 
Gokhman  unpubl.  obs. 
Gokhman  1991 
Gokhman  1989 
Gokhman  1989 
Gokhman  1990b 
Gokhman  1991 


48 


Journal  of  Hymenoptera  Research 


Table  1.    Continued 


Taxon 

n* 

2n* 

Reference(s)+ 

T.  ophthalmicus  (Wesmael) 

22 

Gokhman  1990a 

T.  osculator  (Thunberg) 

22 

Gokhman  1989 

T.  suspicax  (Wesmael) 

22 

Gokhman  1987 

Ventiiria  canescens  (Gravenhorst) 

22 

Speicher  1937 

Virgichneumon  digrammus  (Gravenhorst) 

34 

Gokhman  1990a 

V.  f annus  (Gravenhorst) 

22 

Gokhman  1990a 

Vulgichneumon  snturatorius  (L.) 

9 

18 

Gokhman  1987 

Chrysidoidea 

Bethylidae 

Epyris  niger  Westwood 

14 

28 

present  paper 

Laelius  utilis  Cockerell 

10 

20 

present  paper 

*  In  papers  which  only  quote  n  or  2)i,  the  other  value  has  been  surmised  and  is  given  in  italics, 
t  Data  appearing  in  works  before  1930  should  be  considered  with  great  caution  as  most  resulted  from  his- 
tological rather  than  cytological  protocols,  involving  sectioned  material  rather  than  squash  preparations  and 
also  often  involving  fixation  techniques  not  well  suited  for  the  study  of  chromosomes,  though  some  of  these 
earlier  findings  are  clearly  correct  including  the  oldest  one  (Henking  1892). 

tt  For  some  new  data  we  were  not  able  to  obtain  an  unambiguous  chromosome  number  but  our  best  ap- 
proximation is  presented  (data  indicated  in  table  with  a  "?")  as  in  some  cases  these  still  provide  potentially 
valuable  information. 

'  Hegner  (1915)  did  not  provide  a  definitive  statement  on  chromosome  number  and  the  data  here  come  from 
his  rather  stylized  figures;  such  data  need  therefore  to  be  considered  with  extreme  caution. 
^  Some  workers  consider  H.  juglandis  to  be  a  junior  synonym  of  H.  Iwbetor,  however,  this  is  not  yet  absolutely 
confirmed  and  therefore  we  prefer  to  keep  these  records  separate. 
^  A  single,  probably  aneuploid,  female  specimen  with  25  chromosomes  has  also  been  found. 


3  4  5  6  7  8  9  10         11         12         13         14         15         16         17 

Haploid  chromosome  number 

Fig.  2.     Histogram  of  haploid  chromosome  numbers  of  parasitic  Hymenoptera,  data  from  Table  1  based  on 
n  values  of  each  species. 


Volume  4,  1995 
30  T 


25     - 


49 


20      - 


15      - 


10 


7  8  9  10         11         12         13 

Haploid  chromosome  number 


14 


15 


16 


17 


Fig.  3.     Histogram  of  haploid  ciiromosome  numbers  of  parasitic  Hymenoptera,  data  from  Table  1,  each  n 
value  occurring  in  a  genus  being  represented  only  once. 


two  other  eupelmid  species  in  the  genera 
Macroneura  and  Anastatiis,  which  were 
both  found  to  have  haploid  numbers  of  5 
in  common  with  Nasonia.  Although  the  re- 
sults of  Rasch  et  al.  (1975, 1977)  and  of  Big- 
ot et  al.  (1991)  don't  quite  agree  in  the  or- 
der of  magnitude  of  base  pairs  they  esti- 
mate, probably  due  to  differences  in  pro- 
cedure, they  did  both  show  the  chalcidoid 
to  have  a  larger  genome  than  the  ichneu- 
monid.  Thus,  if  these  values  are  roughly 
representative  of  other  members  of  their 
superfamilies,  then  the  chromosomes  of 
most  chalcidoids  would  be  expected  to 
have  between  four  and  twenty  times  as 
much  DNA  on  average  than  chromosomes 
of  ichneumonids. 

CHROMOSOME  NUMBER  IN 

RELATION  TO 
HYMENOPTERA  PHYLOGENY 

In  karyological  studies  it  is  common 
practice  to  interpret  modal  chromosome 
numbers  as  representing  the  initial  (ances- 
tral) number,  though  to  many  cladists  this 


would  be  interpreted  as  the  application  of 
the  much  decried  commonality  principle. 
The  'common  equals  primitive'  associa- 
tion is  of  course  probabilistic  rather  than 
deterministic  (Watrous  and  Wheeler  1981, 
Frohlich  1987,  Quicke  1993).  Some  further 
insight  into  whether  modal  chromosome 
number  is  likely  to  reflect  the  ancestral 
number  can  come  from  the  comparison  of 
modal  and  median  numbers.  If  these 
numbers  coincide,  and  the  whole  distri- 
bution may  be  approximated  to  a  normal 
one,  then  the  data  provide  no  evidence  of 
a  directed  change,  though  it  must  be  re- 
membered that  there  is  always  the  possi- 
bility that  an  evolutionary  change  in  chro- 
mosome number  early  in  the  evolution  of 
the  group  could  lead  to  the  same  distri- 
bution. In  the  Ichneumonoidea  and  the 
Cynipoidea  both  the  median  and  modal 
chromosome  numbers  are  the  same  11  and 

10  respectively;  in  contrast,  for  example, 
ants  (Formicidae)  have  a  modal  number  of 

11  but  the  median  is  15.  However,  cladis- 
tic  analyses  based  on  independent  char- 


50 


Journal  of  Hymenoptera  Research 


acter  systems,  via  outgroup  comparisons, 
provide  the  most  reliable  means  of  deter- 
mining plesiomorphic  chromosome  num- 
bers and,  where  possible,  this  is  the  ra- 
tional that  we  have  employed. 

According  to  currently  accepted  views 
of  Hymenoptera  phylogeny,  the  sawflies 
('Symphyta')  form  a  paraphyletic  group 
with  respect  to  the  Apocrita  with  the  latter 
being  the  sister  group  of  the  Orussidae, 
and  the  Apocrita + Orussidae  in  turn  most 
probably  being  the  sister  group  of  the 
Xiphydriidae  (Konigsmann  1977,  Rasnit- 
syn  1980, 1988,  Gibson  1985).  Unfortunate- 
ly, chromosome  numbers  are  not  known 
either  for  the  Orussidae  or  for  the  Xiphy- 
driidae, although  they  are  known  for 
members  of  two  other  sawfly  families 
with  claims  for  a  close  relationship  with 
the  Apocrita,  viz  the  Siricidae  and  Cephi- 
dae  (Konigsmann  1977,  Basibuyuk  & 
Quicke  1994,  1995).  In  the  Cephidae  n 
ranges  from  9  to  22-26  (Mackay  1955,  Cro- 
zier  and  Taschenberg  1972),  whilst  in  the 
Siricidae,  according  to  Sanderson  (1932, 
1970),  the  haploid  number  varies  between 
8  and  18.  As  regards  other,  less  derived 
sawflies,  haploid  chromosome  number 
ranges  from  5  to  22  in  the  Tenthredino- 
idea,  with  three  quarters  of  species  having 
an  n  value  ranging  between  7  and  10  (Nai- 
to  1982).  Taking  the  Siricidae  and  Cephi- 
dae as  the  two  sawfly  families  closest  to 
the  ancestral  lineage  of  the  Apocrita  (i.e. 
putative  sister  groups  of  the  Apocrita  + 
Orussidae)  for  which  chromosome  num- 
bers are  available,  it  seems  reasonable  to 
conclude  that  the  plesiomorphic  haploid 
chromosome  number  in  the  latter  was  at 
least  8  and  possibly  rather  higher. 

Within  the  Apocrita  there  is  a  picture 
emerging  from  independent  investiga- 
tions of  phylogenetic  relationships  (Ras- 
nitsyn  1988,  Johnson  1988,  Gibson  1985, 
Mason  1983,  Quicke  et  al.  1993,  1994,  Her- 
aty  et  al.  1994)  that  the  group  divided  rel- 
atively early  in  its  history  into  a  lineage 
giving  rise  to  the  Ichneumono- 
idea+Aculeata  and  a  second  comprising 


the  bulk  of  the  taxa  currently  regarded  as 
'Microhymenoptera'  including  Chalcidoi- 
dea,  Cynipoidea,  Scelionoidea,  Diaprioi- 
dea  and  Proctotrupoidea  s.s.  (Fig.  4). 

Our  data  show  that  the  modal  n  value 
in  the  Ichneumonoidea,  the  probable  sister 
group  of  the  Aculeata,  is  11.  Further,  our 
limited  data  for  the  less  derived  aculeates 
of  the  family  Bethylidae  {Epyris  and  Lae- 
liiis;  Fig.  10),  whilst  demonstrating  some 
degree  of  variation  in  haploid  number  be- 
tween 10  and  14,  when  considered  togeth- 
er with  data  for  other  aculeates  suggest 
the  ancestral  aculeate  may  have  had  a 
haploid  number  around  11,  as  was  also 
concluded  by  Hoshiba,  Matsuura  and 
Imai  (1989).  Similarly,  available  values  for 
three  other  parasitoid  superfamilies,  the 
Diaprioidea  (Fig.  5),  Scelionoidea  and  Cy- 
nipoidea, are  similar.  According  to  Ras- 
nitsyn's  (1988)  phylogenetic  hypothesis 
(see  Fig.  4),  the  Scelionoidea  are  putatively 
the  sister  group  of  the  Chalcidoidea  s.l., 
and  thus  taking  the  former  as  the  out- 
group, the  plesiomorphic  haploid  chro- 
mosome number  for  the  Chalcidoidea 
may  be  postulated  as  being  approximately 
10.  Therefore  the  data  collectively  support 
the  hypothesis  that  the  small  values  of  n 
(from  3  to  7)  found  in  the  majority  of  Chal- 
cidoidea are  likely  to  be  apomorphous. 
Unfortunately,  there  are  no  well  founded 
views  of  relationships  within  the  Chalci- 
doidea (Trjapitzin  1978,  LaSalle  1987,  Bou- 
cek  1988a,  Woolley  1988,  Noyes  1990,  Gib- 
son 1990),  largely  perhaps  because  of  the 
considerable  plasticity  in  adult  morphol- 
ogy displayed  by  many  of  the  families, 
which  may  result  because  of  the  undoubt- 
edly polyphyletic  natures  of  some  family 
level  taxa.  Trjapitzin  (1978)  made  few  pro- 
posals about  higher  level  relationships, 
and  only  suggested  two  possible  group- 
ings, his  'pteromaloid'  group  comprising 
Pteromalidae,  Tanaostigmatidae,  Eupel- 
midae  and  Encyrtidae,  and  a  'tetracam- 
poid'  group  comprising  Tetracampidae, 
Eulophidae,  Elasmidae  and  Aphelinidae. 
A  relationship  between  the  Aphelinidae 


Volume  4,  1995 


51 


Chalckjidae 
Apheiinidae 


Ormyridae 

Pteromalidae 
Torymidae 

Trichogrammatidae 
Encyrtidae 


Cynipoidea 


EvantOKJea 
CeraphronoKiaa 
Trigonatyoidea 
Megatyroidea 
Stephanoidea 


Diaprioidea 

Ichneumonidae 

Braconidae 


Chrysidoidea 


(lO-l'^ 


Vespoidea 
+  Apoidea 


Cephoidea 


Tenthredinoide 


Megaiodontoidea 


Xyeloidea 


Fig.  4.  Chromosome  number  and  evolution  of  the  parasitic  Hymenoptera,  with  haploid  values  overlain  on 
a  cladogram  in  which  superfamilial  relationships  are  those  proposed  by  Rasnitsyn  (1988)  but  with  relation- 
ships within  the  Chalcidoidea  based  on  current  karyological  evidence  for  clarity.  Ranges  and  (in  parentheses) 
modal  values. 


and  the  Encyrtidae  has  also  been  pro- 
posed by  a  number  of  workers,  but  Gib- 
son (1986)  and  WooUey  (1988)  considered 
that  the  supposed  synapomorphies  might 
be  better  regarded  as  resulting  from  con- 
vergence. LaSalle  (1987)  upheld  Trjapit- 
zin's  view  that  the  Tanaostigmatidae,  Eu- 


pelmidae  and  Encyrtidae  form  a  mono- 
phyletic  group,  citing  two  putative  syna- 
pomorphies. Boucek  (1988b)  suggested 
that  four  families,  the  Chalcididae,  Eury- 
tomidae,  Torymidae  and  [some]  Pterom- 
alidae were  relatively  'ancient',  partly  be- 
cause of  their  5  segmented  tarsi,  compar- 


52 


Journal  of  Hymenoptera  Research 


© 


© 


® 


»<^\ 

^'^Z 


*v> 


© 


® 


I 


¥ 


r 


Fig.  5-10.  Photomicrographs  of  chromosomes  of  adult  female  Hymenoptera  revealed  by  Giemsa  staining  of 
ovarian  tissue.  5,  Behjia  depressa  (Diapriidae),  meiosis,  diakinesis  in  mature  egg  {2n  =  16);  6,  7,  Anastatus 
catalonicus  (Eupelmidae),  6,  meiosis,  diplotene  figures  in  developing  oocyte,  7,  metaphase  mitosis  (2«  =  10); 
8,  Diaretiella  rapae  (Braconidae,  Aphidiinae),  metaphase  mitosis  {2n  =  12);  9,  Meteorus  versicolor  (Braconidae, 
Meteorinae),  mitosis,  one  cell  at  pro-metaphase  and  one  at  metaphase  {2n  =  16);  10,  Laelius  utilis  (Bethylidae), 
metaphase  mitosis  {2«  =  20). 


Volume  4,  1995  53 

atively  large  size  and  thoracic  structure,  they  do  not  agree  well  with  the  hypothesis 
and  he  agreed  with  Trjapitzin  and  LaSalle  of  a  close  relationship  between  Eupelmi- 
that  at  least  the  Eupelmidae  and  Encyrti-  dae  and  Encyrtidae  (Trjapitzin  1978,  La- 
dae  might  form  a  closely  related  group.  Salle  1987,  Boucek  1988a,  Woolley  1988, 
However,  this  view  was  not  supported  by  Noyes  1990)  which  have  haploid  numbers 
Gibson  (1989,  1990)  who  considered  both  of  5  (Figs  6,  7)  and  8-11  respectively,  be- 
that  the  Eupelmidae  might  not  be  mono-  cause  if  a  modal  haploid  chromosome 
phyletic  and  that  the  characters  used  to  number  of  5  represents  a  synapomorphy, 
unite  them  with  the  Encyrtidae  and  Tan-  then  this  relationship  would  require  either 
aostigmatidae  are  ".  .  .  either  primitive  two  separate  reductions  (from  c.  10)  or  a 
features  or  apparently  were  derived  in-  reversal.  However,  a  close  relationship  be- 
dependently  several  times".  tween  Eupelmidae  and  Encyrtidae  is  not 
The  present  data  could  suggest  that  universally  accepted  (Gibson  1989,  1990), 
there  has  been  an  approximate  halving  of  and  chromosome  number  should  be  in- 
chromosome  numbers  within  the  Chalci-  corporated  in  future  cladistic  analyses  of 
doidea,  from  around  a  modal  number  of  the  superfamilies  as  an  independent  char- 

10  as  shown  by  the  Eurytomidae  and  En-  acter. 

cyrtidae  to  5  or  6  in  Aphelinidae  (but  see  Potentially  significant  variation  occurs 

below),  Chalcididae,  Eulophidae,  Leucos-  within   the   Braconidae  and   the   Ichneu- 

pidae,  Ormyridae,  Pteromalidae,  Torymi-  monidae.   In   the   former   family,   several 

dae  and  Trichogrammatidae.  It  should  be  subfamilies    have    haploid    numbers    be- 

noted  that  within  the  Aphelinidae,  whilst  tween  8  and  11,  for  example,  the  Bracon- 

most  taxa  examined  have  haploid  values  inae  (Habrobrncon),  Meteorinae  {Meteonis; 

of  5,  Pteropterix  orientalis  has  an  n  value  of  Fig.  9),  and  Miracinae  {Mirnx).  However, 

11  (referred  to  as  Archenomus  orientalis  by  rather  higher  numbers  from  12  to  17  are 
Baldanza  et  al.  1991a).  However,  P.  orien-  found  in  the  Doryctinae  (Heterospilus), 
talis  is  an  highly  derived  species  within  Alysiinae  {Phaenocarpa,  Dacnusa)  and  Opi- 
the  Coccophaginae,  being  either  a  sister  inae  (Biosteres),  whereas  substantially  low- 
group  of  Coccophagoides  or  representing  a  er  numbers  {n  =  4  to  7)  are  found  in  the 
derived  branch  within  Encarsia  itself  (A.  Aphidiinae  {Aphidius,  Diareticlla  (Fig.  8), 
Polaszek  personal  communication).  In  ei-  Ephedrus,  Praon),  in  the  exothecine  genus 
ther  case,  the  high  haploid  chromosome  Rln/sipolis  (n  =  6),  and  in  the  unrelated 
number  in  this  taxon  is  clearly  derived  Charmontinae  {Charmon;  n  =  5).  The  pos- 
with  respect  to  the  values  of  5  (and  6)  dis-  session  of  low  values  within  the  four  gen- 
played  by  the  other  aphelinids  studied  era  of  Aphidiinae  examined  provides  a 
(collectively  representing  both  the  Cocco-  potential  synapomorphy  for  the  group, 
phaginae  and  Aphelininae),  and  these  which  otherwise  seems  an  heterogeneous 
lower  values  may  therefore  be  taken  as  assemblage  in  which  members  are  united 
representing  the  ancestral  range  of  values  by  few  characters  other  than  biology,  and 
for  the  family  as  a  whole.  The  n  value  of  more  taxa  will  need  to  be  examined  before 
11  in  Pteropterix  orientalis,  being  nearly  this  can  be  confirmed.  The  low  n  values 
twice  that  found  in  the  other  aphelinids  found  in  Rln/sipolis  might  be  an  autapo- 
investigated,  is  further  suggestive  that  this  morphy,  and  it  would  be  interesting  to 
taxon  could  have  originated  through  a  know  something  about  chromosome  num- 
polyploidy  event.  bers  in  the  apparently  closely  related  Cli- 

The  karyological  data  summarized  here  nocentrini  and  other  Rogadinae  s.s..  The 

lend  some  support  to  Boucek's  view  that  haploid  number  of  5  found  in  Charmon 

the  Eurytomidae  are  a  relatively  ancient  was  especially  surprising  as  the  subfamily 

and  underived  family  of  Chalcidoidea  but  seems  to  be  close  to  the  Macrocentrinae 


VARIATION  AND  IMPLICATIONS  FOR 
HYMENOPTERA  SYSTEMATICS 


54  Journal  of  Hymenoptera  Research 

(Quicke  and  Achterberg  1990,  Quicke  et  any  great  extent  in  studies  of  Hymenop- 

al.  1994)  for  which  we  obtained  a  value  of  tera  systematics. 

^^Within  the  Ichneumonidae,  most  data  ,, .  J^J^l  ^^D  INTEFSPECIFIC 
available  up  until  now  were  for  the  sub- 
family Ichneumoninae  but  a  few  chromo- 
some numbers  for  the  Campopleginae  Only  with  the  advent  of  techniques  for 
(Venturia),  Cryptinae  {Agr other eutes,  Mas-  examining  chromosome  number  and  mor- 
triis),  Orthocentrinae  (Orthocentrus),  and  phology  in  adult  Hymenoptera  has  it  be- 
Orthopelmatinae  (Orthopelma)  had  also  come  possible  to  study  intraspecific  vari- 
been  published.  The  overwhelming  major-  ation  in  nature  and  thus  even  to  reveal  the 
ity  of  the  species  had  haploid  chromo-  presence  of  hitherto  unsuspected  cryptic 
some  numbers  modal  at  11.  We  extended  species  or  species  complexes.  As  will  be 
this  data  set  by  making  chromosomal  apparent  from  Table  1,  chromosome  num- 
preparations  also  for  the  Banchinae  {Glyp-  ber  is  often  relatively  consistent  within  a 
ta,  Lissonotn),  Pimplinae  {Ephialtes  and  Po-  single  genus.  For  example,  in  the  Ichneu- 
lysphincta),  and  Tryphoninae  (Dyspetes).  monidae,  the  5  species  of  Diadromus  for 
These  additional  data  generally  support  which  chromosome  numbers  are  available 
the  earlier  findings,  although  the  ephial-  all  have  a  haploid  value  of  11.  Even  for 
tine  genus  Ephialtes  had  an  n  value  of  15,  the  large  genus  Ichneumon,  18  of  the  25 
rather  higher  than  appears  typical  for  the  species  examined  have  n  =  12. 
family,  while  the  polysphinctine  genus  Po-  Although  chromosome  numbers  are 
lysphincta  had  the  more  typical,  even  low,  usually  considered  as  differentiating  char- 
haploid  value  of  9.  acters,  serving  to  help  distinguish  between 
In  addition  to  chromosome  number,  closely  related  forms,  they  may  also  be 
chromosome  size  and  structure  have  been  used  in  an  integrative  fashion,  providing 
used  extensively  for  cytotaxonomic  pur-  evidence  for  uniting  related  forms  if  the 
poses  in  other  groups  of  organisms.  Dis-  chromosome  number  represents  a  syna- 
cussion  of  size  and  centromere  position  in  pomorphy.  For  example,  the  discovery 
the  parasitic  Hymenoptera  is  currently  se-  that  all  members  of  the  cynipid  genus  Di- 
verely  hampered  by  the  relative  paucity  of  plolepis  have  n=9,  whereas  the  haploid 
data;  in  fact,  in  many  illustrations,  and  es-  values  for  all  members  of  the  other  cyni- 
pecially  among  the  earlier  ones,  centro-  pid  genera  investigated  to  date  for  which 
meres  are  hardly  (if  at  all)  discernable.  reliable  figures  are  available  is  10,  pro- 
However,  if  we  look  at  the  karyotypes  of  vides  additional  evidence  for  the  mono- 
the  best  studied  groups  (i.e.  Ichneumoni-  phyly  of  Diplolepis. 

dae,  Torymidae,  Cynipidae),  bi-armed  Chromosome  numbers  are  fixed  in  the 

(metacentric  in  the  broad  sense)  chromo-  great  majority  of  species   of  parasitic 

somes  predominate  in  most  cases  (Figs  7-  wasps  whose  populations  have  been  stud- 

10).  Some  Hymenotera,  for  example  sev-  ied  in  detail  in  the  field.  Two  possible  ex- 

eral  Diplolepis  species  (Sanderson  1988),  ceptions,  however,  are  the  ichneumonines 

may  also  have  numerous  acrocentric  chro-  Icheumon    extensorius    and    I.    suspiciosus, 

mosomes.   Even  less  can  be  said   about  both  of  which  were  revealed  by  Gokhman 

chromosome  size,  though  in  general  in  the  (1993a)  to  comprise  individuals  with  two 

parasitic  Hymenoptera  it  is  inversely  re-  different    diploid    numbers,    24    and    26. 

lated  to  chromosome  number.  Much  more  Since    specimens    with    the    intermediate 

by  way  of  comparative  and  quantitative  chromosome    number    were    not    found 

data  will  have  to  be  assembled  before  it  (with  one  possible  exception  in  /.  extenso- 

will  be  possible  to  use  these  features  to  rius)  the  possibility  that  these  represent 


Volume  4,  1995 


55 


Table  2.    Sibling  species  in  the  parasitic  Hymenoptera  detected  by  karyological  features 


Family 


Species 


Chromosomal 
characteristics 


Reference 


Ichneumonidae 
Ichneumonidae 


Ichneumonidae 
Ichneumonidae 

Encyrtidae 


Pteromalidae 
Torymidae 

Torymidae 


Aethecerus  dispar  Wesmael 
Aethecerus  ranuii  Gokhman 

Tycherus  australogemiiius 

Gokhman 
Tycherus  ischionieliniis 

(Gravenhorst) 

Ichneumon  extensorius  L. 

Ichneumon  suspiciosus  Wes- 
mael 

Copidosoma  "tnincatellum 
(Dalman)'" 

Copidosoma  floridanum  (Ash- 
mead) 

C.  floridamim  (Ashmead) 


Nasonia  vitripennis  (Walker) 
Nasonia  vitripennis  (Walker) 

Torymus  californicus  (Ash- 
mead) 

Totrymus  warreni  (Cocker- 
ell)^ 

Monodontomerus  obscurus 

(Westwood)'* 
Monodontomerus  obscurus 

(Westwood)^ 


2n  =  24 

2»  =  22 

2n  =  22 
2«  =  18 

2n  =  24,  26' 
2n  =  24,  26 

n  =  10,  2n  =  20 
n  =  11,  2n  =  22 
n  =  8,2n  =  16 


n  =  5  +  0  -  IB,  2n  =  10 
n  =  6,  2«  =  12 

n  =  6(6M),  2n  =  12 

n  =  6(5M  +  lA),  2n  =  12 


n  =  6,  2n  =  12 


Gokhman  1991a 
Gokhman  1991a 


Gokhman  1993a 
Gokhamn  1993a 

Hunter  &  Bartlett  1975 

Strand  &  Ode  1970 

Leiby  1922;  Patterson  1917, 
1921;  Patterson  &  Porter 
1917 

Nur  ct  al.  1988  and  others 
Goodpasture  1974 

Goodpasture  &  Grissell 
1975 


Goodpasture  &  Grissell 

1975 
McDonald  &  Krunic  1971 


Abbreviations:  A  =  acrocentric;  M  =  metacentric  (in  a  broad  sense). 

'  A  single,  probably  aneuploid,  female  specimen  with  25  chromosomes  has  also  been  found. 

^Apparently  belongs  to  C.  floridamim. 

■  Apart  from  T.  californicus,  the  second  chromosome  pair  in  T.  ivarreni  has  secondary  constrictions. 

■*  Apparently  belongs  to  M.  laticornis  Grissell  &  Zerova. 


two  sibling  species  rather  than  chromo- 
somal races,  seems  much  more  likely, 
though  the  reverse  cannot  at  present  be 
discounted.  Despite  intensive  effort,  in 
neither  case  was  it  possible  to  detect  any 
external  morphological  criteria  to  permit 
delineation  of  these  putative  taxa  in  the 
absense  of  karyological  evidence.  Further, 
in  one  instance,  a  chromosomal  polymor- 
phism in  terms  of  C-banding  pattern  has 
been  detected  in  the  ichneumonine  spe- 
cies, Dirophanes  invisor  (Thunberg).  The  C- 
banding  patterns  of  the  two  homologous 
chromosomes  of  the  second  pair  of  meta- 
centrics do  not  differ  from  one  another  in 
some  individuals,  but  in  others,  this  pair 


is  obviously  heteromorphic,  one  member 
of  the  pair  being  substantially  longer  than 
the  other,  and  its  segment  of  pericentric 
heterochromatin  is  also  much  more  devel- 
oped (Gokhman  1993b). 

Several  other  recently  discovered  ex- 
amples of  apparent  interspecific  variation 
within  other  parasitic  Hymenoptera  are 
summarized  in  Table  2  and  discussed  be- 
low in  more  detail.  The  torymid  chalci- 
doids  Torymus  californicus  and  T.  ivarreni 
were  considered  to  form  a  single  species 
by  Grissell  (1973a).  However,  karyological 
analysis  has  shown  that  despite  the  fact 
that  these  two  wasps  have  the  same  chro- 
mosome number  (2n  =  12),  T.  ivarreni  has 


56 


Journal  of  Hymenoptera  Research 


a  pair  of  acrocentric  chromosomes  and  has 
secondary  constrictions  on  the  second 
largest  pair  of  submetacentrics,  whilst  in 
T.  californicus  all  the  chromosomes  are  bi- 
armed  and  the  second  pair  has  no  con- 
strictions (Goodpasture  and  Grissell  1975). 
Thus  these  two  may  well  be  best  inter- 
preted as  sibling  species.  Other  cases  of 
possible  karyologically-detected  sibling 
species  in  the  Chalcidoidea  are  more  prob- 
lematical. Goodpasture  (1975a)  and  Mc- 
Donald and  Krunic  (1971)  reported  n  =  4 
and  n  =  6  respectively  for  apparently  the 
same  torymid  species,  Monodontomerus  ob- 
scurns.  The  most  likely  explanation  for  this 
difference  is  that  one  of  these  works  in- 
volved a  misidentified  species  with  the 
specimens  examined  by  McDonald  and 
Krunic  actually  belonging  to  the  very  sim- 
ilar species,  M.  laticornis  Grissell  &  Zerova, 
described  14  years  later  (Zerova  and  Gris- 
sell 1985).  Two  new  ichneumonid  species 
of  the  subfamily  Ichneumoninae,  Tycherus 
australogeminus  and  Aetheceriis  ranini,  were 
originally  detected  on  the  basis  of  karyo- 
logical  evidence,  but  in  each  case  reliable 
morphological  differences  were  also 
found  (Gokhman  1991a;  see  also  Table  2). 
Application  of  karyology  may  also  be 
important  in  laboratory  cultures  as  avail- 
able evidence  suggests  that,  at  least  in 
some  instances,  strains  that  were  believed 
to  belong  to  a  single  species  may  in  fact 
represent  more  than  one,  with  different 
laboratories  working  on  different  entities. 
For  example.  Hunter  and  Bartlett  (1975) 
working  with  what  they  referred  to  as 
Copidosoma  truncatellum,  reported  it  as 
having  a  haploid  number  of  10.  C.  trun- 
catellum was  subsequently  partly  synony- 
mized  with  C.  floridanum,  but  Strand  & 
Ode  (1990)  reported  n  =  11  for  apparently 
the  same  species.  Several  earlier  workers 
had  reported  the  haploid  number  for  C. 
floridanum  (as  Paracopidosomopsis  floridan- 
us)  to  be  8  (Leiby  1922,  Patterson  1917, 
1921,  Patterson  and  Porter  1917),  but  their 
findings  have  to  be  treated  with  consid- 
erable caution  as  the  techniques  for  fixa- 


tion and  preparation  used  in  pre-1930 
studies  are  often  unreliable.  Also  in  the 
genus  Copidosoma,  Hegner  (1915)  reported 
n  =  11  and  Patterson  (1921)  reported  n  = 
10  for  C.  gelechiae.  Again  these  data  may 
not  be  fully  reliable  for  technical  reasons. 
However,  it  is  harder  to  interpret  the  ap- 
parent conflict  in  reported  numbers  for  the 
widely  studied  pteromalid,  Nasonia  vitri- 
pennis.  Many  workers  (e.g.  Gershenzon 
1946,  1968,  Pennypacker  1958,  Whiting 
1960,  1968)  have  reported  an  n  value  of  5, 
but  Goodpasture  (1974)  working  on  the 
University  of  California  at  Davis  culture 
found  n  =  6.  As  with  the  case  of  the  ich- 
neumonines  discussed  below,  intensive 
morphological  investigation  of  these  and 
other  strains  by  Goodpasture  failed  to  re- 
veal any  differences.  Therefore,  the  possi- 
bility that  the  Davis  culture  had  devel- 
oped as  a  unique  chromosomal  race  must 
be  considered.  Such  variants  are  not  un- 
common in  cultures  of  other  organisms. 
However,  it  should  be  noted  that  Darling 
and  Werren  (1990)  recently  discovered 
two  cryptic  species  of  Nasonia  in  North 
America,  and  the  karyological  results 
could  also  reflect  a  sibling  species  com- 
plex. 

The  discovery  of  sibling  species  that  can 
only  reliably  be  separated  by  karyotype 
may  pose  a  considerable  nomenclatural 
problem,  since  the  current  Code  of  the 
Zoological  Nomenclature  requires  new 
taxa  to  be  differentiated  from  existing 
ones.  As  it  is  not  normally  possible  to  ob- 
tain karyological  data  from  the  type  spec- 
imens of  the  species  that  have  already 
been  described,  it  would  not  possible  to 
give  scientific  names  to  both  of  them 
(Gokhman  1993a)  unless  it  were  possible 
(for  instance  through  geographic  distri- 
bution) to  infer  the  karyotype  of  the  de- 
scribed taxon. 

Intraspecific  karyotypic  variation  may 
also  be  of  interest  for  population  cytoge- 
netics. This  type  of  variation  is  favoured 
in  Hymenoptera  by  some  characteristics  of 
their  genetic  system,  which  allows  the  sur- 


Volume  4,  1995 


57 


vival  and  comparatively  high  viability  of 
aneuploids  (Imai  et  al.  1984).  The  data  ob- 
tained, for  example,  for  the  ichneumonid 
Tycherus  heUicornis  suggest  the  long-term 
persistence  of  a  chromosomal  population 
polymorphism,  probably  induced  by  a 
translocation  and  subsequent  non-disjunc- 
tion of  chromosomes  (Gokhman  1989). 
Occasional  aneuploid  specimens  were  also 
found  in  Ichneumon  extensor ius  and  /.  gra- 
cilentiis.  Perhaps  the  most  interesting  case 
of  numerical  chromosomal  polymorphism 
is  described  by  Nur  et  al.  (1988)  and  Wer- 
ren  (1991)  and  occurs  in  the  pteromalid, 
Nasonia  vitripennis.  This  type  of  polymor- 
phism implies  the  existence  of  a  particular 
B  chromosome.  Being  transmitted  pater- 
nally into  the  diploid  zygotes,  this  chro- 
mosome eliminates  all  other  chromosomes 
of  the  paternal  set  from  the  zygote,  thus 
converting  the  originally  diploid  zygote 
into  a  haploid  one.  Therefore  this  B  chro- 
mosome may  be  considered  as  the  most 
selfish  genetic  element  ever  known. 

FUNCTIONAL  IMPLICATIONS 

Reduced  chromosome  numbers  will 
generally  be  associated  with  reduced  lev- 
els of  recombination  (Vorontsov  1966, 
White  1973).  Under  many  circumstances, 
a  reduction  in  recombination  will  be  mal- 
adaptive. However,  situations  that  favour 
high  levels  of  inbreeding,  for  example,  ei- 
ther parasitisation  of  aggregated  hosts  or 
gregarious  parasitism  combined  with  mat- 
ing near  the  emergence  site,  may  lead  to 
selection  in  favour  of  low  intrinsic  levels 
of  genetic  diversity.  Such  situations  may, 
for  example,  favour  parthenogenesis  or  in 
the  case  of  the  parasitic  Hymenoptera, 
thelytoky.  Similarly,  under  such  circum- 
stances, high  levels  of  recombination,  or 
large  numbers  of  separate  linkage  groups, 
will  not  be  favoured,  and  consequently, 
mutations  leading  to  reduced  recombina- 
tion will  not  necessarily  be  so  deleterious. 
Such  circumstances  will  permit  a  reduc- 
tion in  chromosome  number  more  readily 
than  will  situations  favouring  high  levels 


of  genetic  diversity.  Consideration  of  the 
data  currently  available  for  parasitic  Hy- 
menoptera may  be  relevant  in  this  respect. 
For  example,  in  the  Aphidiinae  the  chro- 
mosome number  is  markedly  lower  than 
in  the  rest  of  the  Braconidae.  Aphidiines 
parasitise  aphids  which  in  turn  often  form 
clonal  patches.  In  chalcidoids,  many  spe- 
cies are  also  gregarious  or  attack  clustered 
hosts,  perhaps  in  part  as  a  result  of  their 
small  size  and  dispersal  capabilities.  It 
would  be  difficult  to  quantify  this,  but  the 
possibility  exists  that  inbreeding  may  be 
more  common  in  the  Chalcidoidea  as  a 
whole  than  in  the  Ichneumonoidea  or  Cy- 
nipoidea  (Askew  1968).  Information  on  re- 
combination levels  in  parasitic  Hymenop- 
tera is  extremely  limited  to  date  (Crozier, 
1975),  being  based  on  linkage  data  {Habro- 
bracon  hebetor)  and  chiasmata  per  bivalent 
arm  {Aphytis  mytilaspidiis).  Further  studies 
taking  into  account  cross-over  and  levels 
of  heterozygosity  in  parasitic  wasps  with 
different  biologies  and  different  chromo- 
some numbers  and  morphologies  might 
provide  additional  evidence  in  this  re- 
spect. 

FURTHER  PERSPECTIVES 

Though  we  have  managed  for  the  first 
time  to  obtain  cytogenetic  information  for 
several  major  groups  of  parasitic  Hyme- 
noptera, the  overwhelming  majority  of 
these  insects  still  remains  untouched  by 
karyological  investigation,  and  this  in- 
cludes some  entire  superfamilies  such  as 
the  Stephanoidea,  Megalyroidea,  Trigon- 
alyoidea,  Evanioidea  and  Ceraphronoidea 
(Fig.  4).  The  data  available  at  present, 
however,  suggest  that  the  new  chromo- 
somal evidence  may  be  a  substantial  help 
in  future  phylogenetic  and  taxonomic 
studies.  As  regards  the  higher  level  phy- 
logeny  of  the  parasitic  Hymenoptera,  new 
karyological  evidence  is  especially  needed 
for  various  groups  of  Proctotrupoidea  sen- 
su  lato  and  of  Chalcidoidea,  especially  of 
such  apparently  underived  taxa  as  the 
Mymaridae,  Tetracampidae  and   Rotoiti- 


58 


Journal  of  Hymenoptera  Research 


dae  (Noyes  1990)  and  the  pteromalid  sub- 
family Cleonyminae  (Boucek  personal 
communication). 

Recent  investigations  (e.g.  Gokhman 
1991b,  1994,  Costa  et  al.  1993,  Odierna  et 
al.  1993)  also  show  that  other  karyological 
data,  especially  those  obtained  using  dif- 
ferential chromosome  staining  (e.g.  C- 
banding),  can  still  provide  useful  infor- 
mation even  in  the  absence  of  differences 
in  chromosome  number.  More  detailed 
morphological  investigations  are  therefore 
particularly  likely  to  be  of  use  in  future 
species  level  work. 

ACKNOWLEDGEMENTS 

We  would  like  to  thank  Efi  Kazantzidou  (Horti- 
cultural Research  International,  Wellesbourne),  Prof. 
Helmut  van  Emden  (University  of  Reading)  and 
Robert  Belshaw,  Peter  Mayhew  and  Patricia  Reader 
(Imperial  College)  for  providing  living  material  for 
original  investigation,  Lynn  Kimsey  for  enabling  us 
to  see  C.  Goodpasture's  Ph.D.  thesis,  Andy  Polaszek 
for  helpful  discussions  pertaining  to  the  Aphelini- 
dae,  and  Zdenek  Boucek,  John  LaSalle,  and  Mark 
Shaw  for  identifying  various  specimens.  This  work 
was  partly  supported  by  grants  from  the  British 
Council  and  International  Science  Foundation  to 
VEG;  DLJQ  is  supported  by  the  NERC  Initiative  in 
Taxonomy. 

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APPENDIX  I.  METHODOLOGY  FOR 

STAINING  CHROMOSOMES  IN 

ADULT  PARASITIC  WASPS 

General  comments.  Chromosomes  can  be  stained 
in  many  adult  parasitic  wasp  females  especially  in 
recently  eclosed  individuals  using  ovarian  tissue.  As 
always  in  insects,  care  should  be  taken  to  count  mul- 
tiple cells  so  that  the  occasional  polyploid  cell  can 
be  recognised  and  discounted.  In  common  with  Cro- 
zier  (1975)  we  recommend  counting  at  least  ten  in- 
dividual metaphase  plates  though  occasionally  this 
may  not  be  possible.  If  fewer  plates  are  available, 
one  may  gain  extra  confidence  in  the  results  if  it  is 
possible  to  identify  through  size  and  morphology, 
particular  pairs  of  chromosomes  (see  for  example. 
Fig.  9).  Usually,  mitotic  divisions  are  most  evident, 
but  in  some  individuals  and  taxa,  meiosis  may  also 
be  observed,  sometimes  with  very  clear  spreads.  The 
number  of  plates  may  also  be  increased  if  the  wasp 
is  fed  on  honey  water  containing  colchicine  for  a  few 
hours  before  preparation,  though  beyond  about  5 
hours  one  stands  the  risk  of  increasing  the  propor- 
tion of  polyploid  cells.  The  general  method  de- 
scribed below  is  modified  after  that  of  Imai  et  al. 
(1988). 

Method.  Metasomas  of  adult  female  wasps  are 
dissected  in  hypotonic  sodium  citrate  solution  with 
colchicine  (Solution  A).  Ovaries  are  incubated  in  this 
solution  at  room  temperature  (c.  25°C)  for  20  min- 
utes (optimal  times  for  different  taxa  may  vary 
slightly).  Following  incubation,  ovaries  are  trans- 
ferred to  a  thoroughly  cleaned  microscope  slide,  ex- 
cess citrate  solution  pipetted  off  and  the  slide  is 
gently  flushed  with  Fixative  I  taking  care  not  to 
wash  off  the  ovaries.  Whilst  still  moist  with  Fixative 
I,  the  ovaries  are  disrupted  (e.g.  using  fine  mounted 
needles)  and  their  cells  spread  gently  over  the  mid- 
dle part  of  the  slide.  One  or  two  drops  of  Fixative 
II  are  then  applied  to  the  centre  of  the  area  of  spread 
cells  and  the  more  aqueous  phase  which  is  displaced 
to  the  edge  of  the  slide  is  blotted  off.  The  same  pro- 


Volume  4,  1995 


63 


cedure  is  then  performed  with  Fixative  III.  The  slide 
is  then  air  dried  before  staining  for  at  least  20  min- 
utes. Excess  stain  should  be  washed  off  with  dis- 
tilled water  and  the  slide  examined  dry  or  under 
emersion  oil  (do  not  apply  mounting  media).  Slides 
can  be  restained  if  the  initial  result  was  insufficient- 
ly intense,  or  they  can  be  destained  with  alcohol  in 
the  reverse  situation. 


Solutions 

(A)  Hypotonic  sodium  citrate  with  colchicine 
Ig  Na  citrate.2H,0 


5mg  colchicine 

100  ml  distilled  water 

(B)  Fixative  I 

3  parts  Ethanol 

3  parts  Glacial  acetic  acid 

4  parts  distilled  water 

(C)  Fixative  II 

1  part  Ethanol 

1  part  Glacial  acetic  acid 

(C)  Fixative  III 
Glacial  acetic  acid 

(D)  Stain 

2  ml  Giemsa  solution 
50  ml  0.089M  Na.HPO, 
50  ml  0.066M  KH>0, 


J.  HYM.  RES. 
Vol.  4,  1995,  pp.  64-76 

Euryischomyia  Girault  (Hymenoptera:  Chalcidoidea:  Aphelinidae: 

Eriaporinae:  Euryischiini) 

Mary  Carver 
Division  of  Entomology,  CSIRO,  G.P.O.  Box  1700,  Canberra,  A.C.T.  2601,  Australia 


Abstract. — Types  of  some  Australian  Euryischiini  described  by  A.  A.  Girault  have  been  exam- 
ined, and  compared  with  other  material  identified  by  Girault,  together  with  other  reared  and 
collected  material  of  Euryischomyia  Girault.  Euryischomyia  saintpverrei  Girault,  1915,  syn.  nov.; 
Myiocnema  marmorativentris  Girault,  1915,  syn.  nov.;  E.fasciata  Girault,  1916,  syn.  nov.;  and  £.  setosa 
Girault,  1929,  syn.  nov.;  are  placed  in  synonymy  with  £.  flaznthorax  Girault  &  Dodd,  1915;  and  £. 
alami  Shafee,  1975,  syn.  nov.,  (described  fom  India),  is  placed  in  synonymy  with  E.  washingtoni 
Girault,  1914.  A  key  to  the  2  remaining  species  of  Euryischomyia  is  provided.  £.  washingtoni  and 
£.  flavithorax  are  redescribed.  £.  zuasJiingtoni  is  known  as  a  parasite  of  pseudococcids,  and  £. 
flavithorax  as  a  hyperparasite  of  psyllids,  aphids,  coccids  and  pseudococcids. 


INTRODUCTION  compressed  hind  coxae,  large,  bristle-like 

np,        ,              ill.           J                 i.-  setae  on  the  lees  and  elsewhere,  postaxil- 

The  placement,  status  and  composition  ,              ,, 

of  the  subfamily  Eriaporinae  (Chalcidoi-  ^^^^^  ^^'^^  scutellum  and  a  large  propo- 
dea:  Aphelinidae)  have  long  been  a  matter 

of  uncertainty.  The  subfamily  was  erected  '^^^  nomenclatural  history  of  the  Eu- 

in  the  family  Aphelinidae  by  Ghesquiere  ^yischiini  is  complicated  by  the  fact  that 

(1955).   Members  have  been  variously  ^^^^^^  ^^^^^^  ^^^  ^^'"l^^^  ^^^^^^^  ^^^  f^^" 

placed  in  the  Aphelinidae  (e.g.  Ashmead  ^^y  Euryischidae  {sic)  to  contam  Euryischia 

1900;  Compere  1947;  Ferriere  1965)  and  ^^^'  subsequently,  he  included  Myiocnema 

Elasmidae  (e.g.  Girault  1914;  Smith  and  ^^^  Euryischomyia  in  the  family  also  (Sha- 

Compere   1928;   Nikol'skaya   1963).   Cur-  ^^^    1^75)    but,    later,    Shafee   and    Rizvi 

rently,  the  Eriaporinae  are  included  in  the  (^^^O)  retained  only  Euryischia  in  the  fam- 

Aphelinidae  (e.g.  Hayat  1994),  albeit  with  i^Y-  transferring  the  other  2  genera  to  the 

some   reservation   (Goulet   and   Ruber  Aphelinidae.  The  elevation  of  these  genera 

1993).  Contained  taxa  of  Eriaporinae  pos-  ^  family  status,  and  their  separation,  are 

sess  an  unbroken,  posteriorly  expanded  "^o*  accepted  herein. 

parastigma  in  the  fore  wing,  which  is  usu-  Although  Euryischia  is  well  character- 
ally  continued  posterobasally  and  bears  1-  ised,  the  limits  of  the  other  genera  of  Eu- 
3  coarse,  bristle-like  setae.  ryischiini  have  not  been  understood  and 
Hayat  and  Verma  (1980)  created  2  tribes  their  history  is  also  confusing.  Essentially, 
within  the  Eriaporinae,  namely,  the  Eria-  Girault  (1914)  described  the  new  genus 
porini  to  contain  the  genera  Promuscidea  and  species  Euryischomyia  washingtoni  and, 
Girault  and  Eunotiscus  Compere;  and  the  in  1917,  erected  the  genus  Paramyiocnema 
Euryischini  (sic)  for  4  genera:  EuryiscJiia  for  Myiocnema  marmorativentris  Girault  but 
Riley,  1889;  Myiocnema  Ashmead,  1900;  did  not  describe  or  diagnose  the  genus. 
Euryischomyia  Girault,  1914;  and  Para-  Girault  (1930)  placed  Euryischomyia  in  syn- 
myiocnema  Girault,  1917.  Euryischiini  are  onymy  with  Myiocnema.  Hayat  and  Verma 
characterised  by  possession  of  enlarged,  (1980)  transferred  Euryischomyia  flavithorax 


Vt^LUME  4,  1995 


65 


Girault  &  Dodd  to  Parnmyiocnema.  They 
also  opined  that  E.  saintpierrei  Girault,  E. 
fasciata  Girault  and  E.  setosa  Girault  might 
also  belong  in  Paramyiocnema.  Shafee  et  al. 
(1985)  placed  Paramyiocnema  in  synonymy 
with  Myiocnema  and  Hayat  (1994)  placed 
it  in  synonymy  with  Eiiryischomx/ia. 

The  types  of  Eiiryischomyia  washingtoni, 
E.  flavithorax,  E.  saintpierrei,  Myiocnema 
marmorativentris,  E.  fasciata,  and  £.  setosa 
have  been  examined  by  the  author,  to- 
gether with  other  material  identified  by 
Girault,  and  more  recently  reared  and  col- 
lected material  of  EuryiscJioirn/ia.  As  a  re- 
sult, Eiiryischomyia  saintpierrei,  Myiocnema 
marmorativentris,  E.  fasciata,  and  £.  setosa 
are  placed  in  synonymy  with  £.  flavithorax. 

In  addition,  £.  alami  Shafee,  1975,  de- 
scribed from  India,  is  proposed  as  a  junior 
synonym  of  £.  washingtoni,  thus  reducing 
Eiiryischomyia  to  2  species,  namely,  £. 
washingtoni  and  £.  flavithorax. 

METHODS 

All  museum  material  was  examined  as 
provided,  as  dry-mounts,  or  as  uncleared 
or  cleared  slide-mounts.  Girault's  material 
was  recognizable  from  information  pro- 
vided by  Dahms  (1984,  1986),  and  by  rec- 
ognition of  Girault's  or  Mrs  E.J.  Girault's 
handwriting  on  the  labels. 

Whenever  feasible,  reared  material  was 
obtained  by  caging  live,  field-collected 
material  to  allow  maximal  mummification 
of  already-parasitized  hosts.  Aphid  mum- 
mies (mummified,  parchment-like,  some- 
times silk-lined,  aphid  skins  containing 
immature  parasites)  were  removed  and 
placed  individually  in  gelatin  capsules. 
Except  when  slide-mounted,  each  emer- 
gent parasite  was  preserved  in  association 
with  its  mummy.  Reared  material  was 
dry-mounted,  or  preserved  in  80%  etha- 
nol,  or  cleared  and  slide-mounted  using 
10%  potassium  hydroxide,  chloral-phenol, 
and  gum-chloral  mounting  media. 

Micrometer  eyepieces  were  used  for 
measurements,  all  of  which  are  of  maxi- 
mal values;  those  of  the  scape  do  not  in- 


clude the  radicle;  those  of  the  metasoma 
include  the  ovipositor.  Length  of  mesoso- 
ma  +  metasoma  is  given  rather  than  body 
length  because  the  head  of  most  of  Gi- 
rault's specimens  is  separated  from  the 
trunk  and  slide-mounted  fronto-occipital- 
ly  and,  in  those  instances  of  two  or  more 
mountings  per  coverslip,  head  and  trunk 
could  not  be  matched  with  confidence. 

Girault  customarily  enumerated  the 
number  of  specimens  he  used  in  a  descrip- 
tion of  a  new  species,  and  then  implicitly 
designated  only  1  or  2  types  from  that  ma- 
terial. Dahms  (1983)  considered  as  types 
all  specimens  stated  by  Girault  to  have 
been  used  by  him  in  a  description.  Dahms' 
practice  is  followed  herein. 

Dahms  (1984,  1986)  and  Hayat  and  Ver- 
ma  (1980)  catalogued  Girault's  material  of 
Eiiryischomyia  in  detail,  obviating  the  ne- 
cessity to  do  so  herein.  The  information 
contained  in  the  lists  of  specimens  exam- 
ined has  therefore  been  simplified.  Where 
appropriate,  label  data  are  enclosed  in 
quotation  marks;  added  or  interpreted  in- 
formation is  enclosed  in  square  brackets. 
Latitudes  and  longitudes  have  in  most 
cases  been  provided  by  the  author. 

The  terminology  adopted  is  that  of  Nau- 
mann  (1991),  except  for  the  following:  An- 
tennal  formula:  representation  of  the  num- 
ber of  segments  comprising,  respectively, 
the  scape,  pedicel,  anellus,  funiculus,  club. 
Postaxilla:  a  small  triangular  sclerite  pos- 
terior to,  and  apparently  a  division  of,  the 
mesonotal  axilla;  a  term  proposed  by  Sha- 
fee (1975).  Parastigmal  spur:  a  posterobas- 
ally  orientated  extension  of  the  parastig- 
mal vein,  bearing  1-3  stout,  bristles. 

Abbreviations  used  include  the  follow- 
ing: ANIC,  Australian  National  Insect  Col- 
lection, Canberra;  BMNH,  British  Museum 
(Natural  History),  London;  QM,  Queens- 
land Museum,  Brisbane;  SAM,  South  Aus- 
tralian Museum,  Adelaide;  USNM,  United 
States  National  Museum  of  Natural  His- 
tory, Washington  DC;  WADA,  Western 
Australian  Department  of  Agriculture, 


66 


Journal  of  Hymenoptera  Research 


South  Perth;  ZMA,  Zoological  Museum, 
Aligarh. 

Euryischomyia  Girault 

Euryischomyia    Girault,    1914:    285. — Girault 

1915a:  178;  Girault  1930:  1;  Shafee  1975:  27; 

Hayat  &  Verma  1980:  35;  Hayat  1983:  87; 

Dahms  1986:  689;  Viggiani  1988:  101;  Shafee 

&  Rizvi  1990:  103. 
Euryisdiomyiia  [sic]  Girault,  1929:  331. 
Paramyiocnema  Girault,  1917:  93. — Hayat  1994: 

85. 

Type  species  Euryischomyia  washingtoni  Gi- 
rault, by  monotypy  and  original  designation. 

The  following  characters,  in  combina- 
tion, are  diagnostic  for  Euryischomyia: 

Head  with  7  pairs  of  long,  stiff  setae 
near  ocelli  and  inner  orbital  margins;  max- 
illary and  labial  palps  2-segmented;  man- 
dibles tridentate  (or,  depending  on  inter- 
pretation, bidentate,  the  wide  dorsal  tooth 
being  apically  concave  and  truncate);  se- 
tose. Antennae:  scape  unexpanded  in  both 
sexes;  1-2  anelli;  funicle  3-segmented;  club 
3-segmented. 

fronotum  with  row  of  long,  stiff  setae, 
3  on  each  side,  posterosublaterally.  Me- 
soscutum  between  notauli  with  2-3  rows 
of  shorter,  finer  setae  anterior  to  row  of  4 
and  row  of  2  longer,  stiffer  setae;  scutel- 
lum  wider  than  long,  with  2  pairs  of  long, 
stout  setae,  otherwise  bare.  Postaxillae 
present,  bare.  Phragma  evenlv  tapered, 
apically  incised.  Metanotum  not  produced 
posteromedially.  Propodeum  wide,  not 
excavated  posterolaterally. 

Fore  wings  hyaline  or  medially  infus- 
cated;  parastigmal  spur  little-  or  well- 
developed,  with  1-2,  rarely  3,  long,  bris- 
tle-like setae  of  similar  size  to  those  on 
submarginal  vein;  0-2  bristle-like  setae 
posterobasal  to  parastigma,  or  area  pos- 
terior to  submarginal  vein  and  proximal 
of  parastigmal  spur  with  6-13  shorter, 
stiff  setae  in  2-3  irregular  rows;  this  area 
of  disc  otherwise  bare;  discal  setation 
proximally  regularly  or  irregularly  mar- 
gined. 


Legs:  hind  coxa  enlarged,  elongate- 
ovate,  bilaterally  compressed  ventrally; 
hind  femur  elongate,  broad,  laterally  com- 
pressed, laminose  and  arcuate  in  section; 
fore  and  mid  femora  also  broad  and  lat- 
erally compressed.  Only  one  hind  tibial 
spur.  Long,  bristle-like  setae  on  legs  most 
commonly  disposed  in  differently  sized 
pairs  (as  subapically  on  coxae)  or  in  lon- 
gitudinal rows  (as  on  mid  and  hind  tibi- 
ae). 

Petiole  wide,  boomerang-shaped.  Bases 
of  cerci  low,  tuberculate. 

Notes. — Euryischomyia  is  most  closely  re- 
lated to  (the  monotypic)  Myiocnema,  which 
differs  in  possessing  2  hind  tibial  spurs,  3- 
segmented  maxillary  palps,  3  anelli,  and  a 
distinctive  fore  wing  chaetotaxy;  the  costal 
cell  bears  numerous  short,  stiff  setae,  and 
the  basal  discal  area  bears  both  a  very 
long  bristle-like  seta  posterior  to  the  par- 
astigmal spur  and  several  irregular  rows 
of  shorter,  stiff  setae  in  the  angle  between 
the  submarginal  vein  and  the  parastigmal 
spur.  The  chaetotactic  differences  may  be 
of  specific  value  onlv.  Studies  in  progress 
by  J.B.  Woolley  and  M.  Hayat  show  that 
Myiocnema  differs  significantly  from  Eu- 
ryischomyia in  not  possessing  a  looped  me- 
sofurca  (Hayat  1994). 

Euryischia  is  very  distinctive  in  possess- 
ing almost  circular,  disc-like  hind  coxae,  a 
posterolaterally  excavated  propodeum  to 
accommodate  the  coxae,  and  a  postero- 
medial metanotal  process  ('elasmid'  char- 
acteristics). Euryischia  is  further  distin- 
guishable from  Euryischomyia  in  possess- 
ing 2  hind  tibial  spurs,  4-segmented  max- 
illary palps,  and  cylindrical  cereal  bases. 
The  scape  of  the  male  of  some  species  of 
Euryischia  is  greatly  enlarged,  and  the 
chaetotaxy  of  the  mesonotum  and  fore 
wing  of  Euryischia  is  interspecifically  vari- 
able. Hayat  and  Verma  (1980)  provided 
keys  to  the  genera. 

Taxa  of  Euryischiini  clearly  demonstrate 
different  degrees  of  development  of  eria- 
porine  and  euryischiine  features,  namely, 
development  of  the  parastigmal  spur;  spi- 


Volume  4,  1995 


67 


nosity  of  the  legs,  fore  wing  and  body;  en-  thorax,  Myiocnema  comperei  Ashmead  and 
largement  and  compression  of  the  hind  Euryischia  spp.  are  successively  more  en- 
legs  and  concomitant  modifications  of  the  dowed  in  these  respects, 
mesosoma.  Euryischia  washingtoni,  E.  flavi- 


KEY  TO  SPECIES  OF  EURYISCHOMYIA  GIRAULT 

1.  Parastigmal  spur  well-developed,  bearing  2,  rarely  3,  long,  bristle-like  setae.  Area  imme- 
diately posterior  to  submarginal  vein  and  proximal  to  parastigmal  spur  with  2-3  irregular 
rows  of  6-13  relatively  small,  stiff  setae.  Subapical  margin  of  costal  cell  with  4-7  short, 
stiff  setae  anterior  to  cluster  of  fine,  ventral  setae;  otherwise  bare.  Fore  wings  hyaline,  not 
narrow.  Mesoscutum  with  11-21  setae  in  2-3  irregular  rows  anterior  to  row  of  4  and  row 
of  2  bristle-like  setae.  Body  coloration  yellow,  orange,  brown  and  black.  Distribution: 
Australia £.  flavithorax  Girault 

-  Parastigmal  spur  scarcely  developed,  bearing  1-2  bristle-like  setae.  Area  posterobasal  to 
parastigma  bearing  0-2  large,  bristle-like  setae.  Costal  cell  ventro-subapically  with  some 
pale,  fine  setae;  otherwise  bare.  Forewings  narrow,  3.5  times  longer  than  width;  medially 
with  broad  band  of  infuscation.  Mesoscutum  with  only  8  setae  anterior  to  row  of  4  and 
row  of  2  setae.  Body  coloration  predominantly  black,  legs  predominantly  lemon-yellow. 
Distribution:  Australia  and  India E.  washingtoni  Girault 


Euryischomyia  washingtoni  Girault 

Euryischomyia  washingtoni  Girault,  1914:  285. — 
Girault  1915a:  178;  Hayat  &  Verma  1980:  37; 
Dahms  1986:  649. 

Euryischomyia  alami  Shafee,  1975:  27. — Hayat  & 
Verma  1980:  37.  Syn.  nov. 

Type  Material  Examined. — Euryischomyia 
washingtoni  Girault:  Lectotype  9,  paralec- 
totype  6  (both  on  1  slide,  also  parts  of  ho- 
lotype  of  Pleurotropomyia  seditiosus  Gi- 
rault; all  separate);  paralectotype  9,  (on 
tag,  minus  head).  (QM  4060;  [Hy]2742). 
QUEENSLAND:  20°32'S  145°24'E  Cape- 
ville  (Pentland),  by  sweeping  in  forest 
along  the  banks  of  Cape  River,  8. i. 1913 
(Girault  1914). 

Other  Specimen  Examined;  Identified  by 
A. A.  Girault. — "Euryiscliomyiia  [sic]  wash- 
ingtoni" 9  (on  tag  and  on  slide;  slide  also 
contains  a  paratype  of  Amicromelus  gran- 
daevus  Girault).  QUEENSLAND:  17°23'S 
145°19'E  Watsonville  (QM). 

Other  Records. — Euryischomyia  washing- 
toni: QUEENSLAND:  19°15'S  146°48'E 
Townsville,  sweeping  in  forest,  19. i. 1913, 
9  (Girault  1915a). 


Euryischomyia  alami:  ex  Nipaecoccus  viri- 
dis  (Newstead)  on  Mangifera  indica,  MY- 
SORE: Bangalore,  Channapatna,  2.xii.l968, 
holotype  9,  paratype  9,  S.A.  Shafee  (Sha- 
fee 1975)  (ZMA). 

Euryischomyia  alami:  ex  Nipaecoccus  viri- 
dis,  TAMIL  NADU:  Shencottah,  2  9  9,  M. 
Hayat  (Hayat  and  Verma  1980)  (ZMA). 

Redescription. — Coloration  of  9 :  Body 
shiny,  blackish  brown  to  black;  mesono- 
tum  obscurely  dark  green  metallic.  Body 
setae  mostly  pale.  Legs  whitish  except  for 
following:  fore  coxae  dusky  basally;  mid 
and  hind  coxae  black  except  apically;  mid 
and  hind  femora  black  except  apically  and 
basally;  tarsi  brown  apically;  sockets  of 
hind  tibial  bristles  black.  Fore  wings  each 
with  broad  band  of  pale  to  dark  infusca- 
tion extending  from  anterior  to  posterior 
margins  in  region  of  marginal  and  post- 
marginal  veins,  otherwise  hyaline. 

Morphology:  Length  mesosoma  +  me- 
tasoma:  0.90  mm  (paralectotype  9);  1.05 
mm  (lectotype  9);  1.40  mm  (9  t'.v  Watson- 
ville); 0.84  mm  (paralectotype  6).  All  bris- 
tle-like setae  smaller  and  finer  than  cor- 
responding ones  of  £.  flavithorax. 


68 


Journal  of  Hymenoptera  Research 


Head:  submedian  frontal  setae  (of  6): 
(48|JLm);  interocellar  setae:  (68)  ixm.  Max- 
illary and  labial  palps  longer  and  more 
slender  than  in  E.  flavithorax.  Dorsalmost 
tooth  of  mandibles  pointed. 

Antennae  of  9  ex  Watsonville  (and  S): 
antennal  formula:  1:1:1-2:3:3;  relatively 
longer  and  more  slender  than  in  E.  flavi- 
thorax. Scape:  length  5.56  (4.44)  X  maximal 
width;  1.82  (1.73)  X  length  of  pedicel;  1.22 
(1.11)  X  length  of  club.  Pedicel:  2.5  (2.28) 
X  maximal  width.  Anellus  small,  wedge- 
shaped;  a  2nd  anellus  may  be  partly  de- 
limited from  it.  Funicular  segments  mostly 
as  long  as  or  longer  than  wide;  segment  1 
(fl):  length,  1.18  (1.0)  X  width;  f2:  length, 
1.41  (0.94)  X  width;  f3:  length  1.04  (1.14) 
X  width.  Club:  length:  2.56  (3.56)  X  width. 
Lengths  of  segments:  scape,  150  (107)  |xm; 
pedicel,  83  (62)  fxm;  fl:  30  (20)  fxm;  f2:  47 
(21)  |xm;  f3:  41  (30)|jLm;  club:  123  (96)  jim. 

Mesosoma  of  lectotype  9  (and  6):  Pro- 
notum:  6  bristle-like  setae  postero-subla- 
terally,  68  (75)  fxm  long.  Mesoscutum  be- 
tween notauli  with  row  of  2,  and  row  of 
6,  fine,  bristle-like  setae,  33  (26-29)  |xm 
long;  row  of  4  (38)  iJim  long;  and  row  of  2 
thicker,  bristle-like  setae  (51)  ixm  long.  2 
midsublateral  and  2  posterolateral  scutel- 
lar  setae,  (45  ixm)  and  90  (93)  fxm  long, 
respectively.  Axilla  with  2  setae;  posterior 
seta  (34)  ixm  long. 

Legs  of  lectotype  9  (and  6):  Hind  coxa, 
0.21  (0.22)  mm  long,  1.87  (1.74)  X  width, 
slightly  shorter  than  hind  femur,  which  is 
shorter  than  hind  tibia;  subapical  setae  of 
hind  coxa:  30  (36)  and  48  (54)  |xm  long; 
hind  femur,  0.23  (0.26)  mm  long,  2.53 
(2.74)  X  width,  subapical  seta,  26  (30)  |xm; 
hind  tibia,  0.31  mm  long,  gradually  broad- 
ening apicad,  preaxial  row  of  setae  maxi- 
mally 50  |xm  long,  subapical  setae  44  and 
62  fxm  long,  spur  48  jxm  long. 

Fore  wings  of  lectotype  9  (and  6 )  nar- 
row, 3.5  X  width.  Parastigmal  spur  hardly 
developed,  bearing  1-2  bristle-like  setae, 
45  |JLm  long;  1  or  2  bristle-like  setae  pos- 
terobasal of  parastigma,  31  (35)  ixm  and 
(18)  fxm  long,  respectively.  Subapical  area 


of  costal  cell  devoid  of  marginal  setae. 
Discal  setation  with  regular  margin. 

Metasoma  in  dry-mounted  9  9  up- 
turned at  an  angle  of  approximately  60°, 
strongly  tapered  apicad. 

Ovipositor  sheaths  0.18  mm  long,  0.59 
X  hind  tibial  length,  slightly  protrusive 
beyond  apex  of  metasoma,  densely  and 
evenly  furnished  with  short,  bristle-like 
setae. 

Notes. — Based  on  a  knowledge  of  the 
chaetotactic  variability  within  the  tribe,  £. 
alami  is  hereby  placed  in  synonymy  with  E. 
washingtoni,  despite  the  fact  that  the  types 
of  E.  alami  have  not  been  seen  by  the  pres- 
ent author.  £  alami  reportedly  differs  from 
£.  zvashingtoni  in  not  bearing  discal  setae 
posterior  to  the  parastigma,  and  in  possess- 
ing 3  setae  anteriorly  on  the  axilla  (Shafee 
1975;  Hayat  and  Verma  1980).  These  differ- 
ences are  too  small  to  warrant  separation  at 
the  species  level.  Hayat  and  Verma  (1980), 
who  studied  the  above-listed  specimens 
from  Tamil  Nadu,  are  also  of  the  opinion 
that  the  differences  between  the  2  taxa  ap- 
pear to  be  of  doubtful  specific  value. 

The  above  description  of  £.  zuashingtoni 
supplements  those  of  Girault  (1914, 1915a) 
and  Hayat  and  Verma  (1980)  of  £.  wash- 
ingtoni and  that  of  Shafee  (1975)  of  £.  ala- 
mi. Because  of  the  paucity  and  poor  con- 
dition of  the  material  available  for  study, 
a  composite  and  comparative  method  of 
description  is  adopted.  £.  flavithorax,  the 
standard  of  comparison,  is  conventionally 
described  below  in  greater  detail.  The  lem- 
on yellow  coloration  described  by  Girault 
(1914,  1915a)  can  be  presumed  to  have 
faded  with  time  to  white.  The  antenna  of 
the  paralectotype  9  (on  slide)  is  com- 
pressed; measurements  and  ratios  of  the 
9  ex  Watsonville  are  given  instead. 

£.  washingtoni  has  less  developed  eu- 
ryischiine  features  than  the  other  taxa  of 
Euryischiini,  having  a  scarcely  developed 
parastigmal  spur,  and  shorter,  finer  bris- 
tle-like setae  on  the  body,  limbs  and  fore 
wings. 


Volume  4,  1995 


69 


,.J^. 


I  I'tnAniri 


Fig.  1.     Euryischomyia  flavithorax,  male  (X67).  Cleared,  slide-mounted  specimen. 


Biology. — Parasite  of  unknown  status  of 
Nipaecoccus  viridis,  Pseudococcidae. 

Distribution.— AXJSTR ALIA:  Queens- 
land; possibly  also  Beenleigh,  Victoria 
(Viggiani  1988);  INDIA:  Mysore,  Tamil 
Nadu. 

Euryischomyia  flavithorax 

Girault  &  Dodd 
(Figure  1) 

Euryischomyia  flavithorax  Girault  &  Dodd,  1915 
in  Girault  1915a:  178.— Dahms  1984:  614. 

Euryischomyia  saintpierrei  Girault,  1915a:  178. — 
Hayat  &  Verma  1980:  37;  Dahms  1986:  495. 
Syn.  nov. 

Myiociicma  iimnuorativentris  Girault,  1915b: 
64.— Dahms  1984:  795.  Syn.  nov. 

Euryiscliomyia  fasciata  Girault,  1916:  212. — Hay- 
at &  Verma  1980:  37;  Dahms  1984:  579.  Syn. 
nov. 

Paramyiocnema  marmorativentris  (Girault). — Gi- 
rault 1917:  93;  Hayat  &  Verma  1980:  35. 

Euryischomi/i[i]a  sctosa  Girault,  1929:  331.— Hay- 
at &  Verma  1980:  37;  Dahms  1986:  522.  Syn. 
nov. 


Paramyiocnema  flavithorax  (Girault  &  Dodd). — 
Hayat  &  Verma  1980:  33. 

Type  Material  Exmuiued. — Eiin/isdioun/ia 
flavithorax  Girault  &  Dodd:  Lectotype  9 
(on  slide);  paralectotype  9  (on  tag;  head 
and  hind  tibia  on  same  slide  as  lectotype, 
herein  called  paralectotype  1);  paralecto- 
type 9  (also  on  same  slide  as  lectotype, 
herein  called  paralectotype  2).  QUEENS- 
LAND: 17°05'S  145°47'E  "Gordonvale 
(Cairns)"  (QM  3903;  Hy2744). 

Euryischomyia  saintpierrei  Girault:  Holo- 
type  9  (torso  and  1  fore  wing  on  tag,  fore 
wing  separate,  not  listed  by  Dahms  (1986); 
head,  funicular  segments  2  and  3  and 
club,  both  hind  legs  on  slide)  (QM  3905; 
Hy2743).  QUEENSLAND:  27°28'S  153° 
02'E  Brisbane  (Girault  1915a). 

Myiocnenia  marmorativentris  Girault:  Ho- 
lotype  9  (on  slide)  QUEENSLAND: 
17°05'S  145°47'E  "Gordonvale  (Cairns)" 
(QM  3907;  Hy2959). 

Euryischomyia  fasciata  Girault:  Holotype 


70 


Journal  of  Hymenoptera  Research 


6  (not  ?,  as  described  by  Girault;  on  tag 
minus  antennae,  right  wings  and  leg 
parts;  antenna  minus  scape,  fore  wing, 
hind  femur  and  hind  tibia  on  slide;  anten- 
na not  listed  by  Dahms  (1984)).  WESTERN 
AUSTRALIA:  31°57'S  115°51'E  "Perth,  W. 
Austr.,  G.  Compere,  Collector,  837" 
(USNM  19679). 

Euryischomyiia  [sic]  setosa  Girault:  Holo- 
type  9  (on  tag,  minus  head,  with  1  fore 
wing  on  slide)  SOUTH  AUSTRALIA: 
32°49'S  138°11'E  "Melrose,  Oct.,  A.M. 
Lea"  (SAM). 

Other  Specimens  Examined:  identified,  as 
follows,  by  A.A.  Girault. — "Myiocnema  flav- 
ithorax  Gir.  [sic],  brown  scale  [Coccus  hes- 
peridum]  parasites,  [WESTERN  AUSTRA- 
LIA:] 9,  [2]6S,  coll.  L.J.  Newman,  70". 
(On  1  slide.  Host  identified  as  Coccus  hes- 
peridum  in  F.  Wilson  (I960)).  (WADA). 

Euryischomyiia  [sic]  flavithorax  Gir.  [sic], 
9 .  Host  Lecanidae  [Coccidae]  sp.  Ent.  Div. 
Dep.  Ag.  &  Stk  Qld."  (QM  Hy  10). 

"Myiocnema  saintpierrei  Gir.,  S ,  9 .  Host 
Saissetia  sp.,  Sydney  [33°53'S  151°12'E], 
N.S.W.,  June  19-31,  S.E.  Flanders.  Univ. 
Cal.  Citrus  Exp.  Sta.  Ace.  No.  27)."  (On 
slide;  only  2  of  6  specimens  easily  observ- 
able). (QM). 

"Myiocnema  setosa  Gir.,  9,  Myiocnema 
saintpierrei  Gir.,  6 ,  9,  on  lime  scale  [Coc- 
cidae?], [QUEENSLAND:  26°45'S  150°- 
38'E]  Chinchilla,  Aug.  14,  1931,  S.E.  Flan- 
ders". (On  slide;  only  29  9,  26  S  easily 
observable)  (QM). 

Other  Specimens  Examined. — QUEENS- 
LAND: 15°39'S  144°31'E  Split  Rock,  28.v.- 
26.vi.1993,  9,  P.  Zborowski,  I.D.  Nau- 
mann  (ANIC). 

WESTERN  AUSTRALIA:  33°45'S  122° 
32 'E  Condingup,  55km  E  of  Esperance, 
30.xii.l986,  6,  J.S.  Noyes  (BMNH). 

ex  Psyllaephagus  sp.  /  Cardiaspina  albitex- 
tura  Taylor  on  Eucalyptus  blakelyi,  VICTO- 
RIA: nr  36°21'S  146°19'E  Wangaratta, 
ll.iLl955,  39  9,76  6,  E.  Lewis  (ANIC). 

ex  Cardiaspina  sp.  on  Eucalyptus  blakelyi, 
AUSTRALIAN  CAPITAL  TERRITORY: 
35°18'S    149°08'E    Canberra    (site    15a), 


3.ii.l954,  2  9  9,  A.  Magassy.  NEW  SOUTH 
WALES:  35°50'S  147°15'E  Woomargana, 
3.xii.l954,  9,  E.  Lewis  (all  in  ANIC). 

ex  Creiis  sp.,  WESTERN  AUSTRALIA: 
31°59'S  115°52'E  Western  Australian  Insti- 
tute of  Technology,  South  Perth,  1982,  9, 
S.J.  Curry  (ANIC). 

ex  Aphidius  colemani  Viereck  /  Br  achy - 
caudus  helichrysi  (Kaltenbach)  on  safflow- 
er,  NEW  SOUTH  WALES:  29°14'S 
149°51'E  25km  N.  of  Moree,  12.ix.l993, 
3  9  9,  Id,  J.  Stanley  (ANIC). 

ex  Aphidius  colemani  /  Myzus  persicae 
(Sulzer)  on  nectarine,  AUSTRALIAN 
CAPITAL  TERRITORY:  35°15'S  149°05'E 
Aranda,  Canberra,  15.xi.l991,  29  9,  P.J. 
Hart  (ANIC). 

ex  Trioxys  complanatus  Quilis  /  Therioa- 
phis  trifolii  (Monell)  f.  maculata  on  lucerne, 
VICTORIA:  36°25'S  145°56'E  Goorambat, 

1979,  9,  P.  Ridland;  36°33'S  145°59'E  Ben- 
alla,  9.ii.l978,  9,  26  6,  L.T.  Woolcock; 
SOUTH  AUSTRALIA:  34°40'S  138°34'E 
Virginia,  21.ii.l980,  6,  C.  Wilson;  34°56'S 
138°36'E  Northfield,  Adelaide,  20.iii.l978, 
8.ii.l979,  49  9,  36  6,  L.T.  Woolcock; 
35°41'S  139°20'E  Meningie,  27.ii.1979,  6, 
C.  Wilson  (all  in  ANIC). 

ex  Aphelinus  mali  (Haldeman)  /  Eriosoma 
lanigerum  (Hausmann)  on  apple,  NEW 
SOUTH  WALES:  30°31'S  151°40'E  Armi- 
dale,  December  1992,  6  9  9,  S.  Asante 
(ANIC). 

ex  Pseudococcus  sp.,  on  Acacia  melanoxy- 
lon,  VICTORIA:  37°42'S  145°04'E  LaTrobe 
University,  Bundoora,  Melbourne,  l.xii.- 

1980,  29  9,26  6,  G.  Farrell  (BMNH). 
Other  records.— QUEENSLAND:  15°28'S 

145°15'E  Cooktown,  in  forest,  2.iii.l914,  9, 
A.P.  Dodd  (Girault  1915a;  Dahms  1984). 

WESTERN  AUSTRALIA:  27°41'S  114°- 
33 'E  Kalbarri  National  Park,  December 
1986,  9,  J.S.  Noyes;  31°36'S  116°13'E  Avon 
Valley  National  Park,  50km  NE  of  Perth, 
25.xii.1986,  9,  J.S.  Noyes;  31°43'S  116°04'E 
Walyunga  National  Park,  40km  NE  of 
Perth,  24.xii.1986,  6,  J.S.  Noyes  (BMNH) 
(Identified  by  L.D.  Coote). 

ex  Ceroplastes  rubens  Maskell  on  Schef- 


Volume  4,  1995  71 

flera  actinophi/lla,  QUEENSLAND:  27°30'S  Morphology:  Length  mesosoma  +  me- 

153°01'E   Brisbane,  May   1980,  46  6,  J.S.  tasoma,  0.7-L3  mm  long  (mean,  LOl  mm; 

Noyes  (BMNH)  (Recorded  as  Paramyioc-  n  =  2Q;   lectotype,    L06   mm).    Integument 

nema  sp.  by  Viggiani  (1988)).  variously  reticulate-imbricate,  except  that 

Redescription  of  Female. — Coloration:  of  most  of  metanotum  and  propodeum. 
Most  body  setae  dark.  Head  yellow,  or-  which  is  smooth;  imbrication  on  tibiae 
ange  or  tawny  except  for  following:  finely  setose  at  interstices.  Large  setae  Ion- 
brown,   linear  macula  suborbitally  on  gitudinally  ridged. 

gena;  small,  brown  macula  surrounding  Head:  2  pairs  of  large,  stiff,  bristle-like 

posterior  half  of  median  ocellus  and  an-  setae  on  frons  near  margins  of  eyes;  1  sub- 

terior  half  of  each  lateral  ocellus;  exten-  median  pair  between  these  (lOOixm  long  in 

sive,  broad,  brown  to  black,  inverted  U-  lectotype);  1  pair  lateral  to  median  ocellus; 

shaped  macula  in  region  adjacent  to  pro-  1  pair  near  margin  of  eyes  anterolateral  to 

thorax;  mandibles  apically  testaceous.  An-  lateral  ocelli;  1  pair  between  lateral  ocelli 

tennae  yellow,  scape  may  be  dusky.  Eyes  (113  ixm);  1  pair  posteromedial  to  lateral 

red;  margin  and,  internally,  medial  half  of  ocelli.  Dorsalmost  tooth  of  mandible  usu- 

eye  socket  darkly  sclerotized  (conspicuous  ally  slightly  blunt, 

only  in  cleared  specimens).  Ocelli  red.  Antennae  inserted  not  far  distant  from 

Mesosoma,  orange  or  tawny  except  for  clypeal  margin,  the  distance  subequal  to 

following   dark   sclerotization:   extensive,  distance  between  insertions;  setae  short, 

brown  to  black  macula  on  prothorax  ad-  with  stouter,  stiffer  hairs  concentrated  on 

jacent  to  occipital  region  of  head;  notauli  one    side;    antennal    formula:    1:1:1-2:3:3. 

anteriorly,  boundaries  of  anterior  angles  Scape:  length,  2.9-3.4  X  width  (mean,  3.2); 

of  axillae,  anterior  and  posterior  bound-  1.7-2.3  (2)  x  length  of  pedicel;  1-1.2  (1.1) 

aries  of  pronotum,  scutellum  and  meta-  X  length  of  club.  Pedicel:  length,  1.3-2.5 

notum  mav  be  brown;  1-2  small,  brown  (1.8)    X   width;  0.6-1   (0.75)    X   length  of 

to  black  maculae  near  base  of  fore  wing;  anellus(i)    +    funicle.    Anellus    1    large 

sometimes  1  macula  at  base  of  posterolat-  enough  to  be  considered  funicular,  finely 

eral  setae  of  scutellum  and  submarginally  setose  but  non-sensoriated;  a  2nd,  very 

on  metanotum;  propodeum  light  to  dark  small,  anellus  sometimes  variouslv  devei- 

brown  anteriorly  and  on  each  lateral  third  oped  on  half  of  apex  of  anellus  1,  mav  be 

except  for  spiracles,  lateral  margins  may  setose.  Funicular  segments  all  wider  than 

be    darker,    or    whole    segment    may    be  long,  becoming  progressivelv  wider  and 

brown  and  laterally  black.  Ventrally  and  longer  apicad;  segment  1  (fl)  length,  0.58- 

sometimes   pleurally,   mesosoma   light  0.87  (0.7)   X   width;  f2:  length,  0.47-0.82 

brown  to  dark  brown.  (0.74)  X  width;  f3:  length,  0.64-0.9  (0.79) 

Legs  yellow  except  for  following:  fore  X  width;  length  of  anellus  (i)  +  funicle, 

femur  with  linear  macula  postaxially;  mid  0.6-0.9  (0.7)    x    club.   Club  3-segmented; 

coxa  may  be  slightly  dusky  basally;  hind  segments   separable   by    segmentally   ar- 

coxa  brown  to  dark  brown  except  usually  ranged  setal  and  sensorial  patterns,  not  bv 

apically,  sometimes  also  dark  brown  mar-  sutures;  length,  1.5-2.7  (2)  X  width;  width 

ginally;  hind  femur  sometimes  streakily  1-1.8  X  width  of  f3.  Lengths  of  segments: 

dusky;  sockets  of  hind  tibial  bristles  black,  scape,  99-135  fim  (mean,  117  |xm;  n  =  20); 

Metasoma  light  brown  to  black;  petiole  pedicel,  45-68  (58)   |xm;  fl,   14-24  (17.5) 

usually  brown,  often  black  on  lateral  mar-  |xm;  f2,  12-37  (24)  |xm;  f3,  24^1  (31)  |jim; 

gins;  base  of  gaster  medially  and  gastral  club,  93-123  (107)  |xm. 

tergites  2-4  usually  darker  than  rest  of  Mesosoma:  Pronotum  with  row  of  3+3 

gaster,  and  fasciate,  especially  under  high  long  setae  (115  ixm  long  in  lectotype)  pos- 

magnification.  terosublaterally.    Mesoscutum    anteriorly. 


72 


Journal  of  Hymenoptera  Research 


between  notauli,  with  11-21  fine,  stiff  se- 
tae (41  |jLm)  in  2-3  irregular  rows  ;  a  mid- 
dle transverse  row  of  4  long,  stout  setae 
(98  fxm),  and  posterior  row  of  2  long,  stout 
setae  (90  |xm).  Scutellum  with  2  long  setae 
anterosublateral  of  midpoint  (87  ixm)  and 

2  very  long,  stout  setae  posterolaterally 
(168  |xm).  Axillae  each  with  long,  stout 
seta  (84  |xm)  anteriorly  and  2-3,  rarely  4, 
shorter,  differently  sized  setae  (26-36  ixm) 
near  posterior  angle.  Mesosomal  setae  of 
smaller  specimens  shorter  and  less  stout. 

Legs  bearing  long,  stiff  setae,  mostly  in 
rows  or  in  differently  sized  pairs;  other  se- 
tae mostly  disposed  dorsoapically  on  fe- 
mur and  longitudinally  on  tibia;  Fore  coxa 
(of  1  Northfield  specimen):  pair  of  long, 
stiff  setae  subapically  (39  and  57  ixm  long); 
fore  femur:  subapically,  1  seta  ventrally 
(29|jLm),  1  dorsally  (45  lam);  fore  tibia  ap- 
pearing fusiform,  calcar  plumose  (57  ixm), 

3  small,  stiff  setae  apically  (Slfxm),  and 
row  of  7  setae  (maximally  31  ixm)  preaxi- 
ally.  Mid  coxa:  pair  of  setae  subapically 
(42  and  57  lam);  mid  femur:  very  long  pair 
subapically  (65  (xm  ventrally,  78  |xm  dor- 
sally);  mid  tibia:  spur  long  (87  |xm),  plu- 
mose, 1  smaller  seta  near  spur  (18  |xm),  1 
subapically  (53  lam),  preaxial  row  (maxi- 
mally 32  (xm)  hardly  differentiated  from 
other  setal  rows.  Hind  coxa  (0.22  mm 
long),  shorter  than  hind  femur  (0.28  mm), 
which  is  slightly  shorter  than  hind  tibia 
(0.32  mm)  and  subequal  to  hind  tarsus 
(0.27  mm);  hind  coxa  large,  elongate- 
ovate,  length  1.44-1.69  X  width,  bilater- 
ally compressed  ventrally,  pair  of  setae 
subapically  (44  and  66  ixm);  hind  femur 
broad,  approximately  parallel-sided, 
length  2.83-3.14  X  width,  1  seta  subapi- 
cally (58  |JLm);  hind  tibia,  basally  narrow, 
gradually  and  slightly  widening  apicad, 
spur  almost  smooth  (54  |xm  long),  pair  of 
very  long,  stout  setae  subapically  (68  and 
89  |jLm)  and  1  longitudinal  row  of  8  very 
long,  stout  setae  (maximally  75  iJim).  Mid 
basitarsus  longer  than  fore  or  mid  basitar- 
sus,  but  not  conspicuously  long. 

Fore  wings:  6-13  (mean,  10)  moderately 


long  and  fine,  stiff  setae  (30^0  |xm  long 
in  1  Northfield  specimen)  present  in  2-3 
rows  below  submarginal  vein  and  proxi- 
mal of  parastigmal  spur;  2-3  parastigmal 
setae  (70-76  |xm);  costal  cell  in  distal  third 
with  4-7  short,  stiff,  marginal  setae  (25 
fxm)  anterior  to  cluster  of  fine,  stiff,  ventral 
setae;  margin  of  discal  setation  irregular. 

Metasoma  not  upturned  or  strongly  ta- 
pered: Petiole  boomerang-shaped  except 
for  truncated  ends.  Variously  sized,  stiff 
setae  metamerically  disposed  laterad  and 
apicad  on  dorsum  of  gaster. 

Ovipositor  sheaths  0.10-0.11  mm  long, 
0.23-0.33  X  hind  tibial  length,  each  fur- 
nished with  about  10  mostly  subapical  se- 
tae. 

Description  of  Male. — Similar  to  female. 
Colour  pattern  similar  but  yellower  be- 
cause of  less  extensive  dark  sclerotization. 
Linear  macula  usually  present,  anterome- 
dially,  on  vertex;  suborbital,  linear  macula 
of  gena  absent  and  U-shaped  macula  of 
occipital  region  absent.  Mesosoma  later- 
ally pale.  Gaster:  dark  sclerotization  con- 
fined to  narrow  area  adjacent  to  petiole 
and  to  medially  coalesced  fascia  on  gastral 
tergites  2-4,  extending  medially  on  gastral 
tergite  5  and  apicolaterad. 

Antennae  (1  specimen):  scape:  length, 
111  |xm,  3.7  X  width,  2.18  X  length  of  ped- 
icel, 1.19  X  length  of  club;  pedicel:  length, 
51|xm,  1.7  X  width,  0.71  X  length  of  anel- 
lus  +  funicle;  funicular  segment  1  (fl): 
length,  12|jLm,  0.47  X  width;  f2:  length, 
21|jLm,  0.66  X  width;  f3:  length,  27  |am, 
0.66  X  width;  anellus  +  funicle:  length, 
0.77  X  length  of  club;  club:  length,  0.93 
|jLm,  1.32  X  width. 

Hind  coxa  0.24  mm  long,  twice  as  long 
as  wide;  hind  femur  0.24  mm  long,  3.2  X 
width;  hind  tibia  0.3  mm  long. 

Mesosoma  +  metasoma  0.9  mm  long. 
Anterior  mesoscutal  setae,  33^5  juim  long; 
mid  and  posterior  setae,  70  and  81  ixm  long 
respectively.  Anterior  and  posterior  scutel- 
lar  setae  64  and  121  \xm,  respectively;  only 
1  posterior  axillar  seta,  24  |jLm  long. 

Notes. — The  type  material  of  £.  flavitlior- 


Volume  4,  1995  73 

ax  comprises  the  slide-mounted  lectotype  as  possessing  2  anelli,  which  may  have 

designated  by  Hayat  in  1979  (Hayat  and  been  the  reason  why  he  considered  it  a 

Verma  1980),  and  paralectotype  1,  with  species  distinct  from  the  other  species  of 

parts  on  a  tag  and  parts  on  the  same  slide  Euryischomyia,  and  why  he  identified  spec- 

as  the  lectotype  but  mounted  separately,  imens  from  'lime  scale'  as  2  species,  M. 

inbetween  the  lectotype  and  paralectotype  saintpierrei  and  M.  setosa  (see  above  list  of 

2  (see  list  of  type  material  examined).  Par-  specimens   identified   by   Girault).   These 

alectotype  2   is   the  specimen   stated  by  specimens,  on  1  slide,  have  antennae  each 

Hayat  and  Verma  (1980)  to  be  a  species  with  either  1  or  2  anelli  but  are  otherwise 

different  from  the  lectotype,  resembling  similar  to  one  another.  Most  of  the  speci- 

marmorativentris,  and  probably  not  part  of  mens  of  £.  flavithorax  examined   in   this 

the    original    type    material.    Consequent  study  have  both  antennae  with  1  anellus; 

upon  the  above  statements,  the  specimen  others  have  2;  or  1  antenna  may  bear  1 

was  referred  to  by  Dahms  (1984)  as  an  un-  anellus  and  the  other  bear  2;  or  1  small 

identified  chalcidoid.  Paralectotype  2  is,  anellus  may  be  partly  delimited  from  a 

without  doubt,   Euri/ischoniyia  flavithorax.  larger  one. 

Smaller  and  yellower  than  the  lectotype,  it  Distribution. — AUSTRALIA:  Queens- 
does  indeed  resemble  the  holotype  Myioc-  land.  New  South  Wales,  Australian  Capi- 
nema  marmorativentris,  which  was  collect-  tal  Territory,  Victoria,  South  Australia, 
ed  in  the  same  locality  a  week  or  so  later.  Western  Australia  (Figure  2). 

The  female  collected  by  A. P.  Dodd  at  Biology. — E.   flavithorax    is    not    highly 

Cooktown  has  still  not  been  located  (see  host-specific,  its  recorded  hosts  being  chal- 

list  of  other  records  above).  It  may  be  part  cidoid  and  ichneumonoid  Hymenoptera, 

of  Dodd's  collection.  and  psylloid,  aphidoid  and  coccoid  He- 

£.  flavithorax  was  originally  described  miptera  Sternorrhyncha  (Table  1).  E.  flav- 
from  3  swept  specimens,  and  each  of  the  ithorax  was  reared  occasionally  from  im- 
synonymized  species  was  described  from  mature  Aphidiinae  and  Aphclimis  mali,  en- 
single  specimens.  The  relevant  parts  of  the  doparasitic  in  aphidine  and  drepanosiphi- 
descriptions  are  based  mainly  on  colora-  ne,  and  pemphigine  aphids,  respectively, 
tion.  Other  characters  described  by  Girault  as  part  of  a  long-term  study  by  the  author 
are  supra-specific  or  are  not  interspecifi-  of  aphids  and  their  insect  natural  enemies; 
cally  comparable.  However,  all  of  them  fit  and,  also  occasionally,  from  Psyllacphagus 
into  the  above  conception  of  £.  flavithorax.  sp(p).    parasitic    in    lerp-forming    psyllid 

£.  flavithorax  is  variable  in  coloration.  In  nymphs,  as  part  of  an  ecological  study  by 

general,  smaller  females  are  yellower  and  L.R.  Clark  in  1950-74  of  psyllids  associat- 

less  darkly  sclerotized  than  larger  ones,  ed  with  eucalypts  (Riek  1962). 

and  males  are  smaller  and  yellower  than  £.  flavithorax,  however,  is  not  a  true  hy- 

females.  The  differences  are  a  reflection  of  perparasite  of  aphids  if,  like  certain  other 

differences  in  extent  of  dark  sclerotization.  chalcidoids  and  proctotrupc^ids  {e.g.  Pachy- 

The  pattern  of  coloration,  however,  is  cc:)n-  neuron  spp.  (Pteromalidae)  and  Dcndroce- 

stant,  and  is  best  expressed  in  the  lighter-  rus  spp.  (Megaspilidae)  (Takada  1973a,b)), 

coloured  specimens.  The  lectotype  of  £.  it  does  not  parasitize  the  hymenopterous 

flavithorax,  for  instance,  belongs  near  the  host  until  after  the  latter  has  consumed  all 

large,  dark  end  of  the  scale  of  variability,  of  its  aphid  host  except  for  the  dead,  dry 

whereas  paralectotype  2  of  £.  flavithorax  aphid  skin  (mummy);  i.e.  if  it  parasitizes 

and  the  holotype  M.  marmorativentris  be-  the  hymenopterous  contents  of  a  capsule 

long  near  the  small,  pale  end.  The  small,  it  is  strictly  a  primary  parasite.  Such  a 

pale  holotype  £.  fasciata  is  a  male.  probability  is  supported  by  its  broad  host 

Girault  (1913a)  described  £.  saintpierrei  spectrum;  parasitizing  live  hosts  of  such 


74 


Journal  of  Hymenoptera  Research 


■     y 

c 

rt 

-<j 

^-N^ 

■           1 

■ 

'IT'   / 

■  / 

V 

Fig.  2.     Known  distribution  of  Euryischomi/ia  flavithomx. 


Table  1.    Hosts  of  Euryischomyia  flavithorax 

Hemiptera:  Sternorrhyncha 

Psylloidea:  Psyllidae: 

Spondyliaspidinae:  Cardiaspina  albitextum  

Cardiaspina  sp. 
Creiis  sp. 
Aphidoidea:  Aphididae: 

Aphidinae:  Brachycaudus  helichrysi 

Myzus  persicae 

Drepanosiphinae:  Therioaphis  trifolii  f.  maculata 

Pemphiginae:  Eriosoma  lanigerum  

Coccoidea:  Pseudococcidae: 

Pseudococcus  sp. 
Coccoidea:  Coccidae: 

Ceroplastes  rubens 
Coccus  hesperidum 
Saissetia  sp. 


Hymenoptera 


Chalcidoidea:  Encyrtidae: 

Psyllaepliagus  sp. 


Ichneumonoidea:  Braconidae: 
Aphidiinae:  Aphidius  colemani 
Aphidius  colemani 
Trioxys  coniplanatus 
Chalcidoidea:  Aphelinidae: 
Aphelinus  mali 


Volume  4,  1995 


75 


morphological  and  behavioral  diversity 
would  demand  a  high  degree  of  repro- 
ductive adaptability. 

Its  status  as  a  parasite  of  Coccoidea  is 
uncertain.  Female  coccids,  being  sessile 
and  capsule-like,  could  conceivably  func- 
tion as  either  primary  or  secondary  hosts. 

E.  flavithorax  is  exclusively  Australian, 
as  far  as  known.  The  original  hosts  are 
likely  to  be  native  sternorrhynchous  He- 
miptera  and  Hymenoptera,  e.g.  spondy- 
liaspidine  psyllids  and  their  Psi/llncpliagiis 
parasites.  The  recorded  aphid  hosts  and 
their  primary  parasites,  and  the  coccoid 
hosts,  are  all  exotic  species  introduced  into 
Australia. 

E.  flavithorax  was  collected  throughout 
the  year  but  most  commonly  in  summer 
(December  and  February).  £.  flavithorax 
(recorded  as  Myiocnema  sp.)  comprised  2% 
of  the  emergent  parasites  of  Trioxys  coni- 
planatus  /  Therioaphis  trifolii  f.  maculata  in 
several  sites  in  South  Australia  between 
1977  and  1980  (C.G.  Wilson  and  Swincer 
1984). 

Females  were  commoner  than  males,  es- 
pecially among  collected  rather  than 
reared  material. 

ACKNOWLEDGEMENTS 

I  am  indebted  to  the  above-listed  collectors;  to  K.J. 
Lambkin  and  G.B.  Monteith,  Queensland  Museum, 
South  Brisbane;  Jan  Forrest  OAM,  South  Australian 
Museum,  Adelaide;  Frangoise  Berlandier,  Western 
Australian  Department  of  Agriculture,  Perth;  M.E. 
Schauff  and  G.F.  Hevel,  United  States  National  Mu- 
seum, Washington  DC;  and  L.D.  Coote,  Royal  Ontar- 
io Museum,  Toronto,  for  the  loan  of  specimens  in 
their  care;  to  S.O.  Shattuck,  CSIRO  Entomology,  for 
production  of  Figure  2;  and  to  S.O.  Shattuck;  l.D. 
Naumann,  CSIRO  Entomology;  L.D.  Coote;  and  an 
unknown  referee,  for  valuable  comments  on  the 
manuscript. 

LITERATURE  CITED 

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the  Aphelininae.  Canadian  Entomologist  32:  349. 

Compere,  H.  1947.  A  new  genus  and  species,  Eiiry- 
nn/iocnt'iua  aphcUnoides  (Hymenoptera,  Aphelini- 
dae),  and  a  history  of  the  genera  Eun/ifichia  Riley 
and  Myiocnema  Ashmead.  Bulletin  of  Eiitoinologi- 
cal  Research  38:  381-388. 


Dahms,  E.C.  1983.  A  checklist  of  the  types  of  Aus- 
tralian Hymenoptera  described  by  Alexandre  Ar- 
sene  Girault:  II.  Preamble  and  Chalcidoidea  spe- 
cies A-E  with  advisory  notes.  Memoirs  of  the 
Queensland  Museum  21:  1-255. 

Dahms,  E.C.  1984.  A  checklist  of  the  types  of  Aus- 
tralian Hymenoptera  described  by  Alexandre  Ar- 
sene  Girault:  111.  Chalcidoidea  species  F — M  with 
advisory  notes.  Memoirs  of  the  Queensland  Muse- 
um 21:  579-842. 

Dahms,  E.C.  1986.  A  checklist  of  the  types  of  Aus- 
tralian Hymenoptera  described  by  Alexandre  Ar- 
sene  Girault:  IV.  Chalcidoidea  species  N — Z  with 
advisory  notes  plus  addenda  and  corrigenda. 
Memoirs  of  the  Queensland  Museum  22:  319-739. 

Ferriere,  C.  1965.  Hymenoptera  Aphelinidae  d'Eu- 
rope  et  du  Bassin  Mediterraneen.  Faune  de 
I'Europe  et  du  Bassin  Mediterraneen,  1.  206  pp. 
Masson:  Paris. 

Ghesquiere,  J.  1955.  Contribution  a  I'etude  du  genre 
Eriaporus  Waterston  et  genres  affins  (Hym.,  Chal- 
cidoidea, Aphelinidae)  Memoires  de  la  Societe  Roy- 
ale  d'Entomologie  de  Belgique  27:  217-238. 

Girault,  A. A.  1914.  The  third  genus  of  the  family 
Elasmidae  (Hymenoptera).  Canadian  Entomologist 
46:  285-286. 

Girault,  A.A.  1915a.  Australian  Hymenoptera  Chal- 
cidoidea III.  Second  Supplement.  Memoirs  of  the 
Queensland  Museum  3:  170-179.  (Issued  28  Jan. 
1915). 

Girault,  A.A.  1915b.  Australian  Hymenoptera  Chal- 
cidoidea VII.  Metnoirs  of  the  Queensland  Museum 
4:  1-184.  (Issued  4  June  1915). 

Girault,  A.A.  1916.  Australian  Hymenoptera,  Chal- 
cidoidea. General  Supplement.  Memoirs  of  the 
Queensland  Museum  5:  205-230. 

Girault,  A.A.  1917.  New  Australian  chalcid-flies  (Hy- 
menoptera, Chalcidoidea).  Insecutor  Inscitiae 
Menstruus  5:  92-96. 

Girault,  A.A.  1929.  Notes  on,  and  description  of, 
chalcid  wasps  in  the  South  Australian  Museum. 
Concluding  paper.  Transactions  of  the  Royal  Soci- 
ety of  South  Australia  53:  309-346. 

Girault,  A.A.  1930.  New  Pests  from  Australia,  IX.  1  p. 
Girault:  Brisbane.  Pri\'atelv  published  29  Decem- 
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Goulet,  H.  and  Huber  J.T.  (Eds)  1993.  Hymenoptera 
of  the  World:  An  Identification  Guide  to  Fami- 
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Hayat,  M.  1983.  The  genera  of  Aphelinidae  (Hyme- 
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63-102. 

Hayat,  M.  1994.  Notes  on  some  genera  of  the  Aphel- 
inidae (Hymenoptera,  Chalcidoidea),  with  notes 
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Hayat,  M.  and  Verma,  M.   1980.  The  aphelinid  sub- 


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family  Eriaporinae  (Hym.:  Chalcidoidea).  Orien- 
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Naumann,  I.D.  1991.  Hymenoptera.  Pp.  916-1000  in 
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2  volumes.  Melbourne  University  Press:  Carlton. 

Nikols'skaya,  M.N.  1963.  The  Chalcid  Fauna  of  the 
U.S.S.R.  593  pp.  Israel  Program  for  Scientific 
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Riek,  E.F.  1962.  The  Australian  species  of  Psyllaepha- 
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684-757. 

Riley,  C.V.  1889.  Report  of  the  Entomologist.  Report 
of  the  United  States  Department  of  Agriculture  for 
1888,  page  92,  plate  9,  figure  2. 

Shafee,  S.A.  1974.  A  new  family  of  Chalcidoidea  (Ln- 
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Shafee,  S.A.  1975.  A  new  family  of  Chalcidoidea  (In- 
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Shafee,  S.A.  and  Rizvi,  S.  1990.  Classification  and 
phylogeny  of  the  family  Aphelinidae  (Hymenop- 
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(Hymenoptera:  Chalcidoidea).  Indian  Journal  of 
Entomology  2:  27-29. 

Smith,  H.S.  and  Compere,  H.  1928.  A  preliminary 
report  on  the  insect  parasites  of  black  scale,  Sais- 
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Takada,  H.  1973a.  Studies  on  aphid  parasites  of  Ja- 
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1-37. 

Takada,  H.  1973b.  Studies  on  aphid  hyperparasites 
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di  Portici  45:  99-102. 

Wilson,  C.G.  and  Swincer,  D.E.  1984.  Hyperparasi- 
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Aphididae)  in  South  Australia.  Journal  of  the  Aus- 
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Control,  Ottawa,  Canada,  Technical  Communication 
1:  V  +  102. 


J.  HYM.  RES. 
Vol.  4,  1995,  pp.  77-79 


Male  Nest  Defense  in  the  Digger  Wasp  Cerceris  binodis 

(Hymenoptera:  Sphecidae) 

David  Banks 

Snow  Entomological  Museum,  Snow  Hall,  Department  of  Entomology, 
University  of  Kansas,  Lawrence,  KS  66045  USA 


Abstract.— Male  Cerceris  binodis  (Hymenoptera:  Sphecidae:  Philanthinae)  defend  territories  that 
contain  nesting  females  from  conspecific  males.  A  female  which  is  provisioning  a  nest  will  mate 
when  she  returns  to  the  nest  and  a  male  is  present.  This  mating  system  represents  the  first  de- 
scription of  male  nest  defense  in  the  genus  Cerceris.  Presumably,  this  behavior  occurrs  in  C.  binodis 
because  females  nest  in  aggregations,  and  mate  multiply. 


INTRODUCTION 

The  cosmopolitan  genus  Cerceris  con- 
tains over  800  species  (Bohart  and  Menke 
1972),  more  than  any  other  sphecid  genus, 
but  mating  systems  of  relatively  iew  Cer- 
ceris species  have  been  studied.  Non-re- 
source based  territoriality  has  been  re- 
ported in  all  Cerceris  mating  systems 
v^hich  have  been  described  (Alcock  1975a, 
Evans  1971,  Evans  and  O'Neill  1985, 
Evans  and  O'Neill  1988),  and  in  four  other 
genera  in  the  sphecid  subfamily  Philan- 
thinae: Aphilanthops,  Philantliiis,  Cli/f>eadon, 
and  Eiicerceris  (Evans  and  O'Neill  1988; 
O'Neill  1990).  In  this  mating  system, 
males  apply  volatile  secretions  originating 
from  mandibular  glands  to  a  substrate 
with  clypeal  brushes,  and  defend  these 
non-resource  based  territories  from  con- 
specific  males.  Philanthine  females  typi- 
cally nest  in  dispersed  conditions,  mate 
once  at  the  beginning  of  the  nesting  period 
away  from  the  nest,  presumably  are  at- 
tracted to  mandibular  gland  secretions, 
and  enter  territories  to  copulate  (Evans 
and  O'Neill  1988).  The  mating  system  of 
C.  binodis  departs  from  these  conditions. 
Females  nest  in  aggregations,  and  mate 
multiply  at  the  nest.  Resident  C.  binodis 
males  stand  on  or  directly  adjacent  to 
nests,    exclude    conspecific    non-resident 


males  from  nests,  and  do  not  scent  mark 
territories.  Of  the  eight  philanthine  gen- 
era, male  nest  defense  has  been  described 
only  in  Philanthiis,  and  presumably  is  de- 
rived within  that  genus  (Evans  and 
O'Neill  1988).  Male  nest  defense  is  de- 
scribed here  for  the  first  time  in  the  genus 
Cerceris,  and  also  presumably  represents  a 
derived  condition  within  this  taxon. 

METHODS 

An  aggregation  of  nesting  C.  binodis  fe- 
males was  discovered  on  Barbour  Lathrop 
trail  on  Barro  Colorado  Island,  Republic  of 
Panama  on  July  8,  1994.  A  group  of  five 
nests  that  were  close  enough  together  to 
be  observed  simultaneously  were  marked 
with  numbered  tags.  Male  and  female 
wasps  were  individually  marked  with 
enamel  paint  on  the  thorax,  and  head 
widths  were  measured  to  the  nearest  0.05 
mm  with  Manostat  calipers.  The  nests 
were  observed  on  two  days  after  individ- 
uals were  marked.  Observations  were 
made  from  1 355-1 525h  on  the  first  day, 
and  from  1515-1650h  on  the  second  day. 
The  behaviors  recorded  were  the  amount 
of  time  that  males  spent  standing  on  or 
adjacent  tc^  nests,  defensive  interactions 
between  males,  and  copulations. 


78 


Journal  of  Hymenoptera  Research 


RESULTS 

When  a  female  returned  to  a  nest  with 
prey,  and  a  resident  male  was  present,  she 
would  land  on  the  nest  tumulus,  and  cop- 
ulate. However,  a  female  would  fly  di- 
rectly into  her  nest  if  no  male  was  present. 
This  behavior  is  also  described  in  the 
sphecid  genus  Oxybelus  (Crabroninae) 
(Peckham  et.  al.  1973).  Females  were  pro- 
visioning nests  in  all  cases  where  males 
defended  nests  and  mated.  Not  only  did 
males  defend  nests  while  females  were  on 
provisioning  trips,  but  on  two  occassions, 
a  male  was  observed  to  defend  a  nest 
while  the  female  was  inside  the  nest. 
These  males  were  observed  to  bite  at  the 
females,  and  attempted  to  pull  them  out 
of  the  nest  with  their  mandibles.  One  of 
these  males  mated  with  the  female  when 
she  exited  the  nest. 

Nest  defense  duration  was  determined 
by  the  length  of  time  a  resident  male  was 
either  in  contact  with  a  nest,  directly  ad- 
jacent to  a  nest,  or  defending  a  nest  from 
non-resident  males.  On  day  one,  males 
YB,  RB  and  BY,  spent  a  total  of  66.7,  21.1, 
and  12.2  percent  of  the  observation  period 
defending  nests,  respectively.  On  day  two, 
males  WRB,  YB,  RY,  BB  and  RB  spent  a 
total  of  67.3,  41.1,  16.8,  1.2  and  0.8  percent 
of  the  observation  period  defending  nests, 
respectively.  Males  YB  and  WRB  obtained 
71%  (5/7)  of  the  observed  copulations, 
and  there  is  a  positive  correlation  between 
the  duration  of  time  which  males  defend 
nests  and  the  number  of  copulations  that 
males  obtain;  r  =  0.86. 

At  times,  multiple  males  were  simulta- 
neously defending  different  nests  in  the 
aggregation.  Male-male  interactions  were 
considered  a  territorial  defense  when  a 
resident  male  flew  from  his  nest  to  chase 
a  non-resident  male.  These  chases  rarely 
ended  in  grappling  on  the  ground;  other- 
wise, there  was  no  physical  contact.  Four- 
ty  eight  aggressive  interactions  were  re- 
corded between  resident  and  non-resident 
males.  Resident  males  won  significantly 


more  (95%)  interactions  than  non-resident 
males  (Chi-square  =  40.1,  df  =  2,  P< 
0.001).  In  11  of  the  48  interactions,  identity 
and  size  of  both  males  were  known;  resi- 
dent males  won  7  of  these  11  interactions. 
Mean  head  widths  of  winner  and  looser 
males  were  3.83mm  and  3.76mm,  respec- 
tively. The  difference  in  mean  headwidths 
between  winner  and  looser  males  is  not 
statistically  significant  (t-test  =  .82,  df  = 
20,  P  >  0.4). 

DISCUSSION 

Size  has  been  demonstrated  to  deter- 
mine dominance  in  territorial  sphecids 
such  as  Philanthus  (O'Neill  1983a,  b),  and 
in  many  other  insect  mating  systems 
(Thornhill  and  Alcock  1983).  In  C.  binodis, 
however,  territory  ownership,  not  size,  ap- 
parently determines  dominance  (one 
should  note  the  small  sample  size  (n=ll) 
where  identity  of  both  males  was  known). 
This  phenomenon  has  also  been  demon- 
strated in  other  insect  mating  systems  (Da- 
vies  1978,  Eickwort  and  Ginsberg  1980, 
Wang  and  Greenfield  1991).  Data  also  in- 
dicate that  as  the  total  time  invested  in 
nest  defense  per  male  increases,  the  num- 
ber of  copulations  obtained  per  male  also 
increases.  This  is  not  surprising,  since  fe- 
males mate  with  the  resident  male  which 
is  defending  her  nest. 

In  many  Cerceris  species,  females  nest  in 
aggregations,  but  males  do  not  defend 
nests  from  conspecific  males.  The  require- 
ment which  is  likely  responsible  for  the 
difference  between  the  mating  systems  of 
these  species  and  C.  binodis  is  that  C.  bin- 
odis  females  retain  their  receptivity  after 
the  initial  mating.  This  might  maintain 
male  territoriality,  particularly  if  there  is 
last  male  sperm  precedence  (Hook  and 
Matthews  1980,  Brockmann  and  Grafen 
1989).  However,  males  in  several  species 
of  Philanthus  defend  territories  which  con- 
tain nesting  females  that  do  not  display 
continuous  receptivity  (Gwynne  1980, 
Evans  and  O'Neill  1988).  Data  collected 
from  this  study  do  not  indicate  that  fe- 


Volume  4,  1995 


79 


males  are  continuously  receptive  through- 
out the  nesting  period,  only  that  they  mate 
multiply.  In  addition,  last  male  sperm  pre- 
cedence is  not  demonstrated  in  this  spe- 
cies, but  often  occurs  in  insects  (Thornhill 
and  Alcock  1983). 

CONCLUSION 

This  is  the  first  description  of  male  nest 
defense  in  Cerceris.  This  derived  mating 
system  shows  striking  convergence  with 
the  mating  systems  of  both  Oxyhehis 
(Peckham  et.  al  1973,  Hook  and  Matthews 
1980),  and  Philanthus  (Evans  and  O'Neill 
1988)  species.  Moreover,  C.  binodis  males 
do  not  scent  mark  territories,  and  their 
clypeal  brushes  are  reduced.  Since  clypeal 
brushes  are  a  synapomorphy  of  Philanthi- 
nae,  and  are  present  in  most  species  of 
Cerceris  (Alexander  1992),  reduction  of  the 
clypeal  brushes  in  C.  binodis  represents  a 
derived  condition.  Evans  and  O'Neill 
(1988)  state  that  scent  marking  occurs  in 
all  territorial  philanthines  where  the  mat- 
ing systems  have  been  described.  There- 
fore, this  mating  system  represents  an  ex- 
ception to  their  statement  (see  also  Alcock 
and  Gamboa  1975). 

ACKNOWLEDGEMENTS 

Helpful  comments  and  suggestions  were  provided 
by  B.A.  Alexander,  W.  T.  Wcislo,  D.  Windsor,  R.  L. 
Minckley,  and  four  anonymous  reviewers.  This  re- 
search was  funded  by  a  Smithsonian  Tropical  Re- 
search Institute  10  week  graduate  fellowship,  and  by 
a  Sigma  Xi  research  grant. 

LITERATURE  CITED 

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lanthine  wasps(Hymenoptera:  Sphecidae).  Jour- 
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Alcock,  J.  1975b.  Territorial  behavior  by  males  of  Phi- 
lanthus  multimaculatus  (Hymenoptera:  Sphecidae) 
with  a  review  of  territoriality  in  male  Sphecidae. 
Animal  Behavior  23:  889-895. 

Alcock,  J.  and  G.  Gamboa.  1975.  Home  ranges  of 
male  Cerceris  simplex  macrosticta  (Hymenoptera: 
Sphecidae).  Psyche  81:  528-533. 

Alcock,  J.,  C.E.  Jones  and  S.L.  Buchmann.  1977.  Male 
mating  strategies  in  the  bee  Centris  pallida  Fox 
(Hymenoptera:  Athophoridae).  American  Natu- 
ralist 111:  145-155. 


Alexander,  B.A.  1992.  A  cladistic  analysis  of  the  sub- 
family Philanthinae  (Hymenoptera:  Sphecidae). 
Systematic  Entomolog]/  17:  91-108. 

Bohart,  R.M.  and  A.S  Menke.  1976.  Sphecid  Wasps 
of  the  World:  A  Generic  Revision.  Berkely:  Uni- 
versity of  California  Press.  695  pp. 

Brockman,  H.  J.  and  A.  Grafen.  1989.  Mate  conflict 
and  male  behavior  in  a  soitary  wasp.  Trypoxylon 
(Trypargilum)  polifum  (Hymenoptera:  Sphecidae). 
Animal  Behavior  37:  232-255. 

Davies,  N.B.  Territorial  defense  in  the  speckled  wood 
butterfly  (Pararge  aegeria):  the  resident  always 
wins.  Animal  Behavior  26:  137-147. 

Eickwort,  G.C.  and  H.  S.  Ginsberg.  1980.  Foraging 
and  mating  behavior  in  Apoidea.  Annual  Review 
in  Entomology  25:  421-426. 

Evans,  H.E.  1971.  Obser\'ations  on  the  nesting  be- 
havior of  wasps  of  the  tribe  Cercerini.  journal  of 
the  Kansas  Entomological  Society  44:  500-523. 

Evans,  H.  E.  and  K.  M.  O'Neill.  1985.  Male  territorial 
behavior  in  four  species  of  the  tribe  Cercerini 
(Sphecidae:  Philanthinae).  journal  of  the  New  York 
Entomological  Society  93(3):  1033-1040. 

Evans,  H.  E.  and  K.  M.  O'Neill.  1988.  The  Natural 
History  and  Behavior  of  North  American  Bee- 
wolves.  Comstock  Publishing  Associates. 

Hook,  A.  W.  and  R.  W.  Matthews.  1980.  Nesting  bi- 
ology of  Oxybelus  sericeus  with  a  discussion  of 
nest  guarding  by  male  sphecid  wasps  (Hyme- 
noptera). Psyche  87:  21-37. 

Gwynne,  D.T.  1980.  Female  defense  polygyny  in  the 
bumblebee  wolf  Pliilanthus  hicinctus  (Hymenop- 
tera: Sphecidae).  Behavioral  Ecolology  and  Socio- 
btology  7:  213-225. 

O'Neill,  K.M.  1983a.  Territoriality,  body  size,  and 
spacing  in  males  of  the  beewolf  Phihvithus  basi- 
laris  (Hymenoptera:  Sphecidae).  Behavior  86:  295: 
321. 

O'Neill,  K.M.  1983b.  The  significance  of  body  size  in 
territorial  interactions  of  male  beewolves  (Hy- 
menoptera: Sphecidae,  Philanthus).  Animal  Behav- 
ior 3\A0i-AU. 

O'Neill,  K.M.  1990.  Female  nesting  behavior  and 
male  territoriality  in  Aphilanthops  subfrigidus 
Dunning  (Hymenoptera:  Sphecidae).  Pan-Pacific 
Entomologist  66:  19-23. 

Peckham,  D.J.,  F.E.  Kurczewski  and  D.B.  Peckham. 
1973.  Nesting  behavior  of  Nearctic  species  of  Ox- 
ybelus (Hymenoptera:  Sphecidae).  Annals  of  the 
Entomological  Society  of  America  bb:  647-661. 

Peckham,  D.  J.  1977.  Reduction  of  miltogrammine 
cleptoparasitism  by  male  Oxybelus  subulatus  (Hy- 
menoptera: Sphecidae).  Annals  of  the  Entomologi- 
cal Society  of  America  70:  823-828. 

Thornhill,  R.'and  J.  Alcock.  1983.  The  Evolution  of 
Insect  Mating  Systems.  Harvard  Press. 

Wang,  G.  and  M.D.  Greenfield.  1991.  Effects  of  ter- 
ritory ownership  on  dominanace  in  the  desert 
clicker  (Orthoptera:  Acrididae).  Animal  Behavior 
42:  579-587. 


J.  HYM.  RES. 
Vol.  4,  1995,  pp.  80-98 

Revised  Key  to  the  Nearctic  Species  of  Chrysocharis  Forster 
(Hymenoptera:  Eulophidae),  Including  Three  New  Species 

Christer  Hansson 
Department  of  Systematic  Zoology,  Lund  University,  Helgonavagen  3,  S-223  62  Lund,  Sweden 


Abstract. — The  Nearctic  species  of  Chrysocharis  are  keyed,  altogether  63  species.  The  paper  in- 
cludes the  description  of  three  new  species:  waJili,  cornigera,  sentenaca  and  two  new  combinations 
from  Chrysocharis  to  Omphale  Haliday:  varia  Hansson,  gracilicornis  Hansson. 


The  Nearctic  and  Neotropical  species  of  The  page  number  after  each  species  in 

Chrysocharis    were    revised    by    Hansson  the  revised  key  refers  to  Hansson  (1987). 

(1987).  Sixty  one  (61)  species  were  treated  The   descriptions,   diagnoses,   distribution 

and  18  of  the  species  were  described  as  and  hosts  of  Nearctic  species  of  Chrysochar- 

new.  From  the  Nearctic  region  57  species  is  are  found  in  Hansson  (1985,  1986,  1987). 
were  recognized.  However,  the  kev  for  the 

identificatton  of  the  species  is  difficult  and  RECOGNIZING  CHRYSOCHARIS 
laborious  to  use,  mainly  because  too  many  To  facilitate  the  recognition  of  Chryso- 
characters  are  used  under  each  item,  and  charis,  following  should  be  helpful.  Anten- 
characters  used  in  the  two  alternatives  un-  na  with  apical  two  segments  fused,  or 
der  each  item  are  not  always  exclusively  with  all  flagellar  segments  free  (exception: 
the  same.  To  improve  the  key  I  have  re-  imhrasiis  with  three  apical  segments  fused, 
arranged  it  so  that  the  most  obvious  char-  recognized  as  a  Chrysocharis  through  the 
acters  are  used  first,  and  under  each  item  long  postmarginal  vein — 1.5X  stigmal 
only  1-2  characters  are  used — only  occa-  vein — and  antennal  scrobes  join  below 
sionally  are  more  than  two  characters  frontal  suture  in  female).  Clypeus  not  de- 
used.  The  characters  selected  are  those  limited  (exception:  flaviclypeus,  recognized 
easy  to  see  and  that  vary  as  little  as  pos-  as  a  Chrysocharis  through  female  antennal 
sible.  In  some  cases  a  species  varies  in  scrobes  that  join  below  frontal  suture), 
characters  used,  and  this  species  is  found  Postmarginal  vein  longer  than  stigmal 
under  both  alternatives.  vein   (exception   zvahli   Sacornigera,   recog- 

In  the  revised  key  I  have  also  included  nized  as  members  of  Chrysocharis  through 
changes  that  has  taken  place  since  the  the  possession  of  the  three  abovemen- 
publication  of  the  previous  key:  Entedon  tioned  characters).  Without  a  sulcus  sur- 
imbrasus  Walker  has  been  transferred  to  rounding  ocellar  triangle  (this  separates 
Chrysocharis  from  Neochrysocharis  Kurdju-  Chrysocharis  from  Derostemis). 
mov  (Hansson  in  press);  the  species  treat- 
ed as  Zaommomyia  Girault  by  Hansson 
(1986)  have  been  transferred  to  Chryso- 
charis (Schauff  1991).  The  three  species  de-  The  terms  used  in  the  key  and  in  the 
scribed  as  new  in  this  paper  are  also  in-  text  are  indicated  on  Figs  1-8. 
eluded.  Furthermore,  two  species  regard-  Abbreviations  of  collections:  CNC=  Ca- 
ed  as  Chrysocharis  by  Hansson  (1987)  are  nadian  National  Collection  of  Insects  and 
hereby  transferred  to  Omphale  Haliday:  Arachnids,  Ottawa;  LUZM=  Lund  Uni- 
gracilicornis  Hansson,  varia  Hansson,  both  versify  Zoological  Museum;  TAMU=  Tex- 
n.  comb.  as  A&M  University,  College  Station. 


TERMINOLOGY  AND 
ABBREVIATIONS 


Volume  4,  1995  81 


KEY  TO  NEARCTIC  SPECIES  OF  CHRYSOCHARIS 

1.  Flagellum  with  all  five  segments  free  (e.g.  Figs  35,  39)     2 

-  Flagellum  with  apical  two  or  three  segments  fused,  i.e.  the  constrictions  between  the 
basal  funicular  segments  are  narrower  than  the  constrictions  between  the  apical  seg- 
ments (e.g.  Figs  37,  38)    19 

2.  Forewing  with  a  complete  row  of  setae  on  underside  of  costal  cell  (Figs  48,  49)   3 

-  Costal  cell  without  row  of  setae     4 

3.  Forewing  speculum  closed  (Fig.  49);  5th  flagellar  segment  (including  the  narrow  tip) 
1.3X  as  long  as  4th  segment  (Fig.  35) C.  chilo  (Walker)  (female,  male)  p.  30 

-  Forewing  speculum  open  laterally  (Fig.  48);  5th  flagellar  segment  0.9-l.Ox  as  long  as 
4th  segment  (Fig.  36)     C.  pilosa  Delucchi  (female,  male  unknown)  p.  30 

4.  Pronotal  collar  with  a  transverse  carina — at  least  on  median  pronotum — parts  of  pro- 
notum  behind  carina  smooth  and  shiny  (Fig.  87)    5 

-  Pronotum  without  transverse  carina,  hind  margin  of  pronotum  at  most  with  a  very 
narrow  smooth  strip  (Fig.  86)     7 

5.  Petiole  longer  than  wide  (Fig.  62)     C.  acoris  (Walker)  (female,  male)  p.  64 

-  Petiole  at  most  as  long  as  wide  (Figs  56,  59)     6 

6.  Scape  and  pedicel  bright  orange-yellow;  clypeus  pale  yellow    

C.  iUnstris  Graham  (male)  p.  59 

-  Scape  whitish  or  pale  brown,  pedicel  brown;  clypeus  metallic  bluish-green     

C.  occidentalis  (Girault)  (female,  male)  p.  54 

7.  Propodeal  callus  with  2  setae     8 

-  Propodeal  callus  with  at  least  3  setae    14 

8.  Petiole  pale;  male  flagellar  segments  with  a  single  whorl  of  setae  at  the  base  of  each 
segment  (Fig.  42)     C.  beckeri  Yoshimoto  (female,  male) 

-  Petiole  dark;  male  flagellar  segments  also  with  setae  in  middle  and  at  apex    9 

9.  Propodeum  with  2  complete  and  parallel  submedian  grooves  (Fig.  84)    

C.  sulcata  (Hansson)  (female,  male) 

-  Propodeum  without  such  grooves   10 

10.  Forewing  speculum  open  below  (Fig.  47)    C.  vonones  (Walker)  (female,  male) 

-  Forewing  speculum  closed  below    11 

11.  Scape  bright  orange-yellow;  reticulation  on  thoracic  dorsum  fine  and  engraved    

C.  fulviscapus  Hansson  (male,  female  unknown)  p.  45 

-  Scape  whitish  or  brown;  reticulation  on  thoracic  dorsum  raised  and  strong     12 

12.  Occipital  margin  with  a  complete  (reaching  from  eye  to  eye),  high  and  sharp  carina  (Fig. 
91);  frontal  suture  raised     C.  liriomyzae  Delucchi  (male)  p.  26 

-  Occipital  margin  without  a  complete  carina;  frontal  suture  not  raised    13 

13.  All  femora  predominantly  dark    C.  phijtomyzivora  Hansson  (male)  p.  20 

-  Femora  pale     C.  cerodonthae  Hansson  (male)  p.  21 

14.  Petiole  longer  than  length  of  median  propodeum  (Fig.  85)    .  .  C.  viridis  (Nees)  (male)  p.  29 

-  Petiole  at  most  as  long  as  length  of  median  propodeum   15 

15.  Postmarginal  vein  3.0-3.5  x  as  long  as  stigmal  vein   16 

-  Postmarginal  vein  2.0-2.5X  as  long  as  stigmal  vein   17 

16.  Petiole  about  as  long  as  median  propodeum;  dorsellum  usually  excavated  and  divided 
in  two  parts  by  a  median  carina  and  hind  part  with  a  median  incision  (Fig.  69)     

C.  entedonoides  (Walker)  (female,  male)  p.  28 

-  Petiole  shorter  than  median  propodeum;  dorsellum  convex  to  flat,  without  median  in- 
cision and  carina  (Fig.  63)     C.  amyite  (Walker)  (female,  male)  p.  28 

17.  Hind  coxa  conspicuously  long  and  slender,  about  2.5 X  as  long  as  wide  (Fig.  52);  petiolar 
foramen  triangular    C.  longicoxa  Hansson  (male)  p.  26 

-  Hind  coxa  stouter,  about  1.5X  as  long  as  wide;  petiolar  foramen  semicircular  to  qua- 
drangular          18 


82 


Journal  of  Hymenoptera  Research 


Figs.  1-8.  Terminology.  1,  beckeri,  female:  cc=  costal  cell;  dm=  dorsellum;  m=  marginal  vein;  ml=  midlobe; 
msc=  mesoscutum;  om=  occipital  margin;  OOL=  ocell-ocular  line;  pc=  propodeal  callus;  pet=  petiolus;  pm  = 
postmarginal  vein;  POL=  postocellar  line;  POO=  distance  between  hind  ocelli  and  occipital  margin;  prm= 


Volume  4,  1995  g3 

18.  Petiole  as  long  as  wide  (as  in  Fig.  77);  femora  usually  pale — in  a  few  specimens  pre- 
dominantly dark    C.  avia  Hansson  (female,  male)  p.  28 

-  Petiole  transverse  (Fig.  80);  femora  always  predominantly  dark    

C.  phytomyzivora  Hansson  (male)  p.  20 

19.  Flagellum  with  apical  three  segments  fused  (Fig.  41)   

C.  imbrasus  (Walker)  (female,  male  unknown) 

-  Flagellum  with  apical  two  segments  fused    20 

20.  Forewing  with  a  complete  row  of  setae  on  underside  of  costal  cell  (as  in  Figs  48,  49)  .  . 

C  robusta  Yoshimoto  (female,  male)  p.  62 

-  Costal  cell  without  row  of  setae     21 

21.  Pronotal  collar  with  transverse  carina— at  least  on  median  pronotum  (Fig.  87)    22 

-  Pronotum  without  transverse  carina  (Fig.  86)    55 

22.  Flagellum  yellow;  frontal  suture  missing  (Fig.  26)    C.  lualleyi  Yoshimoto  (male)  p.  60 

-  Flagellum  brown;  frontal  suture  present    23 

23.  Petiole  at  least  1.5x  as  long  as  wide    24 

-  Petiole  at  most  as  long  as  wide    26 

24.  Malar  space  very  narrow,  1/15  the  width  of  mouth  opening  (Fig.  92);  frontal  suture 
smoothly  curved;  with  a  procession  between  antennal  toruli     

C.  prodice  (Walker)  (female,  male)  p.  65 

-  Malar  space  not  as  narrow,  1/7  the  width  of  mouth  opening  (Fig.  93);  frontal  suture 
more  straight;  without  procession  between  toruli     25 

25.  Occipital  margin  with  a  low  sharp  carina;  propodeal  callus  with  2  (3)  setae;  female  frons 
above  fork  usually  with  rather  strong  reticulation;  male  frons  smooth  or  with  weak 
reticulation     C.  acoris  (Walker)  (female,  male)  p.  64 

-  Occipital  margin  rounded  and  smooth  (in  a  few  cases  with  a  very  weak  carina);  pro- 
podeal callus  with  4-5  (3)  setae;  female  frons  above  fork  smooth  or  with  very  weak 
reticulation  (male  unknown)     C.  amasis  (Walker)  (female)  p.  65 

26.  Propodeum  with  a  strong  and  complete  median  carina  (Fig.  58) 

C.  walleyi  Yoshimoto  (female)  p.  60 

-  Propodeum  without  a  complete  median  carina     27 

27.  Posteromedian  part  of  propodeum  with  two  submedian,  slightly  curved  carinae  (Fig. 
70);  male  pedicel  bright  orange-yellow    C.  gemma  (Walker)  (female,  male)  p.  59 

-  Posteromedian  propodeum  without  carinae,  or  with  2-6  short  and  straight  carinae  (e.g. 
Fig.  73);  male  pedicel  brown  or  whitish    28 

28.  Anteromedian  part  of  propodeum  with  a  wide  and  strong  median  carina — reaching  half 
the  length  of  propodeum  and  then  dividing  into  2  weaker  carinae  which  diverge  towards 
the  hind  edge  of  propodeum  (Fig.  73)     C.  ilhistris  Graham  (female)  p.  59 

-  Median  carina  on  anteromedian  part  of  propodeum  weaker  or  missing     29 

29.  Clypeus  partly  or  completely  pale  yellow 30 

-  Clypeus  completely  dark  and  metallic     31 

30.  Reticulation  on  thoracic  dorsum  very  dense  and  strong,  almost  like  punctulation;  pro- 
podeal callus  with  5  setae     C.  flaviclypeiis  Hansson  (female,  male  unknown)  p.  63 


propodeum;  prn=  pronotum;  s=  stigmal  vein;  scu=  scutellum;  sl=  sidelobe;  spe=  speculum;  sul=  spiracular 
sulcus.  2,  coUaris,  female:  pcl=  pronotal  collar.  3,  Antenna,  coinpresiiconiis,  female:  an=  anelli;  cla=  clava;  fla  = 
flagellum;  fun=  funiculus;  MPS=  multiporous  plate  sensilla;  pcl=  pedicel;  sca=  scape.  4,  Head,  side  view, 
phytoniyzh'om,  female.  5,  Head,  front  view,  clarknc  male:  cly=  clypeus;  fs=  frontal  suture;  he=  height  of  eye; 
mo=  width  of  mouth  opening;  ms=  width  of  malar  space;  scr=  antennal  scrobes;  tor=  antennal  toruli.  6, 
Mesothorax,  side  view,  longicoxa,  female:  b=  borderline  between  upper  and  lower  mesepimeron.  7, 
Meso+metathorax,  side  view,  tristis,  female:  b=  as  in  Fig.  6;  C2=  mid  coxa;  C3=  hind  coxa;  pc=  propodeal 
callus.  8,  iiicdiivia,  female:  pf=  petiolar  foramen. 


84 


Journal  of  Hymenoptera  Research 


10 


11 


Figs.  9-15.  Chrysocharis  n.spp.  9-12.  cornigera.  9,  Head,  side  view,  female.  10,  Head,  front  view,  female.  11, 
Head,  dorsal  view,  female.  12,  Antenna,  male.  13-15.  wahli.  13,  Head,  side  view,  female.  14,  Head,  front  view, 
female.  15,  Antenna,  male. 


Volume  4,  1995 


85 


Figs.  16-26.     Head,  front  view.  16-17,  ainslici,  female.  18,  Ditto,  male.  19,  oaciiiidi^,  female.  20,  Ditto,  male.  21, 
chromatomyiae,  female.  22,  Ditto,  male.  23,  perditor,  female.  24,  Ditto,  male.  25,  amasis,  female.  26,  wallcyi,  male. 


Figs.  27-40.  27,  Head,  front  view,  tristis,  female.  28,  Ditto  male.  29,  Head,  lateral  view,  coptodiscae,  male.  30, 
Ditto,  pallidigaster,  female.  31,  Ditto,  submiitica,  male.  32,  Head,  dorsal  view,  occidentalis,  female.  33,  Ditto, 
coptodiscae,  female.  34-40.  Antennae.  34,  occidentalis,  female.  35,  chilo,  female.  36,  Apical  2  flagellar  segments, 
pilosa,  female.  37,  coptodiscae,  female.  38,  liriomyzae,  female.  39,  Ditto,  male.  40,  Flagellum,  compressicornis,  male. 


Figs.  41-56.  41-45.  Antennae.  41,  /»/^ras/<s,  female.  42,  heckeri,  male.  43,  /'j/fv»s,  male.  44,  polyzo,  female.  45, 
prodice,  female.  46,  Forewing,  paradoxa.  47^9.  Base  of  forewing.  47,  voiioiics.  48,  pilosa.  49,  c"/i;7o.  50-52.  Hindleg 
from  coxa  to  basal  tibia.  50,  occidentalis.  51,  coptodiscae.  52,  longicoxa.  53,  Hindleg,  polita.  54,  Gaster,  laomedon, 
female.  55,  Ditto,  longigaster,  female.  56,  Petiole + gaster,  illuslris,  male. 


Figs.  57-71.  57,  Body,  dorsal  view,  acutigaster,  female.  58,  Propodeum+ petiole +gaster,  loaUeyi,  female.  59- 
61.  Mesosoma  + petiole,  female.  59,  occidentalis.  60,  mediana.  61,  polita.  62-71.  Propodeum+ petiole,  female.  62, 
acoris.  63,  amyite.  64,  assis.  65,  cerodonthae.  66,  chromatomyiae.  67,  clarkac.  68,  crassiscapus.  69,  entedonoides.  70, 
gemma.  71,  gimulti.  71,  Petiole,  side  view,  clarkac. 


Figs.  73-85.     Propodeum+ petiole,  female.  73,  iUiistris.  74,  laricinellae.  75,  nephereus.  76,  nitetis.  77,  orbicularis. 

78,  oscinidis.  79,  pciithcus.  80,  pJiytouiyzivora.  81,  poh/zo.  82,  pubicornis.  83,  purpurea.  84,  sulcata.  85,  viridis. 


90  Journal  of  Hymenoptera  Research 

-  Reticulation  on  thoracic  dorsum  weak;  propodeal  callus  with  2  setae     

C.  aluta  Yoshimoto  (female,  male)  p.  58 

31.  Anteromedian  part  of  propodeum  with  a  round  to  triangular  fovea  bordered  by  a  raised 
edge  (Fig.  83),  fovea  not  divided  by  a  median  carina;  propodeal  callus  with  3-4  setae 

C.  purpurea  Bukowski  (female,  male)  p.  58 

-  Anteromedian  part  of  propodeum  either  with  carinae  shaped  like  a  V,  Y  or  a  T  (turned 
upside-down)  (e.g.  Figs  74,  76),  or  with  two  submedian  pits  (e.g.  Fig.  75);  propodeal 
callus  with  2  setae     32 

32.  Females,  i.e.  gaster  widest  close  to  base  or  in  the  middle  and  with  apex  pointed  or 
rounded  (e.g.  Figs  1,  54),  scape  comparatively  narrow,  4.3-5.1  X  as  long  as  wide    33 

-  Males,  i.e.  gaster  narrow  at  base  and  gradually  becoming  wider  towards  apex  (Fig.  56), 
scape  comparatively  wide,  3.1-3.3X  as  long  as  wide    45 

33.  Gaster  long — ratio  length  of  mesosoma/ length  of  gaster=  0.6-0.7 — more  or  less  parallel- 
sided  and  with  hind  part  pointed  (Fig.  54)    34 

-  Gaster  shorter — ratio  m/g  at  the  lowest  0.8  but  usually  1.0 — and  oval-shaped  with  hind 
part  rounded    35 

34.  Malar  space  very  narrow — about  1/8  the  width  of  mouth  opening  (Fig.  97);  fore  coxa 
predominantly  pale     C.  laomedon  (Walker)  p.  57 

-  Malar  space  wider — about  1/4  the  width  of  mouth  opening;  fore  coxa  dark  and  metallic 

C.  elongata  (Thomson)  (male  unknown)  p.  56 

35.  Scutellum  with  engraved  reticulation;  flagellum  stout,  flagellar  segments  gradually  be- 
coming shorter  and  wider  towards  apex  (Fig.  37)     C.  coptodiscae  Yoshimoto  p.  53 

-  Scutellum  with  raised  reticulation;  flagellum  slender  (e.g.  Fig.  34)    36 

36.  Scutellum  flattened;  anteromedian  part  of  propodeum  with  two  submedian  pits  (Fig.  75) 

C.  nephereus  (Walker)  p.  52 

-  Scutellum  convex;  anteromedian  part  of  propodeum  with  carinae  shaped  like  a  V,  Y  or 

a  T    37 

37.  Transverse  pronotal  carina  weak  and  present  only  on  median  part  of  pronotum    38 

-  Transverse  pronotal  carina  strong  and  present  along  the  major  part  of  pronotum     ....     40 

38.  Hind  femur  conspicuously  stout,  2.5  x  as  long  as  wide  (Fig.  50);  malar  space  narrower 
than  width  of  scape    C.  occidentalis  (Girault)  p.  54 

-  Hind  femur  slender,  4.0 x  as  long  as  wide  (as  in  Fig.  51);  malar  space  at  least  as  wide  as 
width  of  scape   39 

39.  Malar  space  as  wide  as  width  of  scape  (Fig.  30);  frons  below  suture  golden-red     

C.  pallidigaster  Hansson  p.  49 

-  Malar  space  1.5-2.0  x  as  wide  as  width  of  scape  (Fig.  31);  frons  below  suture  usually 
purple    C.  submutica  Graham  p.  49 

40.  Malar  space  narrower  than  the  width  of  scape     41 

-  Malar  space  at  least  as  wide  as  width  of  scape     42 

41.  Petiole  as  long  as  wide,  with  protruding  forecorners  (Fig.  64)     ....  C.  assis  (Walker)  p.  64 

-  Petiole  transverse,  without  protruding  forecorners  (as  in  Fig.  79)    

C.  paradoxa  Hansson  p.  58 

42.  Pronotal  collar  long  (Fig.  2);  occipital  margin  with  a  carina  behind  ocellar  triangle     .  .  . 

C.  collaris  Graham  p.  52 

-  Pronotal  collar  shorter  (Fig.  87);  occipital  margin  without  carina     43 

43.  Meshes  of  reticulation  with  about  the  same  size  over  entire  frons  (Fig.  95);  scutellum 
distinctly  elongate — ratio  length/width=  1.2    C  laricinellae  (Ratzeburg)  p.  50 

-  Reticulation  on  frons  below  suture  usually  with  larger  meshes  than  on  frons  above 
suture  (as  in  Fig.  96);  scutellum  about  as  long  as  wide    44 

44.  Raised  surface  of  petiole  small  (Fig.  79);  hind  femur  completely  white  or  very  pale  brown 

at  base    C.  pentheus  (Walker)  p.  51 


Volume  4,  1995  91 

-  Raised  surface  of  petiole  larger  (Fig.  76);  hind  femur  usually  predominantly  dark    

C.  nitetis  (Walker)  p.  51 

45.  Third  anellus  large  (e.g.  as  in  Fig.  37)    C.  assis  (Walker)  p.  64 

-  Third  anellus  small  and  discoid   46 

46.  Gaster  with  a  pale  subbasal  spot   C.  pallidigaster  Hansson  p.  49 

-  Gaster  without  a  pale  subbasal  spot    47 

47.  Malar  space  narrow — about  0.3 x  as  wide  as  width  of  scape     48 

-  Malar  space  wider — at  least  0.5  x  as  wide  as  width  of  scape     49 

48.  Propodeum  with  a  complete  median  carina  or  with  2  weak,  complete  and  parallel  carinae 

C.  laomedon  (Walker)  p.  57 

-  Propodeum  without  longitudinal  carinae    C.  paradoxa  Hansson  p.  58 

49.  Transverse  carina  along  pronotum  present  and  complete    50 

-  Transverse  carina  along  pronotum  weak  and  incomplete    54 

50.  Scutellum  flattened  and  about  as  long  as  wide;  reticulation  on  scutellum  small-meshed 
(Fig-  88)    C.  nephereus  (Walker)  p.  52 

-  Scutellum  distinctly  longer  than  wide   51 

51.  Forewing  subtruncate  (as  in  Fig.  46)    C.  paradoxa  Hansson  p.  58 

-  Forewing  rounded  apically     52 

52.  Eyes  4.5x  as  high  as  width  of  malar  space  (Fig.  94) 

C  laricinellae  (Ratzeburg)  (spring  generation)  p.  50 

-  Eyes  5-7x  as  high  as  malar  space  (as  in  Fig.  96)    53 

53.  Entire  frons  with  small-meshed  reticulation  (as  in  Fig.  95) 

C.  laricinellae  (summer  generation)  p.  50 

-  Frons  below  suture  with  larger  meshes  than  above  suture  (Fig.  96)    

C.  pentheiislC.  nitetis  (inseparable)  p.  51 

54.  Malar  space  as  wide  as  width  of  scape;  temples  comparatively  large  and  eyes  compar- 
atively small,  width  of  temples  at  lower  edge  of  eyes  0.15  x  the  height  of  an  eye  (Fig. 

31)    C.  siibmutica  Graham  p.  49 

-  Malar  space  only  0.5  x  as  wide  as  width  of  scape;  temples  smaller  and  eyes  larger,  width 
of  temples  at  lower  edge  of  eyes  0.07X  the  height  of  an  eye  (Fig.  29)    

C.  nephereus/C.  coptodiscae  (inseparable)  p.  52-53 

55.  Propodeum  with  two  complete  parallel  submedian  grooves  (Fig.  84)     

C.  sulcata  (Hansson)  (female,  male) 

-  Propodeum  without  complete  longitudinal  grooves 56 

56.  Petiole  distinctly  longer  than  wide  (at  least  1.4x  as  long  as  wide)    57 

-  Petiole  usually  at  most  as  long  as  wide     63 

57.  Frontal  suture  absent  (Figs  23,  24)   C.  perditor  Hansson  (female,  male)  p.  38 

-  Frontal  suture  present     58 

58.  Petiole  with  a  pair  of  medio-lateral  horns  (Fig.  85);  propodeum  with  2-5  setae  inside 
spiracular  sulci     C.  viridis  (Nees)  (female)  p.  29 

-  Petiole  with  a  pair  of  antero-lateral  horns  (Fig.  71);  propodeum  without  setae  inside 
spiracular  sulci    59 

59.  Scape  completely  pale;  postmarginal  vein  only  1.3x  as  long  as  stigmal  vein 

C.  gibsoni  Hansson  (female,  male)  p.  38 

-  Scape  with  at  least  apical  part  infuscate;  postmarginal  vein  2x  as  long  as  stigmal  vein      60 

60.  Frontal  suture  down-curved  (Figs  27,  28);  speculum  open  below  (as  in  Fig.  47)     

C.  tristis  Hansson  (female,  male)  p.  41 

-  Frontal  suture  curved  upwards  or  straight;  speculum  closed  below  (a  few  specimens 
have  an  open  speculum)     61 

61.  Petiole  at  least  2X  as  long  as  wide    C.  ignota  Hansson  (female,  male)  p.  39 

-  Petiole  less  than  2X  as  long  as  wide    62 

62.  Dorsellum  comparatively  short  and  wide,  6.5x  as  wide  as  long  (Fig.  71);  dorsal  surface 

of  petiole  concave  in  sideview    C.  giratilti  Yoshimoto  (female,  male)  p.  35 


92  Journal  of  Hymenoptera  Research 

-  Dorsellum  longer,  3X  as  wide  as  long  (Fig.  78);  dorsal  surface  of  petiole  straight  in 
sideview     C.  oscinidis  Ashmead  (female,  male)  p.  34 

63.  Petiole  pale,  as  long  as  wide    64 

-  Petiole  dark,  as  long  as  wide  to  transverse   65 

64.  Frontal  suture  almost  straight;  thoracic  dorsum  with  weak  reticulation    

C.  minuta  (Hansson)  (female,  male) 

-  Frontal  suture  V-shaped;  thoracic  dorsum  with  strong  reticulation     

C.  beckeri  Yoshimoto  (female,  male) 

65.  Clypeus  pale  yellow    C  flaviclypeiis  Hansson  (female,  male  unknown)  p.  63 

-  Clypeus  dark  and  metallic    66 

66.  Mouth  opening  wider  than  height  of  an  eye — ratio  width  of  mouth/height  of  eye=  1.1 

C.  kimamaensis  Hansson  (female,  male  unknown)  p.  45 

-  Mouth  opening  narrower  than  height  of  an  eye   67 

67.  Lateral  parts  of  frontal  suture  raised  to  form  two  conspicuous  horns  (Figs  9-11) 

C.  comigera  n.sp.  (female,  male) 

-  Frontal  suture  not  raised,  or  in  one  species  (liriomyzae)  with  entire  frontal  suture  slightly 
raised   68 

68.  Anteromedian  part  of  propodeum  without  carinae,  a  fold  or  a  pit  (e.g.  Fig.  8)    69 

-  Anteromedian  part  of  propodeum  with  carinae,  a  fold  or  a  pit     72 

69.  Propodeum  with  a  complete  median  carina    C.  vonones  (Walker)  (female,  male) 

-  Propodeum  smooth,  without  median  carina   70 

70.  Forewing  with  an  infuscate  spot  below  stigmal  vein;  postmarginal  vein  as  long  as  stigmal 
vein;  male  pedicel  pale   C.  zvahli  n.sp.  (female,  male) 

-  Forewing  hyaline;  postmarginal  vein  twice  as  long  as  stigmal  vein;  male  pedicel  dark       71 

71.  All  femora  usually  predominantly  dark  and  metallic;  hind  femur  comparatively  slender, 
3.5  X  as  long  as  wide;  mesosoma  comparatively  slender,  1.6X  as  long  as  wide  (Fig.  60) 

C.  mediana  Forster  (female,  male)  p.  48 

-  Femora  completely  pale;  hind  femur  conspicuously  stout,  3.2  X  as  long  as  wide  (Fig.  52) 
mesosoma  comparatively  stouter,  1.4X  as  long  as  wide  (Fig.  61)    

C.  polita  (Howard)  (female,  male)  p.  48 

72.  Anteromedian  part  of  propodeum  with  a  single  median  pit  (e.g.  Figs  57,  67,  78)    73 

-  Anteromedian  part  of  propodeum  with  carinae  shaped  like  a  Y  or  a  T  (turned  upside- 
down)  (e.g.  Fig.  68),  a  fold  (e.g.  Fig.  59),  two  submedian  pits  (e.g.  Fig.  75),  or  a  raised 
peak   88 

73.  Petiole  very  short,  less  than  half  as  long  as  wide  (Figs  8,  57)    74 

-  Petiole  longer,  at  least  as  long  as  wide  (e.g.  Figs  78,  81)    76 

74.  Hind  part  of  anteromedian  fovea  on  propodeum  considerably  raised;  female  with  an- 
tennal  clava  distinctly  wider  than  first  2  funicular  segments   

C.  acutigaster  Hansson  (female,  male)  p.  22 

-  No  part  of  anteromedian  fovea  on  propodeum  conspicuously  raised;  female  with  entire 
flagellum  uniformly  slender 75 

75.  All  femora  usually  predominantly  dark  and  metallic;  hind  femur  comparatively  slender, 
3.5  X  as  long  as  wide;  mesosoma  comparatively  slender,  1.6x  as  long  as  wide  (Fig.  60) 

C.  mediana  Forster  (female,  male)  p.  48 

-  Femora  completely  pale;  hind  femur  conspicuously  stout,  3.2 X  as  long  as  wide  (Fig.  52); 
mesosoma  comparatively  stouter,  1.4x  as  long  as  wide  (Fig.  61)    

C.  polita  (Howard)  (female,  male)  p.  48 

76.  Anteromedian  fovea  on  propodeum  very  wide — several  times  wider  than  long  (as  in 
Fig.  1)     C.  vonones  (female,  male) 

-  Anteromedian  fovea  on  propodeum  about  as  long  as  wide     77 

77.  Borderline  between  upper  and  lower  mesepimeron  straight  (Fig.  7);  petiolar  foramen 
with  a  large  membrane  in  upper  part  (Fig.  77)   78 


Volume  4,  1995  93 

-  Borderline  between  upper  and  lower  mesepimeron  at  least  slightly  curved  (Fig.  6);  pet- 
iolar  foramen  with  a  very  small  membrane  in  upper  part,  or  without  membrane 81 

78.  All  coxae  pale — fore  coxa  sometimes  infuscate  in  upper  half    

C.  sjibcirculnris  Yoshimoto  (female,  male)  p.  33 

-  At  least  some,  usually  all  coxae  dark  and  metallic   79 

79.  Forewing  with  speculum  open  below  (as  in  Fig.  47);  female  gaster  long — ratio  length  of 
mesosoma/ length  of  gaster=  0.8,  with  hind  part  pointed  (Fig.  55)   

C.  longigaster  Hansson  (female,  male  unknown)  p.  32 

-  Forewing  with  speculum  closed;  female  gaster  shorter — ratio  length  of  mesosoma/ 
length  of  gaster  =  1.2-1.3,  with  hind  part  less  pointed   80 

80.  Eyes  comparatively  small,  height  of  eye  4.0-7.0  x  the  width  of  malar  space  in  female, 
2.6X  in  male  (Figs  16-18);  femora  usually  predominantly  dark,  but  occasionally  pale; 
postmarginal  vein  frequently  less  than  2x  as  long  as  stigmal  vein 

C.  ainsliei  Crawford  (female,  male)  p.  31 

-  Eyes  comparatively  large,  height  of  eye  9.3x  the  width  of  malar  space  in  female,  6.3X 
in  male  (Figs  19,  20);  femora  pale;  postmarginal  vein  2x  as  long  as  stigmal  vein    

C.  oscinidis  Ashmead  (female,  male)  p.  34 

81.  Frontal  suture  raised;  with  an  interantennal  process  between  toruli  (Fig.  89);  mouth 
opening  with  an  incision  below  eye  (Fig.  90)    C.  liriomyzae  Delucchi  (female)  p.  26 

-  Frontal  suture  not  raised;  without  process  between  toruli;  mouth  opening  without  inci- 
sion         82 

82.  Petiole  with  raised  surface  quadrangular  (Fig.  81)  (shape  varying  from  trapezoid,  qua- 
dratic to  rectangular)     C.  polyzo  (Walker)  (female,  male)  p.  22 

-  Raised  surface  of  petiole  not  quadrangular  (e.g.  Figs  66,  67)   83 

83.  Hind  coxa  long  and  slender,  about  2.5  X  as  long  as  wide  (Fig.  52);  propodeal  callus  with 
5-8  setae;  petiolar  foramen  triangular    C.  longicoxa  Hansson  (female,  male)  p.  26 

-  Hind  coxa  stouter,  about  1.5  x  as  long  as  wide;  propodeal  callus  with  2-A  (some  speci- 
mens of  clarkae  have  5)  setae;  petiolar  foramen  rounded  or  quadrangular    84 

84.  Flagellum  thick,  about  2x  as  wide  as  width  of  scape  (Figs  3,  40);  each  flagellar  segment 
with  MPS  in  two  transverse  rows    C.  cotnpressicomis  Ashmead  (female,  male)  p.  16 

-  Flagellum  more  slender,  at  most  1.5X  as  wide  as  width  of  scape;  each  flagellar  segment 
with  MPS  in  one  transverse  row    85 

85.  Petiole  with  protruding  forecorners,  usually  as  long  as  wide  (shorter  than  wide  in  males) 
(Figs  66,  67)   86 

-  Petiole  without  protruding  forecorners,  shorter  (Figs  65,  80)   87 

86.  Raised  surface  of  petiole  considerably  raised,  with  coarse  sculpture — which  is  situated 
at  a  much  higher  level  than  protruding  forecorners  (Fig.  72);  propodeal  callus  with  3  (4- 

5)  setae     C.  clarkae  Yoshimoto  (female,  male)  p.  23 

-  Raised  surface  of  petiole  lower,  with  strong  punctures  or  irregular  sculpture  (finer  than 
in  clarkae),  top  of  surface  closer  to  forecorners;  propodeal  callus  with  2  setae    

C.  chromatomyiae  Hansson  (female,  male)  p.  24 

87.  Femora  predominantly  dark     C.  phytouiyzivora  Hansson  (female)  p.  20 

-  Femora  pale     C.  cerodonthae  Hansson  (female)  p.  21 

88.  Anteromedian  part  of  propodeum  raised  in  a  peak;  hind  coxa  long  and  slender,  2.5  x  as 
long  as  wide  (Fig.  52)    C.  longicoxa  Hansson  (female,  male)  p.  26 

-  Anteromedian  part  of  propodeum  with  raised  carinae  (Fig.  82);  hind  coxa  stouter,  1.5X 

as  long  as  wide   89 

89.  With  a  fine  median  groove  on  posterior  part  of  midlobe  of  mesoscutum  and  anterior 
scutellum   C.  crassiscaptis  (Thomson)  (female,  male)  p.  18 

-  Without  median  groove  on  mesoscutum  and  scutellum     90 

90.  Propodeal  callus  with  3-4  setae   C.  piibicornis  (Zetterstedt)  (female,  male)  p.  16 

-  Propodeal  callus  with  2  setae     91 


94  Journal  of  Hymenoptera  Research 

91.  Flagellum  thick,  about  2X  as  wide  as  width  of  scape  (Figs  3,  40);  each  flagellar  segment 
with  MPS  in  two  transverse  rows    C.  cotnpressicomis  Ashmead  (female,  male)  p.  16 

-  Flagellum  more  slender,  at  most  1.5X  as  wide  as  width  of  scape;  each  flagellar  segment 
with  MPS  in  one  transverse  row    92 

92.  Forewing  with  speculum  open  below  (as  in  Fig.  47)    C.  gemina  Hansson  (male)  p.  56 

-  Forewing  with  speculum  closed  below    93 

93.  Females,  i.e.  with  gaster  widest  close  to  base  or  in  the  middle  and  with  apex  pointed  or 
rounded  (e.g.  Fig.  54);  scape  comparatively  narrow,  4.3-5.1  X  as  long  as  wide     94 

-  Males,  i.e.  with  gaster  narrow  at  base  and  gradually  becoming  wider  towards  apex  (Fig. 
56);  scape  comparatively  wide,  3.1-3.3X  as  long  as  wide     100 

94.  Hind  femur  conspicuously  stout,  2.5  X  as  long  as  wide  (Fig.  50);  malar  space  narrower 
than  width  of  scape    C.  occidentalis  (Girault)  p.  54 

-  Hind  femur  slender,  3.9  x  as  long  as  wide  (as  in  Fig.  51);  malar  space  at  least  as  wide 

as  width  of  scape    95 

95.  Malar  space  1.5-2X  as  wide  as  width  of  scape     96 

-  Malar  space  as  wide  as  width  of  scape   98 

96.  Postmarginal  vein  only  1.3x  as  long  as  stigmal  vein   C.  gemina  Hansson  p.  56 

-  Postmarginal  vein  2X  as  long  as  stigmal  vein 97 

97.  Eyes  and  ocelli  larger,  ratio  height  of  eye/width  of  mouth=  2.1,  distance  between  hind 
ocelli  2x  width  of  one  ocellus    C.  submutica  Graham  p.  49 

-  Eyes  and  ocelli  smaller,  ratio  height  of  eye/width  of  mouth=  1.7,  distance  between  hind 
ocelli  3.4X  width  of  one  ocellus C.  sentenaca  n.sp. 

98.  Borderline  between  upper  and  lower  mesepimeron  curved  (as  in  Fig.  6)   

C.  griffithsi  Hansson  p.  17 

-  Borderline  between  upper  and  lower  mesepimeron  straight  (as  in  Fig.  7)    99 

99.  Scutellum  flattened  (Fig.  87)     C.  nephereus  (Walker)  p.  52 

-  Scutellum  convex    C.  pallidigaster  Hansson  p.  49 

100.  Gaster  with  a  pale  subbasal  spot   C.  pallidigaster  Hansson  p.  49 

-  Gaster  without  a  pale  subbasal  spot    101 

101.  Hind  pair  of  ocelli  close,  distance  between  them  0.9  X  the  largest  measure  of  one  ocellus 
(Fig.  32)    C.  occidentalis  (Girault)  p.  54 

-  Hind  pair  of  ocelli  further  apart,  distance  between  them  2X  the  largest  measure  of  one 
ocellus  (as  in  Fig.  33)     102 

102.  Malar  space  as  wide  as  width  of  scape   103 

-  Malar  space  0.5  X  as  wide  as  width  of  scape     104 

103.  Thoracic  dorsum  convex;  ocelli  larger,  distance  between  hind  ocelli  1.8X  width  of  one 
ocellus    C.  submutica  Graham  p.  49 

-  Thoracic  dorsum  flat;  ocelli  smaller,  distance  between  hind  ocelli  3.5  X  width  of  one 
ocellus    C.  sentenaca  n.sp. 

104.  Scutellum  convex  and  1.4X  as  long  as  wide;  borderline  between  upper  and  lower  me- 
sepimeron curved  (as  in  Fig.  6)    C.  griffithsi  Hansson  p.  17 

-  Scutellum  flattened  and  about  as  long  as  wide;  borderline  between  upper  and  lower 
mesepimeron  straight  (as  in  Fig.  7)    C.  nephereus  (Walker)  p.  52 


DESCRIPTIONS  1228W,   sect.   32),   l-7.iii.88,   W.E.   Wahl, 

^,  ,      .        ...  MT",  in  CNC.  Paratvpes:  5  females  with 

Chrysochans  ivahh  n.  sp.  ,  i    ,    i  ,    ,  ^    /■        i 

'^  same   label-data   as   holotype;   3   females 

"USA:  California,  San  Luis  Obispo  Coun- 


Figs  13-15 


Type  material. — Holotype  female  la-  ty,  6  mi  SE  Pozo,  1500',  26.iii-9.iv.90,  W.E. 
belled  "USA:  California,  Santa  Barbara  Wahl";  1  female  2  males  with  same  label 
County,   18   mi.   WNW   Cuyama   (TllN,     as  previous  but  collected  9-21.iv.90;  3  fe- 


Volume  4,  1995 


95 


Figs.  86-91.  86-88.  Head+mesosoma,  dorsal  view.  86,  liriomyzae,  female.  87,  nephereus,  female.  88,  Ditto,  male. 
89-91.  liriomyzae.  89,  Head,  front  view.  90,  Semicircular  incision  in  lateral  part  of  mouth  opening.  91,  Occipital 
margin  (om). 


males  in  LUZM,  remaining  in  CNC;  1 
male  "USA:  California,  San  Bernardino 
County,  Summit  Valley,  2mi.  E  Hwy  15, 
28.V.1981,  J  Woolley  81/025"  (TAMU). 

Etymology. — Named  after  W.E.  Wahl, 
who  ran  malaise-traps  in  California  and 
thereby  collected  a  large  material  of  use  to 
me  and  other  taxonomists. 

Diagnosis. — Forewing  with  an  infuscate 
spot  below  stigmal  vein;  postmarginal 
vein  short,  as  long  as  stigmal  vein;  anter- 
omedian part  of  propodeum  smooth;  male 
pedicel  pale. 

Description. — Length  of  body:  Female  = 
1.2-1.6  mm,  male=  0.9  mm. 

Head:   Scape  pale,   remaining  antenna 


dark;  male  pedicel  pale.  Apical  2  segments 
of  flagellum  fused  (Figs  13,  15).  HE/MS/ 
MO  female:  4.6/1.0/2.2,  male:  3.0/1.0/1.9. 
Malar  space  1.5X  as  wide  as  width  of 
scape  in  both  sexes.  Frons  golden-green, 
with  strong  and  small-meshed  reticula- 
tion. Frontal  suture  V-shaped.  Vertex  me- 
tallic greenish-blue,  with  strong  and 
small-meshed  reticulation.  Inner  orbit  of 
eye  with  1  row  of  setae.  POL/OOL/POO: 
2.2/1.0/1.2.  Occipital  margin  rounded. 
Ratio  width  of  head /width  of  thorax 
across  posterior  mesoscutum=  1.1. 

Mesosoma:  Pronotum  without  trans- 
verse carina.  Mesoscutum  and  scutellum 
metallic   greenish-blue;  with  strong  and 


96 


Journal  of  Hymenoptera  Research 


Figs.  92-97.     Head,  front  view.  92,  prodice,  female.  93,  amasis,  female.  94,  laricinellae,  male,  spring  generation. 
95,  Ditto,  female,  summer  generation.  96,  pentheus,  male.  97,  laomedon,  female. 


small-meshed  reticulation;  scutellum  flat-  weakly  curved.  All  coxae  dark  and  metal- 

tened  and  about  as  long  as  wide.  Dorsel-  lie  (fore  coxa  in  some  specimens  pale  with 

lum  small  and  convex,  3.4  X  as  wide  as  darkmetallic  base)  with  weak  reticulation; 

long,  with  strong  reticulation.  Borderline  femora  and  tibiae  pale;  tarsi  pale  with  4th 

between  lower  and  upper  mesepimeron  segment  dark.  Forewing  with  an  infuscate 


Volume  4,  1995  97 

spot  below  stigmal  vein,  rounded  with  truding;  mouth  opening  with  a  semicir- 

speculum  closed,  and  with  a  narrow  costal  cular  incision  below  eye.  Malar  space  1.5X 

cell.  Ratios  length  of  M/PM/S:  4.4/1.0/  as  wide  as  width  of  scape  in  both  sexes. 

1.0.  Propodeum  golden-green;  anterome-  Frons  below  suture  golden-purple  in  fe- 

dian  part  smooth;  surface  with  weak  retic-  male,  golden-green  in   male,  both  sexes 

ulation;  propodeal  callus  with  2  setae.  Pet-  with    weak    reticulation.    Frontal    suture 

iolar  foramen  semicircular.  straight;  lateral  parts  of  frons,  including 

Metasoma:  Petiole  very  short,  in  dorsal  frontal  suture,  considerably  raised  to  form 

view  visible  only  as  a  narrow  and  smooth  2   conspicuous  horns   (Figs  9-11).   Frons 

strip  (as  in  Fig.  75).  Gaster  golden-green  above    suture   and    vertex    golden-green, 

or  golden-purple;  male  with  a  small  pale  golden-red  or  golden-blue,  with  weak  re- 

subbasal    spot;    with    strong    and    small-  ticulation.  Inner  orbit  of  eye  with  1  row  of 

meshed   reticulation;   oval-shaped    in   fe-  setae.   POL/OOL/POO:  2.6/1.8/1.0.   Oc- 

male.  Ratio  length  of  mesosoma/ length  of  cipital  margin  with  a  weak  edge.  Ratio 

gaster  female=  0.8-0.9,  male=  1.0-1.2.  width  of  head /width  of  thorax  across  pos- 

Distribution. — USA  (California).  terior  mesoscutum=  1.2. 

Remarks. — This  species  belongs  in  the  Mesosoma:  Pronotum  without  trans- 
mediana-group,  it  shares  the  narrow  costal  verse  carina.  Mesoscutum  and  scutellum 
cell  in  forewing  and  the  smooth  antero-  golden-green,  golden-blue  or  golden- 
median  part  of  propodeum  with  the  other  red — scutellum  in  some  specimens  with 
species  in  the  group.  anterior  half  purplish;  with  rather  strong 

to  weak  reticulation.  Dorsellum  convex  to 

Chrysocharis  cornigera  n.  sp.  ^^^  ^^^^^1^  ^^^  ^.^1^  2  anterolateral  fov- 

^§^    ~  eas.  Borderline  between  lower  and  upper 

Type   material. — Holotype  female   la-  mesepimeron  straight.  All  coxae  dark  and 

belled  "Canada:  Alberta,  14  km  S  S.  Sas-  metallic  with  strong  reticulation.  Femora 

katchewan  River,  l.vi.l981,  Thormin,  Re-  with  basal  V2  to  %  dark,  with  apical  part 

aney  &  Brouwer,  sand  dune  complex",  in  pale;  tibiae  pale;  tarsi  pale  with  4th  seg- 

CNC.  Paratypes:  8  females  12  males  with  ment    dark.    Wings    hyaline;    forewing 

same  label-data  as  holotype,  3  females  5  rounded    with    speculum    closed.    Ratios 

males  in  coll.  LUZM,  remaining  in  CNC.  length  of  M/PM/S:   6.1/1.0/1.0.   Propo- 

Etymology. — cornigera    is    latin    and  deum    golden-green;    anteromedian    part 

means  "with  horns".  with  a  small  semicircular  fovea;  surface 

Diagnosis. — Lateral  parts  of  frons,  in-  with  rather  strong  to  weak  sculpture/ re- 
cluding  frontal  suture,  considerably  raised  ticulation;  propodeal  callus  with  3  setae, 
to  form  2  conspicuous  horns  (Figs  9-11);  Petiolar  foramen  semicircular, 
clypeus  semicircularly  protruding;  mouth  Metasoma:  Petiole  slightly  transverse 
opening  with  a  semicircular  incision  be-  (as  in  Fig.  68).  Gaster  golden-green;  oval- 
low  eye-mandibles  are  possible  to  point  shaped  in  female.  Ratio  length  of  meso- 
straight  forward  when  fitted  in  these  in-  soma/length  of  gaster  both  sexes=  0.8- 
cisions;  postmarginal  vein  short,  as  long  0.9. 

as   stigmal   vein;   propodeal   callus   with  Distribution. — Canada  (Alberta), 

three  setae.  Remarks. — This  species  comes  closest  to 

Description. — Length  of  bt^dy:  Female=  piibicornis-group,  but  differs  from  the  spe- 

0.9-1.3  mm,  male=  1.1-1.3  mm.  cies  in  this  group  in  the  short  postmargin- 

Head:  Entire  antenna  dark.   Flagellum  al  vein  and  the  protruding  lateral  parts  of 

with  2  apical  segments  fused  (Figs  9,  12).  frons.   Therefore   I   am   not   prepared    to 

HE/MS/MO    female:    4.8/1.0/3.6,    male:  place  corn/'^^t'rrt  in  this,  or  any  other,  group, 

2.8/1.0/2.3.   Clypeus  semicircularly  pro-  but  regard  it  as  species  sola. 


98 


Journal  of  Hymenoptera  Research 


Chrysocharis  sentenaca  n.  sp. 

Type  material. — Holotype  female  la- 
belled "USA:  California,  San  Diego  Coun- 
ty, Sentenac  Canyon,  22. iv. 1981,  J.  Wool- 
ley  81/011",  in  TAMU.  Paratypes:  1  fe- 
male 4  males  with  same  label-data  as  ho- 
lotype, 1  female  2  males  in  coll.  LUZM, 
remaining  in  TAMU. 

Etymology. — Named  after  place  where 
type  specimens  where  collected,  Sentenac 
Canyon  in  California. 

Diagnosis. — Thoracic  dorsum  flattened; 
ocelli  small. 

Description. — Length  of  body:  Female  = 
1.1-1-3  mm,  male=  0.9  mm. 

Head:  Entire  antenna  dark.  Apical  2 
segments  of  flagellum  fused.  HE /MS /MO 
both  sexes:  3.6/1.0/2.2.  Malar  space  1.5X 
as  wide  as  width  of  scape  in  female  and 
1.0  X  in  male.  Frons  metallic  purple  below 
suture,  metallic  greenish-blue  above,  with 
strong  and  small-meshed  reticulation. 
Frontal  suture  V-shaped.  Vertex  golden- 
green,  with  weak  and  small-meshed  retic- 
ulation. Inner  orbit  of  eye  with  1  row  of 
setae.  POL/OOL/POO:  2.8/1.0/1.3.  Oc- 
cipital margm  rounded.  Ratio  width  of 
head /width  of  thorax  across  posterior  me- 
soscutum=  1.2. 

Mesosoma:  Pronotum  without  trans- 
verse carina.  Mesoscutum  and  scutellum 
golden-green;  with  rather  strong  and 
small-mesned  reticulation  on  mesoscu- 
tum, weaker  on  scutellum;  scutellum 
about  as  long  as  wide.  Dorsellum  large, 
2.9  X  as  wide  as  long,  and  almost  flat,  with 
weak  reticulation.  Borderline  between 
lower  and  upper  mesepimeron  weakly 
curved.  All  coxae  dark  and  metallic  with 
weak  reticulation.  Femora  and  tibiae  pale; 
tarsi  pale  with  4th  segment  dark;  except 
weakly  infuscate  fore  tibiae  and  tarsi. 
Forewing  hyaline,  rounded  with  specu- 
lum closed,  and  with  a  narrow  costal  cell. 


Ratios  length  of  M/PM/S:  4.1/1.8/1.0. 
Propodeum  golden-green;  anteromedian 
part  with  a  small  and  weak  fold;  surface 
with  weak  reticulation;  propodeal  callus 
with  2  setae.  Petiolar  foramen  semicircu- 
lar. 

Metasoma:  Petiole  very  short,  in  dorsal 
view  visible  only  as  a  narrow  and  smooth 
strip  (as  in  Fig.  75).  Caster  golden-green 
or  golden-purple;  oval-shaped  in  female. 
Ratio  length  of  mesosoma /length  of  gas- 
ter  female=  0.8-0.9,  male=  0.9. 

Distribution. — USA  (California). 

Remarks. — C.  sentenaca  belongs  to  pen- 
theus-group  and  hence  has  the  characters 
diagnostic  for  that  group  (Hansson  1987: 
49)  and  is  similar  to  C.  suhmutica. 

ACKNOWLEDGMENTS 

Thank  you  to  JT.  Huber  (CNC)  and  an  anonymous 
reviewer  for  comments  on  the  manuscript.  I  am  also 
indebted  to  J.T.  Huber  and  J.B.  Woolley  (TAMU)  for 
loan  of  material,  and  to  B.  Rasmussen  (Zoological 
Museum,  Copenhagen)  for  SEM-work. 

LITERATURE  CITED 

Hansson,  C.  1985.  Taxonomy  and  biology  of  the  Pa- 
learctic  species  of  Chrysocharis  Forster,  1856  (Hy- 
menoptera: Euiophidae).  Entomologica  Scandinav- 
:ca  Siippiemeiit  No.  26:1-130. 

Hansson,  C.  1986.  A  revision  ot  the  Nearctic  species 
of  the  genus  Zaomviomyia  Ashmead  (Hymenop- 
tera: Eulophidae).  Proceedings  of  the  Entomological 
'^-ocietv  in  Washington  88:244-252. 

Hansson,  C.  1987.  Revision  of  the  New  World  spe- 
cies of  Chrysocharis  Forster  (Hymenoptera:  Eulo- 
phidae). Entomologica  Scandinavica  Supplement 
No.  31:1-86. 

Hansson,  C.  1990.  A  taxonomic  study  on  the  Pale- 
arctic  species  of  Chrysonotomyia  Ashmead  and 
Neochrysocharis  Kurdjumov  (Hymenoptera:  Eu- 
lophidae). Entomologica  Scandinavica  21:29-52. 

Hansson,  C.  In  press.  Revision  of  the  Nearctic  spe- 
cies of  Neochrysocharis  Kurdjumov  (Hymenop- 
tera: Eulophidae).  Entomologica  Scandinavica. 

Schauff,  M.E.  1991.  The  Holarctic  genera  of  Entedon- 
inae  (Hymenoptera:  Eulophidae).  Contributions  of 
the  American  Entomological  Institute  26(4):1-109. 


J.  HYM.  RES. 
Vol.  4,  1995,  pp.  99-109 

Review  of  the  World  Species  of  Sigalphiis 

(Hymenoptera:  Braconidae:  Sigalphinae)  and 

Biology  of  Sigalphiis  romeroi,  New  Species 

Michael  J.  Sharkey  and  Daniel  H.  Janzen 

(MJS)  Agriculture  and  Agrifood  Canada,  Centre  for  Land  and  Biological  Resources  Research, 

K.W.  Neatby  Building,  Carling  Avenue,  Ottawa,  Ontario,  KIA  0C6  CANADA; 

(DHJ)  Department  of  Biology,  University  of  Pennsylvania,  Philadelphia,  PA  19104  U.S.A. 


Abstract. — Two  new  species,  Sigalphus  romeroi  Sharkey  from  Costa  Rica  and  Sigalphiis  masoiii 
Sharkey  from  Nepal,  are  described.  The  world  species  are  reviewed,  a  key  to  the  world  species 
is  presented,  and  the  life  history  of  Sigalphus  romeroi  is  described. 


INTRODUCTION  AND  HISTORICAL 
REVIEW 

Until  recently,  there  has  been  little  re- 
search done  on  the  species  of  Sigalphus. 
The  pre-1973  literature  on  Sigalphus  is  cat- 
alogued in  Shenefelt  (1973).  The  more  re- 
cent literature  is  summarized  here.  Tobias 
(1974)  described  Sigalphus  mongolicus  from 
Mongolia.  You,  Zhou  and  Tong  (1991)  de- 
scribed two  putative  new  species  of  Sigal- 
phus from  Hunan  Province  of  China,  viz. 
S.  hunanus  You  and  Tong  and  S.  anomis 
You  and  Tong.  Achterberg  and  Austin 
(1992)  reviewed  the  world  species  of  Si- 
galphus Latreille  and  revised  the  world 
genera  of  Sigalphinae.  Five  species  of  Si- 
galphus were  recognized,  viz.,  S.  bicolor 
(Cresson),  S.  fulvus  Brues,  S.  irrorator  (Fa- 
bricius),  S.  neavei  (Turner),  S.  testaceus 
Granger.  S.  Hunanus  and  S.  anomis  were 
synonymized  with  S.  irrorator.  Achterberg 
and  Austin  apparently  overlooked  S.  mon- 
golicus Tobias  (1974).  He  and  Chen  (1993) 
described  two  new  species  from  China,  Si- 
galphus flavistigmus  He  and  Chen  and  Si- 
galphus nigripes  He  and  Chen.  You  and 
Zhou  (1994)  revised  the  Chinese  species  of 
Sigalphus.  They  disagreed  with  Achterberg 
and  Austin's  (1992)  synonymies  and  elect- 
ed to  reestablish  S.  hunanus  and  S.  anomis, 


suggesting  that  these  two  may  represent 
only  one  species.  He,  Chen,  and  Ma  (1994) 
synonymized  S.  anomis  and  S.  nigripes  un- 
der S.  hunanus  and  described  tw^o  new 
species,  i.e.,  S.  gyrodontus  He  and  Chen 
and  S.  rufiabdominalis  He  and  Chen. 

In  the  present  paper  the  world  species 
are  reviewed;  a  key  to  the  world  species 
of  Sigalphus  is  presented  incorporating 
two  new  species,  and  the  life  history  of 
Sigalphus  romeroi  n.  sp.  is  described. 

Sigalphus  can  be  distinguished  from  all 
other  genera  of  Braconidae  by  the  pres- 
ence of  the  following  combination  of  char- 
acters: metasomal  terga  1  to  3  forming  a 
carapace,  with  a  free  articulation  between 
metasomal  segments  1  and  2  (Fig.  1);  hind 
wing  with  vein  2CU  present  and  very  pos- 
terior in  position  such  that  the  crossvein 
cu-a  is  less  than  .2  X  the  length  of  vein 
ICU  (Figs  3b). 

Although  this  paper  is  co-authored,  the 
senior  author  is  the  sole  authority  of  the 
new  species  of  Sigalp^hus,  the  junior  author 
is  responsible  for  the  biological  data  of  S. 
romeroi. 

Although  widespread,  Sigalphus  has  few 
species;  intensive  collecting  in  the  New 
World  by  members  and  associates  of  the 
Hymenoptera  Section  of  the  Canadian  Na- 


100 


Journal  of  Hymenoptera  Research 


Fig.  1.     Lateral  aspect  of  Sigalphus  bicolor. 


Volume  4,  1995  101 

tional  Collection  of  Insects,  has  yielded  no  or   characters   and    trivial   morphological 

new  species,  and  none  have  been  found  in  characters.  This  decision  will  await  an  ex- 

the  world's  major  collections.  Generally  amination  of  the  types  and  a  more  com- 

specimens  do  not  appear  to  be  rare,  for  plete  investigatic^n  of  color  variability  in  S. 

example  S.  bicolor  and  S.  irrorntor  are  well  irrorator. 

represented  in  the  best  North  American  He  and  Chen  (1993)  also  described  St- 
and European  collections.  The  discovery  galphus  nigripcs  which  the  senior  author,  in 
of  a  new  species  in  Costa  Rica  (S.  romcwi)  accord  with  He  et  al.  (1994),  considers  to 
may  be  the  last  such  discovery  in  the  New  be  a  junior  synonym  of  S.  irrorator.  He  and 
World.  Chen  (1993)  stated  that  it  is  very  similar 


SYNONYMIES 


to  S.  anomis,  and  indeed,  it  is.  The  char- 
acters of  relative  lengths  of  various  veins 
Having  seen  the  type  species  of  both  S.  and  body  parts  that  were  used  to  separate 
hunnmis  You  and  Tong  and  S.  anomis  You  the  two  supposed  species  are  insignificant 
and  Zhou,  the  senior  author  agrees  with  when  compared  to  the  intraspecific  varia- 
Achterberg  and  Austin  (1992)  that  they  are  tion  found  in  both  S.  irrorator  and  S.  bicol- 
both  junior  synonyms  of  S.  irrorator.  In  a  or.  According  to  He  and  Chen  (1993),  one 
recent  paper  He  and  Chen  (1993)  pro-  specimen  of  S.  nigripcs  was  reared  from 
posed  two  new  species  both  of  which  the  Anomis  flava  (Noctuidae),  the  same  host 
senior  author  also  considers  to  be  junior  species  from  which  the  unique  specimen 
synonyms  of  S.  irrorator.  Although  the  of  S.  anomis  was  reared.  This  shared  host 
type  specimens  have  not  been  made  avail-  further  supports  the  proposed  synonymy 
able  to  the  senior  author  (several  requests  of  S.  nigripes.  In  the  most  recent  of  the  pa- 
were  made),  it  is  clear  from  the  descrip-  pers  published  in  China,  He  et  al.  (1994) 
tion  that  Sigalphiis  flavistigmus  He  and  considered  S.  nigripcs  and  S.  anomis  to  be 
Chen  is  a  junior  synonym  of  S.  mongoliciis.  junior  synonyms  of  S.  hunanus. 
These  two  nominal  species  have  similar  The  senior  author  is  dubious  about  the 
color  patterns  except  that  S.  mongoliciis  has  specific  status  of  the  two  new  species  de- 
the  first  and  second  metasomal  terga  red-  scribed  by  He  et  al.  (1994)  as  these  may 
dish  yellow  whereas  only  the  second  also  be  junior  synonyms  of  S.  irrorator. 
metasomal  tergum  of  S./Znz'/sf/g/?/i/s  is  red-  Both  of  these  nominative  species  share 
dish  yellow.  Based  on  color  variation  in  with  S.  irrorator  (including  those  entities 
members  of  S.  bicolor,  of  which  the  senior  synonymized  with  S.  irrorator)  a  unique 
author  has  seen  a  large  series  of  Individ-  state  of  sculpture  on  the  second  metaso- 
uals,  the  color  difference  between  S.  mon-  mal  tergum.  This  is  a  central  longitudinal 
golicus  and  S.  flavistigmus  is  well  within  groove,  which  is  bordered  by  two  carinae. 
the  probable  range  of  intraspecific  varia-  This  groove  is  divided  medially  by  a  weak 
tion.  Not  only  are  there  no  structural  char-  central  keel  and  the  entire  depression  is 
acters  that  sufficiently  separate  the  two  areolate-rugose.  S.  gi/rodontus  and  S.  rii- 
species,  but  the  unique  specimens  of  each  fiabdominalis  differ  from  specimens  of  S.  />- 
supposed  species  were  found  in  close  rorator  that  the  senior  author  has  seen  in 
proximity  to  each  other,  i.e.,  north  central  that  the  two  spines  at  the  apex  of  meta- 
China  and  Mongolia,  further  supporting  somal  tergum  3  are  not  as  sharp  as  those 
their  conspecific  identity.  In  their  recent  of  S.  irrorator.  The  variation  in  this  char- 
revision  of  the  Chinese  species.  He  et  al.  acter  in  S.  irrorator  does  not  appear  to  be 
did  not  consider  S.  mongoliciis.  It  is  the  se-  great,  but  series  of  specimens  from  the 
nior  author's  opinion  that  S.  mongoliciis  it-  east  Palaearctic  must  be  checked.  These 
self  is  likely  synonymous  with  S.  irrorator  two  species  are  therefore  provisionally  ac- 
because  they  apparently  differ  only  in  col-  cepted. 


^Q2  Journal  of  Hymenoptera  Research 

PHYLOGENETIC  CONSIDERATIONS  is  another  character  that  was  not  included 

in  the  Achterberg  and  Austin  data  set  but 

No  hypothesis  of  the  phylogenetic  re-  ^^^^^^  .^  ^^^  ^^^^^  ^^  ^^^  cladogram  a 

lationships  of  Sigalphus  species  has  been  posteriori.  An  assessment  of  the  distribu- 

published.  Achterberg  and  Austin  (1992)  ^.^^  ^^  ^j^.^  character  state  shows  that  the 

analyzed  the  genera  of  the  Sigalphinae.  ^^^^^i^^j.^  of  the  non-Ethiopian  species 

Despite  this  valiant  first  attempt  at  ap-  ^^  Sigalphus  are  relatively  more  costate 

proximating  relationships  among  the  gen-  ^^^^  ^^^^^  ^f  the  Ethiopian  species  which 

era,  the  monophyly  of  Sigalphus  is  ques-  ^^^  mostly  rugose,  as  in  species  of  Min- 

tionable  due  to  the  inclusion  of  the  Ethi-  ^^^^  ^^^  Acampsis.  Again,  this  character 

opian  species  (S.  fiilvus,  S.  testaceus,  and  S.  ^^^^^  ^^es  not  support  the  monophyly  of 

neavei).  Achterberg  and  Austin  (1992)  in  sigalphus. 

their  cladogram  of  sigalphine  genera  cite  3    -p^j-gt  tergite  with  complete  and 
4  synapomorphies  for  the  genus;  they  are  g^^ong  dorsal  carina."  Although  this  char- 
all  suspect.  These  will  be  treated  here  in  ^^^^^  ^^^^^  -g  included  in  the  data  matrix, 
the  order  they  appear  on  the  cladogram.  ^^^  cladogram  does  not  present  the  most 
1.  "Third   tergite  with   lobes   or   teeth  parsimonious  distribution.  The  data  ma- 
ventro-posteriorly."  This  character  state  is  ^^^^  j^g^g  ^j^g  following  taxa  as  possessing 
not  possessed  by  any  of  the  three  Ethio-  ^^^  derived  character  state:  Sigalphus,  Min- 
pian  species,  conversely  it  is  found  in  all  ^^^^^  Acampsis,  and  Afrocampsis.  This  dis- 
members of  Aposigalphus  and   Notosigal-  tribution  is  correct,  but  on  the  cladogram 
phus  in  a  condition  almost  identical  to  that  Achterberg  and  Austin  indicated  that  the 
of  S.  romeroi  (Fig.  2a).  Although  the  char-  character  state  is  an  autapomorphy  for  Si- 
acter  state  appears  on  the  cladogram  of  galphus  and  ignore  the  rest  of  the  character 
Achterberg  and  Austin  it  is  not  included  g^^te  distribution.  There  are  several  parsi- 
in  their  data  set.  Species  of  Minanga  also  monious  reconstructions  of  the  character 
have  teeth  ventro-posteriorly  on  the  third  \y^^^  ^n  indicate  that  the  presence  of  strong 
tergum  but  since  these  do  not  emanate  dorsal  carina  on  the  first  tergite  is  plesiom- 
from  the  margin  of  the  tergum  as  they  do  orphic  with  reference  to  Sigalphus. 
in  the  aforementioned  groups  their  ho-  4    "Face  with  shallow  semicircular  de- 
mology  is  doubtful.  The  character  state  pression."  This  depression  apparently  re- 
has  three  possible  origins  given  the  phy-  f^j-g  to  the  two  longitudinal  depressions 
logeny  of  Achterberg  and  Austin,  viz.  a)  a  t^gt  define  the  raised  medial  area  of  the 
synapomorphy  for  Sigalphus  +  (Aposigal-  f^^e.  Some  specimens  of  S.  irrorator  appear 
phus   +  Notosigalphus)  with  a  loss  in  the  ^  have  these  depressions  somewhat  ex- 
Ethiopian  species  of  Sigalphus,  b)  a  syna-  panded  laterally  but  in  other  species  of  Si- 
pomorphy  for  the  non-Ethiopian  species  galphus  there  is  no  discernable  difference 
of  Sigalphus  and  a  synapomorphy  (conver-  between  these  depressions  and  those  of 
gence)  for  Aposigalphus  +  Notosigalphus,  or  most  other  genera  of  Sigalphinae  except 
c)  a  synapomorphy  for  Minanga  +  (Sigal-  Afrocampsis  which  does  not  have  them. 
phus    +    (Aposigalphus    +    Notosigalphus))  in  summary,  there  is  no  evidence  for  the 
with  a  loss  in  the  Ethiopian  species  of  Si-  monophyly  of  Sigalphus  as  presently  de- 
galphus.  This  last  hypothesis  assumes  that  fined.  That  all  four  of  the  putative  syna- 
the  condition  in  Minanga  and  the  other  pomorphies  proposed  by  Achterberg  and 
genera  is  homologous.  None  of  these  char-  Austin  are  incorrect  suggests  that  their  en- 
acter  state  distributions  provides  support  tire  analysis  is  suspect.  We  do  not  wish  to 
for  the  concept  of  Sigalphus  including  the  address  generic  concepts  in  this  paper, 
Ethiopian  species.  only  to  draw  attention  to  the  fact  that  care 
2.  "Strongly  costate  metapleuron."  This  must  be   taken  in   interpreting  their  re- 


Volume  4,  1995 


103 


b 


Fig.  2.  a,  posterior  aspect  of  metasoma  of  Sigalphus  nmieroi  showing  ridges  on  ventral  apex  (d  =  denticles); 
b,  dorsal  aspect  of  metasomal  tergum  2  of  Sigalpliiis  masoni;  c,  posteroventral  aspect  of  metasoma  of  Sigalpluis 
nmsoiii  showing  denticles  at  ventral  apex  (d  =  denticles,  o  =  ovipositor,  v.f.  =  ventral  foramen);  d,  dorsal 
aspect  of  metasoma  of  Sigalphus  romcwi. 


104 


Journal  of  Hymenoptera  Research 


^V««Kvv,5*^<^VV*S^'^''"''^'' 


,,<^x>'/''- 


.^yyy-^y-^yyy^ 


,^^- 


m-cu 


U  CU-a  ^^^^^^VVVXVVXVSVVV,V.VNV 


.v>S-^" 


.vV-V---^ 


^v^*- 


.v=N^^ 


*»»-^\\V«^^CSVV»^XVVNXVV^^ 


JJ.'tN"" 


"^"^^^--V-..ssvv.vs^-^-^ 


Fig.  3.     a,  fore  wing  of  Sigalphiis  romeroi;  h,  hind  wing  of  Sigalphus  romeroi;  c,  fore  wing  of  Sigalphus  inasoni. 

suits.  A  phylogenetic  analysis  of  the  spe-  paper.  The  non— Ethiopian  members  of  Si- 
des of  Sigalphus  must  be  preceded  by  a  galpJius  may  be  monophyletic,  based  on 
reexamination  of  the  relationships  of  the  the  presence  of  denticles  at  the  ventropos- 
genera  and  this  is  beyond  the  scope  of  this  terior  margin  of  tergum  3. 


KEY  TO  WORLD  SPECIES  OF  SIGALPHUS 

1.  New  World  species 2 

1.'  Old  World  species 3 

2.  Metasoma  orange;  posterior  flange  of  ventral  foramen  of  metasoma  smooth  apically,  ridg- 
es reduced  to  small  crenulae  not  reaching  apex  of  flange  (south  eastern  Canada  and  east- 
ern U.  S.  A.  south  to  Florida) S.  bicolor  (Cresson) 


Volume  4,  1995 


105 


2'.  Metasoma  black;  posterior  flange  of  ventral  foramen  of  metasoma  with  ridges  extending 
to  apex  (Fig.  2a)  (known  only  from  the  type  locality  in  northwestern  Costa  Rica,  but  the 
two  species  of  noctuid  hosts  occur  throughout  the  Mesoamerican  Pacific  dry  deciduous 
forest  and  this  parasitoid  may  be  found  there  as  well) S.  romeroi  n.  sp. 

3.  Head  and  mesosoma  entirely  black    4 

3.'  Head  and  mesosoma  not  enhrely  black,  rather  with  pale  coloration   9 

4.  Apex  of  ventral  foramen  of  metasoma  lacking  denticles  and  two  rounded  lobes  (southern 
Africa) S.  neavei  (Turner) 

4'.  Apex  of  ventral  foramen  of  metasoma  with  two  denticles  or  two  rounded  lobes  (Fig.  2c) 
(Oriental,  Palaearctic) 5 

5.  Apex  of  ventral  foramen  of  metasoma  with  two  sharp  denticles 6 

5'.  Apex  of  ventral  foramen  of  metasoma  with  two  rounded  lobes 7 

6.  Metasoma  entirely  black;  stigma  black  (Palaearctic)    S.  irrorator  (Fabricius) 

6'.  Metasoma  orange  and  black;  stigma  yellow  (Mongolia  and  northern  China)    

S.  tuougolicus  Tobias 

7.  Tergum  2  with  two  parallel  median  longitudinal  carinae  with  a  median  longitudmal 
groove  between  them 8 

7'.  Tergum  2  without  two  parallel  median  longitudinal  carinae  and  without  a  median  lon- 
gitudinal groove S.  masoni  n.  sp. 

8.  Fore  wing  entirely  melanic-infuscate;  tergum  3  as  wide  as  tergum  2 

S.  gyrodontus  He  and  Chen 

8'.  Fore  wing  melanic-infuscate  in  distal  half  only;  tergum  3  wider  than  tergum  2 

S.  rtifiabdominalis  He  and  Chen 

9.  Vertex  of  head  and  apex  of  hind  tibia  black  (southern  Africa) S.  fiilviis  Brues. 

9'.  Head  and  hind  tibia  entirely  reddish  brown  (Madagascar) S.  testaceus  Granger. 


Sigalphus  romeroi  Sharkey  n.  sp. 
(Figs  2a,  d,  3a,  b) 

MALE. 

Length. — 8.4  mm. 

Color. — Entirely  black  except  fore  tarsus 
brown,  wings  evenly  infuscate. 

Head. — Antenna  with  42-44  flagello- 
meres. 

Mesosoma. — Pronotum  smooth  except 
for  weak  crenulae  along  posterior  border; 
subpronope  present;  notaulus  deeply  im- 
pressed and  with  weak  punctures;  scutel- 
lar  furrow  (sulcus)  smooth  except  for  me- 
dian longitudinal  carina;  posterior  scutel- 
lar  depression  composed  of  two  large  pits; 
median  depression  of  metanotum  semicir- 
cular (rounded  posteriorly);  propodeum 
entirely  areolate,  with  pair  of  strong  lon- 
gitudinal carinae  defining  medial  area; 
sternaulus  complete  to  epicnemial  carina; 
fore  wing  (Fig.  3a)  with  veins  M  and  m- 
cu  not  distinctly  converging  anteriorly; 
crossvein  a  of  fore  wing  not  present;  sec- 


ondary crossveins  r  and  m-cu  of  hind 
wing  present  though  weak  and  not  tubu- 
lar (Fig.  3b). 

Metasoma  (Figs  2a,  d). — All  terga  rugose; 
first  tergum  with  two  precurrent  longitu- 
dinal carinae;  second  tergum  with  median 
longitudinal  carina,  and  without  deep  an- 
terolateral depressions  (as  in  S.  bicolor); 
tergum  3  lacking  long  dense  setae;  ventral 
margins  of  carapace  crenulate;  apex  of 
ventral  metasomal  foramen  with  6  riciges 
forming  weak,  sharp,  denticles  (Fig.  2a); 
ventral  foramen  of  carapace  not  extending 
to  apex  of  metasoma,  i.e.,  tergum  3  curved 
around  apex  of  metasoma  forming  poste- 
rior surface. 

Material  Examined. — Holotype  male: 
COSTA  RICA,  Guanacaste  Province, 
Guanacaste  Conservation  Area,  Sector 
Santa  Rosa;  300  m.  Janzen  rearing  data 
base  voucher  number  90-SRNP-828.  Lar- 
val host  Epithisanotia  saiictijolunuia^i  (Noc- 
tuidae)    collected    24    June    1990,    wasp 


106 


Journal  of  Hymenoptera  Research 


eclosed  26  May  1991.  Institute  Nacional  de 
Biodiversidad  (INBio),  Santo  Domingo  de 
Heredia,  Costa  Rica. 

Paratype  male:  COSTA  RICA,  Guana- 
caste  Province,  Guanacaste  Conservation 
Area,  Sector  Santa  Rosa,  300m.  Janzen 
rearing  data  base  voucher  number  92- 
SRNP-2777.  Larval  host  Erochia  miimmia 
(species  complex)  (Noctuidae)  collected  7 
July  1992,  wasp  enclosed  10  May  1993.  Ca- 
nadian National  Collection,  Ottawa,  Can- 
ada. 

Remarks. — The  female  of  this  species  is 
unknown.  S.  romeroi  is  easily  distin- 
guished from  the  only  other  New  World 
species,  S.  bicolor  by  the  characters  given 
in  the  key. 

Eti/mologi/. — The  species  is  named  in 
honor  of  Sra.  Luz  Maria  Romero  in  rec- 
ognition of  her  outstanding  drive  and  in- 
spiration in  guiding  and  developing  the 
Biological  Education  Program  of  the 
Guanacaste  Conservation  Area  in  north- 
western Costa  Rica. 

Biogeographical  Notes. — This  is  the  first 
record  of  members  of  the  genus  Signlphiis 
in  the  Neotropics  and  only  the  second  spe- 
cies to  be  discovered  in  the  New  World. 
The  genus  is  unknown  in  South  America. 

As  suggested  in  the  section  on  phyloge- 
ny,  the  non- Ethiopian  members  of  Sigal- 
phus  may  be  monophyletic  based  on  the 
presence  of  denticles  at  the  ventroposterior 
margin  of  tergum  3.  If  this  is  so,  the  distri- 
bution of  the  New  World  species  is  inter- 
esting from  a  biogeographic  perspective. 

It  is  generally  agreed  that  the  closest 
phylogenetic  relationships  of  most  tem- 
perate North  American  taxa  are  found  in 
the  northern  hemisphere  of  the  Old  World 
(Lavin  and  Luckow  1993).  In  contrast  the 
usual  hypothesis  put  forward  to  explain 
North  American  tropical  fauna  is  dispers- 
al from  South  America. 

If  the  non-Ethiopian  members  of  Signl- 
phus  are  monophyletic,  it  suggests  a  north- 
ern hemisphere  origin  of  S.  romeroi,  the 
only  Neotropical  member  of  the  genus. 
This  distribution  is  consistent  with  what 


has  been  referred  to  as  the  boreotropics 
hypothesis,  which  suggests  a  historically 
more  widespread  tropical  northern  hemi- 
sphere biota.  (Wolfe  1975;  Tiffney  1985a, 
b;  Lavin  and  Luckow  1993). 

Biology. — The  larval  hosts,  Epithisanotia 
snnctijohannasi  and  Erochia  miimmia  (both: 
Noctuidae,  Agaristinae),  are  common  to 
exceedingly  rare  (depending  on  the  year) 
and  univoltine  in  the  primary  to  early  suc- 
cessional  dry  forests  in  eastern  Sector 
Santa  Rosa  of  the  Guanacaste  Conserva- 
tion Area  in  northwestern  Costa  Rica 
(200-300m  elevation;  about  35  km  north  of 
the  town  of  Liberia,  to  the  west  of  the  In- 
teramerican  Highway)  (see  Janzen  1987, 
1988a,  1988b,  1993  for  a  general  site  de- 
scription of  this  forest  and  its  caterpillars). 
E.  sanctijohannasi  and  E.  miimmia  are  most 
easily  located  when  they  are  feeding  on 
the  large  woody  vine  Tetracera  vohibilis 
(Dilleniaceae,  Dilleniales)  (hundreds  of  re- 
cords), and  £.  sanctijohannasi  has  also  been 
found  feeding  on  Curatela  americana  (4  rec- 
ords). These  larvae  eat  mature  leaves  and 
are  non-gregarious  feeders  in  the  penulti- 
mate and  ultimate  instars  (earlier  instars 
unknown)  on  both  host  plants.  Both  spe- 
cies are  univoltine  (last  instar  caterpillars 
occurring  in  late  June  and  early  July,  the 
second  and  third  months  of  the  rainy  sea- 
son) and  the  pupae  remain  dormant 
through  the  remainder  of  the  six  month 
rainy  season  and  all  of  the  dry  season, 
with  the  moths  eclosing  in  May  at  the  be- 
ginning of  the  rains  (see  Janzen  1987, 
1988a,  1993  for  other  examples  of  the  same 
phenomenon). 

Of  36  £.  sanctijohannasi  31  £.  miimmia 
last  instar  larvae  collected  and  reared  from 
T.  voliibilis  in  1992  and  1993,  84  and  93 
percent  respectively  produced  last  instar 
larvae  of  Sigalphus  romeroi.  This  extremely 
high  percent  of  parasitization  is  not  rep- 
resentative of  caterpillar-parasitoid  inter- 
actions in  this  forest,  where  1-30%  para- 
sitization by  a  given  species  of  parasitoid 
is  usually  recorded  (D.  H.  Janzen,  unpub- 
lished  rearing   records).   The   very   large 


Volume  4,  1995 


107 


number  of  Sigalphiis  romeroi  larvae  did  not 
generate  a  large  paratype  series  because, 
as  explained  below,  the  larvae  proved  to 
be  very  incompetent  at  spinning  cocoons 
and  surviving  subsequently  under  artifi- 
cial rearing  conditions. 

The  parasitized  last  instar  caterpillars  of 
£.  sanctijohannnsi  and  E.  uiiimmia  appear  to 
be  perfectly  normal  in  feeding,  fleeing  and 
walking  behaviour.  The  prepupal  caterpil- 
lar searches  in  the  litter  for  a  piece  of  rot- 
ting wood,  tunnels  into  the  wood,  and 
makes  its  pupal  chamber  there.  Within  1- 
3  days  of  the  caterpillar  entering  the  wood 
the  wasp  larva  eats  nearly  all  of  the  con- 
tents of  the  prepupal  caterpillar  and  exits 
the  cadaver.  The  wasp  larva  is  so  large 
that  there  does  not  appear  to  be  sufficient 
food  for  more  than  one  per  caterpillar  of 
either  host.  The  wasp  larva  then  spins  an 
ovoid  white  cocoon  in  the  caterpillar's  pu- 
pal chamber  (late  June  to  early  July), 
where  it  remains  dormant  as  a  prepupal 
larva  until  the  following  May,  when  it  pu- 
pates and  exits  through  a  round  hole  cut 
in  the  cocoon,  and  then  burrows  out 
through  the  wood  chip-filled  entrance 
tunnel  made  initially  by  the  caterpillar. 

The  biology  described  in  the  previous 
paragraphs  is  based  on  observation  of  all 
of  this  behaviour  with  captive  caterpillars 
individually  confined  in  large  plastic  bags 
and  bottles  in  an  open  air  laboratory  a  few 
meters  from  the  forest  where  the  caterpil- 
lars and  wasps  are  found  naturally.  Dor- 
mancy of  moth  pupae  and  of  wasp  larvae 
in  their  cocoons  occurred  in  the  individ- 
uals confined  under  three  conditions  of 
captivity,  i.e.,  under  moist  conditions 
throughout  the  wet  season  with  a  change 
to  dry  conditions  during  the  dry  season, 
entirely  under  dry  conditions  throught^ut 
the  wet  and  dry  seasons,  and  for  those 
captured  in  the  dry  season  they  remained 
in  a  dry  environment  in  captivity.  Moths 
in  the  pupal  stage  and  wasp  larvae  both 
use  some  environmental  cue  that  is  per- 
ceivable through  the  walls  of  a  plastic  bag 
to    initiate    further   development    in    late 


April  or  early  May,  and  this  cue  is  most 
likely  the  abrupt  drop  in  temperature  that 
accompanies  the  first  rains  (see  Janzen 
1993).  By  eclosing  at  the  same  time  as  the 
moths,  the  adult  wasps  are  presumably 
present  at  exactly  the  same  time  as  are  the 
first  instar  larvae  of  the  next  generation  of 
caterpillars  which,  extrapolating  from  the 
known  biology  of  S.  bicolor,  is  the  stage 
most  likely  to  be  attacked. 

The  larva  of  Sigalphus  romeroi  is  excep- 
tional, among  the  species  of  parasitoid 
wasps  that  the  junior  author  has  reared  in 
this  forest,  in  its  inability  to  spin  a  cocoon 
except  in  the  pupal  chamber  of  its  host.  It 
was  usually  not  able  to  spin  a  cocoon  be- 
tween leaves  in  the  litter,  among  crum- 
pled layers  of  dry  or  moist  toilet  tissue,  or 
in  tubular  chambers  of  toilet  tissue.  Of  ten 
larvae  that  did  succeed  in  spinning  co- 
coons among  toilet  tissue  layers,  all  died 
of  unknown  causes  over  the  following 
seven  months. 

Malaise  traps  run  for  the  equivalent  of 
tens  of  years  in  this  dry  forest  between 
1984  and  1988  (e.g.,  Gauld  1988,  Gaston 
and  Gauld  1993)  have  captured  no  speci- 
mens of  S.  romeroi  (I.  D.  Gauld  and  P.  A. 
Mitchell,  personal  communication).  This 
result  is  particularly  striking  given  that 
the  Malaise  traps  were  located  within  sev- 
eral meters  of  the  host  plants  and  in  all 
ages  of  forest  succession. 

In  contrast  to  S.  romeroi  being  large,  sol- 
itary, and  univoltine,  Signlplnis  bicolor,  the 
only  North  American  member  of  the  ge- 
nus, is  small,  gregarious  and  multivoltine. 
S.  bicolor  oviposits  in  the  first  instar  larvae 
of  a  noctuid  caterpillar,  Acronicta  clarescens 
(=  Apatela  clnrescetis),  and  the  larv^ae  come 
out  of  the  prepupal  caterpillar  and  spin 
their  cocoons  within  the  host  cocoon 
(Cushman  1913,  who  described  the  wasp's 
natural  history  under  the  name  Sphaeropyx 
bicolor).  The  only  western  Palaearctic  spe- 
cies of  the  genus,  S.  irrorator  has  also  been 
reared  from  Acronicta  in  Europe,  but  be- 
cause of  its  large  size  (about  the  same  as  S. 
romeroi)   it   is    inferred    to   be   solitary   (S. 


108 


Journal  of  Hymenoptera  Research 


Ward,  personal  communication).  You, 
Zhou  and  Tong  (1991)  report  S.  irrorator  (as 
S.  anomis)  as  a  parasitoid  of  Anomis  flava 
(Noctuidae).  The  biology  of  all  other  spe- 
cies of  Sigalphus  is  unknown. 

Sigalphus  masoni  Sharkey  n.  sp. 
(Figs.  2b,  c,  3c) 

HOLOTYPE  FEMALE. 

Length. — 10.9  mm. 

Color. — Mostly  black  with  yellow  on 
bases  and  ventral  sides  of  all  tibiae  and  on 
metasomal  sterna,  and  orange  on  meta- 
somal  terga  1,  2,  and  anterior  %  of  tergum 
3;  fore  wing  entirely  infuscate  though 
somewhat  paler  basally;  hind  wing  clear 
in  basal  Vs,  infuscate  in  distal  %. 

Head. — Antenna  broken. 

Mesosoma. — Pronotum  weakly  rugose; 
subpronope  present;  notaulus  deeply  im- 
pressed and  crenulate;  scutellar  furrow 
(sulcus)  smooth  except  for  three  longitu- 
dinal carinae;  posterior  scutellar  depres- 
sion composed  of  many  small  pits;  medi- 
an depression  of  metanotum  sharply  tri- 
angular (acute  posteriorly);  propodeum 
weakly  rugose  laterally  with  pair  of  lon- 
gitudinal carinae  defining  medial  area; 
sternaulus  complete  to  epicnemial  carina; 
fore  wing  with  veins  M  and  m-cu  distinct- 
ly converging  anteriorly;  crossvein  a  of 
fore  wing  present  and  distinct;  hind  wing 
with  secondary  crossvein  r  present  and 
secondary  crossvein  m-cu  absent. 

Metasomn  (Figs  2b,  c). — All  terga  rugose; 
first  tergum  with  two  precurrent  longitu- 
dinal carinae;  second  tergum  with  median 
longitudinal  carina  and  an  equally  strong 
pair  of  lateral  longitudinal  carinae  that 
converge  towards  the  mid-line  posterior- 
ly; second  tergum  with  moderately  deep 
anterolateral  depressions  (as  in  S.  bicolor); 
tergum  3  with  long  dense  setae;  ventral 
margins  of  carapace  crenulate;  apex  of 
ventral  metasomal  foramen  with  two 
rounded  denticles  (Fig.  2c);  ventral  fora- 
men of  carapace  (Fig.  2c)  not  extending  to 
apex  of  metasoma. 

Material   Examined. — Holotype    9:    NE- 


PAL: Katmandu,  Godavari,  5,000  ft.  (1524 
m.),  6. VIII. 1967,  Canadian  Nepal  Expedi- 
tion, [Canadian  National  Collection,  Otta- 
wa, Canada] 

Biology. — Unknown 

Remarks. — Members  of  S.  masoni  are 
similar  to  those  of  S.  irrorator  and  S.  mon- 
golicus  in  some  respects,  particularly  in 
their  general  size  and  the  long  setae  on 
metasomal  tergum  3.  In  S.  masoni,  these 
setae  are  longer  and  denser  than  in  the 
New  World  species  of  Sigalphus,  though 
they  are  not  as  exaggerated  as  those  of  S. 
irrorator,  nor  are  they  gold  colored.  Mem- 
bers of  the  two  species  differ  in  color, 
those  of  S.  masoni  having  the  metasoma 
predominantly  orange  versus  entirely 
black  in  S.  irrorator.  The  species  also  differ 
in  the  shape  of  the  denticles  at  the  apex  of 
the  ventral  foramen,  i.e.,  those  of  S.  masoni 
are  wide  and  blunt  whereas  those  of  S.  ir- 
rorator and  S.  mongolicus  are  relatively  nar- 
row and  acute.  The  most  distinctive  fea- 
ture distinguishing  S.  masoni  from  other 
species  of  Sigalphus  is  the  sculpture  of  the 
second  metasomal  tergum.  All  other  Pa- 
laearctic  and  Oriental  species  of  Sigalphus 
have  a  median  longitudinal  groove  bor- 
dered laterally  by  longitudinal  carinae. 
These  are  not  present  in  S.  masoni  (Fig. 
2d). 

Etymology. — The  species  is  named  after 
Dr.  W.  R.  M.  Mason  for  his  many  contri- 
butions to  Hymenopterology,  and  also, 
though  it  is  not  stated  on  the  label,  be- 
cause he  is  thought  to  be  the  collector  of 
the  unique  specimen. 

ACKNOWLEDGMENTS 

This  study  was  supported  by  NSF  grant  BSR  90- 
24770  and  DEB  94-00829  to  D.  H.  Janzen.  Junhua  He 
is  especially  acknowledged  for  the  loan  of  the  type 
specimens  of  S.  hunanus  and  S.  anoniis.  Sondra  Ward 
and  Ian  Gauld  are  gratefully  acknowledged  for  hav- 
ing expressed  great  excitement  over  the  first  speci- 
men reared.  We  thank  Ian  Gauld  for  having  stimu- 
lated this  dry  forest  parasitoid  study  in  the  first  place. 
We  thank  R.  W.  Poole  for  identifying  the  moths.  Hen- 
ri Goulet  drew  and  Susan  Rigby  inked  the  habitus  of 
S.  bicolor  which  was  first  published  in  the  "The  Hy- 
menoptera Of  The  World"  Goulet  and  Huber  (1993). 


I 


Volume  4,  1995 


109 


Jason  Hardis  drew  the  fore  wing  S.  masoni.  All  other 
drawings  were  done  by  Barry  Flahey.  We  thank  Da- 
vid Althoff,  Paul  Marsh,  Lubomir  Masner,  Eric  Rick- 
ey, Ales  Smetana  and  two  anonymous  reviewers  for 
valuable  comments  on  early  drafts. 

LITERATURE  CITED 

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of  the  genera  of  the  subfamily  Sigalphinae  (Hy- 
menoptera:  Braconidae),  including  a  revision  of 
the  Australian  species.  Zoologische  Veiiuvniclingen 
280:1-44. 

Cushman,  R.  A.  1913.  Biological  notes  on  a  few  rare 
or  little  known  parasitic  Hymenoptera.  Proceed- 
ings of  the  Entomological  Society  of  Washington  15: 
153-161. 

Gaston,  K.  J.  and  I.  D.  Gauld  1993.  How  many  spe- 
cies of  pimplines  (Hymenoptera:  Ichneumoni- 
dae)  are  there  in  Costa  Rica?  Journal  of  Tropical 
Ecology  9:491^99. 

Gauld,  I.  D.  1988.  A  survey  of  the  Ophioninae  (Hy- 
menoptera: Ichneumonidae)  of  tropical  Mesoam- 
erica  with  special  reference  to  the  fauna  of  Costa 
Rica.  Bulletin  of  the  British  Museum  (Natural  His- 
tory), Entomology  Series  57  :l-309. 

Goulet,  H.  and  N.B.  Huber.  1993.  Hymenoptera  of  the 
world:  an  identification  guide  to  families.  Research 
Branch  Agriculture  Canada  Publication  1894 /E. 
668pp. 

He,  J.  and  X.  Chen.  1993.  Descriptions  of  two  species 
of  the  genus  Sigalphus  Latreille  from  China  (Hy- 
menoptera: Braconidae:  Sigalphinae).  Acta  Ento- 
mologica  Sinica  34:225-229.  (In  Chinese  with  En- 
glish summary). 

He,  J.,  Chen,  X.  and  Ma  H.  1994.  Revision  of  Sigal- 
phus species  from  China  with  descriptions  of  two 
new  species.  Journal  Zhejiang  Agricultural  Univer- 
sity 20:441-448. 

Janzen,  D.  H.  1987.  How  moths  pass  the  dry  season 
in  a  Costa  Rican  dry  forest.  Insect  Science  and  its 
Application  8:489-500. 


Janzen,  D.  H.  1988a.  Ecological  characterization  of  a 
Costa  Rican  dry  forest  caterpillar  fauna.  Biotro- 
pica  20:120-135. 

Janzen,  D.  H.  1988b.  Guanacaste  National  Park: 
Tropical  ecological  and  biocultural  restoration. 
In:  J.  Cairns,  Jr.,  ed.:  Rehabilitation  of  damaged  eco- 
systems, Vol.  II,  CRC  Press,  Boca  Raton,  Florida, 
pp.  143-192. 

Janzen,  D.  H.  1993.  Caterpillar  seasonality  in  a  Costa 
Rican  dry  forest,  hi:  N.  E.  Stamp  and  T.  M.  Casey 
eds.:  Caterpillars.  Ecological  and  evolutionary  con- 
straints on  foraging,  Chapman  and  Hall,  New 
York,  pp.  448-477. 

Lavin,  M.  and  M.  Luckow.  1993.  Origins  and  rela- 
tionships of  tropical  North  America  in  the  con- 
text of  the  boreotropics  hypothesis.  American 
Journal  of  Botany  80:1-14. 

Shenefelt,  R.D.  1973.  Braconidae  6  Cheloninae.  Hy- 
menopterorum  Catalogus  (n.e.)  10:813-936. 

Tiffney,  B.  H.  1985a.  Perspectives  on  the  origin  of  the 
floristic  similarity  between  eastern  Asia  and  east- 
ern North  America.  Journal  of  the  Arnold  Arbore- 
tum 66:  73-94. 

Tiffney,  B.  H.  1985b.  The  Eocene  North  Atlantic  land 
bridge:  its  importance  in  Tertiary  and  modern 
phytogeography  of  the  northern  hemisphere. 
Journal  of  the  Arnold  Arboretum  66:  243-273. 

Tobias,  V.I.  1974.  Contributions  to  the  fauna  of  Bra- 
conidae (Hymenoptera)  of  Mongolia.  [In  Rus- 
sian]. The  Insects  Of  Mongolia  2:261-274.  Lenin- 
grad Publishing  House. 

You,  L.,  Zhou,  Z.  and  X.  Tong.  1991.  Two  new  spe- 
cies of  Sigalphus  Latreille,  1802  from  Hunan 
Province  (Hymenoptera:  Braconidae:  Cheloni- 
nae). Acta  Entomologica  Sinca  34:225-229. 

You,  L.  and  Z.  Zhou.  1994.  Revision  of  the  genus 
Sigalphus  Latreille,  1802  of  the  subfamily  Sigal- 
phinae (Hymenoptera:  Braconidae)  in  China. 
Journal  of  Hunan  Agricultural  College  20:363-365. 

Wolfe,  J.  A.  1975.  Some  aspects  of  plant  geography 
of  the  northern  hemisphere  during  the  late  Cre- 
taceous and  Tertiary.  Annals  of  the  Missouri  Bo- 
tanical Garden  62:  264-279. 


J.  HYM.  RES. 
Vol.  4,  1995,  pp.  110-120 

Ovipositor  Steering  Mechanisms  in  Braconid  Wasps 

Donald  L.  J.  Quicke,  Mike  Fitton,  Jane  Harris 

(DLJQ)  Department  of  Biology,  Imperial  College  at  Silwood  Park,  Ascot,  Berks  SL5  7PY,  U.K.; 

(MF,  DLJQ)  Department  of  Entomology,  The  Natural  History  Museum,  London  SW7  5BD,  U.K.; 

(JH)  Department  of  Animal  and  Plant  Sciences,  PO  Box  601,  University  of  Sheffield, 

Sheffield  SIO  2UQ,  U.K. 


Abstract. — Ovipositor  features  which  allow  the  ovipositor  tip  to  be  manipulated  and  which  have 
restricted  distributions  among  the  subfamilies  of  Braconidae  are  described  and  illustrated  for  the 
first  time.  Members  of  the  Amicrocentrinae,  Charmontinae,  Helconinae  and  Macrocentrinae  have 
the  rhachis  (the  tongue-like  part  of  the  mechanism  that  interlocks  the  upper  valve  with  the  lower 
ones)  with  an  increased  density  of  scale-like  sculpture  and  often  also  swollen  pre-apically.  This 
modification  leads  to  increased  friction  and  restricted  movement  between  the  upper  and  lower 
valves  and  is  also  associated  with  an  ovipositor  bending  mechanism  in  which  retraction  of  the 
lower  ovipositor  valves  relative  to  the  upper  one  causes  the  ovipositor  tip  to  bend  ventrally.  Many 
members  of  the  subfamilies  Agathidinae  and  Orgilinae  possess  a  pair  of  pre-apical  boss-like  pro- 
jections on  the  upper  ovipositor  valve  (gonapophyses  9)  and  a  corresponding  structure  on  each 
lower  valve  (gonapophyses  8)  that  together  enable  the  wasps  to  bend  and  manipulate  the  ovi- 
positor apex  by  retraction  of  the  lower  ovipositor  valves  relative  to  the  upper  one.  Most  members 
of  the  Doryctinae  have  the  aulax  (the  groove-like  part  of  the  mechanism  that  interlocks  the  lower 
valve  with  the  upper  one)  constricted  opposite  modified  pre-apical,  dorso-lateral  tooth-like  struc- 
tures. These  constrictions  restrict  the  relative  movement  between  the  upper  and  lower  valves  and 
thus  operate  as  an  ovipositor  bending  mechanism.  The  potential  phylogenetic  significance  of  these 
mechanisms  is  discussed,  and  preliminary  parsimony  analyses  are  described  which  suggest  that 
the  pre-apical  boss-like  projections  of  the  Agathidinae  and  Orgilinae  may  represent  a  synapo- 
morphy  uniting  these  two  subfamilies. 


INTRODUCTION 

Oviposition  behaviour  and,  as  a  conse- 
quence, the  ovipositor  have  been  of  im- 
mense importance  in  the  evolution  of  the 
Hymenoptera,  and  are  central  to  the  par- 
asitic way  of  life  (Gauld  &  Bolton  1988). 
The  commonly  held  vievv^  that  the  ovipos- 
itors of  parasitic  w^asps  are  fairly  simple 
analogues  of  hypodermic  needles  is  there- 
fore likely  to  be  a  considerable  over-sim- 
plification as  has  in  fact  been  shown  by 
several  previous  investigations  of  func- 
tional morphology  (Oeser  1961;  Austin  & 
Browning  1981;  Quicke  et  al.  1992a,b, 
1994). 

Several  braconid  genera  have  recently 
been  shown  to  possess  highly  modified 


ovipositors  whose  tips  are  capable  of  be- 
ing independently  actively  manipulated 
by  the  wasp  even  though  the  ovipositor 
valves  themselves  are  devoid  of  intrinsic 
musculature  and  their  relative  movements 
are  controlled  by  muscles  within  the  me- 
tasoma.  Mechanisms,  as  found  for  exam- 
ple in  the  braconine  genus  Zagli/ptogastra 
Ashmead,  were  apparent  because  of  very 
conspicuous  features  of  the  intact  ovipos- 
itor (Quicke  1991;  Quicke  &  Marsh  1992). 
However,  an  ability  to  manipulate  or  steer 
the  ovipositor  tip  has  been  observed  in  a 
number  of  other  parasitic  Hymenoptera 
whose  ovipositors  appear  more  or  less  un- 
modified externally  (Delanoue  &  Aram- 
bourg  1965;  Compton  &  Nefdt  1988).  Fur- 
ther,   our    own    observations    have    also 


Volume  4,  1995 


111 


shown  that  an  ability  to  manipulate  the 
ovipositor  tip  is  present  in  members  of 
several  subfamilies  of  Braconidae  that 
possess  moderately  long  ovipositors.  We 
have  therefore  examined  ovipositor  tip 
structure  in  detail  across  a  range  of  brac- 
onid  taxa.  Here  we  describe  three  different 
types  of  morphological  specialisation 
found  in  the  Braconidae  that  appear  to  un- 
derlie the  manipulatory  abilities  observed. 
These  modifications  all  utilise  the  same 
mechanical  principle  but  they  achieve  this 
through  modification  of  different  struc- 
tures. In  these  cases,  the  structures  in- 
volved can  often  only  be  revealed  by  ex- 
amination of  separated  ovipositor  valves 
using  scanning  electron  microscopy;  how- 
ever, their  existence  supports  the  view 
that  the  ovipositors  of  parasitic  wasps  are 
highly  specialised  and  complex  structures. 
Two  different  ovipositor  steering  mech- 
anisms, one  involving  a  sinuous  apical  re- 
gion of  the  ovipositor  the  other  involving 
the  development  of  an  aulaciform  rod  be- 
tween the  largely  separated  halves  of  the 
upper  ovipositor  valve  have  been  report- 
ed previously  (Quicke  1991;  Quicke  et  al. 
1994).  The  first  of  these  is  possessed  by 
several  unrelated  taxa  of  Braconidae  and 
Ichneumonidae,  the  second  so  far  is 
known  only  from  members  of  the  Ichneu- 
monidae. During  investigations  of  ovipos- 
itors belonging  to  a  large  number  of  other 
ichneumonid  wasps  collectively  repre- 
senting most  subfamilies,  no  steering  spe- 
cialisations similar  to  any  of  the  three  de- 
scribed here  for  Braconidae  have  been  de- 
tected. Functionally  similar  but  morpho- 
logically different  mechanisms  in  members 
of  two  other  families  of  parasitic  Hyme- 
noptera,  Gasteruptiidae  and  Aulacidae, 
are  being  described  elsewhere  (Quicke  & 
Fitton,  in  press). 

GENERAL  MORPHOLOGY 

The  ovipositors  of  Hymenoptera  are 
comprised  of  components  that  are  be- 
lieved to  be  derived  primitively  from  two 
pairs   of   metasomal   appendages   (Smith 


1968,  1970).  The  two  gonapophyses  of  ab- 
dominal segment  9  are  usually  fused 
along  their  length  and  form  the  single  up- 
per valve,  while  those  of  segment  8  re- 
main separate  and  form  the  two  lower 
valves.  The  upper  valve  is  interlocked 
with  each  lower  valve  by  a  k:)ngitudinal 
tongue  and  groove  joint  referred  to  as  the 
olistheter.  The  tongue  or  rhachis  situated 
ventro-laterally  on  each  side  of  the  upper 
valve  is  T'-shaped  in  transverse  section 
and  in  life  runs  within  the  T'-shaped 
groove  or  aulax  on  the  dorso-lateral  face 
of  the  lower  valve  (Fig.  1). 

MATERIALS  AND  METHODS 

Ovipositors  of  70  species  of  Braconidae, 
collectively  representing  members  of  19  of 
the  40-or-so  currently  recognised  subfam- 
ilies (Quicke  &  van  Achterberg  1990;  Shaw 
&  Huddleston  1991)  have  been  examined 
using  scanning  electron  microscopy.  For 
most  of  these,  the  upper  and  lower  valves 
were  carefully  separated  after  soaking  in- 
tact wasp  metasomas  or  excised  oviposi- 
tors in  either  water  or  dilute  aqueous  mild 
detergent.  In  the  case  of  a  few  specimens 
in  which  one  or  both  lower  ovipositor 
valves  were  protruding  beyond  the  apex 
of  the  dorsal  valve,  the  intact  ovipositor 
was  examined. 

Limited  observ^ations  of  ovipositor  ma- 
nipulation were  made  on  some  hand-held 
living  female  wasps.  In  addition,  experi- 
ments were  carried  out  using  excised  ovi- 
positors from  freshly  killed  wasps.  Ovi- 
positors were  submerged  under  water  and 
the  effects  observed  of  pulling  and  push- 
ing the  cut  ends  of  the  basal  portions  of 
the  upper  and  lower  valves  differentially 
with  fine  forceps. 

The  taxa  investigated  are  listed  below 
according  to  subfamily.  Where  more  than 
one  individual  of  a  species  was  examined 
the  number  is  given  in  parentheses. 

Agathidinae:  Braunsia  sp.,  Cameroun 
(3);  Cnmptothlipsis  sp.,  Kenya;  Mcgngnthis 
sp..  Sierra  Leone;  Microdiis  rufipcs  Nees, 
UK.    Alysiinae:    Phacnocarpa    conspurcator 


112 


Journal  of  Hymenoptera  Research 


2a.  Macrocentrus  type 


o 

< 
< 


1.    Stylized 
TS  ovipositor 


2b.  Agathidine  type 


upper 
valve 


aul 


lower 
valve 


c 
■o 

•D 

n> 

< 
< 

(D 


O 

S 

< 

< 


2c.    Doryctine  type 

upper  valve  retracted 


2d.    Doryctine  type 

upper  valve  protracted 


lower  valve 

upper  valve 
lower  valve 


rh    aul 


Figs.  1,2.  1.  Diagramatic  transverse  section  (TS)  through  mid-region  of  braconid  ovipositor  to  show  olistheter 
mechanism:  (ol)  olistheter,  (rh)  rhachis,  (aul)  aulax.  2.  Diagramatic  representations  of  how  the  three  ovipositor 
bending  mechanisms  are  believed  to  operate,  a,  Macrocentrinae,  Amicrocentrinae,  and  Charmontinae,  lateral 
views  showing  preapical  swelling  of  rhachis;  b,  Agathidinae  and  Orgilinae,  lateral  views  showing  pre-apical 
bosses;  c,  d,  Doryctinae,  dorsal  showing  in  the  postulated  increased  grip  against  the  rhachis  when  the  upper 
valve  is  protracted. 


(Haliday),  Netherlands.  Amicrocentrinae: 
Amicrocentrum  sp.,  Tanzania.  Blacinae:  Ar- 
tiocus  spinarius  Achterberg,  Brazil  (2);  Bla- 
cus  hastatus  Haliday,  Sweden;  B.  longipen- 
nis  (Gravenhorst),  UK;  B.  maculipes  Wes- 
mael,  Austria;  B.  nigricornis  Haeselbarth, 
UK;  B.  paganus  Haliday,  UK;  B.  pallipes 
Haliday,  UK;  B.  ruficornis  Nees,  Hungary, 


Russia  (2);  Dyscoletes  lancifer  (Haliday), 
UK,  Germany  (2).  Braconinae:  Digonogas- 
tra  sp.,  Trinidad;  Eiiurohracon  yokahamae 
Dalla  Torre,  Japan;  Glyptomorpha  sp.,  Gam- 
bia; Nesaiilax  sp.,  Papua  New  Guinea;  Ode- 
sia  sp.,  Kenya;  Virgulihracon  endoxylapha- 
gus  Quicke  &  Ingram,  Australia.  Cardi- 
ochilinae:  Cardiochiles  sp.,  Cameroun.  Gen- 


Volume  4,  1995 


113 


ocoeliinae:  Capitonius  spp.,  Brazil  (2). 
Charmontinae:  Charmon  cruentatus  Haliday, 
UK  (2).  Doryctinae:  Acanthodoryctes  tomen- 
tosa  (Szepligeti),  Australia;  Acropjliasmiis 
fernigineus  (Marsh),  USA;  Allorhogas  pyr- 
alophagus  Marsh,  Trinidad;  Binarea  sp.,  Ar- 
gentina; Doryctes  erythromeles  (Brulle), 
USA;  Liobracon  sp.,  Brazil;  Megaloproctiis 
platyantennus  Marsh,  Brazil;  Monarea  sp., 
Brazil;  Rhaconotus  sp.,  USA;  Rutheia  sp., 
Bolivia;  Syngaster  sp.,  Australia;  Spathiiis 
sp.,  Kenya;  Zombrus  sp.,  Kenya.  Helconi- 
nae:  Austrohelcou  sp.,  Australia;  Aspicolpus 
carinator  Nees,  Hungary;  Aspidocolpus  sp., 
Nigeria;  Baecis  abietis  (Ratzeburg),  Poland; 
Briilleia  latiannulatus  (Cameron),  New 
Guinea;  Calohelcon  obsciiripennis  Turner, 
Australia;  Diospilus  capita  (Nees),  Hungary 
(2);  Diospilus  morosus  Reinhard,  Austria; 
Eubazus  pallipes  Nees,  Romania;  £.  tibialis 
Haliday,  Bulgaria;  £.  (Brachistes)  spp.,  Ar- 
gentina, UK  (2);  Helcon  uunciator  Fabricius, 
Hungary;  Helconidea  tentator  (Fabricius), 
Belgium;  Polydegmon  sinuatus  Foerster, 
Hungary;  Taphaeus  hiator  Thunberg,  Hun- 
gary; Wroiightonia  sp.,  Canada;  unde- 
scribed  genus  A,  Australia;  undescribed 
genus  B,  Australia.  Homolobinae:  Homo- 
lobiis  truncator  (Say),  Bulgaria  (2);  H.  {Oiil- 
ophiis)  sp.,  USA.  Macrocentrinae:  Austroz- 
ele  uniformis  Provancher,  USA;  Macrocen- 
trus  linearis  Nees,  Hungary  (2);  Macrocen- 
triis  sp.,  Poland.  Microgastrinae:  Sathon 
falcatus  (Nees),  Germany.  Microtypinae: 
Microtypus  sp.,  Hungary.  Opiinae:  Doryc- 
tobracon  sp.,  Trinidad.  Orgilinae:  Orgilus 
leptocephnlus  (Hartig),  Germany;  Orgilus 
spp.,  Cameroun,  France  (3);  Orgilonia  sp., 
Cameroun;  Stantonia  sp.,  Cameroun.  Pam- 
bolinae:  Monitoriella  elongatn  Hedqvist, 
USA.  Pselaphaninae:  Pselaphanus  trogoides 
Szepligeti,  Surinam.  Rhysipolinae:  Rhysi- 
polis  sp.,  UK 

RESULTS 

Upper  Valve  Rhachis  Scaling  and 
Expansions 

In   all   Hymenoptera  both   the   rhachis 
and  the  margins  of  the  aulax  are  furnished 


with  scale-like  sculpture  which,  in  the  case 
of  the  parasitic  Apocrita  and  the  Orussi- 
dae,  are  generally  rather  widely  spaced 
(Figs.  16,  17).  In  the  Amicrocentrinae,  Bla- 
cinae,  Charmontinae  and  Helconinae  the 
density  of  scaling  was  found  to  be  consid- 
erably increased  over  a  short  region  pre- 
apically  (Figs.  3-11).  Further,  in  the  Char- 
montinae and  Macrocentrinae  this  is  ac- 
companied by  a  distinct  swelling  of  the 
rhachis  (Figs.  3-6;  arrowed)  and  slight 
widening  of  the  corresponding  length  of 
the  aulax. 

Observation  of  living  Macrocentrus  fe- 
males held  in  the  hand  showed  that  they 
are  capable  of  bending  the  apex  of  their 
ovipositors  ventrally  through  nearly  90°.  It 
seems  likely  that  this  is  achieved  in  an 
analogous  way  to  that  proposed  for  the 
Agathidinae  and  Orgilinae  (see  below), 
with  the  swollen  and  more  coarsely  sculp- 
tured region  of  the  rhachis  providing  in- 
creased resistance  to  relative  movement  of 
the  ovipositor  valves  at  the  apical  part  of 
the  ovipositor.  Thus,  if  the  dorsal  valve  is 
protracted  relative  to  the  ventral  ones, 
they  will  not  be  able  to  slide  past  one  an- 
other freely  at  the  apex,  and  so  the  ovi- 
positor will  be  forced  to  curve  ventrally  in 
order  to  accommodate  the  relative  exten- 
sion of  the  upper  valve  in  a  manner  anal- 
ogous to  the  bending  of  a  bimetal  strip  in 
a  thermostat,  as  illustrated  diagrammati- 
cally  in  Fig.  2a.  The  plausibility  of  this  was 
conclusion  was  substantiated  by  manipu- 
lation of  the  cut  ends  of  an  excised  Macro- 
cent  rus  ovipositor  which  showed  that  pro- 
traction of  the  upper  valve  relative  to  the 
lower  one  causes  a  ventral  curving  of  the 
ovipositor  tip. 

Upper  and  Lower  Valve  Bosses 

In  all  members  of  the  Orgilinae  and 
most  Agathidinae  examined,  the  pre-api- 
cal  lateral  margins  of  both  the  upper  (Figs. 
12,  13,  16,  17)  and  lower  (Figs.  14,  15)  ovi- 
positor valves  possess  an  abrupt  escarp- 
ment-like boss  in  lateral  aspect.  These 
structures  are  located  lateral  tt)  the  olis- 


114 


Journal  of  Hymenoptera  Research 


Figs.  3-6.  Scanning  electron  micrographs  of  apical  parts  of  isolated  upper  ovipositor  valves  showing  preap- 
ical  expansion  of  rhachis  (r)  with  denser  scaling  (arrows).  3,  4  (detail),  Clmrniou  cruoitntus.  5,  Macrocentrus 
linearis.  6,  Macrocentrus  sp.  Scale  line:  3,  5:  50|jim;  4,  6:  25(xm. 


Volume  4,  1995 


115 


Figs.  7-11.  Scanning  electron  micrographs  of  apical  parts  of  isolated  upper  ovipositor  valves  showing  weak 
preapical  expansion  of  rhachis  (r)  with  denser  scaling  (arrows).  7,  8,  Amicrocenirum  sp.  9,  Hckonidea  lentator. 
10,  Wwiightonia  sp.  11,  BniUeia  latmwnilatiis.  Scale  line:  7:  100|xm;  8:  ZOjim;  9,  50n.m;  10,  11:  40^,01. 


theter.  In  the  case  of  the  dorsal  valve,  the 
steep  (scarp)  face  of  the  stop  is  at  the  distal 
end  of  the  modified  region  whereas  it  is 
at  the  proximal  end  in  the  lower  valve. 
The  positioning  of  these  modifications  is 
such  that  when  the  ovipositor  is  in  its  rest- 
ing position  with  neither  upper  nor  lower 
valves  apically  protruding  beyonci  the 
other,  the  scarp  faces  of  both  processes  are 
closely  apposed.  The  presence  of  these 
stops    limits    the    relative    longitudinal 


movement  of  the  upper  and  lower  valves 
such  that  the  upper  valve  cannot  be  pro- 
tracted apically,  relative  to  the  lower 
valves,  beyond  the  resting  position  though 
the  upper  valve  can  be  retracted  basally. 
The  functional  significance  of  this  modifi- 
cation was  revealed  when  living  agathi- 
dines  of  the  genus  Agathis  were  held  in  the 
hand  and  the  apical  part  of  the  ovipositor 
was  observed  to  change  its  degree  of  ven- 
tral curvature  as  the  wasp  attempted  to 


116 


Journal  of  Hymenoptera  Research 


I 


Figs.  12-17.  Scanning  electron  micrographs  of  apical  parts  of  isolated  upper  (12,  13,  16,  17)  and  lower  ovi- 
positor valves  (14,  15)  showing  (arrows)  corresponding  preapical,  bosses  lateral  to  the  rhachis  (r)  and  aulax 
(a)  of  the  upper  and  lower  valves  respectively.  12,  13,  15,  Braunsia  sp.  14,  16,  Megagathis  sp.  17,  Orgilonia  sp. 
Scale  line:  12:  50|jLm;  13,  16:  20|xm;  14,  15:  30|xm;  17:  35|xm. 


free  itself.  Such  bending  appeared  to  be  posing  scarp  faces  of  their  bosses.  Because 

achieved  by  the  wasp  protracting  the  up-  the    valves    are    interlocked    along    their 

per  valve  posteriorly  relative  to  the  lower  whole  length  by  the  olistheter  mechanism, 

valves,  thereby  forcing  together  the  op-  the  ovipositor  is  forced  to  curve  ventrally. 


Volume  4,  1995 


117 


as  illustrated  in  Figure  2b.  The  plausibility 
of  this  was  conclusion  was  substantiated 
by  manipulation  of  the  cut  ends  of  an  ex- 
cised Agnthis  ovipositor  which  showed 
that  protraction  of  the  upper  valve  relative 
to  the  lower  one  causes  a  ventral  curving 
of  the  ovipositor  tip. 

No  other  braconids  examined  possessed 
this  modification  and  since  neither  the 
Agathidinae  nor  the  Orgilinae  are  likely  to 
be  the  most  basal  of  extant  braconids 
(Quicke  &  van  Achterberg  1990;  Wharton 
et  al.  1992;  van  Achterberg  &  Quicke  1992) 
(for  example,  because  of  their  koinobiont 
endoparasitoid  life  history),  it  seems  likely 
that  the  presence  of  a  scarp-like  stop 
mechanism  is  apomorphous.  The  potential 
implications  of  this  are  dealt  with  in  the 
discussion. 

Pre-apical  Constrictions  of  the  Aulax 

Unlike  other  braconid  subfamilies,  most 
members  of  the  subfamily  Doryctinae 
have  the  apex  of  the  ovipositor  much 
darker  (piceous  or  black)  and  much  more 
strongly  sclerotized  than  the  remainder 
(Quicke  et  al.  1992a)  such  that  it  is  usually 
extremely  difficult  to  separate  the  valves 
for  study  even  after  softening  in  aqueous 
potassium  hydroxide;  though  with  persis- 
tent manipulation,  the  three  parts  could 
usually  be  separated.  Scanning  electron 
microscopy  shows  that  in  many  genera 
(e.g.  Binnrea,  Liobracon,  Monarea,  Syngasier, 
Zotnbrus)  the  apical  part  of  the  lower 
valves,  as  well  as  having  ventro-lateral 
serrations,  also  possess  one  or  two  appar- 
ently distinct  dorso-lateral  tooth-like 
structures  (Figs.  18-21;  arrows)  (Quicke  et 
al.  1992a).  The  significance  of  these  in 
terms  of  ovipositor  steering  has  been  re- 
vealed by  examination  of  the  adjacent  part 
of  the  aulax.  The  aulax  is  an  apprc^ximate- 
ly  parallel-sided  groove  along  most  of  its 
length,  in  isolated  ventral  valves  it  is 
strongly  constricted  opposite  these  addi- 
tional teeth  by  an  infolding  of  the  lateral 
margin  (Figs.  22-24).  Together  with  the  in- 
creased sclerotization  of  the  ovipositor  tip 


and  our  observation  that  it  is  very  difficult 
to  separate  the  dorsal  and  ventral  valves 
at  the  tip  region,  we  conclude  that  the  nar- 
rowed region  of  the  aulax  acts  to  grip  the 
rhachis  tightly,  thereby  reducing  the  free- 
dom of  movement  of  the  valves  near  to 
the  ovipositor  tip.  The  asymmetric  ar- 
rangement of  the  constrictions  of  the  aulax 
are  such  that  retraction  of  the  upper 
valves  is  likely  to  afford  less  resistance 
than  protraction  (Figs.  2c  cf  d)  which  will 
cause  the  lateral  wall  of  the  aulax  to  roll 
inwards  thus  increasing  the  friction 
against  the  rhachis.  This  prediction  was 
borne  out  by  observations  of  living  Hcca- 
holiis  females  which  were  found  \.o  be  able 
to  bend  their  ovipositors  ventrally  but  not 
dorsally. 

DISCUSSION 

In  all  the  cases  in  which  we  have  found 
to  possess  probable  ovipositor  steering 
mechanisms,  their  hosts  are  concealed  in- 
sect larvae.  Orgilines  mostly  attack  weakly 
concealed  hosts  such  as  leaf  miners,  and 
they  probe  into  their  hosts'  tunnels  often 
through  frass  holes  rather  than  by  boring 
directly  through  the  plant  material  (Flan- 
ders &  Oatman  1982;  Shaw  &  Huddleston 
1991).  Agathidines,  charmontines  and  ma- 
crocentrines  similarly  attack  weakly  con- 
cealed hosts  such  as  leaf  rollers,  leaf  tiers, 
shoot  borers,  and,  particularly  in  the  case 
of  the  Agathidinae,  leaf  miners /tunnellers. 
The  steering  mechanisms  may  therefore 
enable  the  wasps  to  locate  their  hosts  better 
within  their  concealed  feeding  places.  It 
may  also  enable  the  wasps  to  place  their 
eggs  accurately  on  or  within  their  hosts.  In 
the  case  of  agathidines,  for  example,  it  is 
known  that  eggs  are  usually  placed  very 
precisely  within  or  next  to  the  host's  cen- 
tral nervous  system  (references  in  Shaw  & 
Huddleston  1991).  Given  that  their  hosts 
are  concealed  and  often  relatively  small, 
such  accurate  egg  placement  would  be  fa- 
cilitated by  the  existence  of  some  sort  of 
directional  control  of  the  ovipositor  tip. 

In  contrast  to  the  above  mentioned  sub- 


118 


Journal  of  Hymenoptera  Research 


Figs.  18-24.  Scanning  electron  micrographs  of  apical  parts  of  intact  ovipositors  (18,  19)  and  isolated  lower 
ovipositor  valves  (20-24)  of  doryctine  braconids  showing  dorso-lateral  tooth  like  structures  (arrow)  and  in  24, 
the  corresponding  narrowing  of  the  aulax  (a).  18,  Megalopwctus  plaiyantenniis.  19,  24,  Binarea  sp.  20,  Acroplias- 
mus  ferrugineus.  21,  Spathius  sp.  22,  Syngaster  sp.  23,  Zombrus  sp.  Scale  line:  18:  130(xm;  19:  70|xm;  20,  21:  60|xm; 
22,  23:  30|xm;  24:  10|xm. 


families,  many  members  of  the  Doryctinae 
attack  more  strongly  concealed  insects 
such  as  wood-boring  beetle  larvae,  and 
since  they  are  idiobiont  ectoparasitoids, 
the  exact  placement  of  the  paralysing  sting 
or  of  the  egg  is  usually  likely  to  be  of  little 
importance.  The  process  of  boring 
through  bark  or  other  hard  substrates  may 
not  be  very  accurate  and  is  also  quite  time 
consuming,  which  in  turn  probably  places 
the  wasp  under  an  increased  risk  of  pre- 


dation.  Locating  a  host  tunnel  might 
therefore  be  a  good  first  step  towards 
reaching  a  host  if  the  wasp  can  subse- 
quently manipulate  its  ovipositor  tip. 
Steering  mechanisms  might  be  seen  in 
these  cases  as  an  adaptation  which  over- 
comes some  of  the  potential  problems  of 
reaching  a  concealed  host  directly  through 
a  thick  layer  of  substrate. 

Of  some  surprise  was  that  we  failed  to 
find  any  morphological  modifications  that 


Volume  4,  1995 


119 


could  be  involved  in  ovipositor  steering  in 
a  number  of  taxa  with  long  to  very  long 
ovipositors  such  as  members  of  the  bra- 
conine  genera  Archibracon,  Euurobracou, 
Glyptomorpha,  Nesaidax,  Odesia  and  Virgii- 
libracon.  Nor  were  any  steering  mecha- 
nisms identified  in  the  Alysiinae,  Cardi- 
ochilinae,  or  Cenocoeliinae  examined.  Ob- 
servations of  living  females  of  these  taxa 
would  be  valuable  as  it  is  quite  possible 
that  they  have  some  other,  perhaps  less 
conspicuous,  mechanisms  that  enable 
them  to  manipulate  their  ovipositors  or 
ovipositor  tips.  Certainly  for  many  of  the 
braconines  with  long  ovipositors,  ovipo- 
sition  involves  boring  through  hard  wood 
(van  Achterberg  1986)  and  this  may  pose 
limitations  on  the  bending  mechanisms 
they  might  possess.  Alternatively,  it  may 
be  that  most  braconines  that  attack  deeply 
concealed,  wood-boring  hosts  are  simply 
not  adapted  to  hit  the  host's  tunnel  first 
and  then  to  use  the  ovipositor  to  follow 
the  tunnel  to  the  host,  but  rather  reach 
their  hosts  by  drilling  directly  through  the 
substrate  to  them.  If  this  is  generally  the 
case,  then  the  strong,  possibly  defensive, 
smell  of  most  braconines  (Quicke  1988; 
Quicke  &  Wharton  in  preparation)  could 
be  an  adaptation  to  protect  them  during 
protracted  periods  of  drillng  for  hosts, 
during  which  time  they  may  be  particu- 
larly vulnerable  to  predation. 

Interestingly,  the  Agathidinae  and  Or- 
gilinae  did  not  appear  together  in  the  phy- 
logenetic  analyses  of  Quicke  &  van  Ach- 
terberg (1990)  and  Wharton  et  al.  (1992) 
though  their  relationships  were  rather  la- 
bile. These  two  subfamilies  have  a  similar 
general  appearance  and  perhaps  more  im- 
portantly, have  two  other  putative  syna- 
pomorphies,  a  relatively  narrow  forewing 
marginal  cell  with  an  approximately 
straight  vein  3-SR,  and  in  those  species 
with  a  complete  2nd  submarginal  cell,  this 
is  small  and  usually  triangular  or  at  least 
has  vein  2-SR  very  short.  The  discovery  of 
a  new  apomorphous  character  present  in 
both,  further  suggests  the  possibility  that 


they  could  be  sister  groups,  though  the 
possibility  also  exists  that  the  ovipositor 
bending  mechanisms  possessed  by  mem- 
bers of  the  two  subfamilies  reflect  conver- 
gence as  a  result  of  the  similarities  in  their 
hosts'  biologies.  In  a  preliminary  investi- 
gation, we  have  added  the  presence  or  ab- 
sence of  an  ovipositor  boss  as  an  addition- 
al character  to  a  data  matrix  of  braconid 
subfamilies  based  on  that  of  Quicke  &  van 
Achterberg  (1990)  modified  in  accordance 
with  the  suggestions  of  Wharton  et  al. 
(1992)  and  of  van  Achterberg  &  Quicke 
(1992).  Whereas  the  Agathidinae  and  Or- 
gilinae  did  not  appear  as  sister  groups  in 
the  most  parsimonious  trees  (length  391), 
trees  with  them  so  arranged  were  only 
one  step  longer  than  the  most  parsimoni- 
ous ones  obtained.  The  absence  of  this 
character  in  the  Pselaphaninae  might  be 
taken  to  indicate  that  they  are  not  the  sis- 
ter group  of  the  Agathidinae  (van  Achter- 
berg 1985),  however,  it  should  be  noted 
that  in  Pselaphanus,  the  ovipositor  is  rather 
robust  and  is  probably  unsuited  to  "steer- 
ing" and  that  such  a  mechanism  could 
have  been  secondarily  lost.  In  this  context, 
it  should  also  be  noted  that  members  of 
some  genera  of  Agathidinae  that  have 
shorter  and  more  robust  ovipositors  were 
also  found  to  lack  the  bosses,  and  there- 
fore this  feature  may  be  readily  lost  in  spe- 
cies whose  ovipositors  evolve  to  become 
less  flexible.  We  therefore  conducted  a  sec- 
ond phylogenetic  analysis  on  the  data  ma- 
trix described  above  but  this  time  scoring 
the  Pselaphaninae  as  unknown  for  the 
new  ovipositor  character,  thus  allowing 
the  possibility  of  a  reversal  in  that  group 
without  adding  to  tree  length.  In  this  case 
the  most  parsimonious  trees  included  a 
subset  containing  the  clade  (Orgilinae  -I- 
(Agathidinae  +  Pselaphaninae)).  We 
therefore  suggest  that  the  newly  discov- 
ered ovipositor  features  should  be  incor- 
porated in  future  phylogenetic  analyses  of 
the  Braconidae. 

ACKNOWLEDGMENTS 

We  would  like  to  th.ink  Rachel  Kruft  and  Laraine 
Ficken  for  their  assistance  with  scanning  electron  mi- 


120 


Journal  of  Hymenoptera  Research 


croscopy,  and  Kees  van  Achterberg,  Paul  Marsh  and 
Jeno  Papp  for  providing  some  of  the  material  exam- 
ined. Our  thanks  also  to  Mark  Shaw  and  Robert  Bel- 
shaw  who  provided  many  helpful  comments  on  an 
earlier  version  of  the  manuscript.  The  work  was  part- 
ly supported  by  a  NERC  research  grant  (GR3/8060) 
to  Donald  Quicke  and  Mike  Fitton  and  by  the  NERC 
Initiative  in  Taxonomy. 

LITERATURE  CITED 

Achterberg,  C.  van.  1985.  Notes  on  Braconidae.  V. 
The  systematic  position  of  the  genera  Ecnomios 
Mason  and  Pselaphanus  Szepligeti  (Hymenop- 
tera: Braconidae).  Zoologische  Verhandeliiigen,  Lei- 
den 59:  341-348. 

Achterberg,  C.  van.  1986.  The  oviposition  behaviour 
of  parasitic  wasps  with  very  long  ovipositors 
(Ichneumonoidea:  Braconidae).  Eutomologische 
Berichten  46:  113-115. 

Achterberg,  C.  van  and  D.  L.  J.  Quicke.  1992.  Phy- 
logeny  of  the  subfamilies  of  the  family  Braconi- 
dae: A  reassessment  assessed.  Cladistics  8:  237- 
264. 

Austin,  A.  D.  and  T.  O.  Browning  1981.  A  mecha- 
nism for  movement  of  eggs  along  insect  ovipos- 
itors. International  Journal  of  Insect  Morphologx/  and 
Emhryologi/  10:  93-108. 

Compton,  S.  and  R.  Nefdt.  1988.  Extra-long  ovipos- 
itors in  chalcid  wasps;  some  examples  and  ob- 
servations. Antenna  12:  102-105. 

Delanoue,  P.  and  Y.  Arambourg.  1965.  Contribution 
a  I'etude  en  laboratoire  d'Eupelmus  urozonus 
Dalm.  (Hym.  Chalcidoidea  Eupelmidae).  Annales 
de  la  Societe  Entomologique  de  France  (N.S.)  1:  817- 
842. 

Flanders,  R.  V.  and  E.  R.  Oatman.  1982.  Laboratory 
studies  on  the  biology  of  Orgiliis  jenniae  (Hyme- 
noptera: Braconidae),  a  parasitoid  of  the  potato 
tuberworm,  Phthorimaea  operculella  (Lepidoptera; 
gelechiidae).  Hilgardia  50:  1-33. 

Gauld,  I.  D.  &  B.  Bolton  (eds).  1988.  The  Hymenoptera. 
British  Museum  (Natural  History/Oxford  Uni- 
versity Press,  Oxford.  332pp. 

Oeser,  R.  1961.  Vergleichend-MorphologischeUnter- 
suchungen  iiber  den  Ovipositor  der  Hymenop- 
teren.  Mitteihmgen  aiis  dem  Zoologische  Museum  in 
Berlin  37:  1-119. 

Quicke,  D.  L.  J.  1988.  Host  relationships  in  the  Bra- 
coninae  (Hymenoptera:  Braconidae) — How  little 
we  know!  Entomological  Society  of  Queensland 
News  Bulletin  16:  85-92. 


Quicke,  D.  L.  J.  1991.  Ovipositor  mechanics  of  the 
braconine  wasp  genus  Zaglyptogastra  and  the  ich- 
neumonid  genus  Pristomerus.  Journal  of  Natural 
History  25:  971-977. 

Quicke,  D.  L.  J.  and  C.  van  Achterberg.  1990:  Phy- 
togeny of  the  subfamilies  of  the  family  Braconi- 
dae (Hymenoptera).  Zoologische  Verhandelingen 
258:  1-95. 

Quicke,  D.  L.  J.,  L.  C.  Ficken,  and  M.  G.  Fitton.  1992a. 
New  diagnostic  ovipositor  characters  for  doryc- 
tine  wasps  (Hymenoptera,  Braconidae).  Journal  of 
Natural  History  26:  1035-1046. 

Quicke,  D.  L.  J.,  L.  C.  Ficken,  and  M.  G.  Fitton.  1992a. 
New  diagnostic  ovipositor  characters  for  doryc- 
tine  wasps  (Hymenoptera,  Braconidae).  Journal  of 
Natural  History  26:  1035-1046. 

Quicke,  D.  L.  J.  and  M.  G.  Fitton.  In  press.  Ovipositor 
steering  mechanisms  in  parasitic  wasps  of  the 
families  Gasteruptiidae  and  Aulacidae  (Hyme- 
noptera). Preceedings  of  the  Royal  Society. 

Quicke,  D.  L.  J.,  M.  G.  Fitton,  J.  R.  Tunstead,  S.  N. 
Ingram,  and  P.  V.  Gaitens.  1994.  Ovipositor 
structure  and  relationships  within  the  Hymenop- 
tera, with  special  reference  to  the  Ichneumono- 
idea. Journal  of  Natural  Histoty  28:  635-682. 

Quicke,  D.  L.  J.  and  P.  M.  Marsh.  1992.  Two  new 
species  of  Neotropical  parasitic  wasps  with  high- 
ly modified  ovipositors  (Hymenoptera,  Braconi- 
dae: Braconinae  and  Doryctinae).  Proceedings  of 
the  Entomological  Society  of  Washington  94:  559- 
567. 

Quicke,  D.  L.  J.  and  R.  A.  Wharton.  In  prep.  Antero- 
lateral scent  glands  of  braconine  wasps  (Hyme- 
noptera: Braconidae). 

Shaw,  M.  R.  and  T.  Huddleston.  1991.  Classification 
and  biology  of  braconid  wasps.  Handbooks  for  the 
Identification  of  British  Insects.  Vol.  7,  Part  11.  Roy- 
al Entomological  Society,  London. 

Smith,  E.  L.  1968.  Biosystematics  and  morphology  of 
Symphyta.  I.  Stem-galling  Euura  of  the  California 
region,  and  a  new  female  genitalic  nomenclature 
(Hymenoptera:  Tenthredinidae).  Annals  of  the  En- 
tomological Society  of  America  61:  1389-1407. 

Smith,  E.  L.  1970.  Evolutionary  morphology  of  the 
external  insect  genitalia.  2.  Hymenoptera.  Annals 
of  the  Entomological  Society  of  America  63:  1-27. 

Wharton,  R.  A.,  S.  R.  Shaw,  M.  J.  Sharkey,  D.  B.  Wahl, 
J.  B.  Woolley,  J.  B.  Whitfield,  P.  M.  Marsh  and  J. 
W.  Johnson  1992.  Phylogeny  of  the  subfamilies 
of  the  family  Braconidae  (Hymenoptera:  Ichneu- 
monoidea): a  reassessment.  Cladistics  8:  199-235. 


J.  HYM.  RES. 
Vol.  4,  1995,  pp.  121-136 


The  Wasp  Genus  Tachytella  Brauns,  1906  (Hymenoptera:  Sphecidae) 

WOJCIECH  J.  PULAWSKI 
California  Academy  of  Sciences,  Golden  Gate  Park,  San  Francisco,  California  94118,  USA 


Abstract.— The  southern  African  genus  Tachytella  Brauns  includes  aurcopilosa  Brauns,  1906  (west- 
ern South  Africa)  as  well  as  two  new  species:  heliophila  (western  Namibia)  and  nama  (southern 
Namibia,  northwestern  South  Africa).  A  lectotype  is  designated  for  Tachytella  aureopilosa  var.  nana 
Arnold,  1936,  the  latter  name  being  synonymized  with  Tachytella  aureopilosa  Brauns,  1906.  The 
genus  is  redescribed  using  previously  known  and  newly  discovered  characters,  a  key  to  the  spe- 
cies is  provided,  and  phylogenetic  relationships  between  the  species  are  discussed. 


INTRODUCTION 

General. — Tachytella  Brauns,  a  little 
known  genus  from  southern  Africa,  was 
thought  to  be  monotypic  until  now.  Spec- 
imens are  rarely  collected  (Bohart  and 
Menke  1976,  for  example,  knew  of  only 
six).  A  decade  ago,  I  discovered  an  unde- 
scribed  species  in  materials  from  the  State 
Museum  Windhoek  that  Ole  Lomholdt, 
then  at  the  Zoological  Museum,  Copen- 
hagen, kindly  had  transshipped  to  me. 
During  our  expedition  to  Namibia  in  1990, 
Maximilian  Schwarz  and  I  collected  spec- 
imens that  represented  another  new  spe- 
cies. These  findings  prompted  me  to  re- 
vise the  genus.  I  follow  the  format  used 
for  my  revisions  of  Kohliella  Brauns,  Holo- 
tacfn/spliex  de  Beaumont,  and  Gastrosericus 
Spinola  (Pulawski  1991,  1992,  1995). 

Technical  Terms. — Most  morphological 
terms  follow  Bohart  and  Menke  (1976), 
but  the  mandibular  terms  are  taken  from 
Michener  and  Fraser  (1978).  The  following 
terms  are  here  redefined  for  convenience: 

Clypeal  lobe:  the  projecting  medioventral 
portion  of  the  clypeus. 

Mandible: 

— adductor  ridge:  extends  distad  from  the 
adductor  swellings  (on  the  inner  side) 
and  gradually  becomes  visible  from  the 
outside,  constituting  the  distal  part  of 


the  mandibular  posterior  margin;  the 
two  portions  differ  in  size,  the  distal  one 
being  higher  than  the  basal  one; 

— condylar  ridge:  arises  from  the  condyle, 
extends  distad,  and  forms  the  basal  por- 
tion of  the  posterior  mandibular  mar- 
gin; it  is  angulate  distally  in  many  Lar- 
rinae; 

— condyle:  mandibular  articulation  on  the 
occipital  side  of  the  head  capsule; 

— notch:  an  emargination  on  the  posterior 
margin,  delimited  basally  by  the  con- 
dylar ridge  and  distally  by  the  expand- 
ed portion  of  the  adductor  ridge; 

— posterior  margin:  extends  between  the 
condyle  and  mandibular  apex;  called 
externoventral  margin  by  Bohart  and 
Menke  (1976)  and  lower  margin  by 
Michener  and  Fraser  (1978);  it  actually 
consists  of  two  components:  the  con- 
dylar ridge  basally  and  the  adductor 
ridge  distally  (the  term  posterior  is  pre- 
ferred because  the  head  is  hypognath- 
ous,  and  this  edge  is  thus  oriented  pos- 
terad); 


Sternum,    tergum:    shortened    terms 
gastral  sternum,  gastral  tergum. 

Vertex: 


for 


— length:  the  distance  between  a  hind- 
ocellus  and  an  imaginary  line  connecting 
eye  hindcorners  (i.e.,  the  point  where  the 


122 


Journal  of  Hymenoptera  Research 


inner  and  the  posterior  portions  of  the  or- 
bit meet); 

— width:  the  shortest  interocular  distance 
in  the  ocellar  region. 


specimens 
that   have 


Origin  of  Material. — The  29 
examined  apparently  are  all 
been  collected  so  far.  Institutional  or  per- 
sonal collections  in  which  the  material  is 
deposited  are  abbreviated  in  the  text  as 
follows  (names  of  contact  persons  are  giv- 
en in  parentheses): 

AMG:     Albany    Museum,    Grahams- 
town,    South    Africa    (Friedrich 
W.  Cess). 
BMNH:     British   Museum   (Natural   His- 
tory), London,  current  nonstat- 
utory name:  The  Natural  Histo- 
ry Museum  (Loraine  Ficken). 
CAS:     California  Academy  of  Sciences, 
San  Francisco,  California  (Woj- 
ciech  J.  Pulawski). 
FSCA:     Florida  State  Collection  of  Ar- 
thropods, Gainesville,  Florida 
(Lionel  A.  Stange,  James  R.  Wi- 
ley). 
MS:     Maximilian  Schwarz,  Ansfelden 
bei  Linz,  Austria  (personal  col- 
lection). 
SAM:     South   African   Museum,   Cape 
Town,  South  Africa,  including 
G.  Arnold  collection  previously 
housed  in  Bulawayo,  Zimbabwe 
(Margie  A.  Cochrane,  Vincent  B. 
Whitehead). 
FMP:     Transvaal  Museum,  Pretoria, 
South  Africa  (Ilema  Fourie). 
USNM:     Smithsonian   Institution,   Na- 
tional Museum  of  Natural  His- 
tory, Washington,  D.C.  (Arnold 
S.  Menke). 

Genus  Tachytella 

Tachytella  Brauns,  1906:56.  Type  species:  Tach- 
ytella aureopilosa  Brauns,  1906,  by  monotypy. 

Diagnosis. — Tachytella  is  a  member  of 
Larrini  (Bohart  and  Menke  1976)  and  thus 
has  the  hindocelli  modified  to  flat,  elon- 


gate scars  (Figs.  8,  12,  20),  the  only  rem- 
nant of  a  lens  being  a  narrow,  translucent 
band  along  the  scar's  outer  margin  (the 
band  is  broadly  interrupted  on  the  scar's 
posterolateral  or  lateral  side,  depending 
on  its  orientation).  Within  the  tribe,  Tach- 
ytella can  be  recognized  by  the  following 
four  characters: 

1.  The  frons  has  a  median,  slightly 
raised,  flat  area  that  is  delimited  by  a  lat- 
eral sulcus  (Figs.  6,  10,  18,  22,  28,  31),  the 
sulci  being  subparallel  and  close  to  the  or- 
bits. Similar  sulci  are  found  in  some  An- 
cistromma  W.  Fox,  e.g.,  capax  W.  Fox,  and 
in  less  specialized  Parapiagetia  Kohl  such 
as  genicularis  (F.  Morawitz)  and  odontosto- 
ma  (Kohl),  in  which  however,  the  frons  is 
not  raised. 

2.  The  pronotal  collar  is  rounded  and 
reaches  the  scutum  level,  a  condition  also 
found  in  some  Gastrosericiis,  e.g.  baobabiciis 
Pulawski  and  eurypus  Pulawski.  Tachytel- 
la, however,  has  three  submarginal  cells, 
whereas  Gastrosericiis  has  two. 

3.  The  submarginal  cell  III  in  the  fore- 
wing  is  essentially  rhomboidal  (Fig.  1),  a 
previously  unused  character,  with  the  an- 
terior and  posterior  margins  equal  in 
length  or  nearly  so;  the  proximal  and  dis- 
tal margins  are  weaklv  sinuous,  almost 
straight,  nearlv  parallel  to  each  other. 

4.  The  hindwing  jugal  lobe  is  short  and 
ends  well  before  crossvem  cu-a;  conse- 
quently, the  jugal  and  anal  excisions  are 
widely  separated  (Fig.  1).  A  similar  con- 
dition is  found  in  some  specialized  Gastro- 
sericiis such  as  braunsi  Arnold,  herero  Pu- 
lawski, and  pulchellus  R.  Turner  (Pulawski 
1995). 

Description. — Bohart  and  Menke  (1976) 
regarded  Tachytella  as  monotypic  and, 
consequently,  considered  as  generic  some 
structures  that  are  found  only  in  aureopi- 
losa but  not  in  the  other  two  species  (e.g., 
such  male  structures  as  the  edentate  inner 
mandibular  margin,  dentate  clypeal  free 
margin,  and  sterna  VI-VIII  with  dense, 
erect  setae).  They  thought  that  the  prono- 
tal collar  not  closely  appressed  to  scutum 


Volume  4,  1995 


123 


0.5  mm 


0.25  mm 


Figs.  1-3.  Some  of  the  generic  characters  of  Tachytella.  1,  fore  and  hindwing  of  aureopilosa  (drawn  from  the 
holotype,  the  model  Bohart  and  Menke  1976,  used  for  their  illustration);  arrows  indicate  third  submarginal 
cell,  jugal  excision,  and  anal  excision.  2,  anterior  part  of  thorax  of  aureopilosa  (arrow  indicates  pronotum).  3, 
foretarsus  of  female  nama. 


124 


Journal  of  Hymenoptera  Research 


also  was  a  generic  character,  but  in  fact  it 
is  only  individual.  The  following  rede- 
scription  takes  into  account  the  three  spe- 
cies now  recognized  and  also  some  struc- 
tures not  considered  previously.  Included 
are  those  features  that  vary  within  other 
Larrini  but  are  universal  in  Tachytella. 

Labrum  flat,  not  emarginate  apically, 
not  or  barely  protruding  beyond  clypeal 
free  margin;  prementum  convex  but  not 
compressed  laterally;  stipes  nearly  flat. 
Mouthparts  not  elongate.  Paramandibular 
process  broadly  separated  from  back  side 
of  clypeus  (mandibular  socket  open).  Pos- 
terior mandibular  margin  notched;  notch 
delimited  proximally  by  angulate  apex  of 
condylar  ridge,  and  distally  by  marked, 
roundly  expanded  distal  portion  of  ad- 
ductor ridge.  Clypeus  produced  into  mes- 
al  lobe,  free  margin  of  lateral  section  con- 
cave. Antennal  socket  narrowly  separated 
from  frontoclypeal  suture.  Inner  orbits 
convergent  above.  Frons  flat,  with  no  gla- 
brous swelling  above  each  antennal  sock- 
et. Hindocellar  scars  elongate,  broadly  di- 
verging anterolaterad  (their  long  axes 
forming  an  angle  of  about  120-140°), 
shorter  than  distance  that  separates  each 
one  from  midocellus.  Occipital  carina  ef- 
faced just  short  of  hypostomal  carina.  Pro- 
notum  anterodorsally  with  transverse 
groove;  collar  swollen,  reaching  level  of 
scutum.  Postspiracular  carina  evanescent, 
about  as  long  as  midocellar  diameter.  Ep- 
isternal  sulcus  originating  near  middle  of 
subalar  fossa,  ending  well  before  reaching 
anteroventral  mesopleural  margin;  scrobal 
sulcus  absent;  subalar  fossa  not  margined 
below;  metapleural  flange  not  expanded. 
Axilla  the  usual  shape,  i.e.,  not  expanded 
or  carinate.  Propodeum  short,  distance  be- 
tween spiracle  and  metanotum  less  than 
spiracle's  length;  dorsum  setose  through- 
out. No  additional  sclerites  between  me- 
tasternal  apex  and  propodeum  (no  "pro- 
podeal  sternum").  Forewing  with  three 
submarginal  cells,  none  petiolate;  recur- 
rent veins  separate  or  (some  aureopilosa) 
forming  short  petiole.  Hindwing  crossvein 


cu-a  vertical  (not  inclined).  Forecoxal  apex 
not  expanded  into  process.  Hindcoxal  dor- 
sum: inner  margin  not  carinate.  Hindtibia 
not  margined.  Forebasitarsus  and  apical 
tarsomeres  without  ventral  spines.  Claws 
in  each  pair  equal  in  size.  Hindtarsomere 
II  long  (0.6-0.7  X  hindtarsomere  I).  Base 
of  tergum  I  variable:  with  or  without 
short,  oblique  carina  that  extends  from 
each  anterolateral  corner.  Lateral  carina  of 
tergum  I  complete,  reaching  tergal  hind- 
margin.  Tergum  II  not  carinate  laterally. 
Body  without  erect  setae  (except  setae  on 
apical  sterna  in  male  of  aureopilosa);  those 
on  propodeal  dorsum  inclined  obliquely 
toward  head. 

Female.  Inner  mandibular  margin  with 
tooth.  Forecoxa  not  carinate  anteriorly, 
venter  slightly  convex;  outer  surface  of 
foretibia  without  preapical  spines.  Foretar- 
sus  with  rake  that  consists  of  stiff,  flat- 
tened, widely  spaced  spines  (four  on  for- 
ebasitarsus, two  on  foretarsomere  II);  tar- 
someres IV:  length  more  than  apical 
width,  apicoventral  margin  emarginate; 
tarsomeres  V:  apicoventral  margin  slightly 
arcuate.  Claws  without  teeth.  Tergum  VI 
fairly  convex:  angle  between  lateral  mar- 
gin of  tergum  and  lateral  margin  of  pygid- 
ial  plate,  in  side  view  about  30-40°.  Pygid- 
ial  plate  well  defined  (margined  laterally), 
rounded  apically,  sparsely  punctate  and 
with  thin,  inconspicuous  setae  (Figs.  9,  21, 
30).  Sting  sheaths  somewhat  flattened  dor- 
soventrally. 

Male.  Foretrochanter  and  forefemur  not 
emarginate;  forebasitarsus  1  with  one  rake 
spine  (in  basal  half);  outer,  apical  spine  of 
foretarsomere  II  shorter  than  foretarso- 
mere III;  sterna  without  velvety  patches; 
tergum  VII  carinate  apicolaterally,  not  de- 
pressed apically;  sterna  III- VI  impunctate 
at  least  mesally;  sternum  VIII  rounded 
apically  or  (some  aureopilosa)  minimally 
emarginate.  Venter  of  penis  valve  finely 
denticulate  subapically  (Figs.  17,  27,  35). 

Character  Polarization. — The  tribe  Larri- 
ni, of  which  Tachytella  is  a  member,  con- 
sists of  two  lineages,  the  subtribes  Larri- 


Volume  4,  1995 


125 


heliophila         aureopilosa         nama         aureopilosa        heliophila 


nama 


Figs.  4a  and  b.     Cladograms  showing  hypothetical  cladistic  relationships  of  the  three  species  of  Tachytella;  2: 
modified  cleft  of  inner  mandibular  margin,  8:  penis  valve  with  medioventral  tooth. 


na  and  Tachytina  (Bohart  and  Menke 
1976).  The  autapomorphy  of  Larrina  is  a 
unique  frons:  the  midocellus  is  placed  in 
a  depression,  and  the  transverse  sweUing 
delimiting  the  depression  anteriorly  joins 
a  linear  swelling  along  the  inner  orbit. 
The  hindocelli  are  oriented  along  a  trans- 
verse line,  and  the  transparent  lens  rem- 
nant is  horseshoe-like,  with  the  two 
branches  parallel  and  close  to  each  other, 
apparently  the  plesiomorphic  condition 
within  the  tribe.  In  Tachytina,  the  frons  is 
the  usual  plesiomorphic  condition,  with 
no  depression  or  swelling,  but  the  hin- 
docelli are  oriented  in  a  V-shaped  pattern 
and  the  lens  remnant  is  clearly  derived: 
the  anterior  branch  is  longer  than  the  pos- 
terior or  (in  Parapiagetia)  the  branches  are 
roughly  circular  (open  on  the  lateral 
side).  Because  of  the  shape  of  the  lens 
remnant,  Tacln/tella  is  a  member  of  Tach- 
ytina, but  its  relationships  within  the  sub- 
tribe  are  unclear.  Bohart  and  Menke 
(1976)  noted  that  Tacln/telln  resembled 
Ancistrotnma  and  Larropsis  Patton  in  the 
shape  of  the  mandible,  ocellar  scar,  fe- 
male foretarsal  rake,  and  male  sternum 
VIII.  None  of  these  similarities,  however, 
appears  to  be  a  synapomorphy.  I  have 
found  no  single  derived  character  linking 
Tacln/telln  with  any  of  the  other  nine  gen- 
era of  the  subtribe.  For  the  purpose  of  this 
analysis,  I  regard  the  Larrina  and  the  re- 


maining Tachytina  as  the  outgroup.  The 
characters  found  in  both  Tacln/telln  and 
the  outgroup  are  considered  plesiom- 
orphic, unless  there  is  strong  evidence 
that  they  have  been  acquired  indepen- 
dently. The  characters  present  only  in 
some  Tncln/tella  but  nc^t  the  outgroup  are 
regarded  as  apomorphic.  The  character 
codings  are:  0.  ancestral,  1  and  2.  derived. 
The  following  characters  have  been  con- 
sidered: 

1.  Female  mandible  inner  margin:  sub- 
basal  tooth:  0.  well  defined,  1.  inconspic- 
uous, obtuse  (as  in  Tnclnjtelln  nureopilosa). 
The  tooth  is  present  in  most  members  of 
the  outgroup,  but  absent  in  Lnrrn  Fabri- 
cius,  the  subgenus  Motes  Kohl  of  Liris  Fa- 
bricius,  in  Parapiagetia,  several  specialized 
Gastrosericus,  and  in  two  Tachysphex  Kohl 
{nefarius  Pulawski  and  ramses  Pulawski). 
Because  of  their  many  specializations, 
none  of  these  are  likely  to  share  a  unique 
common  ancestor  with  Tacln/telln,  and 
consequently  I  regard  the  presence  of  a 
well  defined  mandibular  tooth  in  Tacliy- 
tella  as  ancestral. 

2.  Female  mandible  inner  margin:  cleft: 
0.  narrow,  1.  shallow,  obtusely  angulate 
(as  in  Tacln/tella  aureopilosa ,  2.  deep,  broad 
(as  in  Tacln/tella  nama).  A  cleft  is  present 
and  narrow  in  most  Larrinae  (including 
Tacln/tella  heliophila),  but  absent  in  Larra, 
the  subgenus  Motes  of  Liris,  Dalara  Ritse- 


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Journal  of  Hymenoptera  Research 


9  aureopilosa 
Q  heliophiia 


nama 


( 


Fig.  5.     Collecting  localities  of  Tachytella. 


ma,  Paraliris  Kohl,  Holotachysphex,  many 
specialized  Gastrosericus,  Parnpiagetia  geni- 
cularis  (F.  Morawitz),  Tachysphex  nefarius 
and  ramses.  None  of  these  is  likely  to  share 
a  unique  common  ancestor  with  Tachytel- 
la,  and  I  therefore  accept  that  a  narrow 
cleft  is  ancestral  in  the  latter  genus.  The 
broad,  deep  cleft  is  unique  within  the 
tribe. 

3.  Male  clypeus:  0.  without  longitudinal 
swellings,  1.  swellings  present.  The  clyp- 
eal  swellings  of  aureopilosa  are  unique 
within  Larrinae  and  thus  apomorphic. 


4.  Propodeum:  0.  without  V-shaped  im- 
pression, 1.  with  V-shaped  impression. 
The  impression  is  unique  to  Tachytella  au- 
reopilosa and  thus  apomorphic. 

5.  Male  foretarsus:  0.  tarsomere  II  not 
expanded,  1.  tarsomere  II  expanded.  The 
expanded  tarsomere  of  Tachytella  aureopi- 
losa is  unique  within  the  tribe  and  thus 
apomorphic. 

6.  Tergum  I:  0.  with  a  pair  of  short,  bas- 
al carinae,  1.  carinae  absent.  Within  Lar- 
rini,  the  carinae  are  absent  in  Gastrosericus, 
Holotachysphex,  Kohliella,  Parapiagetia,  Tach- 


Volume  4,  1995 


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Figs.  6-9.     Tachytella  aiireopilosa,  female.  t>,  head  frontally.  7,  clypeus.  S,  head  dorsally.  9,  pygidial  plate. 


i/sphex,  manv  Tachvtes  Panzer,  and  Larrop- 
sis  chilopsidis  (Cockerell  and  Fox).  They  are 
present  m  all  other  Larrini  and  other 
Sphecidae  (those  with  a  petiolate  gaster 
are  obvious  exceptions).  1  therefore  regard 
the  absence  of  carinae  in  Tachytella  aiireo- 
pilosa as  derived. 

7.  Male  gaster:  0.  sternal  setae  all  short, 
1.  setae  of  sterna  VI-VIII  long,  erect.  Ster- 
nal setae  are  short,  appressed  in  most  Lar- 
rinae,  but  long,  erect  on  apical  sterna  in 
many  Liris  and  in  Tachytella  aiireopilosa. 
These  species  are  not  likely  to  share  a  sin- 
gle common  ancestor.  In  addition,  the  se- 
tal  configuration  c^f  T.  aiireopilosa  is  unique 
and  I  regard  it  to  be  derived. 

8.  Penis  valve:  0.  without  medioventral 


tooth,  1.  with  medioventral  tooth.  This 
tooth,  found  m  Tachuteila  heliophila  and 
nama,  is  unique  within  Larrinae  and  thus 
apomorpnic. 

The  following  5  characters  cannot  be 
polarized  because  both  states  are  com- 
monly found  in  the  outgroup: 

9.  Gena:  a.  thin  in  dorsal  view,  b.  thick 
in  dorsal  view. 

10.  Mesopleural  vestiture:  a.  not  con- 
cealing integument,  b.  concealing  integu- 
ment. 

11.  Hindtarsomere  IV:  a.  length  about 
1.0  X  apical  width,  b.  length  about  1.2-1.3 
X  apical  width. 

12.  Pygidial  plate  of  female:  a.  apex 
with    a    row   of   setigerous   punctures. 


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13 


Figs.  10-13.     Tachytella  aureopilosa,  male.  10,  head  frontally.  11,  clypeus.  12,  head  dorsally  with  outline  of 
vertex  showing  variation.  13,  scape. 


b.  apex  without  a  row  of  setigerous  punc- 
tures. 

13.  Male  mandible:  a.  inner  margin  an- 
gulate,  b.  inner  margin  simple. 
Cladistic  Analysis. — Distribution  of  the  8 
polarized  characters  is  given  below: 


Number: 
ancestor: 

aureopilosa: 
heliophila: 


nama: 


12345678 
00000000 
11111110 
00000001 
02000001 


Only  two  derived  characters  are  found 
in  more  than  one  species:  character  2  (a 


modified  cleft  on  inner  mandibular  mar- 
gin) is  shared  by  aureopilosa  and  nama,  and 
character  8  (penis  valve  with  medioventral 
tooth)  by  heliophila  and  nama.  The  clado- 
gram  resulting  from  character  2  (Fig.  4a) 
is  heliophila  +  {aureopilosa  +  nama),  the 
cladogram  based  on  character  8  (Fig.  4b) 
is  aureopilosa  +  {heliophila  +  nama).  I  see 
no  reason  to  prefer  one  over  the  other. 

Life  History. — Unknown.  The  presence, 
in  the  female,  of  a  foretarsal  rake  and  a 
pygidial  plate  suggests  ground  nesting. 

Geographic  Distribution  (Fig.  5). — Namib- 
ia and  western  half  of  South  Africa. 


Volume  4,  1995  129 


KEY  TO  THE  SPECIES 

1.  Propodeal  dorsum  with  V-shaped  impression  that  begins  behind  each  spiracle  and  ends 
at  hindmargin  (impression  inconspicuous  in  smallest  individuals);  tergum  I  basally  without 
pair  of  short,  oblique  carinae.  Female:  clypeus  (except  basally)  with  finely,  closely  punctate 
median  impression  that  separates  two  unsculptured,  longitudinal  areas  (Fig.  6);  imier  man- 
dibular margin  with  inconspicuous  subbasal  tooth  (Fig.  6).  Male:  clypeus  with  pair  of 
glabrous,  longitudinal  swellings,  free  margin  with  small,  median  tooth  (Fig.  10);  sterna  VI- 
VIII  with  dense  rows  or  tufts  of  erect  setae  (Fig.  15);  penis  valve  without  ventral  tooth 
(Fig.  17)    aurcopilosa  Brauns 

-  Propodeal  dorsum  without  such  impression;  tergum  I  basally  with  pair  of  short,  oblique 
carinae.  Female:  clypeal  bevel  without  median  row  of  small  punctures;  inner  mandibular 
margin  with  well-defined  subbasal  tooth  (Figs.  18,  28).  Male:  clypeus  with  no  longitudinal 
swellings,  free  margin  without  small,  median  tooth;  sterna  without  dense  rows  or  tufts  of 
erect  setae;  penis  valve  with  ventral  tooth  (Figs.  27,  35) 2 

2.  Mesopleural  vestiture  conspicuous,  fully  concealing  integument.  Length  of  hindtarsomere 
IV  1.2-1.3  X  apical  width.  Female:  free  margin  of  clypeal  lobe  weakly  arcuate,  corner  well 
defined  but  not  prominent  (Fig.  18);  inner  mandibular  margin  with  usual,  narrow  cleft 
(Fig.  18).  Male:  clypeal  lobe  broad,  nonprominent  (Fig.  22);  inner  mandibular  margin  sim- 
ple (Fig.  22) heliophila  sp.  n. 

-  Mesopleural  vestiture  inconspicuous,  integument  easily  visible.  Length  of  hindtarsomere 
IV  about  1.0  X  apical  width.  Female:  free  margin  of  clypeal  lobe  sinuate,  with  prominent 
corner  (Fig.  28);  inner  mandibular  margin  with  broadly  expanded  cleft  (Fig.  28).  Male: 
clypeal  lobe  prominent,  markedly  narrowing  anterad  (Fig.  31);  trimmal  margin  obtusely 
dentate  (Fig.  31) nama  sp.  n. 


DESCRIPTIONS  OF  SPECIES  (TMP),  present  designation,  examined.  New 

synonym. — As    Tachytella    aiireopilosa    nana: 

Diagnostic  characters  for  each  species  ^^^^^^  ^^^  ^^^^^  ^^7^^253  (illustration  of 

are  given  m  the  key  and  are  not  repeated  ^^^^  ^ead),  256  (new  status,  listed), 
individually  in  the  descriptions  below. 

Synonymy. — Arnold  (1936)  described  a 
Tachytella  aiireopilosa  Brauns  yar.  nana  for  specimens  that  had  red  gas- 
Figures  6-17  tral   segments   I-III,   shallower  V-shaped 

„    ,    ^  ,,                ,1,            inn^  err        i     •  imprcssion  on  thc  propodcal  dorsum,  and 

Tnan/tella  aiireo-vilosa  Brauns,  1906:57,  male,  in-  '^                ,,,,,,                 •      . 

.  :     ,    ,      .        ..         u  1  .  ^  truncate  rather  than  shallowly  emarginate 

correct    original    hyphenation.    Holotype:  -^            " 

male.  South  Africa:  Cape  Province:  Willow-  male  sternum  VIII.  Because  these  charac- 
more  (TMP),  examined.— Arnold  1923:218  ters  vary  individually,  I  regard  var.  nana 
(revision),  1930:5  (listed);  Bohart  and  Menke  as  a  synonym  of  aiireopilosa. 
1976:231  (illustration  of  ocelli),  253  (illustra-  Description. — Gena  thin  in  dorsal  view 
tion  of  male  head),  255  (illustration  of  (Figs.  8,  12).  Vertex  with  triangular,  gla- 
wings),  257  (listed),  258  (illustrations  of  pro-  brous,  impunctate,  dull  area  that  extends 
notum  and  scutum),  268  (illustration  of  fe-  f^.^^  Q^bit  toward  midline.  Head,  thorax, 
male  foretarsus),  279  (illustration  of  male  ^^^^^^^  ^^^  femora  minutely  punctate, 
sternum  VIII),  280  (illustration  of  male  geni-  ,  i.u  i  ^-  I^  .*.  ,^  ^^^..f 
talia);  Gess  1981:19  (South  Africa:  nefting  punctures  no  more  than  1  diameter  apart, 
probably  in  friable  soils).  hence  integument  mat  except  for  shiny 
Tachytella  aureo-pilosa  race  nana  Arnold,  1936:29,  clypeal  swellings  in  male.  Propodeal  dor- 
female,  male.  Lectotype:  male.  South  Africa:  sum,  in  most  specimens  studied,  with 
Cape  Province:  Resolution  in  Albany  District  pair   of   impressions   that   meet   postero- 


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Figs.  14-17.     Tachytella  aiireopilosa,  male.  14,  foretarsomere.  15,  gastral  apex  in  profile.  16,  volsella.  17,  penis 
valve. 


mesally  and  delimit  triangular  area  re-  throughout.  Length  of  hindtarsomere  IV 

sembling   propodeal   enclosure   of   most  about  1.0   X    apical  width.  Gastral  seg- 

Sphecidae  (impressions  barely  visible  in  ment  I  without  basolateral,  oblique  cari- 

smallest  specimens,  apparently  a  result  of  na. 

allometric   growth).   Propodeal  hindface  Mesopleural  vestiture  concealing  integ- 

not    ridged.    Femora    and    tibiae    setose  ument. 


i 


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Figs.  18-21.     Tacln/tt'lla  heliophila,  female.  18,  head  frontally.  19,  clypeus.  20,  head  dorsally.  21,  pygidial  plate. 


Head  and  thorax  black  but  the  follow- 
ing are  reddish:  mandible  (apical  third 
dark),  scape,  tegula,  and  humeral  plate. 
Gastral  segment  I  to  I-III  red,  remainder 
black.  Mid-  and  hindfemora  black,  forefe- 
mur  (except  dorsally),  tibiae  and  tarsi  red. 
Wings  nearly  hyaline.  Terga  I-V  silvery 
fasciate  apically. 

Female. — Mandible:  inner  margin  di- 
vided by  an  obtuse  angle  into  basal  and 
distal  portions,  with  shallow,  obtusely  an- 
gulate  cleft.  Clypeus  with  median,  finely, 
densely  punctate  impression  that  sepa- 
rates two  unsculptured,  longitudinal  areas 
(Figs.  6,  7);  lobe  free  margin  arcuate  except 
markedly  concave  near  corner,  which  is 
prominent  (Figs.  6,  7).  Length  of  flagello- 
mere  I  2.0  X  apical  width.  Vertex  width 
1.4  X  length.  Tergum  V  punctate  and  se- 


tose throughout.  Pygidial  plate  with 
preapical  row  of  setigerous  punctures 
(Fig.  9).  Length  7.3  mm 

Male. — Mandible:  inner  margin  simple, 
not  angulate  (Fig.  10).  Clypeus  (Figs.  10, 
11):  lobe  free  margin  with  median  tooth, 
corner  well  defined  but  not  prominent; 
distance  between  corners  about  0.8  X  dis- 
tance between  corner  and  orbit;  middle 
section  with  a  pair  of  longitudinal  swell- 
ings that  are  either  microsculptured,  dull, 
or  unsculptured,  shiny.  Scape  elongate, 
swollen  apicoventrally  (Fig.  13).  Length  of 
flagellomere  I  1.7-1.8  X  apical  width.  Ver- 
tex width  1.6  X  length.  Foretarsomere  II 
enlarged  (Fig.  14).  Tergum  VII:  punctures 
averaging  about  1  diameter  apart,  some  of 
them  about  3  diameters  apart.  Apex  of 
sternum  VI,  sternum  VII,  and  VIII  with 


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23 


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25 


Figs.  22-25.     Tachytella  heliophila,  male.  22,  head  frontally.  23,  clypeus.  24,  head  dorsally.  25,  scape. 


erect  setae;  longest  setae  equal  to  basal 
mandibular  width.  Length:  6.1-8.5  mm. 
Volsella:  Fig.  16.  Penis  valve  without  ven- 
tral tooth  (Fig.  17). 

Geographic  Distribution. — Western  South 
Africa. 

Records.— SOUTH  AFRICA:  Cape  Prov- 
ince: Colesberg  (1  female,  SAM),  Gra- 
hamstown:  Hilton,  sand  pit  (1  male, 
AMG),  NE  Ouberg  Pass  31  road  km  NE 
Montagu  at  33°39'S  20°18'E  (1  male, 
AMG),  Resolution  in  Albany  District  (1  fe- 
male, 1  male,  AMG;  1  male,  SAM;  1  head- 
less female,  1  male  lectotype  of  nana, 
TMP),  10  km  W  Steinkopf  (1  female, 
FSCA),  Willowmore  (1  male,  TMP,  holo- 
type  of  aureopilosa),  Worcester  (3  males, 
BMNH;  1  male,  USNM). 


Tachytella  heliophila  Pulawski,  new  species 
Figures  18-27 

Name  Derivation. — Heliophila,  a  Neolatin 
feminine  adjective  derived  from  two 
Greek  words:  helios,  sun;  and  philos,  a 
friend;  with  reference  to  the  open,  sunny 
habitat  where  the  species  was  found. 

Description. — Gena  thin  in  dorsal  view 
(Figs.  20,  24).  Vertex  without  impunctate, 
triangular  area.  Head,  thorax,  coxae,  fem- 
ora and  tibiae  minutely  punctate,  punc- 
tures no  more  than  1  diameter  apart, 
hence  integument  mat  except  for  the  un- 
sculptured,  shiny  clypeal  bevel.  Propodeal 
hindface  not  ridged.  Femora  and  tibiae  se- 
tose throughout.  Length  of  hindtarsomere 
IV  1.2-1.3  X  apical  width.  Gastral  segment 
I  basolaterally  with  short,  oblique  carina. 


Volume  4,  1995 


133 


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Figs.  26  and  27.     Tachytella  heliophila.  26,  volsella.  27,  penis  valve. 


Mesopleural  vestiture  concealing  integ- 
ument. 

Head  and  thorax  black  but  the  follow- 
ing are  reddish:  mandible  (apical  third 
black),  clypeal  bevel  anteriorly,  scapal 
venter  apically,  pronotal  lobe,  tegula,  and 
humeral  plate.  Gaster  red.  Coxae  and  fem- 
ora black  (femora  red  apically),  tibiae  and 
tarsi  red.  Wings  hyaline.  Terga  I-V  silvery 
fasciate  apically. 

Female. — Mandible:  inner  margin  with 
small  subbasal  tooth  and  narrow  cleft  (Fig. 
18).  Clypeus  (Figs.  18,  19):  free  margin  of 
lobe  arcuate,  corner  angulate  but  not 
prominent;  apicomedian  portion  of  mid- 
dle section  shiny,  practically  unsculptured 
(with  only  a  few  microscopic  punctures). 
Length  of  flagellomere  I  2.75  X  apical 
width.  Vertex  width  1.7  X  length.  Tergum 
V  punctate  and  setose  throughout.  Pygid- 
ial  plate  with  preapical  row  of  setigerous 
punctures  (Fig.  21).  Foretibia  without 
preapical  spines.  Length  6.6-7.5  mm. 

Male. — Mandible:  inner  margin  simple. 


not  angulate  (Fig.  22).  Clypeus  (Figs.  22, 
23):  lobe  free  margin  arcuate,  corner  ob- 
tusely angulate;  surface  finely  punctate 
and  setose  throughout  except  for  narrow 
apical  lip;  distance  between  corners  about 
0.9  X  distance  between  corner  and  orbit. 
Scape  swollen  ventrally  (Fig.  25).  Length 
of  flagellomere  I  2.1  X  apical  width.  Ver- 
tex width  3.0  X  length.  Tergum  VII:  punc- 
tures averaging  less  than  one  diameter 
apart.  Length  6.4  mm.  Volsella:  Fig.  26.  Pe- 
nis valve  with  ventral  tooth  near  mid- 
length  (Fig.  27). 

Geographic  Distribution. — Known  from  a 
single  area  in  Namib  desert,  western  Na- 
mibia. 

Records. — Holotype:  female,  Namibia: 
Karibib  District:  65  km  SW  Usakos,  24  Feb. 
1990,  W.J.  Pulawski  (CAS). 

Paratypes:  NAMIBIA:  Karibib  District: 
55  km  SW  Usakos,  1  Mar.  1990,  W.j.  Pu- 
lawski (1  female,  CAS);  65  km  SW  Usakos, 
M.  Schwarz,  24  Feb.  1990  (1  female,  MS), 
1  Mar.  1990  (1  female,  MS;  1  male,  CAS). 


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Figs.  28-30.     Tachytella  nama,  female.  28,  head  frontally.  29,  head  dorsally.  30,  pygidial  plate. 


Tachytella  nama  Pulawski,  new  species 
Figures  28-35 

Name  Derivation. — Nama,  a  Hottentot 
tribe  that  immigrated  from  southern  Af- 
rica into  the  central  Namib;  a  noun  in  ap- 
position to  the  generic  name. 

Description. — Gena  thick  in  dorsal  view 
(Figs.  29,  32).  Vertex  without  impunctate, 
triangular  area.  Scutal  punctures  varying 
from  mostly  about  1  diameter  apart  (fe- 
male from  Goegap)  to  many  diameters 
apart  (most  specimens).  Mesothoracic  ven- 
ter with  minute  punctures  that  are  several 
diameters  apart.  Propodeal  hindface  mi- 
croscopically, densely  ridged  and  with  ev- 
anescent punctures.  Fore-  and  midfemo.ral 
venter  with  a  few,  scattered  punctures,  as- 
etose.  Outer  side  of  foretibia  as  well  as 
mid-  and  hindtibial  dorsum  impunctate, 
asetose.  Length  of  hindtarsomere  IV  about 


1.0  X  apical  width.  Gastral  segment  I  ba- 
solaterally  with  short,  oblique  carina. 

Mesopleural  vestiture  inconspicuous, 
not  concealing  integument. 

Head  and  thorax  black  except  mandible 
yellowish  basally  and  reddish  distally, 
middle  clypeal  lobe  of  male  reddish,  male 
flagellum  yellowish  brown  ventrally.  Gas- 
tral segments  I-III  red,  remainder  black. 
Wings  almost  hyaline.  Terga  I-III  silvery 
fasciate  apically,  but  fasciae  evanescent  in 
most  specimens. 

Female. — Mandible:  inner  margin  with 
subbasal  tooth  and  broadly  expanded 
cleft  (Fig.  28).  Clypeus  (Fig.  28):  lobe  free 
margin  sinuous,  corner  prominent;  sur- 
face all  or  largely  unsculptured  (at  most 
punctate  basally).  Length  of  flagellomere 
I  2.0  X  apical  width.  Vertex  width  2.4  X 
length.  Tergum  V  with  impunctate,  gla- 


Volume  4,  1995 


135 


0.5  mm 


32 


0.5  mm 


0.1  mm 


Figs.  31-35.     Tachytella  nama,  male.  31,  head  frontally.  32,  head  dorsally.  33,  scape.  34,  volsella.  35,  penis  valve. 


brous  apical  depression.  Pygidial  plate 
with  no  preapical  row  of  setigerous  punc- 
tures (Fig.  30).  Foretibia  with  two  spines 
between  inner  surface  and  dorsum:  one 
at  midlength,  the  other  at  two  thirds  of 
length.  Length  7.5  mm. 

Male. — Mandible:  inner  margin  obtuse- 
ly angulate  (Fig.  31).  Clypeus  (Fig.  31): 


middle  section  markedly  convex;  lobe 
prominent,  markedly  narrowing  anterad, 
with  unsculptured  apical  part  and  shal- 
lowly  emarginate  anterior  margin.  Scape 
minimally  swollen  ventrally  (Fig.  33). 
Length  of  flagellomere  I  1.5  X  apical 
width.  Vertex  width  2.4  X  length.  Tergum 
VII:  punctures  averaging  more  than  one 


136 


Journal  of  Hymenoptera  Research 


diameter  apart.  Length  7.2  mm.  Volsella: 
Fig.  34.  Penis  valve  with  ventral  tooth 
(Fig.  35)  near  midlength. 

Geographic  Distribution. — Southern  Na- 
mibia and  northwestern  South  Africa. 

Records. — Holotype:  female.  South  Afri- 
ca, Cape  Province:  Hester  Malan  [now 
Goegap]  Nature  Reserve  near  Springbok, 
15-21  Oct.  1987,  F.W.  and  S.K.  Gess 
(AMG). 

Paratypes:  NAMIBIA:  Liideritz  District: 
Namuskluft  88,  2716  Dd  [=  between 
27°45'  and  28°00'S  and  16°45'  and 
17°00'E],  collector  unknown,  12-15  Sept. 
1973  (1  female,  SMW),  7-15  Oct.  1970  (1 
male,  SMW). 

SOUTH  AFRICA:  Cape  Province: 
Anenous,  29°14'30"S,  17°34'45"E,  11-13 
Oct.  1988,  D.W.  Gess  (2  females,  AMG); 
Dikbome,  Merweville  Koup,  [South  Afri- 
can] Museum  Expedition,  Oct.  1952  (1 
male,  SAM);  Hester  Malan  [now  Goegap] 
Nature  Reserve  near  Springbok,  15-21 
Oct.  1987,  F.W.  and  S.K.  Gess  (1  female, 
CAS);  same  data  but  10-12  Oct.  1988  (1 
male,  AMG);  between  Kamieskroon  and 
Springbok,  [South  African]  Museum  Staff, 
Oct.  1952  (2  females,  CAS,  SAM). 

ACKNOWLEDGMENTS 

Illustrations  were  prepared  by  Mary  Ann  Tenorio 
on  a  Macintosh  computer  using  the  Adobe  Illustrator 
program.  Fieldwork  in  Namibia  in  1989,  where  I  col- 
lected specimens,  was  supported  by  the  National  Sci- 
ence Foundation  (grant  number  BSR-8722030)  and 
museum  work  in  South  Africa  the  same  year  by  the 
California  Academy  of  Sciences  through  its  In-House 


Research  Fund.  I  sincerely  thank  persons  who  sent 
specimens  for  study.  I  am  indebted  to  Arnold  S.  Men- 
ke  and  Friedrich  W.  Gess  for  their  critical  reviews  of 
the  manuscript.  I  sincerely  thank  my  friend  and  trav- 
el companion  to  Namibia,  Herr  Maximilian  Schwarz, 
who  generously  donated  the  unique  male  of  Tachy- 
tella  heliophila  to  the  California  Academy  of  Sciences. 

LITERATURE  CITED 

Arnold,  G.  1923.  The  Sphegidae  of  South  Africa.  Part 
III.  Annals  of  the  Transvaal  Museum  9:191-253. 

Arnold,  G.  1930.  A  check-list  of  the  Sphegidae  of  the 
Ethiopian  Region.  University  Press,  Cambridge, 
England.  21  pp. 

Arnold,  G.  1936.  New  African  Hymenoptera  No.  3. 
Occasional  Papers  of  the  Rhodesian  Museum  5:1-38, 
pi.  I. 

Bohart,  R.M.,  and  A.S.  Menke.  1976.  Sphecid  zoasps  of 
the  world.  A  generic  revision.  University  of  Cali- 
fornia Press,  Berkeley,  Los  Angeles,  London.  1 
color  plate,  IX  +  695  pp. 

Brauns,  H.  1906.  Zur  Kenntnis  der  siidafrikanischen 
Hymenopteren.  Verhandlungen  der  Zoologisch-Bo- 
tanischen  Gesellschaft  in  Wien  56:43-59. 

Gess,  F.  1981.  Some  aspects  of  an  ethological  study 
of  the  aculeate  wasps  and  the  bees  of  a  karroid 
area  in  the  vicinity  of  Grahamstown,  South  Af- 
rica. Annals  of  the  Cape  Provincial  Museums  (Nat- 
ural History)  14:1-80. 

Michener,  CD.  and  A.  Fraser.  1978.  A  comparative 
anatomical  study  of  mandibular  structures  in 
bees.  The  University  of  Kansas  Science  Bulletin  51: 
463^82. 

Pulawski,  W.J.  1991.  A  revision  of  the  wasp  genus 
Kohliella  (Hymenoptera:  Sphecidae).  Proceedings 
of  the  California  Academy  of  Sciences  47:289-302. 

Pulawski,  W.J.  1992.  World  species  of  the  wasp  ge- 
nus Holotachysphex  de  Beaumont  (Hymenoptera: 
Sphecidae).  Proceedings  of  the  Etitomological  Society 
of  Washington  94:223-242. 

Pulawski,  W.J.  1995.  The  wasp  genus  Gastrosericus 
Spinola,  1839  (Hymenoptera:  Sphecidae).  Mem- 
oirs of  the  California  Academy  of  Sciences  (in  press). 


J.  HYM.  RES. 

Vol.  4,  1995,  pp.  137-193 

Revision  of  the  Ant  Genus  Gnantptogenys  in  the  New  World 

(Hymenoptera:  Formicidae) 

John  E.  Lattke 

Institute  de  Zoologia  Agricola,  Universidad  Central  de  Venezuela, 
Apartado  4579,  Maracay,  Venezuela  2101-A 


Abstract. — Ants  of  the  genus  Gnmnptogenys  Roger  in  the  New  World  are  revised.  The  study  is 
based  mostly  on  worker  morphology;  73  species  are  recognized,  fifteen  of  which  are  new:  G. 
andina,  G.  hoUviensis,  G.  brunnea,  G  ciineiforma,  G.  ejuncida,  G.  extra,  G.  gentri/i,  G.  latkephala,  G. 
ilimaui,  G.  nigrivitrea,  G.  pilosa,  G.  siapensis,  G.  stellae,  G.  transversa  and  G.  volcano.  G.  ericae  is 
revalidated.  In  the  present  work  sixteen  names  have  been  synonymized,  and  a  morphological 
synopisis  of  the  genus  is  given.  The  species  range  from  southern  U.S.  to  northern  Argentina,  but 
most  are  found  in  Central  America  and  tropical  South  American  forests.  The  evolutionary  history 
of  the  genus  is  studied  using  phylogenetic  systematics  criteria.  Six  species  groups  are  recognized: 
striatula  group  (22  species);  niiniita  group  (9  species);  rastrata  group  (14  species);  sulcata  group  (8 
species);  nwrdax  group  (17  species),  and  the  concinna  group  (3  species).  A  key  for  the  identification 
of  workers  is  presented.  Illustrations  of  morphological  features  supplement  the  key  and  descrip- 
tions. 


INTRODUCTION 

The  genus  Gnamptogem/s  Roger  is  a  di- 
verse group  of  ponerines  with  a  distribu- 
tion that  covers  the  Neotropics,  Indoma- 
laysia  and  parts  of  the  Nearctic  and  Aus- 
tralia. The  genus  as  treated  here  is  mostly 
the  result  of  extensive  synonymy  by 
Brown  (1958)  in  his  revision  of  the  Tribe 
Ectatommini.  In  this  treatment,  however, 
the  minuta  group  species  (=  Alfarin  Forel 
group  of  previous  authors),  are  excluded 
and  treated  elsewhere  (Brandao  and  Latt- 
ke 1990;  Lattke  1992).  Since  Brown  (1959), 
research  on  Gnamptogem/s  has  consisted 
mostly  of  the  description  of  new  forms 
and  scattered  bits  of  ecological  informa- 
tion. The  only  known  fossils  of  the  genus 
are  twc^  species  reported  from  Dominican 
Amber  by  Baroni  Urbani  (1980).  Lattke 
(1990)  reviews  the  Venezuelan  species  and 
their  natural  history.  Due  to  the  growing 
number  of  specimens  in  collections,  and 
the  additional  information  published  c^r 
accumulated  in  notes,  a  revision  of  the 
New  World  species  seemed  desirable. 


MATERIALS  AND  METHODS 

The  revision  is  based  primarily  on 
worker  morphology  and  the  species  have 
been  defined  conservatively  because  of 
considerable  interpopulation  variation. 
The  criteria  for  judging  allopatric  popula- 
tions as  conspecific  are  as  in  Ward  (1984: 
131):  if  they  are  at  least  as  dissimilar  as 
closely  related,  sympatric  species  and  are 
not  connected  by  known  intermediate 
forms.  For  a  phylogenetic  analysis,  the  ge- 
nus Ectatomma  Fr.  Smith  is  used  as  an  out- 
group  in  order  to  help  determine  the  char- 
acter state  polarities.  This  genus  is  consid- 
ered very  close  to  Gnamptogemjs  and  pre- 
sents a  greater  number  of  primitive 
character  states  than  either  Gnamptogemjs 
or  Rhytidoponera  Mayr  (Lattke  1994).  Spe- 
cies groups  were  determined  and  then 
compared  using  a  character  matrix  (Table 
1)  and  Farris's  Hennig  '86  v.  1.5  phyloge- 
netic inference  program.  The  "ie-"  option 
was  used  to  generate  the  most  parsimo- 
nious tree  (Fig.  19)  and  a  nelsen  consensus 


138 


Journal  of  Hymenoptera  Research 


Table  1.    Character  state  matrix 


123456789 

0123456789 

012345 

Ectatomma 

000000000 

0000000000 

000000 

LEMQ 

G.  haenschi 

110001101 

0010000000 

012110 

G.  concinna 

100011111 

1110000001 

000001 

G.  schmitti 

100001021 

1010000011 

001010 

strigata  sbgp. 

010010020 

0000011201 

000100 

porcata  sbgp. 

000010020 

0000011201 

000100 

LNKD 

striatuln  sbgp. 

000010020 

0000011200 

000100 

moniax  group 

111111121 

111-011102 

000011 

MCZC 

sulcata  group 

110111121 

1111011122 

001011 

rastrata  group 

110011120 

0010111202 

000010 

miuiita  group 

100011000 

0010010001 

110110 

MHNG 

MIZA 

tree  calculated  from  the  13  most  parsi- 
moniuos  trees  calculated  by  the  "ie"  op- 
tion. Due  to  logistical  difficulties,  it  was 
not  possible  to  examine  some  types,  this 
is  especially  true  for  European  specimens. 
Thus  this  revision  depends,  in  such  cases, 
upon  Brown's  concept  of  the  species  in- 
volved, either  through  examining  species 
determined  by  him  during  the  course  of 
his  studies,  reading  the  notes  he  took 
while  visiting  European  collections  in 
1963  or  from  his  1958  revision.  When  type 
material  was  examined  it  is  followed  by  a 
"T"  in  the  species  list. 

Specimens  were  examined  from  or  de- 
posited in  the  following  collections: 

BMNH     The  Natural  History  Museum, 
London,  U.K. 

CMLB  Coleccion  Martha  Lucia  Baena, 
Cali,  Colombia 

CASC  California  Academy  of  Sciences, 
San  Francisco,  California,  U.S.A. 

CFFC  Coleccion  Fernando  Fernandez, 
Bogota,  Colombia 

CKWJ  Coleccion  Klaus  Werner  Jaffe, 
Universidad  Simon  Bolivar,  Ca- 
racas, Venezuela 

CNIC  Cornell  University  Insect  Col- 
lection, Ithaca,  New  York,  U.S.A. 

CUNB     Coleccion  Universidad  Nacion- 
al,  Bogota,  Colombia 
JTLC     John  T.  Longino  Collection,  Ev- 
ergreen College,  Olympia,  Wash- 
ington, U.S.A. 


LACM  Los  Angeles  County  Museum 
of  Natural  History,  Los  Ange- 
les, California,  U.S.A. 
Lyman  Entomological  Muse- 
um, McGill  University,  Ste. 
Anne  de  Bellevue,  Quebec, 
Canada. 

Landessammlungen  fiAr  Natur- 
kunde,  Karlsruhe,  Germany 
Museum  of  Comparative  Zool- 
ogy, Harvard  University,  Cam- 
bridge, Massachusetts,  U.S.A. 
Museum    d'Histoire    naturelle, 
Geneva,  Switzerland. 
Museo  del  Instituto  de  Zoologia 
Agricola,   Universidad   Central 
de  Venezuela,  Maracay,  Vene- 
zuela 
MUSP     Museu    Universidade    de    Sao 
Paulo,  Sao  Paulo,  Brazil 
NHMW     Naturhistorisches  Museum,  Vi- 
enna, Austria. 
PSWC     Philip  S.  Ward  Collection,  Uni- 
versity of  California,  Davis, 
California,  U.S.A. 

USNM  United  States  National  Muse- 
um, Washington,  D.C.,  U.S.A. 

WPMC  William  P.  MacKay  Collection, 
University  of  Texas,  El  Paso, 
Texas,  U.S.A. 

Measurements  and  Indices 

Morphological  measurements  were 
made  at  magnifications  of  up  to  100  X  with 
an  ocular  micrometer  on  a  Wild  M5  ster- 
eoscope. Since  ratios  derived  from  the 
measurements  are  conveniently  scaled  in- 
dices by  themselves,  the  orthodox  proce- 
dure of  multiplying  them  by  100  was  not 
followed.  The  measurements  follow  those 
of  Lattke  (1990). 

HL  Head  length:  midline  length  of 
head  proper,  measured  in  full-face 
(dorsal)  view,  from  the  anterior 
clypeal  margin  to  the  midpoint  of 
a  line  drawn  across  the  vertexal 
margin. 

ML     Mandibular    length:     length    of 


Volume  4,  1995 


139 


closed  mandibles,  from  the  anteri- 
or clypeal  margin  to  the  apices  of 
the  mandibles,  measured  in  the 
same  plane  as  HL. 

SL  Scape  length:  length  of  the  first  an- 
tennal  segment,  excluding  the  neck 
and  basal  condyle. 

ED  Eye  Diameter:  diameter  of  eye, 
measured  along  its  long  axis  in  lat- 
eral view 
HW  Head  width:  maximum  width  of 
head,  measured  in  full-face  (dor- 
sal =  frontal)  view,  excluding  the 
eyes. 
WL  Weber's  length  of  the  mesosoma 
(alitrunk):  diagonal  length  mea- 
sured in  lateral  view,  from  the  an- 
terior margin  of  the  pronotum  (ex- 
cluding collar)  to  the  posterior  ex- 
tremity of  the  metapleural  lobe. 

CI     Cephalic  index:  HW/HL 

MI     Mandibular  index:  ML/HW 

SI     Scape  index:  SL/HW 

OI  Ocular  index:  ED/HW 
SSC  Scape  setal  count:  the  number  of 
standing  hairs  (not  pubescence), 
i.e.  those  forming  an  angle  of  45° 
or  more  with  the  cuticular  surface, 
visible  in  outline  on  the  upper  sur- 
face of  the  scape,  with  the  line  of 
view  at  right  angles  to  the  plane  of 
funicular  inflexion. 

New  World  Gnamptogenys  Roger 

Generic  Synopsis  (Worker). — Sculpture 
consisting  mostly  of  even  parallel  costae, 
costulae  or  strigae,  occasionally  rugose; 
distinct  dorsomedian  cephalic  carinae 
usually  absent,  at  most  never  extending 
more  than  half-way  between  clypeus  and 
vertex;  eyes  at  midlength  of  head  or  pos- 
terad,  surrounded  by  a  fine  groove;  lobes 
of  frontal  carinae  broadly  convex  and  par- 
tially raised,  incompletely  covering  anten- 
nal  condyles  (except  miniitn  group);  fun- 
niculus  filiform  or  incrassate,  never  with  a 
distinct  club;  anterior  clypeal  border  with 
narrow  lamella  of  variable  length;  mese- 
pisternum    anteroventrally    bordered    by 


narrow  lamella;  pronotum  unarmed, 
without  protuberances;  propodeal  spira- 
cles round  or  slightly  ovoid,  never  slit- 
like; transverse  sutures  and  grooves  on 
mesosomal  dorsum  present  or  absent; 
mesonotum  never  prominently  convex 
and  bulging;  anterior  prostemal  process 
bidentate;  anterior  mesepisternal  process 
produced  as  thin  triangular  lobe  with 
pointed  or  bluntly  pointed  apex;  metepis- 
ternum  with  deep  posterior  cleft  for  peti- 
olar  insertion  and  continuous  with  open 
metacoxal  fossae;  metepisternal  process 
located  anterad  of  cleft  and  of  variable  de- 
velopment; metacoxal  dorsum  usually 
with  denticle,  lobe  or  tubercle,  absent  in 
some  species;  helcium  protruding  medi- 
anly  on  anterior  postpetiolar  face;  tergite 
of  helcium  much  larger  than  sternite;  for- 
tibial  apex  lacking  stout  moveable  setae; 
outer  border  of  foretarsal  comb  (opposite 
calcar)  usually  with  single  prominent  seta; 
meso-  and  metatibial  spurs  one  or  two 
(weakly  developed),  barbulate  or  simple; 
empoida  lacking.  In  most  species  the  pre- 
tarsal  claws  are  bidentate  on  all  legs,  and 
the  median  tooth  may  vary  in  its  position 
among  species,  and  the  claws  may  not 
necessarily  be  alike  on  all  legs.  In  small 
species  the  claws  may  be  hard  to  observe. 

Malpighian  tubule  number  six  (Brown 
1988).  The  genus  is  so  diverse  that  char- 
acterizing it  can  be  difficult.  The  following 
characters  can  be  considered  synapomor- 
phies  of  the  genus  which  distinguish  it 
from  its  closest  relatives:  Ectatomnia  and 
Rhi/tidoponera:  the  single  stout  moveable 
seta  on  the  foretibial  apex  and  a  spine  or 
tubercle  on  the  metacoxal  dorsal  surface. 
This  former  trait  is  lacking  in  some  spe- 
cies. The  following  combination  of  char- 
acters are  synapomorhic  for  the  genus  if 
one  excludes  the  problem  species  dis- 
cussed further  on  in  the  text:  inconspicu- 
ous metanotum,  disappearance  of  the  me- 
dian cephalic  carina,  lack  of  row  of  stout 
setae  on  foretarsal  base  opposite  the  strig- 
il,  leaving  only  one  seta. 

Characters. — The  choice  of  characters  for 


140 


Journal  of  Hymenoptera  Research 


1 


8 


Figs.  1-8.  Lateral  view  of  mandibles  and  anterior  section  of  head  of  Ectatomma  and  Gnmnpiogemjs.  Transverse 
section  of  mandible  depicted  in  black  with  chewing  border  to  the  left  and  frontal  surface  below.  1,  E.  niidum; 
2,  G.  porcata;  3,  G.  moellerius;  4,  G.  liaenschi;  5,  G.  triangularis;  6,  G.  mordax;  7,  G.  tortuolosa;  8,  G.  concinna. 


use  is  complicated  by  convergence  in  sev- 
eral instances.  Some  character  states  con- 
sidered apomorphic  have  apparently  been 
reached  independently  by  one  or  two  spe- 
cies in  different  lineages.  Examples  are  the 
smooth  and  shining  scapes  of  G.  rastrata 
and  of  G.  mordax,  or  the  conspicuous  me- 
tanotal  groove  of  G.  bisulca  and  G.  brimnea. 
There  are  also  some  apparent  reversals,  as 
well  as  retention  of  plesiomorphies  pres- 
ent in  the  outgroup.  But  these  cases  are 
few,  isolated  and  do  not  represent  the  ma- 
jor trends  of  their  respective  species 
groups.  Therefore,  they  were  not  taken 
into  account  for  defining  group  character 


states.  Mandibular  shape  is  an  apparent 
useful  indicator  of  phylogenetic  trends, 
with  the  general  direction  being  from  ple- 
siomorphic  triangular  mandibles  to  apo- 
morphic forms  as  subtriangular  or  subfal- 
cate.  Subtriangular  mandibles  have  been 
independently  derived  in  the  haenschi,  sul- 
cata, and  mordax  groups,  as  indicated  by 
other  mandibular  traits  such  as  the  config- 
uration of  the  chewing  border,  sculpture 
and  cross  section.  In  the  last  two  groups 
(Figs.  6-8),  the  mandibular  cross  section 
reveals  thick,  robust  mandibles  as  op- 
posed to  the  slim  cuneiform  section  of  G. 
haenschi,  which  is  the  plesiomorphic  state 


Volume  4,  1995 


141 


11 


Figs.  9-14.     Lateral  view  of  propodeum  of  Ectatomma  and  Gnamptogenys.  9,  E.  lugeus;  10,  £.  quadridens;  11, 
E.  tuberculatum;  12,  G.  bispinosa;  13,  G.  perspicax;  14,  G.  triangularis. 


exhibited  by  the  outgroup  and  the  striatula 
and  rastrata  groups  (Figs.  1-5).  The  sub- 
petiolar  process  in  its  primitive  state  is  tri- 
angular, lobe-like  and  anteriorly  protrud- 
ing. Modification  has  been  towards  a  sub- 
quadrate  shape,  but  some  species,  such  as 
G.  striatula,  exhibit  both  conditions,  and  in 
the  derived  mordax  subgroup  the  triangu- 
lar subpetiolar  porocess  is  probably  a  re- 
versal. 

Some  characters  are  shared  by  the  out- 
group and  certain  groups  or  subgroups  of 
Gnamptogenys.  Two  of  these  are  particu- 
larly important:  the  propodeal  denticles 
and  the  setae  on  the  foretarsal  base.  The 


16 


Figs.  15-16.  Outer  lateral  view  of  apex  of  tibia  and 
base  of  tarsus  of  foreleg.  15,  typical  configuration  in 
Ectatomma  with  single  calcaria  on  tarsal  apex  and  row 
of  setae  in  foretarsal  concavity;  16,  same  in  most 
Gnamptogenys  with  only  one  stout  seta  on  foretarsal 
concavity. 


propodeal  denticles  of  Ectatomma  may  be 
absent  in  some  species  and  when  present, 
they  are  quite  modest,  usually  lobe-like 
and  directed  dorsally.  The  declivitous  pro- 
podeal face  runs  evenly  and  directly  to  the 
denticular  apex  (Figs.  9-11).  The  denticles 
in  the  rastrata  group  are  stout  and  inclined 
posterad,  and  their  base  forms  a  more 
abrupt  separation  from  the  propodeal  de- 
clivity (Figs.  12-14).  With  the  exception  of 
a  few  highly  derived  species,  propodeal 
denticles  are  lacking  in  Rhytidoponcra  and 
the  striatula  group  of  Gnamptogenys.  This 
suggests  that  the  presence  of  propodeal 
denticles  is  probably  independently  de- 
rived in  Ectatomma  and  Gnamptogenys  and 
consequently  was  lacking  in  the  ancestral 
stock  of  this  group  of  genera.  For  this  rea- 
son this  state  has  been  given  value  0  in  the 
character  matrix  for  Ectatomma.  The  base 
of  the  foretarsus,  in  the  concavity  oppos- 
ing the  strigil,  has  an  outer  row  of  stout 
setae  in  Ectatomma,  but  only  one  promi- 
nent seta  in  most  Gnamptogenys  (Figs.  15- 
16).  The  exceptions  are  all  the  minuta- 
group  species  besides  three  other  species, 
and  each  of  these  exceptions  are  unique 
within  the  genus  as  will  be  discussed  later 
on.  The  brief  anterior  peduncle  of  the  pet- 
iole, typical  of  Ectatomma  is  considered 
plesiomorphic  for  the  porcata  subgroup. 


142 


Journal  of  Hymenoptera  Research 


0^ 


17 


18 

Figs.  17-18.     Ventral  view  of  the  anterior  postpetiolar  process.  17:  a,  G.  triajjgularis;  b,  G.  haenschi;  c,  G.  striatiila. 
18,  G.  sulcata. 


but  autapomorphic  for  the  mordax  sub- 
group. The  following  characters  were 
taken  into  account  for  all  the  species 
groups  or  individual  species  in  the  matrix, 
but  have  been  restricted  to  the  minuta  sub- 
group of  the  minuta  group  (Lattke  1991). 
Characters  8,  17,  18  were  considered  non- 
additive. 

The  following  character  states  were 
taken  into  account: 

1.  Head  shape  in  frontal  view:  0= wider 
posterad  than  anterad,  l=subquadrate/ 
other. 

2.  Funiculus  shape:  0  =  filiform  to  subfi- 
liform,  l  =  incrassate. 

3.  Scape  length:  0= surpassing  vertexal 
border  (Figs.  39,  41),  l  =  does  not  surpass, 
at  most  reaching  the  vertexal  border  (Fig. 
43). 

4.  Scape  sculpture:  0= scapes  with  ap- 
preciable sculpture,  1  =  smooth  and  shin- 
ing. The  aforementioned  sculpture  usually 
consists  of  punctulae,  punctae  or  striae. 

5.  Development  of  the  clypeal  lamella: 
0= poorly  developed,  very  short,  l=well 
developed.  This  character  refers  to  how 
far  anterad  the  lamella  extends.  In  poorly 
developed  cases  the  longitudinal  cephalic 
costulation  is  not  clearly  delimited  from 
the  lamella. 

6.  Shape  of  anterior  clypeal  margin: 
0= convex  clypeus  (Figs.  33,  37,  39,  41), 
1  =  straight/concave  (Figs.  34-36,  62,  64). 

7.  Shape  of  lateral  extremities  of  ante- 
rior clypeal  lamella:  O=softly  curves  (Figs. 


33,  37,  39,  41),  1= angular /pointed  (Figs. 
34-35). 

8.  Presence  of  median  clypeal  carinae: 
0= present,  1=  variable,  2  =  absent  or  indis- 
tinct. 

9.  Mandibular  shape:  0  =  triangular 
(Figs.  39,  41),  l=subtriangular/subfalcate 
(Figs.  43,  54,  69,  62). 

10.  Shape  of  cross  section  at  apical  one 
third  of  mandible:  0  =  thin  and  cuneiform 
(Figs.  2-5),  l=wide  and  robust  (Figs.  6,  7). 

11.  Sculpture  of  mandibular  dorsum: 
0=striae/rugulae  (Fig.  54),  l=smooth  and 
shining  (Fig.  43). 

12.  Development  of  promesonotal  su- 
ture: 0  =  deeply  impressed,  cleaving  sculp- 
ture (Figs.  46,  49,  52),  1=  weakly  im- 
pressed, effaced  (Fig.  55). 

13.  Development  of  metanotal  groove: 
0= deeply  impressed  (Fig.  55),  l=weak  to 
effaced  (Figs.  46,  49,  52). 

14.  Development  of  propodeal  denti- 
cles: 0= absent  (Figs.  46,  49,  52),  1= present 
(Figs.  12-14). 

15.  Position  of  spiracle  on  propodeal 
side,  in  lateral  view:  0  =  anterad,  well  sep- 
arated from  declivitous  propodeal  face, 
1=  posterad,  close  to  declivitous  propo- 
deal face. 

16.  Presence  of  setae  on  foretarsal  base 
opposite  the  strigil:  O  =  row  of  stout  setae 
(Fig.  15),  l=one  prominent  seta  only  (Fig. 
16). 

17.  Development   of   dorsal   metacoxal 


Volume  4,  1995 


143 


process:  0  =  absent,  1  =  variable,  2= well  de- 
veloped (Figs.  46,  49). 

18.  Shape  of  the  metasternal  process: 
0  =  stout  denticles  or  tubercles,  l=acicular, 
straight  denticles,  2=acicular,  arched  den- 
ticles. 

19.  Shape  of  petiolar  node:  0=erect  and 
high  node  (Figs.  46,  49,  52),  l=high  and 
posterad,  2  =  subcylindrical/other  (Figs. 
61,  63,  65). 

20.  Petiolar  spiracles:  0  =  directed 
obliquely;  1=  directed  ventrally. 

21.  Petiolar  spiracles:  0  =  -at  the  same 
level  as  the  surrounding  integument;  l=in 
a  distinct  depression. 

22.  Shape  of  postpetiolar  process  in  lat- 
eral view:  O=broad  (Fig.  17),  l=bluntly 
pointed  (Fig.  18). 

23.  A  brief  median  carina  on  the  dor- 
sum of  the  postpetiolar  process:  0= absent, 
1=  present.  This  is  a  low  carinae  that  starts 
at  the  anterior  margin  of  the  process  and 
briefly  extends  posterad. 

24.  Abdominal  segment  IV:  0=with  a 
ventral  stridulitrum,  1=  without  a  ventral 
stridulitrum.  Usually  recognizable  as  a 
longitudinal  band  that  refracts  light  into 
rainbow  colors. 

25.  Shape  of  the  fourth  abdominal  seg- 
ment: 0  =  ventrally  reflexed  (Fig.  55), 
l=straight  (Fig.  44). 

Species  Groups. — Within  the  New  World 
Gnamptogenys  six  species  groups  are  de- 
fined here:  striatula,  sulcata,  concinna,  min- 
uta,  mordax  and  rastrata.  The  internal  clas- 
sification of  the  genus  is  depicted  in  Table 
2.  The  characters  used  in  defining  the  fol- 
lowing informal  groupings  are  diverse, 
thus  they  may  or  may  not  be  synapomor- 
phies.  A  question  mark  following  the  spe- 
cies name  means  that  its  position  within 
the  group  is  not  clear. 

\.  striatula  group:  mandibles  triangular 
with  rugulae  or  striae  on  frontal  surface; 
convex  anterior  clypeal  margin;  scapes 
usually  surpassing  vertexal  margin,  sculp- 
turing punctate  or  vermiculate,  never 
smooth  and  shining;  eyes  slightly  behind 
cephalic  midlength;  head  wider  posterad 


Table   2.    Internal   classification   of   Gnamptogenys 
(except  minuta  group) 

striatula  group 
porcata  subgroup:  acuta,  brunnea,  ejuncida,  extra, 

gentryi,  gracilis,  nigrivitrea, 
pilosa,  pleurodon,  porcata 
striatula  subgroup:  ammophila,  moelleri,  striatula 

strigata  subgroup 
strigata  complex:  andina,  bisulca,  ilimani,  pitti- 

eri,  strigata 
haytiana  complex:         haytiana,  inina,  reichenspergi, 

rclicta 

rastrata  group 

rastrata  subgroup: 
hispinosa  complex:        bispinosa,  perspicax 
rastrata  complex:  cuneiforma,  ingeborgae,  lanei, 

lineolata,  lucaris,  mecotyle, 
meiiozzi,  rastrata,  triangu- 
laris 
banksi  subgroup: 
banksi  complex:  laticephala,  mediatrix 

semiferox  complex:       semiferox 

sulcata  group 

sulcata  subgroup: 

fernandezi  complex:  fernandezi 

sulcata  complex:  acuminata,  curvoclypeata,  tor- 

tuolosa 
ericae  subgroup: 

ericae  complex:  ericae 

lucaris  complex:  lucaris  siapensis 

volcano  complex:  volcano 

mordax  group 

regularis  subgroup: 
hartmaui  complex:        bruchi,  hartmani,  transversa 
regularis  complex:        horni,  nana,  pristina,  regularis, 

rimulosa 
ainndata  complex:        annulata,  kempfi 
mordax  subgroup:  continua,  boliviensis,  interrup- 

ta,  mordax,  stellae 
alfaroi  subgroup:  alfaroi 

levinates  subgroup:         levitjates 

concinna  group 
concinna,  haenschi,  schmitti 


than  anterad;  propodeal  spiracle  close  to 
declivity;  anterolateral  propodeal  declivity 
without  spines  or  tubercles;  anterior  pros- 
ternal  process  broadly  concave  medianly; 
metacoxal  dorsum  always  with  denticle  or 
lobe;  high  petiolar  node;  relatively  wide 
anteroventral  postpetiolar  process;  second 


144  Journal  of  Hymenoptera  Research 

gastric   segment   ventrally   arched.    Most  view;    anterior    clypeal    margin    usually 

seem  to  be  generalist  predators,  but  noth-  straight;  mandibular  front  usually  striate 

ing  is  yet  known  about  the  diet  of  the  stri-  or   rugulose,   sometimes   smooth;   scapes 

gata  subgroup  species.  usually  surpassing  vertex,  sometimes  with 

Three  subgroups  are  recognizable:  1)  longitudinal  rugulae,  vermiculate  or 
striatula  subgroup:  petiolar  node  erect  and  smooth  and  shining;  promesonotal  suture 
sessile;  subpetiolar  process  variably  feebly  impressed  to  absent,  never  totally 
shaped,  either  subquadrate  or  triangular  dividing  sculpture;  metanotal  suture  well 
and  projecting  anterad  in  lateral  view.  2)  impressed;  propodeum  usually  armed 
porcata  subgroup:  petiolar  node  posterior-  with  denticles  or  spines;  petiolar  node 
ly  inclined,  with  short  anterior  peduncle;  low;  subpetiolar  process  shape  variable, 
subpetiolar  process  lobe-like,  projecting  usually  projecting  anterad  but  sometimes 
anterad  in  lateral  view.  3)  strigata  sub-  subquadrate;  metacoxal  teeth  always  pres- 
group:  petiolar  node  variably  shaped;  sub-  ent,  usually  acicular;  second  gastric  seg- 
petiolar  process  subquadrate  with  a  cu-  ment  ventrally  arched.  The  ants  of  this 
neiform  ventral  edge  (Fig.  30)  as  opposed  group  are  specialized  millipede  predators 
to  the  uniformly  parallel  sides  of  the  other  and  can  be  subdivided  into  two  sub- 
striatula  group  species  (Fig.  29).  Many  of  groups.  1)  rastrata  subgroup:  mandibles 
the  species  have  their  propodeal  spiracles  triangular.  Within  the  subgroup  two  com- 
elevated  on  small  prominences.  Most  spe-  plexes  are  defined:  a)  bispinosa  complex: 
cies  are  smaller  in  size  than  those  of  other  large  species,  anterior  clypeal  margin  con- 
subgroups  and  all  are  cryptobiotic  leaflit-  vex,  propodeal  teeth  above  level  of  spira- 
ter  inhabitats.  Two  species  complexes  can  cles;  anterior  lobe  of  subpetiolar  process 
be  pointed  out  in  this  subgroup:  a)  strigata  prominent,  b)  rastrata  complex:  smaller 
complex:  cephalic  vertex  sculptured;  no  species;  anterior  clypeal  margin  straight; 
distinct  propodeal  lobes;  petiolar  node  not  propodeal  teeth  at  same  level  as  spiracles; 
parallel-sided  in  lateral  view,  anterior  subpetiolar  process  subquadrate.  2)  banksi 
margin  usually  convex  (Figs.  40,  57).  All  subgroup:  mandibles  subfalcate  to  falcate, 
species  inhabitat  cloud  forests,  mostly  of  Two  species  complexes  are  defined,  a) 
the  Andes,  with  the  exception  of  one  banksi  complex:  metacoxae  and  propo- 
found  in  the  Cordillera  de  la  Costa  of  Ven-  deum  with  denticles,  b)  semiferox  complex: 
ezuela.  b)  haytiana  complex:  cephalic  ver-  metacoxae  and  propodeum  lacking  denti- 
tex  mostly  smooth  and  shining;  small  pro-  cles.  In  a  study  of  the  sting  apparatus  of 
podeal  lobes  present;  petiolar  node  several  species  of  Gnamptogenys  Kugler 
strongly  compressed  with  subparallel  an-  (1991)  grouped  triangularis  and  bispinosa, 
terior  and  posterior  faces  (Fig.  21);  some  the  only  members  of  the  rastrata  group  as 
species  have  modified  sculpturing  from  defined  above,  into  one  group, 
the  usual  costulate  patterns  of  the  genus.  IIL  mordax  group:  antennal  scapes  usu- 
Most  species  inhabitat  the  Amazon-Ori-  ally  do  not  reach  vertex  and  are  com- 
noco  river  basin,  except  for  one  species  pressed,  smooth  and  shining;  mandibles 
found  on  Hispaniola  Island.  Kugler  (1991)  subtriangular  to  subfalcate  with  front 
obtained  similar  results  studying  the  sting  smooth  and  shining;  head  subquadrate  to 
apparatus  of  brunnea,  nr.  strigata,  porcata,  elongate;  mesosoma  mostly  parallel  sided; 
moelleri  and  gracilis.  They  were  the  only  small  propodeal  lobes  frequently  present, 
members  of  the  striatula  group  included,  denticles  absent  (except  one  species);  me- 
as  defined  above,  and  were  grouped  to-  tacoxal  denticles  or  lobes  frequently  lack- 
gether.  ing;  second  gastric  segment  straight,  with- 

II.  rastrata  group:  head  subquadrate  or  out  ventral  arching.   Considerable  inter- 
wider  anterad   than  posterad   in   frontal  population  variability  in  the  average  size 


Volume  4,  1995 


145 


is  observed  in  species  of  this  group.  The 
group  can  be  divided  into  4  subgroups:  1) 
regulciris  subgroup:  mandibles  subtrian- 
gular;  anterior  clypeal  border  straight  and 
usually  with  laterally  acutely  pointed  la- 
mella; no  transverse  dorsal  mesosomal  su- 
tures; petiolar  node  lacking  anterior  pe- 
duncle; subpetiolar  process  subquadrate. 
Within  this  subgroup  three  species  com- 
plexes are  recognizable,  a)  hartfiiaiii  com- 
plex: mandibles  triangular  to  subtriangu- 
lar,  frontal  surface  partially  sculptured; 
vertical  sculpturing  on  declivitous  propo- 
deal  face;  b)  regiilaris  complex:  mandibles 
subtriangular,  their  frontal  sides  smooth 
and  shining;  vertical  sculpturing  on  de- 
clivitous propodeal  face;  c)  annulata  com- 
plex: mandibles  subtriangular,  front 
smooth  and  shining;  horizontal  sculptur- 
ing on  propodeal  declivity.  2)  mordax  sub- 
group: mandibles  subfalcate,  frontal  sur- 
face smooth  and  shining;  anterior  clypeal 
margin  concave  with  rounded  lamellar 
sides;  well  impressed  metanotal  groove; 
petiolar  node  slightly  pedunculate;  sub- 
petiolar process  projects  anterad.  3)  alfaroi 
subgroup:  mandibles  triangular,  head 
prominently  elongate;  denticles  on  pro- 
podeum.  The  elongate  head  and  armed 
propodeum  makes  this  subgroup  quite 
distinct  from  the  others.  4)  levinates  sub- 
group: subtriangular  mandibles,  frontal 
surface  sculptured;  scapes  surpassing  ver- 
tex; subpetiolar  process  subquadrate.  Its 
clypeus  projects  anterad  more  than  in  any 
extant  species  of  the  group. 

IV.  sulcata  group:  mandibles  subtrian- 
gular, front  smooth  and  shining;  head  su- 
bquadrate; anterior  clypeal  border  usually 
straight;  lamella  usually  laterally  angular; 
scapes  smooth  and  shining;  metacoxal 
denticles  wanting  in  some  species;  usually 
no  transverse  mesosomal  sutures;  petiolar 
node  low,  lacking  peduncle;  metasternal 
process  acicular  and  arched;  second  gas- 
tric segment  relatively  straight;  anteroven- 
tral  postpetiolar  process  relatively  narrow 
and  bluntly  pointed,  in  contrast  with  wid- 
er  process   of   other   species.    Two    sub- 


groups can  be  pointed  out.  1)  sulcata  sub- 
group: posterior  face  of  petiolar  node  with 
horizontal  costulae.  Two  species  complex- 
es are  recognizable:  a)  fernandezi  complex: 
mandibles  triangular;  clypeal  lamella  con- 
vex, b)  tornata  complex:  mandibles  sub- 
triangular; clypeal  lamella  straight.  2)  sul- 
cata subgroup:  posterior  node  face  with 
longitudinal  costulae.  This  subgroup  is 
made  up  of  three  complexes,  a)  ericae  com- 
plex: clypeal  lamella  laterally  acutely  an- 
gulate;  mandibles  subtriangular.  b)  lucaris 
complex:  clypeal  lamella  laterally  round- 
ed; mandibles  subtriangular.  c)  volcano 
complex:  clypeal  lamella  laterally  obtusely 
angulate;  mandibles  triangular. 

V.  concinna  group:  head  subquadrate; 
scapes  surpassing  vertex  in  frontal  view; 
median  clypeal  carina  frequently  present; 
body  sculpture  densely  striate;  small  pit 
frequently  situated  medianly  on  weakly 
impressed  promesonotal  suture;  metaster- 
nal process  slender  to  acicular;  row  of 
stout  setae  on  base  of  foretarsus  opposite 
strigil  present;  petiolar  node  with  no  an- 
terior peduncle;  metacoxae  and  propo- 
deum unarmed.  All  three  members  of  the 
group  are  large  species.  This  group  is  nev- 
ertheless heterogeneous  and  its  three 
members  were  considered  individually  in 
the  matrix. 

VI.  minuta  group:  head  subquadrate; 
frontal  carinae  broadly  expanded  laterad; 
row  of  stout  setae  on  base  of  foretarsus 
opposite  strigil  present;  petiolar  spiracles 
facing  directly  ventrad  and  sunken  within 
a  pit. 

Phylogenetics. — The  possible  evolution- 
ary history  of  the  six  species  groups  is  de- 
picted in  Fig.  19.  Fig.  19  was  computed  by 
the  "ie-"  option  of  Hennig  '86.  Figure  20 
is  a  nelsen  consensus  tree  calculated  from 
13  trees  generated  by  the  "ie"  option  of 
Henning  '86. 

The  striatula  group  subgroups  are  con- 
sidered a  sister  group  to  the  rest  of  the 
Gnamptogem/s  species  considered.  The  sul- 
cata and  mordax  groups  are  the  most  de- 
rived and  are  considered  sister  groups. 


146 


Journal  of  Hymenoptera  Research 


19 


Ectatomma 
strigata  subgroup 
porcata  subgroup 
striatula  subgroup 
minuta  group 
rastrata  group 
G.  schmitti 
G.  haenschi 
G.  concinna 
mordax  group 
sulcata  group 


20 


Ectatomma 
strigata  subgroup 
porcata  subgroup 
striatula  subgroup 
G.  haenschi 
G.  concinna 
G.  schmitti 
rastrata  group 
minuta  group 
mordax  group 
sulcata  group 


Figs.  19-20.  Cladograms  generated  from  the  data  in  Table  1  by  Hennig  '86.  19,  most  parsimonious  tree  found 
by  the  "ie-"  option,  length  51,  ci  58.  20,  Nelsen  consensus  tree  generated  from  the  13  most  parsimonious  trees 
found  by  the  "ie"  option,  length  51,  ci  58. 


These  results  closely  parallel  Kugler's 
(1991)  findings  based  upon  the  sting  ap- 
paratus of  several  Gnamptogeni/s  species. 
Within  each  of  these  groups  we  find  spe- 
cies that  apparently  represent  the  transi- 
tion from  striatida-\ike  ancestors  and  thus 
give  additional  support  to  the  program  re- 
sults: the  hartmani  complexes  of  the  mordax 
group,  and  the  volcano  and  fernandezi  com- 
plexes of  the  sulcata  group.  The  rastrata 
group  is  probably  closer  to  the  striatula 
group  than  to  the  mordax  or  sulcata  groups 
on  account  of  characters  such  as  triangular 
mandibles,  long  and  usually  sculptured 
scapes,  the  convex  clypeal  lamella  of  the 
bispinosa  complex  and  the  well  developed 


metacoxal  tooth.  The  consensus  tree  rec- 
ognizes two  major  groupings  within  the 
genus:  striatula,  and  the  rest.  The  "rest" 
make  up  a  six-branched  polytomy.  Clearly 
further  studies  will  be  needed  in  the  fu- 
ture in  order  to  gain  a  better  picture  of  the 
phylogenetic  history  of  Gnamptogenys. 

The  concinna  group  is  probably  not 
monophyletic.  It  is  diverse  in  mandibular 
shape,  and  in  other  details  such  as  the  de- 
velopment of  the  clypeal  lamella  or  me- 
dian clypeal  carina.  These  traits  as  well  as 
vestigial  promesonotal  suture  are  unusual 
remnants  of  plesiomorphies  when  com- 
pared with  most  other  members  of  the  ge- 
nus. G.  haenschi  retains  more  plesiomor- 


Volume  4,  1995 


147 


phies  than  the  other  two  species  and  could 
be  considered  closer  to  the  rootstock  of  the 
group.  All  are  apparently  specialized,  ei- 
ther in  feeding  or  nesting  habits.  This 
group  opens  a  number  of  questions  which 
can  not  be  answered  to  satisfaction  at 
present.  Are  the  unarmed  metacoxae  ple- 
siomorphic  or  apomorphic  for  this  group? 
Could  they  represent  relicts  of  an  other- 
wise extinct  lineage  (or  lineages)  in 
Gnamptogenys  history?  Interesting  paral- 
lels can  be  found  in  the  shallow  areolate 
ground  sculpture  and  well-developed  ver- 
texal  lobes  of  mimita  subgroup  members 
and  those  of  several  Old  World  species 
groups.  A  comparative  study  with  Old 
World  species  could  shed  new  light  on  the 
phylogeny  of  the  genus.  The  striate  sculp- 
ture of  the  concinna  group  can  also  be 
found,  in  a  lesser  degree,  in  the  suniilans 
complex. 

The  origins  of  Ectatommini  can  be 
traced  to  the  late  Cretaceous  to  early  Ter- 
tiary, at  least  after  the  separation  of  Africa 
from  Gondwana  (80-90  Ma),  but  before 
the  separation  of  South  America  from  Pa- 
leoantarctica  (36-41  Ma)  (Lattke  1994). 
Given  the  presence  of  Gnamptogenys  in  the 
Indomalaysian  Region  and  their  absence 
in  Africa,  the  origin  of  the  genus  also  ex- 
tends at  least  to  this  time  period  and  geo- 
graphically could  be  related  to  southern 
South  America  -Palaeoantarctica.  The 
spread  of  the  ancestral  fauna  from  South 
America  to  Central  America  probably 
took  place  during  the  mid  Tertiary  (30-38 
Ma)  as  the  Panama  land  bridge  devel- 
oped. Chances  for  an  earlier  exchange 
through  possible  connections  between 
parts  of  the  Greater  Antilles  and  South 
America  could  have  existed  during  the 
late  Cretaceous  (Donelley  1988:26),  but  the 
geological  history  of  the  Caribbean  region 
is  still  fraught  with  controversy  (Rull 
1989).  The  mordax  group  fossils  from  Do- 
minican Amber  show  that  by  the  late  Oli- 
gocene  to  early  Miocene  (15-25  Ma),  di- 
versification of  the  American  Gnamptogen- 
ys fauna  was  well  underway.  Potential  ter- 


restrial connections  between  Central 
America  and  the  Greater  Antilles  could 
have  existed  during  the  Cenozoic  via  the 
presently  submerged  Nicaraguan  rise 
which  lays  between  Honduras  and  Jamai- 
ca (Donnelly  1988:28).  It  offered  possibili- 
ties for  dispersal  until  its  subsidence  dur- 
ing the  middle  Cenozoic.  The  presence  of 
the  Hispaniolan  endemic  species  G.  hayti- 
ana,  G.  schmitti,  and  G.  semiferox  lend  sup- 
port to  such  a  connection  and  to  the  di- 
versification of  Gnamptogenys  lineages 
some  26-34  Ma. 

Ecology. — The  ecology  of  Gnamptogenys 
found  in  Venezuela  is  treated  in  Lattke 
(1990)  and  should  be  consulted  for  more 
information.  With  the  exception  of  a  few 
additional  tidbits  of  information  little  is 
added  here.  Generally  most  species  are 
dwellers  of  mesic,  forested  habitats  from 
Texas  and  Louisiana  to  northern  Argenti- 
na. Nests  are  generally  constructed  in  de- 
composing wood  on  the  ground  and  tend 
to  be  small,  rarely  exceeding  500  adults.  A 
few  species  exhibit  polygyny. 

The  striatula  group  species  are  mostly 
generalist  predators,  but  other  species 
groups  have  developed  varying  degrees  of 
specialization.  This  is  taken  to  an  extreme 
by  the  rastrata  group  species  which  prey 
only  on  certain  diplopods.  Beetles  are  the 
favorite  of  many  members  of  other 
groups,  and  some  species  such  as  G.  hart- 
manni  and  G.  horni  have  also  become  fond 
of  other  ants  as  prey. 

New  World  Gnamptogenys: 
A  Synonymic  List 

When  type  material  of  a  determined 
species  was  personally  studied  a  "T"  fol- 
lows the  distribution,  plus  the  acronym  of 
the  collection  where  the  specimen  is  de- 
posited. Type  material,  in  this  case,  in- 
cludes specimens  labelled  either  holotype, 
paratype,  cotype,  type  series,  or  syntype. 

acuniiiiaia  Emery  1869.  Tropical  S.  America. 
acuta  (Brown  1957a).  Colombia-Bolivia.  T:MCZC 
alfaroi  Emery  1894.  Costa  Rica-Ecuador 
aitimopliila  Lattke  1990.  se  Venezuela.  T:MIZA 


148 


Journal  of  Hymenoptera  Research 


andina  Lattke  new  sp.  sw  Colombia-Ecuador. 
annulata  Mayr  1887.  Costa  Rica-tropical  S.  America. 
hanksi  (Wheeler  1930).  Panama-Ecuador.  T:MCZC 
bispinosa  (Emery  1890).  Costa  Rica-Colombia. 
bisiilca  Kempf  y  Brown  1968.  Costa  Rica-Ecuador.  T: 

MCZC 
bufonis  (Mann  1926).  s  Mexico-Nicaragua. 
boliviensis  Lattke,  new  species,  n  Bolivia. 
briichi  (Santschi  1922).  Argentina.  T:MCZC 
brunnea  Lattke,  new  species,  nw  Colombia. 
caelata  Kempf  1967.  Colombia-se  Brazil. 

=  soror  Kempf  and  Brown  1968 
concinna  (F.  Smith  1858).  Mexico-tropical  S.  America. 
T:BMNH 

=  romani  Wheeler  1923. 

=  semicircularis  Borgmeier  1929. 

=  conica  Borgmeier  1929. 
continiia  Mayr  1887  Mexico-tropical  S.  America. 

=  panamensis  Santschi  1931 

=  exarata  Emery,  1901.  new  synonymy. 
cuneiforma  Lattke,  new  species.  Panama. 
curvoclypeata   Lattke   1990.   Colombia-Venezuela.   T: 

MCZC 
ejuncida  Lattke,  new  species,  s  Colombia. 
ericae  Forel  1912,  revalidated,  n  S.  America. 

=  bufonum  Weber  1938,  new  synonymy. 
extra  Lattke,  new  species.  Colombia-Ecuador. 
falcifera  Kempf  1967.  n  S.  America. 
fernandezi  Lattke  1990.  Colombia-Ecuador.  T:MIZA 
fieldi  Lattke  1990.  n  central  Venezuela.  T:MIZA 
gentryi  Lattke,  new  species,  sw  Colombia. 
gracilis  (Santschi  1929).  Guiana  Shield.  T:MCZC 
laticephala  Lattke,  new  species.  Ecuador. 
haenschi  Emery  1902.  Costa  Rica-tropical  S.  America. 
hartmanni  (Wheeler  1915).  s  United  States-n  S.  Amer- 
ica. T:MCZC 

=  nigrifrons  Borgmeier,  1949.  new  synonymy.  T: 
MUSP 

=  turmalis  Kempf  y  Brown,  1968.  new  synonymy. 
T:MCZC 
haytiana  (Wheeler  and  Mann  1914).  Hispaniola.  T: 

MCZC 
horni  Santschi  1929.  Panama-Bolivia.  T:MCZC 
ilimani  Lattke,  new  species.  Bolivian  Andes. 
ingeborgae  Brown  1992.  e  Colombia.  T:MCZC 
interrupta  Mayr  1887.  Mexico-tropical  S.  America,  Ja- 
maica. 
kempfi  Lenko  1964.  Brazil-Peru. 
lanei  Kempf  1960.  Brazilian  Amazonas. 
levinates  Baroni  Urbani  1980.  Dominican  Amber. 
lineolata  Brown  1992.  Hispaniola.  T:MCZC 
lucaris  Kempf  1968.  s  Brazil. 
mecotyle  Brown  1958.  Panama-tropical  S.  America.  T: 

MCZC 
mediatrix  Brown  1958.  Amazon  drainage.  T:MCZC 
menozzi  Borgmeier  1948.  s  Brazil.  TiMUSP 

=  schubarti  (Borgmeier,  1948)  new  synonymy.  T: 
MUSP 


mina  (Brown  1957).  w  S.  America.  T:MCZC 
minuta  (Emery  1896).  Belize-Brazil. 

=  bufonis  (Mann  1922) 

=  emeryi  (Forel  1901) 

=  scabrosus  (Mann  1922) 

=  mus  (Santschi  1931) 

=  panamensis  (Weber  1940) 

=  carinata  (Weber  1940) 

=  pneodonax  Kempf  1968 
moelleri  (Forel  1912).  Tropical  S.  America. 

=  splendens  (Santschi  1929) 

=  concinna  (Santschi  1929) 

=  teffensis  (Santschi  1929)  new  synonymy.  T:MCZC 
niordax  (F.  Smith  1858).  Mexico-tropical  S.  America. 
T:BMNH 

=  nodosa  (Latreille  1802) 

=  purensis  Forel  1912 

=  sebastiani  Borgmeier  1937 
nana  Kempf  1960.  Brazil. 
nigrivitrea  Lattke,  new  species,  sw  Colombia. 
perspicax  Kempf  and  Brown  1970.  Colombia-Ecuador. 

T:MCZC 
petiscapa  Lattke  1990.  ne  Venezuela.  T:MIZA 
pilosa  Lattke,  new  species,  sw  Colombia. 
pittieri  Lattke  1990.  n  Central  Venezuela.  T:MIZA 
pleurodon  (Emery  1896)  Tropical  S.  America. 

=  emeryi  (Santschi  1929). 

=  vidua  (Santschi  1929). 
porcata  (Emery  1896).  Honduras-Bolivia. 

=  magnifica  (Santschi  1921).  new  synonymy. 
pristina  Baroni  Urbani  1980.  Dominican  Amber. 
rastrata  (Mayr  1866).  s  Brazil. 

=  trigona  Emery  1905.  new  synonymy. 
regularis  Mayr  1870.  Mexico-Paraguay. 

=  splendida  Pergande  1895.  T:CASC 

=  fiebrigi  Forel  1909. 

=  arcuata  (Santschi  1929). 
reichenspergi  (Santschi  1929).  Amazon-Orinoco  drain- 
age. 
relicta  (Mann  1916).  Amazon-Orinoco  drainage.  T: 

USNM 
rimulosa  (Roger  1861).  s  Brazil. 
schmitti  (Forel  1901).  Hispaniola. 

=  minor  (Wheeler  1936).  T:MCZC 
semiferox  Brown  1958.  Hispaniola.  T:MCZC 
siapensis  Lattke,  new  species,  s.  Venezuela. 
simulans  (Emery  1896).  Costa  Rica.  T:USNM 
stellae  Lattke,  new  species.  Costa  Rica. 
striatula  Mayr  1883.  Mexico-Argentina,  Caribbean.  T: 
BMNH 

=  curtula  (Emery  1896)  new  synonymy. 

=  stolli  (Forel  1899)  new  synonymy. 

=  brasiliensis  (Emery,  1902). 

=  angustiloba  (Forel  1908). 

=  paulina  (Forel  1908). 

=  simplicoides  (Forel  1908)  new  synonymy. 

=  pernambucana  (Santschi  1929). 

=  calcarata  (Santschi  1929). 


Volume  4,  1995 


149 


=  antillana  (Santschi  1929).  T:MCZC 

=  hybrida  (Santschi  1929). 

=  mayri  (Santschi  1929). 

=  isthmica  (Santschi  1929). 

=  recta  (Santschi  1929).  T:USNM 

=  rustica  (Santschi  1929)  new  synonymy. 

=  wheeleri  (Santschi  1929)  new  synonymy.  T:MCZC 

=  arcuata  (Santschi  1929)  new  synonymy. 

=  wasmanni  (Santschi  1929)  new  synonymy. 
striolata  (Borgmeier  1957).  se  Brazil.  T:MUSP 
strigata  (Norton  1871).  Mexico-Colombia. 

=  simplex  (Emery  1896). 
sulcata  (F.  Smith  1858).  Mexico-tropical  S.  America.  T: 
BMNH 

=  tornata  (Roger  1861).  new  synonymy. 


=  lineata  Mayr  1870. 

=  cearensis  Forel  1912. 

=  nitens  Mann  1916.  TiUSNM 

=  ypirangensis  Borgmeier  1928. 
tortuolosa  (F.  Smith  1858).  Amazon-Orinoco  drainage. 
T:BMNH 

=  quitensis  Forel  1920. 
transversa  Lattke,  new  species.  Panama. 
triangularis  Mayr  1887.  Panama-Argentina;  se  United 
States. 

=  richteri  (Forel  1920) 

=  aculeaticoxae  (Santschi  1921)  new  synonymy. 
volcano  Lattke,  new  species.  Costa  Rica. 
vriesi  Brandao  &  Lattke,  1990.  Ecuador. 


KEY  TO  THE  WORKERS  OF  THE  NEW  WORLD  SPECIES  OF  GNAMPTOGENYS. 

Note:  In  some  species  the  difference  between  the  promesonotal  suture  and  the  metanotal  groove  may  be 
hard  to  establish  when  only  one  of  the  two  is  present,  which  is  the  usual  case.  In  a  dorsal  view  the  lateral 
margins  of  the  pronotum  converge  posterad  and  at  the  end  of  the  convergence,  when  the  margins  more  or 
less  straighten  out,  the  promesonotal  suture  is  usually  slightly  arched  anteriorly,  while  the  metanotal  groove 
is  usually  straight. 

1.  Petiolar  spiracle  directed  ventrally  and  within  a  depression   ....   mn;Hfrt  group  (see  Lattke  1991) 

-  Petiolar  spiracle  directed  anteroventrally  and  not  depressed;  Pronotum  separated  from 
mesonotum  by  very  distinct  suture  which  completely  cuts  dorsal  sculpture    2 

-  Petiolar  spiracle  directed  ventrolaterally  and  not  depressed;  Promesonotal  suture  absent 

to  well  impressed,  but  never  completely  cuts  sculpture     23 

2.  Petiolar  in  lateral  view  with  subparallel  anterior  and  posterior  margins  sharply  set  off  at 
right  angles  to  relatively  brief  dorsal  margin  (Fig.  21);  vertex  usually  smooth  and  shining, 
occasionally  with  weak  transverse  triae     3 

-  Petiole  not  as  above,  in  lateral  view  with  anterior  margin  either  broadly  curving  into 
dorsal  margin  or  separated  by  broad  angle  (Figs.  22-4,  46,  49,  52);  vertex  strongly  sculp- 
tured           6 

3.  Mesosomal  dorsum  with  extensive  smooth  and  shining  areas     4 

-  Mesosomal  dorsum  totally  sculptured,  usually  rugulose  or  costulate 5 

4.  Metanotal  groove  well  impressed,  breaking  sculpture;  metacoxae  armed;  anterolateral 
lobes  on  declivitous  propodeal  face  lacking  (Amazon-Orinoco  drainage)    relicta 

-  Metanotal  groove  absent;  metacoxae  unarmed;  small  lobes  present  on  anterolateral  de- 
clivitous propodeal  face    (Amazon-Orinoco  drainage)   reichenspergi 

5.  Postpetiolar  dorsum  transversely  rugulose;  body  color  yellowish  (Hispaniola)    ....  haytiana 

-  Postpetiolar  dorsum  longitudinally  costulate;  black  to  dark  brown  (w  South  America)    ....  mina 

6.  Mesosomal  dorsum  with  well  developed  metanotal  groove    7 

-  Metanotal  groove  absent    9 

7.  Dorsum  of  gastric  tergites  I  and  II  smooth  and  shining  (sw  Colombia)    gentryi  n.  sp. 

-  Dorsum  of  gaster  with  longitudinal  costulae    8 

8.  Subpetiolar  process  subquadrate,  with  posterior  angle  (Fig.  21;  Costa  Rica  to  Ecuador, 
Andes)   bisulca 

-  Subpetiolar  process  shaped  as  an  anteriorly  projecting  lobe  with  no  posterior  angle  Fig. 

42)  (nw  Colombia)    brunnea  n.  sp. 

9.  Petiolar  node  in  lateral  view  relatively  erect,  posterodorsal  and  anterodorsal  angles  not 
differing  notably  (Fig.  22)     10 

-  Petiole  in  lateral  view  posteriorly  inclined,  anterior  margin  joins  dorsal  margin  through 
broad  convexity  that  contrasts  with  sharp  angle  between  posterior  and  dorsal  margins 
(Figs.  23,  24,  46,  49,  52)     12 


150  Journal  of  Hymenoptera  Research 

10.  SL  <  1.08  mm   11 

-  SL  >  1.08  mm  (tropical  S.  America)    moelleri 

11.  Declivitous  propodeal  face  with  5-11  longitudinal  costulae  between  spiracles  and  1-3 
transverse  costulae  between  each  spiracle  and  longitudinal  costulae  (Fig.  26);  decumbent 
pubescence  on  scapes  sparse,  <  six  decumbent  and  suberect  hairs  (Mexico  to  Argentina, 
Caribbean) striatula 

-  Declivitous  face  with  13-16  longitudinal  costulae  between  spiracles  and  no  transverse 
costulae  between  each  spiracle  and  longitudinal  costulae  (Fig.  25);  scapes  with  dense  and 
uniform  decumbent  pubescence,  1-3  subdecumbent  hairs  (se  Venezuela)    ammophila 

12.  Posterodorsal  angle  of  petiolar  node  forms  a  sharp  to  bluntly  pointed  apex  that  overhangs 
posterior  margin  (Figs.  23,  24,  46,  49,  52);  subpetiolar  process  in  ventral  view  with  uni- 
formly narrow  ventral  edge,  posteriorly  widening  only  at  junction  with  node  (Fig.  29)     13 

-  Petiolar  node  not,  or  slightly  overhanging  posterior  margin  (Figs.  40,  57);  subpetiolar 
process  in  ventral  view  with  cuneiform  ventral  edge:  anteriorly  thin,  wider  posterad  with 
bifurcate  apex  that  forms  posterior  angle  of  subquadrate  process  (Fig.  30)     20 

13.  Petiolar  node  with  a  blunt  posterodorsal  apex  (Figs.  24,  46,  49,  52)    14 

-  Petiolar  node  with  acute  posterodorsal  point  (Fig.  23;  Colombia  to  Bolivia)     acuta 

14.  Postpetiolar  sternum  totally  costulate  or  striate;  vertex  usually  with  one  (occasionally  up 

to  3)  transverse  costulae  next  to  vertexal  carinae    15 

-  Postpetiolar  sternum  with  a  basal  median  smooth  and  shining  area;  vertex  with  4-5  trans- 
verse costulae  next  to  vertexal  carinae  (Guiana  Shield)    gracilis 

15.  SSC  <  10  (Fig.  27)     18 

-  SSC  >  10  (Fig.  28)     16 

16.  Body  and  legs  brown,  sculpture  costulate  to  striate;  body  with  abundant  decumbent  pu- 
bescence; subpetiolar  process  lobiform  and  without  acute  projection    17 

-  Body  piceous  and  legs  ferruginous;  body  with  abundant  pubescence;  subpetiolar  process 
usually  with  an  acute  anteroventral  projection,  sometimes  blunt,  especially  in  specimens 
from  Central  America  (Honduras  to  Bolivia)    porcata 

17.  Femora  with  abundant  erect  to  suberect  hairs;  ventral  surfaces  of  meso-  and  metacoxae 
with  erect  hairs;  gastric  sculpture  striate  (sw  Colombia)    pilosa,  n.  sp. 

-  Femora  with  decumbent  to  subdecumbent  hairs  and  few  or  no  erect  to  suberect  hairs; 
ventral  surfaces  of  meso-  and  metacoaxe  wirthout  erect  hairs;  gastric  sculpture  costulate 

(s  Colombia)    ejuncida,  n.  sp. 

18.  Five  or  more  standing  hairs  on  scapes  and  tibiae     19 

-  One  or  no  suberect  and  subdecumbent  hairs  on  scapes  and  tibiae  (Colombia-Ecuador) 

extra,  n.  sp. 

19.  Declivitous  propodeal  face  with  transverse  costulae;  petiolar  node  with  anterior  peduncle 
(Fig.  24;  tropical  S.  America)    pleurodon 

-  Declivitous  propodeal  face  with  longitudinal  costulae;  node  without  peduncle  (Fig.  59; 

sw  Colombia)    nigrivitrea,  n.  sp. 

20.  Postpetiolar  sternum  with  well  defined  costulae  or  striae;  dorsal  propodeal  face  not  no- 
tably depressed  below  mesonotum,  dorsal  and  declivitous  faces  confluent    21 

-  Postpetiolar  sternum  with  weak  irregular  rugae;  in  lateral  view  dorsal  propodeal  face 
separated  from  mesonotum  by  notable  depression,  dorsal  and  declivitous  faces  sharply 
separated  by  a  ridge  (n  Central  Venezuela)    pittieri 

21.  Scapes  with  moderate  to  weak  pubescence,  five  or  more  standing  hairs  present     22 

-  Scapes  with  dense,  white  subdecumbent  pubescence,  0-2  standing  hairs  present  (Bolivian 
Andes)    ilimani  n.  sp. 

22.  HW  >  0.84,  WL  >  0.35;  Ol  <  0.16  (sw  Colombia-Ecuador)    andina,  n.  sp. 

-  HW  <  0.84,  WL  <  0.35,  OI  >  0.16  (Mexico-Colombia)    strigata 

23.  With  head  in  frontal  view  antennal  scapes  barely  or  do  not  reach  the  vertexal  margin, 
never  beyond,  when  laid  back  as  straight  as  possible  from  insertions    24 

-  Scapes  clearly  reach  beyond  vertexal  margin    38 


Volume  4,  1995  151 

24.  Mandibular  dorsum  with  continuous  striae  or  costulae  over  the  basal  one-fourth  or  more 

of  length    25 

-  Mandibular  dorsum  entirely  smooth  and  shining,  sometimes  with  scattered  punctures    27 

25.  Distinct  median  carinae  visible  on  clypeus;  eyes  small,  OI  <  0.1;  mandibular  dorsum 
entirely  and  densely  striate  (Costa  Rica-tropical  S.  America)    haenschi 

-  No  distinct  median  carinae  on  clypeus;  eyes  larger,  OI>  0.2;  mandibular  dorsum  without 
dense  striae   26 

26.  Mandibular  dorsum  with  costulae  on  basal  one  fourth  or  less,  rest  smooth  and  shining; 
propodeal  declivity  with  longitudinal  costulae  (s  United  States-n  South  America)    .... 

hartmani  (in  part) 

-  Mandibular  dorsum  mostly  rugulose;  propodeal  declivity  with  transverse  costulae    .  .  . 

transversa  n.  sp. 

27.  Propodeal  dorsum  mostly  transversely  striate    28 

-  Propodeal  dorsum  mostly  longitudinally  costulate  or  striate,  continuous  with  rest  of  notal 
sculpturing     30 

28.  Head  subquadrate,  CI  >  0.78;  body  brown  to  dark  brown   29 

-  Head  notably  rectangular  and  elongate,  CI<  0.78;  body  black  (Costa  Rica-Ecuador)     .  .  .  .alfaroi 

29.  Costulae  on  mesosomal  and  petiolar  dorsum  subopaque;  propodeum  with  small  teeth; 
metacoxal  teeth  (Brazil  and  Peru)    kempfi 

-  Very  fine  striae  on  mesosoma  and  node;  no  teeth  on  propodeum  nor  on  metacoxae  (Costa 
Rica  to  tropical  S.  America)    attnulata 

30.  Clypeal  lamella  medianly  straight  and  laterally  acutely  angulose;  subpetioloar  process 
subquadrate,  usually  with  acute  posteroventral  tooth    36 

-  Clypeal  lamella  usually  concave  and  laterally  rounded,  never  acutely  angular  (rarely  with 
slight  median  projection  and  laterally  bluntly  angular);  subpetiolar  process  usually  pro- 
jecting anteriorly  as  subtriangular  lobe,  sometimes  subquadrate,  but  never  with  acute 
posteroventral  tooth    31 

32.  Mandibles  subtriangular  to  subfalcate;  metanotal  groove  well-impressed     32 

-  Mandibles  triangular;  metanotal  groove  absent  (Brazil)   nana 

32.  Metacoxa  with  dorsobasal  lobe  or  tooth    34 

-  Metacoxa  lacking  tooth  or  lobe,  at  most  small  swelling  or  low  tubercle  present    33 

33.  Body  sculpture  striate;  gastric  tergum  2  smooth  and  shining;  clypeal  lamella  laterally 
bluntly  angulose,  slightly  projecting  anterad  medianly  and  with  small  concavity  in  middle 

of  projection  (Fig.  60;  Costa  Rica)     stellae  n.  sp. 

-  Body  sculpture  costulate;  gastric  tergum  2  longitudinally  costulate;  clypeal  lamella  lat- 
erally rounded  and  medianly  concave  (Fig.  43;  Mexico  to  tropical  S.  America,  Jamaica) 

interrupta 

34.  Metacoxal  dorsum  with  low  triangular  lobe;  HW  <  1.12,  WL  <  1.81  mm 35 

-  Metacoxal  dorsum  with  high,  approximately  parallel-sided  lobe  or  tooth;  HW  usually  > 
1.12,  WL  >  1.81  mm  (Mexico  to  tropical  S.  America)     mordax 

35.  Cephalic  dorsum  striate;  anterolateral  lobes  of  declivitous  propodeal  face  weakly  devel- 
oped (n  Bolivia)     holiviensis,  n.  sp. 

-  Cephalic  dorsum  costulate;  anterolateral  propodeal  lobes  well  developed  (Mexico  to  trop- 
ical S.  America)   continua 

36.  Metanotal  groove  weakly  impressed,  visible  only  with  limited  angles  of  view;  subpetiolar 
process  with  acute  posteroventral  tooth;  anterolateral  angles  of  declivitous  propodeal  face 
bordered  by  raised  carinae  that  cross  over  costulate  sculpture    37 

-  Metanotal  groove  well-impressed,  visible  with  any  angle  of  view;  subpetiolar  process 
without  acute  posterior  tooth;  declivitous  propodeal  face  lacking  carinae  different  from 
costulate  sculpture  (s.  Brazil)    rimiilosa 

37.  Declivitous  propodeal  face  with  longitudinal  costulae  that  converge  posterad;  anterior 
nodal  face  with  transverse  costulae;  meso-metapleural  suture  absent  or  very  weakly  im- 
pressed (Panama  to  Bolivia)     homi 


152  Journal  of  Hymenoptera  Research 

-  Declivitous  propodeal  face  with  longitudinal  parallel  costulae;  anterior  nodal  face  mostly 
longitudinally  costulate,  basal  transverse  costulae  frequently  present  and  rarely  occupy 

all;  meso-metapleural  suture  distinctly  impressed  (Mexico  to  Paraguay)    regularis 

38.  Mandibles  very  elongate:  falcate  or  subfalcate,  only  their  apices  touching  or  crossing  when 
closed  (Figs.  35,  54)     39 

-  Mandibles  triangular  or  subtriangular,  apical  and  basal  margins  separated  by  rounded 

or  angular  basal  angle  (Figs.  33-34,  36-37)    43 

39.  Declivitous  propodeal  face  with  spines  or  denticles;  metacoxal  tooth  present    40 

-  Declivitous  propodeal  face  rounded,  no  trace  of  spines  or  denticles;  coxal  tooth  absent    42 

40.  Inner  basal  mandibular  border  with  short  triangular  tooth  that  represents  basal  angle 
(Fig.  54)    41 

-  Inner  basal  mandibular  border  with  broadly  convex  lobe  or  flange  (Amazon  drainage) 

mediatrix 

41.  Clypeus  with  two  lobes  betwen  antennal  fossae  and  anterior  lamella,  lobe  protrude  over 
lamella  in  front  view;  mesothoracic  spiracles  not  set  in  deep  depressions  (Ecuador)     .  . 

laticephala,  n.  sp. 

-  Clypeus  without  lobes  between  antennal  fossae  and  anterior  lamella;  spiracles  of  meso- 
thorax  set  in  deep  depressions  (Panama  to  Ecuador)   banksi 

42.  Mandibles  long  and  slender  (Fig.  36);  dorsal  propodeal  face  evenly  and  transversely  cos- 
tulate (Hispaniola)     schmitti 

-  Mandibles  shorter  and  wider;  dorsal  propodeal  face  subvermiculately  and  longitudinally 
costulate  (Hispaniola) semiferox 

43.  Mandibles  mostly  smooth  and  shining,  with  scattered  punctures,  subtriangular  to  trian- 
gular, sometimes  striae  or  costulae  present  on  basal  one-third  or  less;  usually  without 
transverse  sutures  on  mesosomal  dorsum  (except  concinna)     44 

-  Mandibles  with  longitudinal  rugulae  or  striae  (sometimes  weakly  impressed),  always 
triangular,  never  subtriangular;  metanotal  suture  distinctly  impressed     55 

44.  HW  >  1.43;  WL  >  2.27  mm     45 

-  HW  <  1.43;  WL  <  2.27  mm     46 

45.  Sculpture  costulate;  metacoxae  with  dorsal  tooth;  petiolar  node  ends  in  blunt  angle;  black 
(Amazon-Orinoco  drainage)     tortuolosa 

-  Sculpture  striate;  metacoxae  without  dorsal  tooth,  with  low  tubercle  at  most;  petiole  ends 

in  acute  point  or  beak;  ferruginous  (Mexico  to  tropical  S.  America)    concinna 

46.  Sculpture  striate;  anterolateral  corners  of  declivitous  propodeal  face  borderd  by  carinae 
that  cross  costulae;  antennal  scape  when  laid  back  does  not  surpass  vertexal  margin  by 
more  than  one  apical  width   47 

-  Sculpture  costate;  declivitous  propodeal  face  without  carinae  distinct  from  costae;  scapes 
surpass  vertexal  margin  by  more  than  an  apical  width    48 

47.  Posterior  face  of  petiolar  node  longitudinally  costulate  (s  United  States  to  n  S.  America) 

hartmani  (in  part) 

-  Posterior  nodal  face  transversely  costulate  (Argentina)    bruchi 

48.  Posterior  face  of  petiolar  node  transversely  costulate     49 

-  Posterior  face  of  node  longitudinally  costulate   51 

49.  Anterior  clypeal  lamella  laterally  acutely  angular;  mandibles  subtriangular,  basal  and 
apical  margins  joined  by  very  broad  continuous  convexity  (Fig.  34)     50 

-  Clypeal  lamella  very  evenly  convex  laterad;  mandibles  triangular,  basal  and  apical  mar- 
gins separated  by  angle  (Fig.  33;  Venezuela  to  Ecuador)   femandezi 

50.  Petiolar  node  with  posterior  pointed  projection;  declivitous  propodeal  face  with  longi- 
tudinal costulae;  body  color  uniformly  brown  (tropical  S.  America) acuminata 

-  Node  sometimes  with  slight  posterior  projection,  but  never  pointed;  declivitous  propodeal 
face  with  transverse  costulae,  rarely  longitudinal;  color  variable,  usually  with  brown  and 
ferruginous  areas,  rarely  totally  brown  (Mexico  to  tropical  S.  America)   sulcata 

51.  Anterior  clypeal  lamella  laterally  acutely  angulate     52 


Volume  4,  1995  153 

-  Clypeal  lamella  laterally  rounded  or  obtusely  angulate   53 

52.  HW  <  1.11,  WL  <  1.72  mm;  anterior  clypeal  lamella  straight  (Fig.  31;  n  S.  America)    .  .  . 

ericae  reval.  sp. 

-  HW  >  1.11,  WL  >  1.72  mm;  anterior  clypeal  lamella  medially  convex  and  laterally  con- 
cave (Fig.  32;  Venezuela  and  Colombia)    ciirvoclypeata 

53.  Declivitous  propodeal  face  with  longitudinal  costae;  clypeal  lamella  straight  anterad  and 
laterally  rounded;  mandibles  subtriangular,  basal  and  apical  margins  meeting  through 
broad  convexity  (Fig.  62)    54 

-  Declivitous  propodeal  face  with  transverse  costae;  clypeal  lamella  excised  mesad  and  with 
two  lateral  convexities;  mandibles  roughly  triangular,  apical  and  basal  margins  separated 

by  notch  (Fig.  64;  Costa  Rica)     volcano,  n.  sp. 

54.  Petiolar  node  with  dorsal  margin  more  or  less  evenly  convex  in  lateral  view;  metacoxal 
tooth  denticle-like;  subpetiolar  process  with  acutely  pointed  angles  (s  Brazil)    lucaris 

-  Petiolar  node  with  dorsal  surface  flat;  metacoxal  tooth  broadly  triangular;  subpetiolar 
process  bluntly  angulate  (s  Venezuela)    siapensis,  n.  sp. 

55.  Antennal  scapes  usually  smooth  and  shining   56 

-  Scapes  with  longitudinal  striae  or  rugulae  (sometimes  partially  effaced)    60 

56.  Scapes  surpass  vertexal  margin  by  two  or  more  apical  widths;  HW  >  1.07,  WL  >  1.64 
mm     59 

-  Scapes  when  laid  back  do  not  surpass  vertexal  margin  by  more  than  one  apical  width; 
HW  <  1.07,  WL  <  1.64  mm     57 

57.  Body  sculpture  striate;  no  propodeal  denticles;  ventral  petiolar  process  in  lateral  view 
projecting  anterad  as  a  lobe,  lacking  a  posterior  angle;  Hispaniola  Island    lineolata 

-  Sculpturing  costulate;  propodeal  denticles  present;  petiolar  ventral  process  with  a  pointed 
posterior  corner;  South  America     58 

58.  Costulae  on  dorsum  of  metanotum  and  propodeum  well  defined;  petiole  in  dorsal  view 
with  concentric  costulae  (s  Brazil)    rastrata 

-  Costulae  on  dorsum  of  metanotum  and  propodeum  mostly  effaced;  petiole  in  dorsal  view 
transversely  costulate  (e  Colombia)   ingeborgae 

59.  Propodeal  teeth  longer  than  diameter  of  propodeal  spiracles;  petiolar  node  longer  than 
wide  and  subpetiolar  process  projects  anterad  as  triangular  lobe  (Brazilian  Amazon  drain- 
age)     land 

-  Propodeal  teeth  shorter  than  diameter  of  propodeal  spiracles;  petiolar  node  wider  than 
long;  subpetiolar  process  subquadrate  (s  Brazil)     menozzi 

60.  Anterior  pronotal  margins  with  at  least  five  transverse  costulae  or  rugulae;  eyes  promi- 
nent and  bulging  (Fig.  37)    61 

-  Anterior  pronotal  margin  with  longitudinal  costulae  that  extend  caudad  across  rest  of 
mesosomal  dorsum;  eyes  not  bulging  and  prominent,  relatively  flat  (Fig.  38;  Florida,  Pan- 
ama to  Argentina)     triangularis 

61.  Vertex  with  4-5  transverse  costulae;  anterior  one-half  of  postpetiolar  tergum  rugose   ...     62 

-  Vertex  with  longitudinal  costulae;  anterior  one-half  of  postpetiolar  tergum  longitudinally 
costulate     63 

62.  Dorsum  of  metanotum  and  all  of  propodeum  with  transverse  costulae  (Panama)     .... 

cuneiforma  n.  sp. 

-  Dorsum  of  metanotum  and  propodeum  longitudinally  costulate;  declivitous  propodeal 
face  with  weak  longitudinal  costulae,  smooth  and  shining  mesad  (Panama  to  tropical  S. 
America)    mecotyle 

63.  Propodeal  spines  at  least  as  long  as  distance  between  inner  sides  of  their  base  (Fig.  12); 
ferruginous  species  (Costa  Rica  to  Colombia)   bispinosa 

-  Propodeal  spines  shorter  (Fig.  13);  black  species  (Colombia  to  Ecuador)    perspicax 


154 


Journal  of  Hymenoptera  Research 


New  World  species  of  Gnamptogenys 

In  this  section  the  species  are  arranged 
in  alphabetical  order.  In  the  synonymy 
listings,  when  more  than  one  citation  fol- 
lows a  junior  synonym,  the  first  author, 
after  the  author  of  the  species,  is  respon- 
sible for  that  synonymy.  The  diagnosis  is 
meant  to  complement  the  results  obtained 
in  the  key.  In  order  to  avoid  repeating  in- 
formation already  published  the  reader 
should  consult  Lattke  (1990)  for  additional 
data  concerning  ecology  and  localities  of 
species  previously  known  from  Venezue- 
la. For  the  sake  of  brevity  the  dates  and 
collector's  names  have  been  omitted. 

Gnamptogenys  acuminata  Emery 

Gnamptogenys  acuminata  Emery  1896:50;  Brown 
1958:228,^236,  299;  Kempf  1961:491;  Kempf 
1972:111;  Kempf  1976:51;  Lattke  1990:9. 

Diagnosis. — Mandibles  subtriangular; 
longitudinal  costulae  on  mesosoma  and 
declivitous  propodeal  face;  transverse  su- 
tures weak,  visible  only  with  certain  an- 
gles of  light;  body  dark  brown  to  black; 
coxae  and  basal  two-thirds  of  femora  tes- 
taceous to  brown-testaceous;  tibiae,  apex 
of  femora,  antenna  and  mandibles  brown. 

Ecology. — Found  in  humid  forests,  nest- 
ing in  rotting  wood. 

Comments. — The   acute   posterior   tooth 


on  the  petiolar  node,  combined  with  lon- 
gitudinal costulae  on  the  declivitous  pro- 
podeal face,  seem  to  provide  the  most  re- 
liable distinction  from  tornata.  Color  in  ac- 
uminata is  not  so  variable  but  light-pig- 
mented  forms  can  occasionally  be  found. 
The  mesometepisternal  suture  is  usually 
absent. 

Specimens  Examined. — BRAZIL,  Amazo- 
nas:  Ig.  Marianil,  Rio  Branco  Rd,  24  km 
NE  Manuas.  COLOMBIA,  Meta:  Cano  EL 
Buque,  S  Villavicenicio,  480  m.  PERU,  Lo- 
reto:  15  km  WSW  Yurimagua,  5°59'S 
76°13'W,  220  m;  Panguana:  9°37'N 
74°56'W,  220  m.  VENEZUELA,  Bolivar: 
Rio  Cuyuni,  66  km  SSE  El  Dorado,  6°  09' 
N  61°  30'W,  250  m;  10  km  E  Icabaru,  700 
m.  Amazonas:  Rio  Baria,  0°50'N  66°10'W, 
140  m. 

Gnamptogenys  acuta  Brown 

Holcoponera  acuta  Brown  1957:491. 
Gnamptogenys  acuta  (Brown);  Brown  1958:300; 
Kempf  1972:111. 

Diagnosis. — Similar  to  striatula;  four  to 
six  transverse  costulae  on  anterior  prono- 
tal  face,  rest  of  mesosoma  with  longitudi- 
nal costulae;  longitudinal  costulae  on  an- 
terior face  of  postpetiole;  body  dark 
brown  to  black,  legs  brown. 


Figs.  21-24.     Lateral  view  of  petiole.  21,  G.  relicta;  22,  G.  inoelleri;  23,  G.  acuta;  24,  G.  pleurodon. 


Volume  4,  1995 


155 


26 

Figs.  25-26.     View  of  declivitous  propodeal  face.  25,  G.  ammophila;  26,  G.  striatiila. 

Figs.  27-28.     Frontal  view  of  scapes,  showing  hairs  and  pilosity.  27,  G.  pleurodon;  28,  G.  porcata. 


Ecology. — Found  in  low  to  premontane 
jungles. 

Comments. — Not  frequently  collected, 
knowledge  of  its  range  has  expanded 
slowly. 

Specimens  Examined. — ECUADOR, 
Napo:  2-8  miles  N  Puyo,  Pastaza,  935  m; 
original  locality  unknown:  Quarantine 
Station,  San  Pedro,  California.  PERU,  Ju- 
nin:  Colonia  Perene,  Rio  Perene,  18  km  NE 
La  Merced;  Loreto:  Boqueron,  500  m;  CO- 
LOMBIA, Meta,  Mesetas,  780  m. 

Gnamptogenys  alfaroi  Emery 

Gnamptogenys  alfaroi  Emery   1894:145;   Brown 
1958:301;'Kempf  1972:111. 

Diagnosis. — Mandibular  dorsum  smooth 
and  shining  with  some  punctulae;  meta- 
coxal  spine  present;  subpetiolar  process 
anteriorly  projected;  gastric  tergum  2  with 
basal  one-half  longitudinally  costulate,  the 
rest  smooth  and  shining;  disk  of  pospetio- 
lar  sternum  smooth  and  shining;  body 
black;  legs  and  antennae  brown. 

Ecology. — Known  from  montane  rain 
forest. 

Comments. — The  Ecuador  record  ex- 
tends its  distribution  from  the  only  pre- 
viously known  localities  in  Costa  Rica. 
This  species  is  a  striking  member  of  the 
mordax  group  due  to  its  elongated,  almost 
cylindrical  body.  Its  relationship  with  oth- 
er members  of  the  mordax  group  is  not 
clear.  It  is  rarely  collected. 

Specimens  Examined. — COSTA   RICA, 


Turrialba.  ECUADOR,  Guayas:  3  km  SW 
Bucay  (MCZC). 

Gnamptogenys  ammophila  Lattke 

Gnamptogenys  ammop>hila  Lattke  1990:9. 

Diagnosis. — Black,  finely  costulate  with 
brown  scapes  and  mandibles,  light  brown 
legs.  Subpetiolar  process  subquadrate,  not 
as  projecting  anterad  as  in  striatula.  Me- 
sosomal  dorsum  with  longitudinal  costu- 
lae  and  erect  hairs  up  to  0.30  mm  in 
length. 

Ecology. — This  species  is  unique  within 
the  genus  due  to  its  savanna  nest  sites  as 
opposed  to  the  forest  preferences  of  most 
species.  Its  range  is  apparently  restricted 
to  high  (>1200m),  cool  savannas  of  the 
Upper  Rio  Caroni  watershed  (La  Gran  Sa- 
bana).  It  has  not  been  found  in  lower  sa- 
vannas, despite  the  fact  that  another  ant, 
Pogonomyrmex  naegeli,  is  widespread  in  sa- 
vannas isolated  in  the  midst  of  great  ex- 
panses of  forest.  The  extent  of  savannas  in 
the  Guiana  Shield  was  much  broader  dur- 
ing the  last  glaciation  (75,000-11,000  B.P.), 
as  were  generally  cooler  temperatures  and 
a  drier  climate  (Schubert  1988;  Clapperton 
1993).  Such  conditions  could  concievably 
have  favored  the  origin  of  this  species 
from  its  apparent  sister  species  G.  striatula 
during  the  course  of  the  last  glaciation. 

Comments. — See  Lattke  1990. 

Specimens  Examined. — VENEZUELA,  Bo- 
livar: via  El  Doradc-)-Santa  Elena,  km  141, 
Rio  Apanwao,  5°52'N  61°26'W,  1500  m. 


156 


Journal  of  Hymenoptera  Research 


^: 


iT 


31 


29 


30 


32 


Figs.  29-30.     Ventral  view  of  petiole  showing  shape  of  subpetiolar  process  (outlined  in  black).  29,  G.  gracilis; 
30,  G.  andina. 

Figs.  31-32.     Frontal  view  of  clypeal  lamella.  31,  G.  sulcata;  32,  G.  curvoclypeata. 


Gnamptogenys  andina  Lattke, 
new  species 
(Figs.  39-40) 

Type  Specimens  Examined. — Holotype 
worker.  COLOMBIA,  Valle:  Parque  Far- 
allones  de  Cali,  El  Topacio,  20  km  NW 
Cali,  76  37'N  2  30'W,  1550  m,  31-XII-1981, 
J.  Lattke  No.  220.  Deposited  in  MIZA. 
Paratypes  (all  from  Colombia,  Valle):  (1). 
22  workers  from  tye  same  nest  series  as 
the  holotype.  (2)  21  workers,  same  data  as 
holotype  except  collection  numbers  220, 
214,  218.  (3).  18  workers,  Cali-Buenaven- 
tura  road,  km  21,  15  km  NE  Cali,  1300- 
200  m,  l-X-1975,  J.  Lattke,  leg.  (4).  6  work- 
ers, CVC  station  near  Pance,  15  km  W 
Cali,  1700  m,  12-XII-1975,  J.  Lattke,  leg. 

Additional  Specimens  (not  paratypes). — 
ECUADOR,  Bolivar,  20  km  de  Palzabam- 
ba,  1500-1800  m,  18-X1-87,  N.  Zavala,  leg. 
Six  paratypes  deposited  in  MCZC  and 
three  in  each  of  the  following  :  LACM, 
BMNH,  CASC,  MUSP.  The  remainder  of 
paratypes  and  other  specimens  deposited 
in  MIZA. 

Worker. — Holotype  (Paratypes)  mea- 
surements: HL  1.04  (1.02-1.08);  ML  0.40 
(0.26-0.36);  HW  0.88  (0.82-0.94);  SL  0.86 
(0.86-0.92);  ED  0.12  (0.10-0.14);  WL  1.36 
(1.36-1.60)  mm;  CI  0.85  (0.83-0.90);  SI  0.98 
(0.93-1.10);  OI  0.14  (0.11-0.17)  n  =  7. 

Typical  striatida  group  member:  Rough- 
ly costulate  sculpture.  Vertexal  margin 
concave  in  frontal  view;  clypeal  costulae 
extend  slightly  onto  anterior  lamella;  man- 


dibles triangular  and  mostly  striate  except 
for  smooth  and  shining  lamellate  basal 
corner  and  margin  of  chewing  border; 
compound  eyes  relatively  small;  in  lateral 
view  dorsal  propodeal  margin  not  notably 
depressed  below  rest  of  mesosomal  dor- 
sum; propodeal  spiracle  slightly  elevated 
above  rest  of  sculpture  and  at  edge  of  de- 
clivitous surface;  petiolar  node  posteriorly 
inclined;  subpetiolar  process  subquadrate, 
typical  of  strigata  subgroup.  Longitudinal 
costulae  present  on  declivitous  postpetio- 
lar  face.  Weak  transverse  costulae  on  post- 
petiolar  ventrum. 

Fairly  abundant  standing  hairs  on  body, 
but  very  sparse  appressed  pubescence. 
Legs  smooth  and  shining,  except  for  nu- 
merous piligerous  punctures.  Metacoxal 
dorsum  with  a  well-developed  denticle. 
Body  dark  brown  to  black  with  yellowish 
legs. 

Male. — Longitudinal  costulae  on  cephal- 
ic dorsum;  mandibles  triangular  and  cos- 
tulate; pronotum  smooth  and  shining  with 
piligerous  punctures;  pronotum  with  pili- 
gerous punctures;  mesonotum  mostly 
smooth  and  shining  except  for  narrow  an- 
terior band  of  transverse  striae;  declivitous 
propodeal  face  with  longitudinal  rugulae 
that  do  not  reach  anterior  dorsal  face;  gas- 
ter  smooth  and  shining. 

Queen. — Unknown. 

Discussion. -This  species  is  near  strigata 
and  though  the  differences  are  not  striking, 
they  seem  consistent  enough  to  define  a 


Volume  4,  1995 


157 


35 


Figs.  33-38.     Frontal  view  of  head.  33,  G.  fernandezi;  34,  G.  sulcata;  35,  G.  haenschi;  36,  G.  schmitti;  37,  G. 
bispinosa;  38,  G.  triangularis. 


species.  G.  strigata  is  smaller  (HW  0.74- 
0.80);  WL  1.20-1.34  mm),  with  relatively 
larger  eyes  (OI  0.17-0.20)  and  shorter 
scapes  (0.70-0.78  mm)  and  dorsal  propo- 
deal  face  in  lateral  view  is  usually  de- 
pressed below  the  rest  of  the  mesosomal 
dorsum.  It  is  probably  sympatric  with  nn- 
dina  in  part  of  its  range,  having  been  found 
only  51  km  away  from  the  andina  type  lo- 
cality. The  male  of  strigata  has  a  median 
area  of  rugosity  on  the  mesonotum  and  to- 
tally rugulose  propodeum.  Individuals  of 
the  Ecuadorean  sample  are  slightly  larger 
than  the  Colombian  specimens. 

Etymology. — The  name  alludes  to  the 
Andes,  a  portion  of  which  is  inhabited  by 
this  species. 

Gnamptogenys  annulata  Mayr 

Ectntomma  {Gnamptogenys)  rimulosa  var.  annulata 
May  1887:543. 


Ectatomma   {Gnamptogenys)   annulatutn   Mayr; 

Mann  1916:408;  Luederwalt  1926:23. 
Gnamptogenys  annulatum  Mayr;  Mann  1922:3; 

Brown    1958:300;    Kempf    1960:390;    Kempf 

1961:491;  Kempf  1969:275;  Kempf  1972:112; 

Kempf  1976:51;  Lattke  1990:11. 

Diagnosis. — Finely  striate  species;  longi- 
tudinal striae  on  head,  promesonotum 
and  gastric  segments  2  and  3;  petiolar  dor- 
sum with  transversely  arched  striae;  man- 
dibles subtriangular;  metacoxal  spine  ab- 
sent; body  reddish  brown;  coxae  ferrugi- 
nous yellow;  tibiae  and  femora  yellow  to 
testaceous  but  apically  ferruginous  yellow 
to  ferruginous. 

Ecology. — J.  Longino  (pers.  comm.)  re- 
ports from  a  nest  midden  chitinous  frag- 
ments of  Coleoptera,  aradid  bug  nymphs, 
an  unidentified  male  ant,  and  a  Cauipotio- 
tus  worker.  An  apparently  recently  killed 


158 


Journal  of  Hymenoptera  Research 


42 


Figs.  39-42.     Frontal  view  of  head,  and  lateral  view  of  mesosoma  and  petiole.  39-40,  G.  andina;  41-41,  G. 
brunnea. 


Pheidole  (cephalica?)  queen  was  also  found 
in  a  nest  chamber. 

Comments. — The  gauge  of  costulation 
can  vary  from  striate  to  finely  costulate, 
but  coloration  is  fairly  stable. 

SPECIMENS  EXAMINED:  BOLIVIA, 
Caranavi:  vie.  Radio,  800  m.  BRAZIL, 
Amazonas:  80  km  NNE  Manaus,  2°25'S 
59°46'W,  80  m.  COLOMBIA,  Valle:  Bajo 
Calima;  Cauca:  Isla  La  Gorgona.  COSTA 
RICA,  vie.  Guapiles,  Rio  Toro  Amarillo; 
Prto.  Limon;  Limon:  Tortuguero,  <  100m, 
10°35'N  83°31'W;  Heredia:  10°20'N 
84°84'W,  500  m;  3  km  S  Puerto  Viejo, 
10°26'N  84°00'N,  50  m;  Pq.  Nac.  Braulio 
Carrillo,  Carrillo,  500  m;  Peninsula  de  Osa, 
Sirena,  8°28'N  83°35'W;  Alajeula:  Rio  Peii- 
as  Blancas,  10°19'N  84°43'W,  800  m.  EC- 
UADOR, Pichincha:  6  km  W  Sto.  Domingo 


de  los  Colorados,  953  m;  Maquipucuna,  5 
km  ESE  Nanegal,  0°70'N  78°38'W,  1250m; 
Sucua.  GUIANA,  Kartabo.  PERU,  Pangu- 
ana:  9°37'S  74°56'W,  220  m;  Tingo  Maria: 
Valle  de  Monson;  Madre  de  Dios:  Res.  Rio 
Tambopata,  30  km  SW  Prto.  Maldonado, 
290  m;  Loreto:  Boqueron,  500m;  San  Mar- 
tin: Davidcillo,  30  km  NNE  Tarapoto, 
6°15'S  76°15'W.  VENEZUELA,  Tachira: 
Uribante-Caparo,  Las  Cuevas,  7°47'N 
71°46'W,  500  m;  Barinas:  Ticoporo,  8°04'N 
70°48'W,  250  m;  Amazonas:  Alto  Mavaca, 
2°01'N  65°07'W,  200  m;  Alto  Siapa,  1°42'N 
64°38'W,  500  m. 

Gnamptogenys  banks!  (Wheeler) 

Emeryella  banksi  Wheeler  1930:10. 
Gnamptogenys   banksi   (Wheeler);   Brown   1958: 
227,233;  Kempf  1972:112. 


Volume  4,  1995  159 

Diagnosis. — Finely  costulate;  head  broad  from  the  first  sighting  to  the  prey  was  6 

with  longitudinally  striated  scapes;  meso-  m.  I  followed  a  returning  worker  as  far  as 

metanotal  suture  not  deeply  impressed,  I  could  into  a  tree  fall  tangle,  10  m  from 

but  distinct.  It  lacks  the  long,  fine  mandib-  prey."    The    second    attack    involved    46 

ular  sulcus  present  in  laticepJmla.  workers  tackling  a  6  cm  long  millipede 

Ecology. — Found  in  rotten  logs  in  low-  within  a  rc^tten  log.  Once  the  millipede 

land  rain  forest,  this  species  is  a  specialist  was  subdued  they  carried  it  back  to  the 

millipede  predator.  nest. 

Comments. — It    is    rarely    collected.    A  Two  millipedes  taken  from  the  nest  by 

specimen  from  Barro  Colorado  has  two  Longino  were  determined  by  R.L.  Hoff- 

deep  fossae  on  the  posterolateral  margin  man   as   Trichomorpha   sp.    (Polydesmida: 

of  the  mesonotum.  The  type  was  exam-  Chelodesmidae),  and   Epinannolene  sp. 

ined.  (Spirostreptida:  Epinannolenidae). 

Specimens         Examined. — ECUADOR,  Comments. — This  unique  ant  was  previ- 

Guayas:  3  km  S  Bucay.  PANAMA,  Isla  ously  only  known  from  a  restricted  area 

Barro  Colorado  (MCZC).  in  Central  America.  Along  with  perspicax, 

it  represents  a  highly  specialized  form  in 

Gnamptogenys  bispinosa  (Emery)  ^^^  lineage  of  millipede  hunters.  The  an- 

Ectatomma   {Poneracantha)   bispinosum   Emery  terior  clypeal  setae  are  more  developed 

1897:547.  than  in  any  other  of  the  rastrata  group  spe- 

Ectatomma  (Holcoponera)  bispinosum  Emery;  Em-  cies  and  an  interesting  autapomorphy  is  a 

ery  1890:40.  brief  anteromedian  longitudinal  sulcus  on 

Holcoponera  bispinosa   (Emery);  Wheeler   1952:  \\^q  clypeus. 

132-133.  Specimens  Examined. — COLOMBIA,  Val- 

Gnamptogenys  bispinosa  (Emery);  Brown  1958:  ^^.    ^QSTA    RICA-     Heredia-     10°20'N 

?g79  i^r^^  ''''^   ^'°'^''   ^^^^''^^^''   ^'""^^  84°04'W,    500m;    Peninsula    de   Osa,    Pq. 

Nac.  Corcovado,  Sirena,  <  100m,  8°28'N 

Diagnosis. — Eyes    subglobulose,    scapes  83°35'W;  Puntarenas:  2  km  S  San  Vito,  Fin- 
long  and  surpassing  vertexal  margin;  ca  Las  Cruces,  1219  m;  Rio  Reventazon,  4 
mandibles  edentate.  Promesonotal  suture  km  E  Turrialba  (MCZC). 
breaks  sculpture  dorsally  but  not  laterally; 

mesometanotal   suture   deep   and   broad.  Gnamptogenys  bisulca  Kempf  and 

Head,  mesosoma  and  postpetiole  rugose;  Brown 

„   ^           ■      .                .                     .        .  Gnamptogenys  bisulca  Kempf  and  Brown  1968: 

£co/o^/.-This   wet   forest   species   is   a  92-3;  ^  Wheeler    and    Wheeler    1971:1202; 

millipede  specialist.  J.   Longino  (pers.  Kempf  1972112 
comm.)  observed  the  following  two  raids 

from  the  same  colony,  both  at  11:00  AM:  Diagnosis. — Longitudinal  costulae  on 

"  A  column  of  20  workers  was  moving  propodeal  declivity,  metacoxal  spines 

along  a  liana  on  the  forest  floor.  They  left  hooked  and  triangular,  not  acicular;  well 

the  liana  and  moved  very  slowly  across  defined  transverse  costulae  on  postpetio- 

the   leaf   litter,    frequently  bunching   up.  lar  sternum;  propodeal  spiracle  as  low  as 

They  all  went  under  a  leaf  and  then  agi-  surrounding  sculpture,  not  raised, 

tated  ants  could  be  seen  coming  out  from  Ecology. — Occurs  in  mesic  forested  hab- 

under  the  leaf,  2  or  3  ants  at  a  time,  clean-  itats.  The  localities  suggest  preference  for 

ing  their  antennae  and  mandibles.  I  re-  relatively  cool  sites  as  the  lowest  altitudes 

moved  the  leaf  to  find  the  ants  attacking  are  found  at  higher  latitudes  (10°N)  and 

a  millipede.  Some  were  stinging  and  some  as  latitude  approaches  the  equator  the  lo- 

were  pulling  on  the  legs.   The  distance  calities  ascend  in  altitude. 


160 


Journal  of  Hymenoptera  Research 


Comments. — Within  the  strigata  complex 
this  species  is  unique  due  to  the  develop- 
ment of  its  metanotal  groove,  and  within 
the  striatula  group  two  other  species  also 
show  this  character  state.  Recent  collec- 
tions show  a  wider  range  than  previously 
known. 

Specimens  Examined. — COLOMBIA, 
Choco:  10  km  SW  San  Jose  del  Palmar,  Rio 
Torito,  Finca  Los  Guaduales,  800  m,  1/4- 
VI-78;  Valle:  vie.  Lago  Calima,  1600  m;  vie. 
Saladito,  1900-2100  m.  COSTA  RICA,  He- 
redia:  17  km  S.  Puerto  Viejo,  10°18'N 
84°02'W,  550  m;  17  km  N  Volcan  Barba, 
10°17'N  84°05'W,  880-1020  m.  ECUA- 
DOR, Pichincha:  25  km  ENE  Alluriquin, 
via  Chiribaga,  1400-1800  m;  Tinalandia, 
16  km  SE  Sto.  Domingo  de  los  Colorados, 
9-VI-76;  3  km  E  Tandapi,  1300  m. 

Gnamptogenys  boliviensis  Lattke, 
new  species 
(Figs.  43,  44) 

Type  Material. — Holotype  worker.  BO- 
LIVIA, Tumupasa,  W.M.  Mann,  leg.  De- 
posited in  the  USNM.  Paratypes:  Seven 
workers  deposited  in  the  USNM;  One 
worker  and  one  male  in  each  of  the  fol- 
lowing: BMNH,  MIZA,  LACM,  MCZC. 
One  worker  in  MUSP.  All  from  same  lo- 
cality and  probably  from  the  same  nest. 
One  specimen  bears  an  additional  label: 
"Mulford  Biological  Expedition,  1921- 
1922."  Consultation  of  maps  locates  Tu- 
mupasa at  14°09'S  67°55'W  in  lowland  for- 
ests of  the  upper  Rio  Beni  watershed. 

Worker. — Holotype  (Paratypes)  mea- 
surements: HL  0.88  (0.83-0.88);  ML  0.36 
(0.39-0.41);  HW  0.70  (0.69-0.70);  SL  0.49 
(0.48-0.50);  ED  0.10  (0.10-0.12);  WL  1.23 
(1.06-1.15)  mm;  CI  0.70  (0.80-0.84);  SI  0.70 
(0.69-0.71);  OI  0.15  (0.15-0.18).  n=4. 

Head  in  frontal  view  elongate,  lateral 
margins  fairly  parallel,  vertexal  margin 
concave;  anterior  lamella  of  clypeus  later- 
ally rounded  and  medianly  convex;  eyes 
situated  at  mid-length;  antennae  smooth 
and  shining,  dorsoventrally  compressed 
widest    apicad;    mandibles    smooth    and 


shining  dorsally,  with  sparse  punctae,  su- 
bfalcate,  laterobasally  with  small  area  of 
longitudinal  rugae,  including  small  sulcus 
never  longer  tha  one-fourth  of  the  man- 
dible; gula  with  longitudinal  costulae, 
front  of  head  with  longitudinal  striae. 

Occasional  transverse  costulae  on  pron- 
otal  collar;  mesosomal  dorsum  with  lon- 
gitudinal costulae,  slightly  thicker  on 
metanotum  and  propdeum  than  on  pro- 
mesonotum;  pronotal  suture  softly  im- 
pressed, visible  only  in  certain  angles  of 
view;  metanotal  suture  deeply  impressed 
but  doesn't  break  longitudinal  sculpture; 
declivitous  propodeal  face  with  oblique  to 
transverse  costulae  and  two  superolateral 
ridges;  pronotum  laterally  with  transverse 
costulae,  smooth  and  shining  along  ven- 
tral margin;  anepisternum  elongate  and 
obliquely  costulate;  katepisternum  and 
metepisternum  with  transverse  costulae; 
costulae  on  metepisternum  have  same  di- 
rection as  on  katepisternum,  but  costulae 
on  lateral  propodeal  face  are  more  hori- 
zontal; propodeal  spiracle  round  and  not 
prominently  higher  than  surrounding 
sculpture;  petiole  slighty  pedunculate, 
dorsally  with  longitudinal  costulae,  ante- 
rior face  with  transverse  costula  or  rugae, 
sometimes  partially  effaced,  laterally  with 
longitudinal  costulae  with  slight  oblique 
tendency;  node  in  lateral  view  with  softly 
convex  dorsal  margin,  anteriorly  concave; 
anterior  and  posterior  faces  fairly  sharply 
separated  from  dorsal  face;  posterior  face 
with  4-5  convex  costulae;  subpetiolar  pro- 
cess varies  from  subquadrate  anteriorly 
and  posteriorly  concave. 

Anterior  postpetiolar  face  smooth  and 
shining;  gastric  terga  I  and  II  with  longi- 
tudinal striae,  sternum  I  laterally  costula- 
te, but  discal  area  smooth  and  shining  as 
is  rest  of  gaster;  procoxa  anteriorly  and 
anterolaterally  smooth  and  shining,  pos- 
terolaterally  costulate;  mesocoxae  with 
transverse  costulae  that  tend  to  be  effaced; 
dorsum  of  metacoxa  with  transverse  cos- 
tulae and  basal,  low  triangular  lobe;  tibia 
and  femora  smooth  and  shining.  Body  fer- 


Volume  4,  1995 


161 


Figs.  43-44.     Micrographs  of  G.  boliviciiisis.  Scale  bars  =  0.5  mm.  43,  head;  44,  thorax  and  gaster. 


162 


Journal  of  Hymenoptera  Research 


ruginous  brown;  legs  and  antennae  testa- 
ceous. 

Queen. — Unknown. 

Male. — Mandibles  triangular,  apical 
edge  serrate,  dorsally  smooth  and  shining; 
cephalic  dorsum  with  longitudinal  striae 
that  tend  to  be  effaced  before  reaching 
mid-ocellus,  faint  transverse  rugae  above 
the  posterior  ocelli  but  rest  of  head 
smooth  and  shining;  propodeum  rugose; 
petiole  with  lateral  rugulae,  dorsum  shin- 
ing, with  slight  roughened  aspect  and  me- 
dian longitudinal  costulae;  gastric  sterna 
and  terga  smooth  and  shining;  vestigial 
arolea  present. 

Discussion. — Several  specimens  were 
found  with  determination  labels,  probably 
written  by  Mann,  indicating  that  he  rec- 
ognized this  species  as  new.  G.  boliviensis 
is  near  to  G.  continua  and  the  possibility 
exists  that  the  collection  and  study  of 
more  material  may  show  them  to  be  con- 
specific.  In  G.  continua  the  sculpture  is 
coarser,  costate-costulate,  and  the  mandi- 
bles are  more  robust,  without  a  concave 
inner  edge. 

Etymology. — The  name  is  derived  from 
the  name  of  the  type  locality  country,  Bo- 
livia. 

Gnamptogenys  bruchi  (Santschi) 

Ectatomma  (Parectatomma)  bruchi  Santschi  1922: 

241. 
Gnamptogenys   bruchi   (Santschi);   Brown   1958: 

227;  Kempf  and  Brown  1968:92;  Kempf  1969: 

275;  Kempf  1972:112. 

Diagnosis. — Striate  body  sculpture  with 
weak  transverse  impressions;  metacoxal 
process  is  shaped  as  a  low,  triangular  lobe; 
clypeal  lamella  broadly  convex  anterad. 

Ecology. — Apparently  a  predator  of  Tra- 
chymyrmex  and  other  leaf  cutting  ants 
(Kempf  and  Brown,  1968:92). 

Comments. — Kempf  (1969)  notes  bruchi 
as  close  to  hartmani  and  that  a  synonymy 
could  be  possible.  He  also  mentions  the 
variability  of  the  extent  of  impression  of 
the  metanotal  groove  and  discards  it  as  an 
indicator  of  species  separation.  This  could 


very  well  be  only  a  variant  of  hartmanni 
but  due  to  differences  of  sculpture  on  the 
posterior  petiole  face,  postpetiolar  sternal 
process  and  clypeal  lamella,  I  have  chosen 
to  conserve  this  name  until  more  material 
can  be  studied.  The  sculpture  on  the  dor- 
sum of  the  petiole  can  vary  from  longitu- 
dinal to  whorled. 

Specimens  Examined. — ARGENTINA, 
Cordoba:  Alta  Gracia  (cotype:MCZC). 
BRAZIL,  Sao  Paulo:  Fazenda  B.  Rico. 

Gnamptogenys  brunnea  Lattke, 
new  species 
(Figs.  41,42) 

Type  Material. — Holotype  worker.  CO- 
LOMBIA, Choco,  10  km  SW  San  Jose  del 
Palmar,  Finca  Los  Guaduales,  760  m,  2-VI- 
1978,  C.  Kugler,  leg.  Deposited  in  MIZA. 
Para  types:  One  queen  and  18  nidotype 
workers.  Queen  and  6  workers  deposited 
in  MIZA.  Two  workers  deposited  in  each 
of  the  following:  MCZC,  LACM,  BMNH, 
MUSP,  USNM. 

Worker. — Holotype  (Paratypes)  mea- 
surements: HL  0.90  (0.88-0.92);  ML  0.30 
(0.30-0.32);  HW  0.74  (0.68-0.80);  SL  0.74 
(0.80-0.84);  WL  1.20  (1.10-1.28)  mm;  CI 
0.82  (0.77-0.87);  SI  1.00  (0.95-1.09);  OI  0.18 
(0.16-0.18)  n=5. 

Cephalic  dorsum  longitudinally  costu- 
late;  in  frontal  view  vertexal  margin  con- 
cave and  eyes  slightly  behind  mid-length; 
mandibles  triangular,  with  dorsal  costu- 
lae, basal  and  apical  margins  with  narrow 
smooth  and  shining  strip  that  is  sharply 
separated  and  lower  than  the  rest  of  man- 
dibular surface;  anterior  pronotal  margin 
with  6-8  transverse  costulae  which  arch 
around  and  become  longitudinal  on  dor- 
sum; mesometanotal  suture  well-im- 
pressed; mesonotum  narrow  and  trans- 
verse; in  lateral  view  dorsal  propodeal 
face  slightly  depressed  below  rest  of  me- 
sosomal  dorsal  margin;  declivitous  pro- 
podeal face  distinctly  separated  from  rest 
of  mesosoma  by  low  lateral  ridges;  mostly 
transverse  costulae  with  longitudinal  cos- 
tulae from  dorsum  extending  only  to  an- 


Volume  4,  1995  163 

terior  one-fourth  of  declivitous  face;  peti-         Diagnosis. — A  large  finely  striate  species 

olar  node  in  lateral  view  transversely  cos-  with  transverse  striae  on  propodeum,  de- 

tulate  and  posteriorly  inclined:  a  broad  clivity    medianly    smooth;    promesonotal 

anterior  convexity  with   sharp   posterior  suture  weakly  impressed  or  absent;  me- 

drop  and  concave  posterior  margin;  sub-  sometanotal  suture  deep  and  wide;  meta- 

petiolar  process  shaped  as  an  anteriorly  coxal  tooth  absent,  low  crest  or  tubercle  at 

projecting  lobe;  anterior  postpetiolar  face  most. 

and  sternum  with  transverse  costulae.  Ion-  Ecology. — Consistently  collected  and  ob- 

gitudinal  on  rest;  costulae  on  the  gastric  served  on  trees,  this  species  is  an  arboreal 

tergum  II  finer  than  on  preceding  seg-  nester.    Longino   found    a    nest   inside   a 

ment;  metacoxal  spine  present.  large,   deep   knothole   and    records   prey 

Body   with   sparse   decumbent   pubes-  items  of  the  following:  Heteroptera:  Pen- 

cence  and  erect  to  suberect  hairs;  pubes-  tatomidae,  Aradidae;  Coleoptera:  Ceram- 

cence  denser  on  legs;  scapes  with  abun-  bycidae,  Histeridae,  Platypodidae,  Chrys- 

dant  decumbent  pubescence,  no  erect  or  omelidae,  Tenebrionidae,  and  Passalidae. 

suberect  hairs.  Body  reddish-brown  and  One  observed  foraging  group  was  made 

legs  brownish-yellow.  up  of  9  workers  and  a  dealate  queen.  Ob- 

Queen. —HL  0.98;  HW  0.86;  ML  0.32;  SL  servations  of  group  foraging  of  up  to  40 

0.84;  ED  0.18;  WL  1.44  mm;  CI  0.88;  SI  single-file    workers,    plus    the    following 

0.97;  Ol  0.21.  Like  workers  with  exception  Longino  observation,  indicate  trail  and /or 

of  usual  caste  differences.  recruiting  pheromone  communication  in 

Male. — Unknown.  this  species.  "The  foragers  walked  with 

Discussion. — This  species  is  close  to  ni-  their  gasters  curled  and  touching  the  sub- 

grivitrea  on  account  of  size,  posteriorly  tilt-  strate.  When  they  were  together  near  the 

ed  petiolar  node,  sleek  appearance,  sub-  prey  I  could  often  make  out  a  tiny  white 

petiolar  process  shape,  sparse  pilosity  and  area  (gland?)  exposed  at  the  very  tip  of  the 

lack  of  any  trace  of  anterior  petiolar  pe-  gaster,  which  other  workers  would  inves- 

duncle.  But  nigrivitrea  is  different  in  its  pi-  tigate". 

ceous  body  coloration  with  dark  brown         Comments. — Color  can  vary  from  ferru- 

legs,   lack  of  metanotal  groove,   slightly  ginous  to  light-brown,  and  the  striae  on 

more  elevated  propodeal  spiracles,  longer  the  posterior  nodal  face  may  be  effaced, 
pilosity,  and  a  sleeker  petiolar  node  with         Specimens  Examined. — BRAZIL,  Amazo- 

a  more  prominent  apical  point.  nas:  Ilha  de  Curari,  (varzea);  Amapa:  Villa 

Etymolog}/. — The  name  comes  from  the  Amazonas.    COLOMBIA,    Meta:    Reserva 

Latin  term  for  brown,  brunneus.  La  Macarena,  Cafio  La  Curia,  580  m.  COS- 
TA RICA,  Peninsula  de  Osa:  Pq.  Nac.  Cor- 

Gnamptogenys  concinna  (Smith)  covado,  Llorona;  Heredia:  Finca  La  Selva. 

PANAMA,   Isla   Barro   Colorado.    PERU, 

Ectatommn  concinnna  F.  Smith  1858:103.  Madre  de  Dios:  15  km  E  Prto.  Maldonado, 

Ectakvnma{Gnamptogenys)concinmim{¥.Smm;  200    m;    Amazonas,    Panguana,    9°37'S 

Mann  1916:406;  Wheeler  1922:2.  ^a^^c^'wi     nnn   ^     x-  \a     ^      \7  n      a 

r.  ,^  74  56  W,  220  m;  Tmgo  Maria:  Valle  de 

tctatomma  (Gnamptogenys)  concinnum  var.   ro-  ^.        .      -roiMTMr-v  a  r^    a    •        \i  w        i  r-^ 

■  AAfu    1     in-^oo  D  ir^ron-^r,  Monson.  TRININDAD,  Arima  Valley,  152 

mnn?  Wheeler  1923:2;  Brown  1958:227.  a/cxtc^ttct  a       a  ai/    r,- 

Lcia\omma{Gnampto^em,s)concmnumv,.r.comca  "^-    VENEZUELA,    Amazonas:    Alto    Rio 

Borgme.er  1928:196;  Brown  1958:227.  Mavaca,  2  02'N  65°06'W,  200m. 

Ectatommn  {GnarnptoQenys)  concimmni  var.  senii-  ^  ^  .•  »,r 

,        D  .      ir.or.ir^r    r,  irvr-r,  Gnamptogenys  contiHua  Mavr 

circularis  Borgmeier  1929:195;  Brown  1958:  r     &      j  / 

227.  Ectatommn    [Gnnmptogenys)    conti)U(uiii    Mayr 

Grwmptogcnys   co}tci)uia   (Smith);    Brown    1958:  1887:544. 

227;  Kempf  1972:112;  Lattke  1990:11.  Ectatommn    {Giinmptogcin/^)    cxaratum    Emery 


164 


Journal  of  Hymenoptera  Research 


1901:50;   Brown   1958:303;   Kempf   1972:112; 

Lattke  1990:13.  NEW  SYNONYMY. 
Ectaomma  {Gnamptogenys)  continuum  var.  pana- 

mensis  Santschi   1931:265;   Brown   1958:228, 

304. 
Gnamptogenys  continua  (Mayr);  Brown  1958:228, 

304;  Kempf  1972:112;  Lattke  1990:12. 

Diagnosis. — Mandibles  with  linear  ten- 
dency; mesometanotal  suture  impressed; 
declivituous  propodeal  face  with  small  su- 
perolateral lobes;  upper  half  with  longi- 
tudinal costulae  and  inferior  half  with 
transverse  costulae;  sternal  disc  of  post- 
petiole  smooth;  anterior  nodal  face  rugose 
with  some  inferior  transverse  costulae. 

Ecology. — Collected  in  leaflitter  samples 
from  humid  forsets. 

Comments. — The  synonymy  of  exarata 
was  inevitable  as  more  material  accumu- 
lated and  it  became  impossible  to  separate 
the  two  forms.  The  gap  between  OD  and 
apical  scape  width  closed.  There  are 
changes  in  the  gauge  of  the  costulation, 
degree  of  anterior  clypeal  convexity,  ef- 
facement  of  the  sculpture,  shape  of  the 
subpetiolar  process,  extent  of  the  petiolar 
peduncle,  and  size  (HW  0.56-0.96;  HL 
0.64-1.16;  WL  0.88-1.62).  The  occiput  has 
a  median  dent  that  is  most  noticeable 
when  the  occiput  is  smooth  and  shining 
as  opposed  to  costulate  forms.  Deposited 
in  the  MCZC  is  a  specimen  from  Vera 
Cruz  that  was  compared  with  the  type  by 
Brown  in  1954. 

Specimens  Examined. — BELIZE,  Caves 
Branch,  4  km  S  Belmopan.  BRAZIL,  Rio  de 
Janeiro:  Itatiara;  Sao  Paulo:  Fazenda  Con- 
quista,  Rio  Preto;  Guquitiba,  BRE  km  76. 
HONDURAS,  La  Lima.  ECUADORA,  Pi- 
chicincha:  Tinalandia,  16  km  SE  Sto.  Do- 
mingo del  los  Colorados,  680  m.  COLOM- 
BIA, Meta:  Quebrada  Susumuko,  23  km 
NW  Villavicencio,  1000  m;  Magdalena: 
Parque  Tayrona,  Pueblito,  360  m.  MEXI- 
CO, Vera  Cruz,  Tuxtla,  Las  Hamacas,  17 
km  N  Santiago.  PERU,  Panguana,  9°37'S 
74°56'W,  220  m;  Huanuco:  16  km  SW  Las 
Palmas,  1000  m;  Pasco:  Pozuzo,  1000,  SU- 
RINAM,  Kartabo,    Bartica   Dam.    VENE- 


ZUELA, Barinas:  17  km  SSW  Ciudad  Bo- 
livia, 8°02'N  70°46'W,  240  m;  Guarico:  24 
km  N  Altagracia  de  Orituco,  300  m. 

Gnamptogenys  cuneiforma  Lattke, 
new  species 

Type  Material. — Holotype  worker:  PAN- 
AMA, Bocas  del  Toro,  8°47'N  82°11'W, 
500m;  16/18-VII-87,  D.M.  Olson,  leg.  De- 
posited in  MCZC.  Paratypes:  One  worker 
deposited  in  MCZC  and  two  workers  de- 
posited in  MIZA,  all  with  same  locality 
data  as  the  holotype. 

Worker. — Holotype  (Paratype)  measure- 
ments: HL  0.90  (0.84-0.88);  ML  0.62  (0.60- 
0.64);  HW  0.84  (0.82-0.84);  SL  0.82  (0.80- 
0.82);  ED  0.20  (0.21-0.22);  WL  1.40  (1.38- 
1.40)  mm;  CI  1.07  (1.02-1.05);  SI  0.98  (0.98); 
OI  0.24  (0.26-0.28),  n=4. 

Head  subquadrate  in  frontal  view,  sides 
very  broadly  convex,  slightly  wider  ante- 
riorly than  posteriorly,  posterior  margin 
broadly  concave,  longitudinally  costulate. 
Anterior  clypeal  margin  with  a  small  la- 
mella, medianly  straight  and  laterally 
rounded.  A  brief  longitudinal  carinae, 
thicker  than  surrounding  sculpture,  pres- 
ent between  frontal  lobes.  Mandibles  tri- 
angular and  elongate,  masticatory  border 
concave,  dorsal  surface  of  blade  convex 
and  with  longitudinal  rugulae.  Scapes 
shining  and  with  low  rugosities.  Scapes 
without  pilosity,  only  decumbent  hairs. 

Anterior  pronotal  face  with  4-5  trans- 
verse costulae,  sometimes  quite  rough  and 
resembling  rugae,  longitudinal  costulae 
on  the  dorsum  posterad  including  the 
mesonotum.  Metanotum  and  propodeum 
with  transverse  costae. 

Promesonotal  suture  well  to  poorly  im- 
pressed, metanotal  groove  well  im- 
pressed. In  lateral  view  the  mesosoma  has 
a  very  broadly  convex  dorsal  margin,  pro- 
podeal denticles  brief  and  triangular,  de- 
clivity concave.  Sides  of  pronotum  with 
regular  longitudinal,  slightly  oblique  cos- 
tae. Anepisternum  fulcrum  shaped.  Kat- 
episternum  with  a  well-developed  flange. 
Mesopleura,  propodeum  and  dorsal  two- 


Volume  4,  1995 


165 


thirds  of  metapleura  with  transverse/ 
oblique  costae;  metapleura  ventrad  of  spi- 
racle with  longitudinal  costae. 

Petiole  in  lateral  view  with  a  low  and 
broadly  convex  dorsal  margin,  ventral 
process  projects  anterad.  Dorsally  with 
transverse  costae,  posterior  margin  ru- 
gose. Gastric  dorsum  with  longitudinal 
costulation;  ventrum  of  postpetiole  with 
transverse  rugae  along  posterior  half;  ir- 
regular rugulae  anterad  to  the  process. 

Base  of  foretarsus  with  a  single  stout  se- 
tae opposite  the  strigil.  Forecoxae  with 
transverse  costulae  on  lateral  face,  as  well 
as  dorsal  faces  of  meso-  and  metacoxae. 
Legs  shining  and  with  abundant  shallow 
punctulae.  Very  sparse  pilosity  on  femora 
and  tibiae.  Color  ferruginous  to  dark 
brown;  mandibles,  antennae  and  legs  fer- 
ruginous. 

Queen,  Male. — Unknown. 

Discussion. — This  species  may  be  con- 
fused with  two  other  small  rastrata  group 
species:  mstrata  and  mecoti/le.  It  is  separa- 
ble from  the  former  on  account  of  the  lon- 
ger scapes  (surpassing  the  vertexal  mar- 
gin), transverse  costulae  on  the  propo- 
deum  as  opposed  to  longitudinal,  and  a 
well-defined  metapleura,  separated  from 
the  propodeum  laterally  by  an  impressed 
line,  lacking  in  rnstratn.  The  costulae  of  ine- 
cotyle  have  a  coarse  aspect  in  contrast  with 
the  smooth  regularity  in  cuneiforma,  tneco- 
tyle  also  has  the  striae  on  the  scapes  and 
dorsal  mandibuar  surface  better  defined 
and  the  masticatory  border  lacks  the  con- 
cavity of  cuneiforma. 

Etymology. — The  name  alludes  to  the 
shape  of  the  anepisternum. 

Gnamptogenys  curvoclypeata  Lattke 
Gnamptogenys  curvoclypeata  Lattke  1990:  12. 

Diagnosis. — Piceous  species  with  ferru- 
ginous antenna,  legs  and  coxae;  prome- 
sonotal  suture  very  weakly  impressed, 
visible  only  with  certain  angles  of  light; 
meso-  and  metacoxae  smooth  and  shining 
on  median  and  basal  lateral  faces;  meta- 


coxae with  weakly  developed  triangular 
dorsal  lobe. 

Ecology.— See  Lattke  (1990). 

Comments. — See  Lattke  (1990). 

Specimens  Examined. — COLOMBIA, 
Amazonas:  7  km  N  Leticia  (MCZC). 

Gnamptogenys  ejuncida,  Lattke, 
new  species 
(Figs.  48,  50) 

Type  Material. — Holotype  worker.  CO- 
LOMBIA, Putumayo,  Mocoa,  610  m,  4-1- 
77,  C.  Kugler,  leg.  Deposited  in  MIZA. 

Worker. — Holotype  measurements:  HL 
1.10;  ML  0.26;  HW  0.88;  SL  1.02;  ED  0.18; 
WL  1.54  mm,  CI  0.80;  SI  1.56;  OI  0.21. 

With  head  in  frontal  view:  vertexal  mar- 
gins slightly  convex,  lateral  margins 
broadly  convex  and  anterior  clypeal  mar- 
gin convex;  mandibles  triangular  and  with 
rugose  dorsum;  anterior  pronotal  face 
with  six  transverse  costulae,  dorsum  with 
longitudinal  costulae;  mesonotum  with 
anteriorly  convex  costulae,  transverse  at 
metanotum  and  posteriorly  convex  on 
dorsal  propodeal  face;  anepisternum  with 
three  transverse  costulae  that  descend 
from  mesonotum;  rest  of  mesosomal  side 
obliquely  costulate,  some  curve  around 
propodeal  spiracles;  declivitous  propodeal 
face  with  longitudinal  costulae;  in  lateral 
view  dorsal  mesosomal  outline  evenly 
convex,  with  no  abrupt  breaks  or  depres- 
sions; petiolar  node  posteriorly  inclined: 
in  lateral  view  transversely  costulate  with 
convex  anterior  margin,  concave  posterior 
margin  and  overhanging  apex;  subpetiolar 
process  triangular  and  projecting  anterad. 
Anterior  face  to  dorsal  two-thirds  of  post- 
petiolar  dorsum  with  transverse  costulae; 
rest  of  gastric  sculpture  longitudinal;  post- 
petiolar  sternum  anteriorly  transversely 
costulate,  posteriorly  divergent;  transverse 
costulae  on  procoxae  and  anterior  faces  of 
meso-  and  metacoxae;  metacoxal  tooth 
well  developed.  Abundant  suberect  hairs 
and  pubescence  on  body  and  extremities. 

Queen,  Male. — Unknown. 

Ecology. — Unknown. 


166 


Journal  of  Hymenoptera  Research 


Figs.  45-53.     Lateral  view  of  posterior  leg,  mesosoma  and  petiole,  and  frontal  view  of  head.  45-47,  G.  extra; 
48-50,  G.  ejuncida;  51-53,  G.  pilosa. 


Discussion. — On  account  of  size,  sculp- 
ture, pilosity  and  posteriorly  inclined  pet- 
iole node  this  species  seems  to  be  most 
closely  related  to  pilosa,  a  very  similar  spe- 
cies found  in  an  Andean  canyon  NW  of 
Cali,  Colombia.  The  sculpture  of  pilosa  is 
much  finer  and  its  pilosity  is  a  bit  denser, 
including  abundant  erect  hairs,  practically 
lacking  in  ejuncida.  The  presence  of  erect 
to  suberect  hairs  in  pilosa  vs.  subdecum- 
bent  to  decumbent  hairs  in  ejuncida  is  es- 
pecially notable  on  the  femora.  In  lateral 
view  the  angle  formed  by  the  junction  of 
the  dorsal  and  declivitous  propodeal  faces 
is  greater  in  ejuncida  than  in  pilosa,  giving 
it  a  more  slender  appearance. 

Etymology. — This  sleek  species  inspired 
the  use  of  the  Latin  word  for  slender,  ejun- 
cida. 


Gnamptogenys  ericae  Forel,  revalidated 

Gnamptogenys  tornata  var.  ericae  Forel  1912:33; 

Brown  1958:229. 
Grtamptogenys  sulcata  (F.  Smith);  Brown  1958: 

329;  Lattke  1989:24. 
Gnamptogenys  sulcatum   bufonum  Weber   1938: 

208.  NEW  SYNONYMY. 

Diagnosis. — Mandibles  subtriangular; 
posterior  mesosomal  dorsum  and  propo- 
deum  with  longitudinal  costulae,  trans- 
verse sutures  barely  visible  under  certain 
angles  of  lighting;  small  metacoxal  tooth. 
Body  black;  antennae,  legs  and  mandibles 
brown  to  dark  brown. 

Ecology. — Can  be  found  nesting  in  dis- 
turbed forests  and  coffee  plantations  as 
well  as  in  undisturbed  forests. 

Comments. — Much  confusion  has  accu- 


Volume  4,  1995 


167 


mulated  regarding  the  identity  of  this  spe- 
cies and  the  similar  sulcata.  Many  previous 
references  to  sulcata  are  actually  ericae,  but 
the  confusion  makes  it  quite  difficult  to 
distinguish  between  each  reference.  Dr. 
Ivan  Lobl  of  the  MHNG  kindly  examined 
the  ericae  type  and  confirmed  the  longitu- 
dinal direction  of  the  costulae  on  the  pos- 
terior face  of  the  node  of  the  specimen.  A 
specimen  in  the  MCZC  from  Guiana  with 
a  determination  label  as  bufonum  in  N. 
Weber's  handwriting  and  another  label 
stating  "in  Bufo  marinus  stomach"  coin- 
cides with  the  information  in  Weber's  de- 
scription. Even  though  it  has  no  label  des- 
ignating it  as  a  type,  it  seems  reasonable 
to  conclude  that  Weber's  bufonum  is  a  syn- 
onym of  ericae.  For  a  broader  discussion 
on  the  separation  of  the  forms  known  as 
tornata  and  sulcata  the  reader  should  turn 
to  the  "Discussion"  for  G.  sulcata.  Color  in 
this  species  is  relatively  constant  but  oc- 
casional ferruginous,  brown  or  mottled 
specimens  will  turn  up.  Size  and  shape  of 
the  petiolar  node  does  not  vary  as  much 
as  in  sulcata.  Workers  seem  to  always  have 
longitudinal  costulae  on  the  propodeal  de- 
clivity, though  queens  may  have  trans- 
verse costulae. 

Specimens  Examined. — BOLIVIA,  Rosa- 
rio.  COLOMBIA,  Magdalena:  Minca,  610 
m;  Guajira:  Serrania  de  Macuira,  7  km  S 
Nazareth,  70-200m;  Meta:  San  Juan  de  Ar- 
ama,  914m;  La  Macarena,  Rio  Guayabero, 
260m;  65  km  E  Puerto  Lopez.  GUIANA, 
Forest  Settlement,  R.  Mazaruni.  TRINI- 
DAD, Port  of  Spain,  Federation  Park. 
VENEZUELA,  Merida:  Santa  Cruz  de 
Mora,  600m;  Guarico:  Hato  Masaguaral; 
Bolivar:  Campamento  Rio  Grande,  8°07'N 
6r48'W,  280  m;  Anzoategui:  20  km  S  El 
Tigre. 

Gnamptogenys  extra  Lattke,  new  species 
(Figs.  45-47) 

T\/pe  Material. — Holotype  worker.  EC- 
UADOR, Pichincha:  Tinalandia,  16  km  SE 
Santo  Domingo  de  Los  Colorados,  VI- 
1976,  S.  &  J.  Peck,  leg.  Deposited  in  MIZA. 


Paratypes:  (1)  One  worker  with  the  same 
data  as  the  holotype.  Deposited  in  MCZC. 
(2)  Two  workers  from  COLOMBIA,  Cho- 
c6:  Lloro,  Vereda  Pefialoza,  26-III-1988,  V. 
Jaimes,  leg.  One  worker  in  BMNH  and 
LACM,  respectively. 

Worker. — Holotype  (Paratypes)  mea- 
surements: HL  1.20  (1.10-1.12);  HW  1.00 
(0.94-1.00);  ML  0.34  (0.30-0.34);  ED  0.16 
(0.14-0.18);  SL  1.26  (1.00-1.20);  WL  1.68 
(1.44-1.50)  mm;  CI  0.83  (0.85-0.98);  SI  1.26 
(1.06-1.20);  lO  0.16  (0.15-0.19)  n=4. 

With  head  in  frontal  view:  vertexal  mar- 
gin straight,  laterally  convex  with  eyes 
slightly  behind  midlength,  and  anterior 
clypeal  margin  convex;  mandibles  trian- 
gular with  dorsal  striae;  scapes  relatively 
thick  in  appearance  and  surpassing  ver- 
texal margin;  vertex  with  2-3  transverse 
costulae;  anterior  pronotal  face  with  5-7 
transverse  costulae,  dorsally  with  anteri- 
orly arching  costulae;  costulae  on  meso- 
notal  concentric  or  longitudinal;  costulae 
on  metanotum  and  dorsal  propodeal  face 
transverse  to  longitudinal,  sometimes 
arched;  declivitous  propt^deal  face  with 
longitudinal  costulae  that  diverge  poste- 
riorly, basally  with  2-3  transverse  costu- 
lae. 

Petiolar  node  in  lateral  view  subquad- 
rate,  with  broad  anterodorsal  convexity 
and  sharp  posterior  drop  with  overhang- 
ing apex;  subpetiolar  apex  is  an  anteriorlv 
projecting  lobe;  anterior  postpetiolar  face 
with  transverse  costulae  that  arch  back 
and  bect")me  longitudinal  posteriorlv;  rest 
of  dorsal  gastric  cc^stulae  longitudinal,  di- 
vergent on  posterior  postpetiolar  sternum; 
ventral  stridulitrum  present  on  fourth  ab- 
dominal segment.  Scapes  with  abdundant 
decumbent  pilosity  and  no  suberect  or 
subdecumbent  hairs;  decumbent  pilosity 
also  present  on  coxae  and  legs,  with  no 
standing  hairs;  sparse  suberect  hairs  on 
rest  of  body.  Body  dark  brown,  legs  and 
antennae  slightly  lighter. 

Male,  Female. — Unknown. 

Discussion. — This  species  has  a  fairly 
pointed  petiolar  node  apex,  but  it  doesn't 


168 


Journal  of  Hymenoptera  Research 


approach  the  extremely  pointed  condition 
of  G.  acuta.  The  node  is  more  robust  and 
subquadrate  (lateral  view)  than  in  other 
porcata  subgroup  species  and  is  reminis- 
cent of  the  condition  in  G.  tortiiolosa.  This 
species  could  be  closely  related  to  G.  stria- 
tula  on  account  of  the  node  shape  (erect) 
and  the  lacking  anterior  peduncle. 

Gnamptogenys  fernandezi  Lattke 

Gnamptogenys  fernandezi  Latike  1990:  14. 

Diagnosis. — Triangular  mandibles  with 
a  smooth  and  shining  dorsum;  anterior  la- 
mella of  clypeus  convex;  promesonotal  su- 
ture lightly  impressed  and  mesometanotal 
suture  absent;  low,  elongate  petiole,  pos- 
terodorsally  bluntly  pointed;  metacoxal 
teeth  absent. 

Ecologif. — A  rarely  collected,  but  fairly 
widespread  species  in  n  South  America.  In 
Venezuela  known  only  from  2  forested  lo- 
calities in  the  Cordillera  de  la  Costa  be- 
tween 650-1100  m. 

Comments. — See  Lattke  (1990). 

Specimens  Examined. — COLOMBIA:  Val- 
le:  50  m  (no  other  data).  ECUADOR.  Mo- 
rora:  Santiago,  Los  Tayos.  PERU,  Madre 
de  Dios:  5  km  SE  Prto.  Maldonado,  Rio 
Tambopata,  260m;  Cuzco  Amazonico,  15 
km  NE  Prto.  Maldonado,  200m.  VENE- 
ZUELA, Carabobo:  Hacienda  Palmichal, 
Canoabo  900  m. 

Gnamptogenys  gentryi  Lattke, 
new  species 

Type  Material. — Holotype  worker.  CO- 
LOMBIA, Valle,  2250  m,  Insp.  Pance,  Re- 
serva  Natural  Hato  Viejo,  J.B.  Hillaire,  leg. 
Deposited  in  MIZA.  Thank  you  to  Fernan- 
do Fernandez  for  making  the  specimen 
available. 

Worker. — Holotype  measurements:  HL 
1.22;  ML  0.32;  HW  0.92;  ED  0.22;  SL  1.04; 
WL  1.64  mm;  CI  0.75;  SI  1.13;  Ol  0.24. 

Head  in  frontal  view  elongate,  anterior 
clypeal  margin  convex,  posterior  margin 
concave,  eyes  relatively  flat,  finely  costu- 
late  throughout,  though  diverging  at  ver- 


tex, with  a  band  of  three  transverse  cos- 
tulae  next  to  the  vertexal  carinae;  mandi- 
bles triangular,  frontal  surface  rugulose. 
Scapes  mostly  smooth  and  shining  except 
for  sparse  punctulae. 

Pronotal  dorsum  with  whorled  costulae 
almost  enclosing  3  brief  longitudinal  cos- 
tulae, rest  of  mesosomal  dorsum  with  lon- 
gitudinal costulae,  metanotal  groove 
deeper  than  pronotal  suture.  In  lateral 
view  the  mesosoma  has  a  convex  pronotal 
profile  that  becomes  an  inclined  and  more 
or  less  straight  slope  that  bends  sharply  at 
the  propodeal  declivity,  which  has  a 
broadly  concave  profile.  Node  in  lateral 
view  tilted  backwards,  with  oblique  to 
longitudinal  rugulae  laterally  and  trans- 
verse costulae  on  the  anterior  and  poste- 
rior faces,  ventral  process  subquadrate, 
with  anterior  corner  projecting  more  than 
posterior  corner.  Gastric  tergite  I  mostly 
smooth  and  shining  except  for  anterior 
face  which  has  a  small  area  of  transverse 
costulae  and  the  sternite  with  transverse 
striae;  the  rest  of  the  gaster  is  smooth  and 
shining.  Color  black. 

Queen,  Male. — Unknown. 

Discussion. — This  striatula  group  species 
is  quite  easy  to  separate  from  all  others 
just  considering  the  presence  of  a  metan- 
otal groove  and  the  mostly  smooth  and 
shining  gaster. 

Etymology. — Named  in  honor  of  the  late 
botanist  Alwyn  Gentry,  he  died  in  the  line 
of  duty  in  the  Andes  of  Ecuador. 

Gnamptogenys  gracilis  Santschi 

Holcoponera  gracilis  Santschi  1929:468. 
Gnamptogenys  gracilis  (Santschi);  Brown  1958: 
228,  232;  kempt  1972:112;  Lattke  1990:17. 

Diagnosis. — Finely  costulae;  pronotal 
costulae  concentric  to  arched;  mesonotum 
frequently  with  longitudinal  costulae  sur- 
rounded by  concentric  costulae;  metano- 
tum  and  propodeal  dorsum  with  trans- 
verse costulae,  and  longitudinal  on  decliv- 
itous propodeal  face;  subpetiolar  process 
anteriorly  triangular  and  posteriorly  an- 
gular. 


Volume  4,  1995 


169 


Ecology. — This  is  an  epigaeic  forager 
known  only  from  primary  and  secondary 
wet  forests  of  the  Guiana  Shield  and  its 
remnants  in  the  Venezuelan  Amazonas, 
mostly  between  altitudes  of  850-1600  m. 
Within  the  range  of  1000-1200  it  can  be 
locally  abundant.  One  record  from  550  is 
next  to  Angel  Falls.  This  suggests  that 
proximity  with  the  higher  forested  talus 
slopes  of  Auyan  Tepuy  could  explain  its 
ocurrence  at  such  a  low  elevation.  This 
species  obviously  prefers  cool,  mesic  hab- 
itats. This  distribution  plus  climatic  evi- 
dence (Schubert  1988)  suggests  that  G. 
gracilis  may  have  been  more  widely  dis- 
tributed in  this  area  during  the  last  glacial 
period. 

Comments. — On  account  of  size,  similar 
sculpture,  posteriorly  inclined  node  and 
fair  amount  of  pilosity,  its  nearest  relative 
appears  to  be  pilosa,  also  an  apparent  en- 
demic, of  the  Colombian  Cordillera  N  and 
NNW  of  Cali.  An  occasional  specimen 
may  have  a  second,  smaller  seta  just  be- 
low the  prominent  seta  of  the  foretarsal 
base. 

Specimens  Examined. — VENEZUELA, 
Bolivar:  Auyan  Tepuy,  Sector  Aonda, 
6°02'N  62°37'W,  1600  m;  Auyan  Tepui, 
Cafion  del  Diablo,  Isla  Raton,  550  m;  San 
Ignacio  de  Yuruani,  5°00'N  61°08'W,  800 
m  (MIZA). 

Gnamptogenys  haenschi  Emery 

Gnamptogcmis  haenschi  Emery  1902:27;  Brown 
1958:302;"Kempf  1972:112. 

Diagnosis. — Vertexal  margin  concave  in 
frontal  view;  mandibles  with  longitudinal 
costulae;  declivitous  propodeal  face  with 
longitudinal  striae  superiorly  and  basally 
transverse  rugae;  petiolar  node  transverse; 
postpetiolar  sternum  mostly  transversely 
strigulate;  no  metacoxal  tooth. 

Ecology. — This  species  inhabitats  forest 
and  apparently  nests  in  rotten  wood  as 
well  as  in  soil.  It  has  also  been  taken  in  oil 
palm  plantations.  Some  specimens  were 
collected  from  carrion  traps  baited  with 


iguana  meat.  Longino  (pers.  comm.)  re- 
ports finding  a  nest  beneath  a  rotten  log; 
the  ants  burrowed  into  the  soil  on  expo- 
sure. 

Comments. — This  species  is  possibly  the 
only  survivor  of  an  otherwise  extinct  lin- 
eage in  the  genus.  See  discussion  on  phy- 
logeny  of  the  different  species  groups. 

Specimens. — BOLIVIA,  Lower  Rio  Mad- 
idi;  El  Montero,  70  km  N  Sta.  Cruz.  CO- 
LOMBIA, Cauca:  Isla  La  Gorgona;  Guaji- 
ra:  Quebrada  Guacoche,  vie.  Don  Diego, 
10  m;  Magdalena:  4  km  N  San  Pablo, 
10°57'N  74°03'W,  550  m.  COSTA  RICA, 
Peninsula  de  Osa:  Corcovado,  8°28'N 
83°35'W;  Heredia:  La  Selva;  3  km  S  Prto. 
Viejo,  10°26'N  84°00'W,  50m.  ECUADOR, 
Napo:  Prto.  Misahualli,  350  m.  PERU,  Tin- 
go  Maria:  Valle  de  Monson;  Puerto  Mal- 
donado,  260m;  Loreto:  Ramon  Castillo,  5 
km  NW  Leticia.  PANAMA,  Darien:  Rio 
Tacaruna,  580m.  VENEZUELA,  Alto  Rio 
Siapa,  1°40'N  64°35'W,  530m. 

Gnamptogenys  hartmani  (Wheeler) 

Ectatoniuw  {Parectatomma)  hartmani  Wheeler 
1915:390. 

Gnamptogenys  txartmani  (Wheeler);  Brown  1958: 
228,  302;  Echols,  1964:137;  Kempf  1972:112; 
MacKay  1988:127. 

Gnamptogenys  nigrifnvis  Borgmeier  1948;  Brown 
1958:  228,  236;  Kempf  1972:114.  NEW  SYN- 
ONYMY. 

Gnamptogenys  turmalis  Kempf  and  Brown  1968: 
93;  Kempf  1972:116.  NEW  SYNONYMY. 

Diagnosis. — Superolateral  corners  of  de- 
clivitous propodeal  face  with  small  lobes 
or  carinae;  mandibles  triangular  to  sub- 
triangular;  metanotal  groove  vaguely  im- 
pressed and  posterior  nodal  face  has  lon- 
gitudinal costulae;  metacoxal  dorsum 
with  lobes;  subpetiolar  process  subquad- 
rate. 

Ecology. — One  record  from  soil  in  a  ba- 
nana farm  and  another  from  soil  in  a  de- 
stroyed Trachymyrmex  nest.  These  ants 
have  on  several  occasions  been  reported 
as  predators  of  Trachymrmex  ants  (Echols, 
1964:137;  Kempf  and  Brown,  1968:94).  J. 


170 


Journal  of  Hymenoptera  Research 


Longino  (pers.  comm.)  reports  from  Costa 
Rican  field  notes  of  Dana  Myer:  "a  nest 
was  found  found  in  leaf  litter  amidst  the 
remains  of  a  Trachymrmex  nest  and  many 
cut  up  workers  and  a  queen  of  the  attines 
were  also  found  along  with  many  wound- 
ed Gnamptogenys  workers."  Longino  has 
observed  this  species  carrying  its  brood  in 
a  loose  3  m  column,  fleeing  from  raiding 
Eciton.  One  specimen  was  taken  from  the 
stomach  contents  of  Dendrobates  lecomelns. 

Comments. — Kempf  and  Brown  de- 
scribed G.  turmalis  as  being  close  to  G.  ni- 
grifrons  but  slightly  larger  and  more  ro- 
bust, with  finer  sculpturing  and  a  uniform 
ferruginous  color.  These  authors  realized 
that  G.  hartmani,  G.  nigrifrons  and  G.  tur- 
malis were  so  close  as  to  constitute  possi- 
ble synonyms,  and  they  were  right.  The 
study  of  specimens  at  hand  show  colors 
vary  enough  to  make  it  an  unreliable  char- 
acter for  separating  species  and  the  same 
is  true  for  irregularities  in  the  sculpture. 
Specimens  from  more  southern  localities 
have  finer  costulation.  Also  found  to  vary 
continously  was  the  length  vs.  width  of 
the  petiolar  node,  as  well  as  other  size  in- 
dicators. 

Specimens  Examined. — BRAZIL,  Bahia, 
Ilheus-Itab.,  km  22,  (CEPEC).  COSTA 
RICA,  Peninsula  de  Osa:  Sirena,  8°28'N 
83°35'W,  50m.  GUYANA,  Kartabo;  HON- 
DURAS, La  Lima,  Zapote  farm.  MEXICO, 
Tamaulipas:  10  km  W  EL  Encino,  23°N 
08'W  99°10'W;  PANAMA,  Isla  Barro  Col- 
orado. PERU,  Valle  de  Chanchamayo,  800 
m.  USA,  Louisiana:  Lucky;  Texas:  Bent- 
son-Rio  Grande  State  Park,  30.  VENE- 
ZUELA, Amazonas,  vie.  Cerro  Ya-Pakana. 

Gnamptogenys  haytiana  (Wheeler  and 
Mann) 

Spaniopone  haytiana  Wheeler  and  Mann  1911:11. 
Gnamptogenys  hai/tiana   (Wheeler  and   Mann); 
Brown  1958:228,  316;  Kempf  1972:113. 

Diagnosis. — Petiole  node  disciform,  with 
anterior  and  posterior  faces  more  or  less 
parallel  to  one  another,  dorsal  and  lateral 


nodal  faces  with  transverse  rugulae;  me- 
tanotal  groove  barely  impressed;  propo- 
deal  spiracles  mounted  on  turrets  at  mid- 
height  of  the  lateral  edge  of  declivitous 
propodeal  face. 

Ecology. — Taken  from  forest  and  coffee 
plantation  leaf  litter.  The  dramatic  defor- 
estation of  Hispaniola  has  considerably  re- 
duced the  range  of  this  species. 

Comments. — Endemic  to  Hispaniola  Is- 
land. On  account  of  the  yellowish  color, 
more  pronounced  curvature  of  the  second 
gastric  segment  and  different  sculpture, 
this  species  is  outstanding  amongst  the 
other  members  of  its  species  complex.  It  is 
possible  that  it  represents  an  independent 
development  from  the  strigata  complex. 
The  holotype  was  examined  in  the  MCZC. 

Specimens  Examined. — DOMINICAN 
REPUBLIC,  Barahana:  2  km  N  Polo, 
1000m. 

Gnamptogenys  horni  Santschi 

Gnamptogenys  regularis  horni  Santschi  1929:475. 
Gnamptogenys  horni  Santschi;  Brown  1958:228, 

235,  316;' Kempf  1961:491;  Kempf  1972:113; 

Lattke  1990:17. 

Diagnosis. — Subtriangular  mandibles; 
dorsum  of  petiolar  node  with  posterome- 
dian longitudinal  costulae,  transversely 
arched  anteriorly;  episternal  costulae 
curve  on  to  declivitous  propodeal  face. 
Body  dark  brown;  legs  and  antennae  fer- 
ruginous. 

Ecology. — A  series  was  taken  from  the 
stomach  of  Colostethus  nubicola.  See  Lattke 
1990. 

Comments. — This  common  species  has  a 
wide  range  in  the  neotropics.  There  is  fair 
size  variation  and  smaller  specimens  usu- 
ally have  more  vermiculate  costulation. 
The  position  of  the  propodeal  spiracle  can 
vary  from  mid-heigth  to  below  mid-heigth 
and  the  metacoxal  tooth  from  triangular  to 
denticulate.  Nests  of  small  forms  can 
sometimes  be  found  in  the  same  locality 
as  larger  forms.  Intermediates  were  found 
in  other  localities.  A  specimen  labelled  as 
a  "cotype"  was  examined  in  the  MCZC. 


Volume  4,  1995 


171 


Specimens  Examined. — BRAZIL,  Amazo- 
nas:  Rio  Taruma;  Benjamin  Constant;  Re- 
serva  Ducke;  via  Manaus-Itacoatiara  km 
50;  Manaus;  Rondonia:  Vilhema;  Para:  Ca- 
rajas;  Utinga  Tract  vie.  Belem.  BOLIVIA, 
Caranavi,  vie.  radio,  800m.  COLOMBIA, 
Choco,  Rio  Napipi.  ECUADOR,  Pichincha: 
Estacion  Rio  Palenque;  Sucua;  Morona: 
Santiago,  Los  Tayos;  Esmeraldas:  48  km  S 
Atacames.  GUYANA,  Kamakusa.  PANA- 
MA, Isla  Barro  Colorado.  PERU,  Pangu- 
ana,  9°37'S  74°56'W,  220m;  Tingo  Maria: 
Valle  del  Monson.  SURINAM,  Tambah- 
redjo;  Dirkshoop;  La  Poolle;  Maripa- 
hueuei,  Vank.  VENEZUELA,  Amazonas: 
Alto  Rio  Mavaca,  2°02'N  65°06'W,  200m; 
Alto  Rio  Siapa,  1°40'N  64°35'W,  600m. 

Gnamptogenys  ilimani  Lattke, 
new  species 

Type  Material. — Holotype  worker.  BO- 
LIVIA, 22  km  N  Caranavi,  Vivero  Ilimani, 
1700  m,  22-VI-81,  C.  Kugler,  leg.  Primary 
forest  clearing  with  Cinchona,  nest  in  rot- 
ten wood.  Deposited  in  the  MIZA.  Para- 
types:  paranidotypic  workers  from  the 
same  nest  as  the  holotype.  Deposited  in 
each  of  the  following:  BMNH,  LACM, 
MUSP,  MCZC,  MIZA. 

Worker. — Holotypes  (Paratypes)  mea- 
surements: HL  0.90  (0.84-0.90);  HW  0.78 
(0.72-0.78);  ML  0.30  (0.26-0.32);  ED  0.14 
(0.14-0.18);  SL  0.70  (0.70-0.72);  WL  1.20 
(1.12-1.20)  mm;  CI  0.87  (0.84-0.88);  SI  0.90 
(0.92-0.97);  OI  0.19  (0.18-0.27)  n=5. 

Coarsely  costulate  strigata  group  spe- 
cies. With  head  in  frontal  view  vertexal 
margin  convex;  anterior  pronotal  face 
with  3-4  costulae  that  sharply  bend  back 
laterally  to  become  longitudinal;  declivi- 
tous propodeal  face  with  longitudinal  cos- 
tulae; node  anteroposteriorly  compressed, 
its  posterior  face  sharply  marginate  later- 
ally with  a  flat  to  slightly  concave  surface 
and  effaced  sculpture;  anterior  postpetio- 
lar  face  with  2-3  transverse  costulae,  rest 
longitudinal;  postpetiolar  sternum  with 
longitudinal  costulae;  lateral  mesocoxal 
face  with  oblique,  rough  costulae  and  that 


of  metacoxae  with  rough  rugulae;  meta- 
coxal  tooth  low  and  triangular.  Body  black 
with  yellow-brown  legs  and  scapes;  abun- 
dant appressed  pilosity  on  legs  and 
scapes,  but  no  standing  hairs. 

Qi/mj.— Measurements:  HL  0.88  (0.88); 
HW  0.72  (0.74);  ED  0.20  (0.18);  SL  0.72 
(0.72);  WL  1.28  (1.20)  n  =  2.  Differences 
from  workers  are  the  usual;  caste  differ- 
ences, though  the  node  is  more  disciform. 

Male. — Unknown. 

Discussion. — This  species  is  nearest  to  G. 
strigata  and  G.  pittieri  but  they  are  sepa- 
rable on  several  on  several  accounts:  the 
posterior  petiolar  node  face  is  not  sharply 
set  off  and  has  raised  costulae;  in  dorsal 
view  their  node  is  anteriorly  convex,  not 
straight;  standing  hairs  on  the  body  and 
especially  the  scapes  are  abundant,  scape 
pilosity  is  longer.  In  G.  pittieri  the  anterior 
transverse  costulae  of  the  pronotum  curve 
around  gently  at  the  sides. 

Etymology. — The  species  name  alludes 
to  the  type  locality. 

Gnamptogenys  ingeborgae  Brown 

Gnamptogenys  ingeborgae  Brown  1992:279. 

Diagnosis. — Mandibular  masticatory 
margin  denticulate,  promesonotal  suture 
present,  at  least  partially,  and  the  longi- 
tudinal costulae  on  the  propodeal  dorsum 
and  postpetiolar  tergite  is  frequently  ill- 
defined  to  effaced. 

Ecology. — A  specialized  millipede  feeder 
from  Colombia.  See  Brown  1992. 

Comments. — See  Brown  1992. 

Specimens  Examined. — COLOMBIA, 
Cundinamarca,  Bogota-Villavicencio  road, 
km  79,  1100  m  (type  series:  MCZC). 

Gnamptogenys  interrupta  Mayr 

Ectatomuia    {Gnamptogenys)    interruptum    Mayr 

1887:543. 
Gnamptogenys  interruptum  (Mayr);  Mann  1922:3; 

Brown  1958:228,  303;  Kempf  1968:377;  Kempf 

1972:  113;  Lattke  1990:18. 

Diagnosis. — Sublinear  mandibles.  Ce- 
phalic dorsum,  mesosoma,  and  gastric  ter- 


172 


Journal  of  Hymenoptera  Research 


gum  I  with  longitudinal  costulae;  gastric 
tergum  II  smooth;  pleura  also  with 
smooth  patches;  metacoxal  tooth  absent. 
Body  reddish  brown;  legs  and  antennae 
ferruginous. 

Ecology. — Found  in  humid  forests  of 
lowlands  and  mountains  (cloud  forest). 
Taken  from  leaf  litter  samples  and  beneath 
bark  of  rotting  logs  on  ground. 

Comments. — Lateral  mesosomal  costulae 
can  be  effaced  to  a  variable  degree  and  the 
second  gastric  segment  can  ocassionally 
have  weak  longitudinal  costulae,  medi- 
anly  effaced.  Declivitous  propodeal  face 
with  longitudinal  costulae  and  weakly  de- 
veloped anterolateral  lobes. 

Specimens  Examined. — COLOMBIA, 
Magdalena:  3  km  SE  Minca,  11°08'N  74° 
06'W,  1050m;  Valle:  Puerto  Merizalde,  10 
m.  COSTA  RICA,  Puntarenas:  Montever- 
de,  10°18'N  84°48'W,  1500m.  HONDU- 
RAS, Lombardia;  JAMAICA,  Mandeville. 

Gnamptogenys  kempfi  Lenko 

Gnamptogenys  kempfi  Lenko   1964:257;   Kempf 
1972:114. 

Diagnosis. — Mandibles  subtriangular, 
dorsally  smooth  and  shining,  transverse 
costulae  on  mesosomal  dorsum  and  node, 
no  apparent  transverse  sutures  on  meso- 
somal dorsum.  Propodeal  spiracle  large 
and  metacoxal  tooth  present. 

Ecology. — Apparently  from  lowland  for- 
est, one  nest  was  found  in  rotten  wood  on 
the  ground. 

Comments. — A  widepread  but  uncom- 
mon species. 

Specimens  Examined. — COLOMBIA, 
Amazonas:  7  km  N  Leticia.  PERU,  Pan- 
guana,  9°37'S  74°56'W,  220m  (MCZC). 

Gnamptogenys  lanei  Kempf 

Gnamptogenys  lanei  Kempf  1960:388-90;  Kempf 
1968:377;  Kempf  1970:  325;  Kempf  1972:114. 

Diagnosis. — Relatively  small  eyes;  clyp- 
eal  lamella  medianly  concave;  promeson- 
otal  suture  vestigial  and  small  denticles  on 


the  propodeum;  node  elongate,  with 
transverse  costulae. 

Ecology. — Its  morphology  puts  it  in  the 
rastrata  group  of  millipede  hunters. 

Comments. — Kempf  1968:377  reports  a 
series  with  transverse  costulae  on  the  an- 
terior pronotal  face,  differing  from  the  lon- 
gitudinal sculpture  of  the  type  series. 

MflfenVi/.— BRAZIL,  Para:  Belem,  Rio 
Guama  (MUSP). 

Gnamptogenys  laticephala  Lattke, 
new  species 
(Figs.  54,  55) 

Type  Material. — Holotype  worker.  EC- 
UADOR, Guayas:  3  km  S  Bucay,  24-VII- 
73,  W.L.  Brown,  leg.  Deposited  in  MCZC. 

Worker. — Holotype  measurements:  HL 
1.71;  ML  1.01;  HW  1.94;  SL  1.53;  ED  0.34; 
WL  2.32  mm;  CI  1.13;  SI  0.79;  OI  0.18. 

Head  in  frontal  view  broad,  vertexal 
margin  fairly  straight,  lateral  margins 
slightly  convex  and  anteriorly  diverging; 
eyes  large  and  situated  at  cephalic  mid- 
length;  anterior  clypeal  lamella  broadly 
concave  with  lateral  triangular  teeth  near 
mandibular  insertions;  clypeus  with  two 
small  anteriorly  projecting  lobes  between 
anterior  border  of  antennal  fossae  and  an- 
terior lamella;  cephalic  dorsum  with  lon- 
gitudinal rugae  that  diverge  slightly  pos- 
terad  on  to  vertex;  rugae  between  frontal 
lobes  and  eyes  more  irregular  than  rugae 
between  and  behind  frontal  lobes;  abun- 
dant piligerous  punctures  present.  Anten- 
nal scapes  smooth  and  shining,  surpassing 
vertexal  margin;  gula  with  median  longi- 
tudinal carinae  and  transverse  costae  at 
each  side;  mandibles  falcate,  longitudinal- 
ly costulate  with  interior  glabrous  sulcus 
that  ends  at  apical  tooth,  basal  tooth 
marks  the  end  of  basal  margin  and  is  fol- 
lowed by  two  more  pre-apical  teeth;  an- 
terior pronotal  face  opaque,  not  as  shiny 
as  rest  of  pronotum  and  with  faint  trans- 
verse rugulae;  mesosomal  dorsm  with 
parallel  longitudinal  costulae,  promeson- 
otal  suture  well  impressed  but  does  not 
break  longitudinal  sculpture,  mesometa- 


Volume  4,  1995 


173 


Fig.  54-55.     Micrographs  of  G.  laticcplmla.  Scale  bars  =1.0  mm.  54,  head;  55,  body. 


174 


Journal  of  Hymenoptera  Research 


notal  suture  deep  and  interrupting  sculp- 
ture except  for  some  ridges  that  join  a  few 
costulae  on  either  side  of  suture;  lateral 
pronotal  face  with  slightly  oblique  parallel 
costulae;  pronotal-mesopleural  suture 
very  broad  and  deep;  meso-  and  metepis- 
terna  with  parallel,  longitudinal  costulae; 
mesometepisternal  suture  well  impressed 
and  metepisternal-propodeal  suture  dis- 
tinct; metepisternal  lobe  well  developed; 
declivitous  propodeal  face  with  longitu- 
dinal costulae;  petiolar  node  in  lateral 
view  low,  anterodorsal  margin  convex 
and  posterodorsal  margin  sharply  convex; 
transversely  costulate;  subpetiolar  process 
anteriorly  triangular,  with  slighty  round- 
ed posterior  lobe. 

Postpetiolar  costulation  longitudinal, 
weakly  roughened  and  becoming  rugulo- 
se  laterally,  ventrally  with  transverse  cos- 
tulae that  tend  to  fade  medially;  gastric 
tergum  II  strongly  vaulted  anteroventrally 
with  dorsal  and  lateral  sculpture  as  on 
preceding  segment;  apical  gastric  seg- 
ments with  faint  transverse  strigulae, 
tending  to  smooth  and  shining;  anterior 
face  of  procoxae  smooth  and  shining  lat- 
erally with  oblique  costulae;  meso-  and 
metacoxae  laterally  smooth  and  shining, 
dorsally  transversely  costulate;  metacoxal 
teeth  well  developed;  body  with  sparse 
suberect  and  decumbent  hairs,  longest  on 
gastric  apex  and  clypeus,  no  appressed  pi- 
losity  on  scapes,  just  suberect  and  decum- 
bent hairs.  Mesosoma  dark  reddish 
brown;  head,  node  and  gaster  darker;  legs 
and  antennae  brown;  mandibles  brown  to 
yellow  brown. 

Queen,  Male. — Uknown. 

Discussion. — The  two  lobes  between  the 
antennal  fossae  and  clypeal  lamella  distin- 
guish this  species  from  all  other  New 
World  members  of  the  group,  which  lack 
such  lobes,  including  its  closest  relative, 
banksi.  Other  characters  in  which  banksi 
differs  are  the  following:  areolate  sculp- 
ture on  the  cephalic  dorsum,  occiput  and 
anterior  pronotal  face  with  transverse  ru- 
gae, deep  lateral  and  mesonotal  pits  for 


the  spiracles  (such  pits  lacking  in  frauda- 
trix),  pronotal  suture  interrupts  sculpture, 
and  the  inner  mandibular  mandibular  sul- 
cus only  reaches  the  basal  angle.  In  gen- 
eral the  sculpture  of  banksi  is  rougher, 
mostly  of  gaster  I,  which  is  reticuate  ru- 
gose and  gaster  II  has  rough,  widely 
spaced,  dorsal  longitudinal  rugae,  becom- 
ing more  irregular  laterally.  The  form  of 
the  pronotal-mesopleural  suture  and  the 
lobes  formed  by  the  pronotum  and  anepi- 
sternum  at  the  base  of  the  suture  are  rem- 
iniscent of  some  minuta  group  species.  The 
possibility  exists  that  this  could  be  an  ab- 
errant specimen  of  banksi  (it  was  found  de- 
termined as  such),  but  given  the  observed 
differences  I  have  opted  for  the  most  con- 
servative course.  It  is  hoped  that  the  cap- 
ture of  additional  specimens  may  throw 
more  light  onto  the  status  of  laticepJiala. 

Etymology. — The  name  alludes  to  the 
relatively  broad  head  when  seen  in  frontal 
view. 

Gnamptogenys  lineolata  Brown 

Gnamptogenys  lineolata  Brown  1992:275. 

Diagnosis. — Head  in  frontal  view  su- 
bquadrate,  scapes  smooth  and  shining; 
mandibles  denticulate;  sculpturing  of 
body  striate,  lacking  propodeal  denticle. 

Ecology. — A  specialized  millipede  feeder 
endemic  to  Hispaniola. 

Comments. — See  Brown  1992. 

Specimens  Examined. — DOMINICAN 
REPUBLIC,  La  Vega,  La  Cienaga,  1100  m. 

Gnamptogenys  lucaris  Kempf 

Ectatotwna  tornatum  Luederwalt  1926:236  (nee 

Roger  1862). 
Gnamptogenys  lucaris  Kempf  1968:379;  Kempf 

1972:114. 

Diagnosis. — Mandibles  subtrianglar; 
clypeal  lamella  laterally  bluntly  rectangu- 
lar, not  acutely  dentate;  propodeum  with 
longitudianal  costulae;  node  low,  its  de- 
clivitous face  narrow. 

Ecology. — Luederwalt   (1926)    reports   a 


Volume  4,  1995 


175 


Specimen  on  a  shrub  devouring  the  re- 
mains of  a  beetle. 

Comments. — This  species  is  very  close  to 
G.  siapensis  and  the  possiblity  exists  that 
they  are  conspecific. 

Specimens  Examined. — None. 

Gnamptogenys  mecotyle  Brown 

Gnauipfogcm/s  mecotyle  Brown  1958:318;  Kempf 
1972:114.' 

Diagnosis. — Promesonotal  suture  and 
metanotal  groove  impressed  but  not 
breaking  sculpture;  propodeal  spiracle 
opening  large  and  round,  right  behind 
teeth;  metacoxal  teeth  long  and  sharp;  oc- 
cipital carinae  wide  and  visible  in  frontal 
view;  anterior  clypeal  lamella  laterally 
rounded  and  anteriorly  straight  to  very 
broadly  convex. 

Ecology. — Millipede  feeder  captured  in 
litter  samples  from  wet  forest.  Some  spec- 
imens found  in  stomach  contents  of  Den- 
drobates  histroniciis. 

Comments. — The  cephalic  sculpture  can 
vary  from  evenly  costulate  to  roughly  ru- 
gose and  the  petiole  length  vs.  width  is 
also  variable. 

Specimens  Examined. — COLOMBIA,  An- 
tioquia,  Providencia,  Estacion  Biologica; 
Choco:  Serrania  de  Baudo,  Camino  de 
Yupe,  500-700m;  Amazonas:  7  km  N  Le- 
ticia.  COSTA  RICA,  Puntarenas:  6  km  SW 
Monteverde.  10°16'N  84°50'W,  900m. 
PANAMA,  Isla  Barro  Colorado. 

Gnamptogenys  mediatrix  Brown 

Gnamptogenys  mediatrix  Brown  1958:362;  Kempf 
1972:n4.' 

Diagnosis. — Mandibles  elongate  and  tri- 
angular, their  basal  %  rugulose  and  apical 
Vi  smooth  and  shining;  clypeal  lamella  me- 
dianly  concave;  scapes  with  no  erect  hairs; 
petiolar  costulation  mostly  transverse, 
node  broader  than  long;  metacoxal  teeth 
very  slender,  propodeal  teeth  short. 

Ecology. — Apparently  of  lowland  for- 
ests, one  series  taken  from  a  varzea  habi- 
tat. Its  morphology  puts  it  within  the  ras- 


trata  group,  and  it  is  quite  probably  a  mil- 
lipede hunter. 

Comments. — Rarely  collected,  it  is  a  close 
relative  of  G.  laticephala. 

Specimens  Examined. — BRAZIL,  Amazo- 
nas: Ilha  de  Curari;  Para;  Igarape  Mana 
(MCZC). 

Gnamptogenys  menozii  (Borgmeier) 

Ectatomma    {Parectatomma)    menozzi   Borgmeier 

1928:32 
Gnamptogenys  menozzi  (Borgmeier);  Brown  1958: 

228,  316,  321;  Kempf  1972:113 
Ectatomma  {Parectatomma)  schubarti  Borgmeier 

1948:198.  NEW  SYNONYMY. 
Gnamptogenys   schubarti   (Borgmeier);   Brown 

1958:316;  kempf  1960a:390;  Kempf  1972:114. 

Diagnosis. — Eyes  convex;  mandibles; 
promesonotal  suture  impressed  but  not 
deep,  metanotal  groove  very  deep;  pro- 
podeal teeth  short;  mesosoma  longitudi- 
nally costulate. 

Ecology. — Found  in  forested  areas,  in- 
cluding montane  forests. 

Comments. — Borgmeier  described  G. 
schubarti  upon  characters  always  "a  bit 
more"  than  for  G.  menozzi  and  it  is  curious 
to  note  how  authors  frequently  refer  to 
these  two  forms  together  when  discussing 
their  differences  from  other  species. 
Brown  (1958:316)  discusses  differences  be- 
tween the  two  forms:  size,  number  of  cos- 
tulae,  color  and  the  shape  of  the  subpetio- 
lar  process.  He  also  admits  that  the  forms 
could  belong  to  one  species.  Size  differ- 
ences between  the  two  forms  overlap  and 
the  color  and  gauge  of  the  costulation  are 
also  continuously  variable.  The  differences 
in  the  shape  of  the  subpetiolar  process  are 
well  within  infraspecific  variation,  at  least 
as  observed  in  other  Gnamptogenys. 

Specimens  Examined. — BRAZIL,  Rio 
Grande  do  Sul  {menozzi  cotype:  MUSP); 
Sao  Paulo:  Monte  Alegre,  900m  {schubarti 
cotype:  MUSP);  Rio  Corocovado;  Espiritu 
Santo:  Res.  Nova  Lombardia,  4  km  N 
Santa  Tereza,  900  m. 


176 


Journal  of  Hymenoptera  Research 


Gnamptogenys  mina  (Brown) 

Holcoponera  mina  Brown  1957:494. 
GnampHogeiJi/s  mina  Brown  1958:220,  228;  Kempf 
1972:114;'Lattke  1990:18. 

Diagnosis. — Mandibular  dorsum  smooth 
and  shining  and  apex  of  scapes  surpassing 
the  vertexal  margin;  declivitous  propodeal 
face  flat  and  separated  from  the  dorsal 
face  by  a  sharp  angle;  propodeal  spiracles 
elevated  at  the  apex  of  cylindrical  tuber- 
cles. 

Ecology. — Found  in  leaf  litter  and  earth 
samples  from  humid  forests. 

Comments. — The  series  from  near  Ka- 
vanayen  show  very  sharply  defined  cos- 
tulae  with  fine  vermiculation  in  contrast 
with  the  glazed  and  effaced  aspect  of  oth- 
er series. 

Specimens  Examined. — COLOMBIA,  Pu- 
tumayo:  Villa  Garzon  (MCZC).  VENE- 
ZUELA, Bolivar:  10  km  E  Kavanayen, 
1200  m  (MIZA). 

Gnamptogenys  moelleri  (Forel) 

Holcoponera  moelleri  Forel  1912:34;  Mann  1916: 

404. 
Holcoponera   teffensis   Santschi   1929:449.   NEW 

SYNONYMY. 
Holcoponera  teffensis  var.  concinna  Santschi  1929: 

449  (nee.  F.  Smith  1858);  Brown  1958:229. 
Holcoponera  moelleri  var.  splendens  Santschi  1929: 

450;  Brown  1958:228. 
Gnamptogenys  moelleri  (Forel);  Brown  1958:329; 

Kempf  1972:113;  Kempf  1976:51. 
Gnamptogenys  teffensis  (Santschi);  Brown  1958: 

329;  Kempf  1972:115;  Lattke  1990:24. 

Diagnosis. — Slightly  roughened  costu- 
lae;  pronotum  with  concentric  costulae; 
posterodorsal  mesosomal  face  with  longi- 
tudinal costulae;  subpetiolar  process  sub- 
quadrate,  sometimes  anteriorly  projecting 
into  a  lobe. 

Ecology. — Epigaeic  forager  of  mostly 
lowland  forested  areas. 

Comments. — Comparison  of  specimens 
determined  as  G.  moelleri  with  G.  teffensis 
forms  reveal  nothing  beyond  normal  in- 
fraspecific  variation.  Santschi  described  G. 


teffensis  as  a  relative  of  G  striatida  differ- 
ing in  gauge  of  costulation,  pilosity,  and 
size.  The  use  of  length  vs.  width  of  the 
funnicular  segments  in  separating  G  moel- 
leri from  G  teffensis  is  useless  as  even  in 
members  of  the  same  nest  series  the  ratio 
will  differ  either  way. 

Specimens  Examined. — ARGENTINA,  50 
km  S.  Oran.  BRAZIL,  Amazonas:  Fazenda 
Esteio,  80  km  NNE  Manaus,  2°25'S 
59°46'W,  80m.  COLOMBIA,  Amazonas:  7 
km  N  Leticia;  Putumayo:  5  km  S  Mocoa, 
610m;  Meta:  Rio  Guayabero,  Isla  Angos- 
tura, 279  m;  Reserva  La  Macarena,  580m; 
Choco:  Finca  Los  Guaduales,  800m;  Nari- 
fio:  28  km  SE  Mocoa,  510m.  ECUADOR, 
Napo:  Limoncocha,  250m;  59  km  E  Quev- 
edo,  Los  Rios,  2000m;  PARAGUAY, 
Neembuca,  Pilar.  PERU,  Panguana,  9°37'S 
74°56'W,  220m;  Puerto  Maldonado,  260m. 
VENEZUELA,  Barinas:  17  km  SW  Ciudad 
Bolivia,  8°04'N  70°48'W,  240  m. 

Gnamptogenys  mordax  (Smith) 

Ponera  mordax  F.  Smith  1858:98. 

Ponera  nodosa  Latreille  1802:217;  Brown  1958: 
228. 

Ectatomma  (Gnamptogenys)  mordax  (Smith);  Em- 
ery 1896:49. 

Gnamptogenys  mordax  var.  purensis  Forel  1912; 
Brown  1958:228. 

Gnamptogenys  mordax  (Smith);  Mann  1922:3; 
Brown  1958:319;  Kempf  1972:113;  Kempf 
1976:51;  Lattke  1990:19. 

Gnamptogenys  mordax  var.  sebastiani  Borgmeier 
1937:220;'Brown  1958:228. 

Diagnosis. — Mandibles  subtriangular; 
declivitous  propodeal  face  transversely 
costulate;  gastric  tergum  II  can  vary  from 
smooth  to  longitudinally  costulate  or  cos- 
tulate-rugose.  Body  very  dark,  brown,  al- 
most black;  legs  dark  brown.  Smaller  in- 
dividuals can  be  confused  with  G.  contin- 
iia. 

Ecology. — A  common  dweller  of  humid 
forests. 

Cojnments. —Size  (HW  1.08-2.08;  WL 
1.66-2.92  mm)  and  sculpture  patterns  of 


Volume  4,  1995 


177 


57 


59 


Figs.  56-59.     Frontal  view  of  head,  and  lateral  view  of  mesosoma  and  petiole.  56-57,  G.  iliniani;  58-59,  G. 
nigrivitrea. 


this  widespread  and  timid  species  is  quite 
variable. 

Specimens  Examined. — COSTA  RICA, 
Heredia:  17  km  N.  Volcan  Barba,  10°17'N 
84°05'W,  880  m.  GUYANA,  Kaietur.  MEX- 
ICO, Chiapas:  Jetja.  PERU,  Panguana, 
9°37'S  74°56'W,  220  m;  Tingo  Maria:  Mon- 
son  Valley;  69  km  E  Tingo  Maria,  1200m. 
VENEZUELA,  Amazonas:  Alto  Rio  Siapa, 
1°40'N  64°35'W,  600m. 

Gnamptogenys  nana  Kempf 

Gnamptogein/s   nana   Kempf   1960b:422;   Kempf 

1972:113.' 
Parectatomma  dina  Kusnezov,  1969:35;  Kempf 

1978:35. 

Diagnosis. — Head  subquadrate,  mandi- 
bles subtriangular  with  striae  at  base, 
clypeal  lamella  straight,  no  transverse  me- 
sosomal  impressions,  propodeal  declivity 
with  vertical  costulae. 


Ecology. — Unknown.  Its  nearest  rela- 
tives, horni  and  regularis  include  beetles  as 
a  significant  part  of  their  diets,  while  horni 
also  preys  on  ants. 

Comments. — A  rarely  collected  species, 
the  specimen  mentioned  below  is  topotyp- 
ic  (MCZC). 

Specimens  Examined. — BRAZIL,  Sao  Pau- 
lo: Agudos. 

Gnamptogenys  nigrivitrea  Lattke, 
new  species 
(Figs.  58,  59) 

Type  Material. — Holotype  worker:  CO- 
LOMBIA, Valle:  Reserva  Forestal  de  Yo- 
toco,  3°50'N  76°25'W,  42  km  NNE  Call, 
1100-1500  m,  6-1-84,  W.P.  MacKay,  leg. 
No.  7221.  Deposited  in  MIZA.  Paratype: 
One  worker  with  same  locality  data  as  ho- 
lotype, deposited  in  LACM. 

VSlorker. — Holotype  (paratype)  measure- 


178 


Journal  of  Hymenoptera  Research 


merits:  HL  1.10  (1.10);  HW  0.86  (0.82);  ML 
0.86  (0.82);  ED  0.14  (0.12);  SL  0.82  (0.84); 
WL  1.40  (1.36)  mm;  CI  0.79  (0.79);  SI  0.95 
(1.02);  OI  0.16  (0.15).  Head  elongate;  fairly 
parallel  sided  in  frontal  view;  posterior 
margin  slightly  concave;  clypeal  lamella 
widest  medianly,  giving  it  a  bluntly  point- 
ed aspect;  eyes  moderately  protuberant. 
Mandibles  triangular  and  shallowly  cos- 
tulate;  no  trace  of  mesometanotal  suture; 
pronotum  anterolaterally  convex;  declivi- 
tous propodeal  face  longitudinally  costu- 
late;  metacoxal  tooth  triangular;  node  pos- 
teriorly inclined,  in  lateral  view  with  a 
broadly  convex  anterior  margin  and  a 
bluntly  pointed  apex  which  slightly  over- 
hangs the  posterior  margin,  in  dorsal  view 
with  longitudinal  costulae;  subpetiolar 
process  is  an  anteriorly  projecting  lobe; 
anterior  face  of  postpetiole  with  3  trans- 
verse costulae,  costulation  longitudinal  on 
dorsal  face;  postpetiolar  sternum  with  dis- 
tinct costulation;  scape  with  few  standing 
hairs  and  moderate  amount  of  decumbent 
pilosity.  Body  black;  antennae,  legs,  me- 
tapleural  gland  area,  and  gastric  apex 
brown.  Coxae  very  dark  brown. 

Female,  Male. — Unknown. 

Discussion. — The  closest  species  to  G.  ni- 
grivitrea  is  G.  hrunnnea,  but  its  well  im- 
pressed mesometanotal  groove,  angulate 
anterolateral  pronotal  area  and  depressed 
apical  and  basal  mandibular  margins  per- 
mit easy  separation. 

Etymology. — The  species  name  is  de- 
rived from  the  Latin  adjectives  for  black, 
nigra,  and  for  glassy,  vitra,  and  alludes  to 
the  black  and  shining  sculpture. 

Gnamptogenys  perspicax 

Kempf  and  Brown 

Gnamptogenys  perspicax  Kempf  and  Brown  1970: 
316;  Kempf  1972:114. 

Diagnosis. — Eye  slightly  behind  cephalic 
mid-length;  scape  longitudinally  striate 
with  abundant  standing  hairs;  petiole 
node  more  or  less  evenly  convex;  postpe- 


tiolar sternum  with  transverse  costulae  or 
rugae. 

Ecology. — A  millipede  feeder  found  in 
humid  forests. 

Comments. — Apparently  a  sister  species 
of  bispinosa. 

Specimens  Examined. — ECUADOR,  Pi- 
chincha,  Estacion  Rio  Palenque  (MCZC). 

Gnamptogenys  pilosa  Lattke, 
new  species 
(Figs.  51-53) 

Type  Material. — Holotype  worker.  CO- 
LOMBIA, Valle:  Calima,  Canon  El  Pital,  F. 
Castano,  leg.  Deposited  in  MIZA.  Para- 
types:  Three  workers  with  same  locality 
data  as  holotype.  One  deposited  in  each  of 
the  following:  LACM,  BMNH,  MCZC. 

The  late  Mr.  F.  Castano  kindly  provided 
additional  information  about  the  type  lo- 
cality. The  site  is  by  a  stream  called  Que- 
brada  El  Pital  (tributary  of  Rio  Calima)  in 
a  gorge  900-1300  m  above  sea  level.  Ap- 
proximate coordinates:  3°08'N  76°40'W. 
The  site  will  soon  be  flooded  due  to  con- 
struction of  a  dam.  The  specimens  were 
found  on  leaf  litter  in  a  cloud  forest. 

Worker. — Holotype  (paratypes)  mea- 
surements: HL  1.44  (1.40-1.48);  HW  1.20 
(1.20-1.22);  ML  0.54  (0.46-0.54);  ED  0.28 
(0.28-0.30);  SL  1.46  (1.50-1.57);  WL  2.00 
(2.00-2.06)  mm;  CI  0.83  (0.81-0.86);  SI  1.21 
(1.25-1.28);  OI  0.23  (0.23-0.25)  n=4. 

With  head  in  frontal  view,  vertexal  mar- 
gin fairly  straight,  sides  broadly  convex; 
anterior  clypeal  margin  convex,  lamella 
well  developed;  eyes  prominent  on  pos- 
terior one  half  of  head;  mandibles  trian- 
gular, apical  margin  denticulate,  dorsum 
striate;  cephalic  dorsum  with  longitudinal 
parallel  striae,  diverging  posterad  at  ver- 
tex; 1-2  transverse  striae  parallel  to  occip- 
ital carina;  anterior  prontal  face  trans- 
versely striate;  dorsal  mesosomal  surface, 
including  all  of  propodeum,  longitudinal- 
ly striate;  striae  curve  around  propodeal 
spiracles  with  no  transverse  striae  be- 
tween spiracles;  lateral  pronotal  surface 
obliquely  to  transversely  striate,  striation 


Volume  4,  1995 


179 


on  pleura  and  lateral  propodeal  face  par- 
allel with  dorsal  striae;  inner  procoxal  fac- 
es punctate,  anterolaterally  transversely 
striae;  lateral  faces  of  meso-  and  metacox- 
ae  weakly  striate;  metacoxa  with  well-de- 
veloped tooth. 

Petiolar  node  transversely  striate,  low; 
viewed  laterally  with  convex  anterior 
margin  and  concave  posterior  margin, 
apex  overhanging;  subpetiolar  process 
with  a  projecting  anterior  lobe  and  su- 
bquadrate  posteriorly;  transverse  striae 
anteriorly  on  gastric  tergum,  up  to  three- 
fourths  length  of  tergum,  rest  of  gaster 
longitudinally  striate;  anteroventral  post- 
petiolar  process  prominent;  sternite  with 
longitudinal  striation,  diverging  caudad; 
abundant  erect  to  suberect  hairs  on  body 
and  extremities;  abundant  pilosity  on  cox- 
ae, propodeum,  petiole,  antennae  and 
legs. 

Queen,  Male. — Unknown. 

Discussion. — This  is  the  hairiest  of  all 
Gnamptogem/s  species.  On  account  of  size, 
slender  habitus,  posteriorly  inclined  peti- 
olar node,  fine  costulation,  and  abundant 
pilosity  its  nearest  reatives  appear  to  be  G. 
gracilis  and  G.  ejuncida.  The  sculpture  of 
ejuncida  is  coarser,  the  pilosity  is  not  as 
dense  and  the  presence  of  erect  hairs  is 
considerably  less.  The  break  between  the 
dorsal  and  declivitous  propodeal  faces  is 
sharper  in  G.  pilosa,  giving  it  a  more  ro- 
bust aspect.  G.  gracilis  is  an  endemic  from 
the  Guiana  Shield  area,  and  has  the  post- 
petiolar  sternal  disc  smooth  and  shining. 

Etymology. — The  species  name  is  de- 
rived from  the  Greek  word  for  hair:  pilos. 

Gnamptogenys  pittieri  Lattke 
Gnamptogenys  pittieri  Lattke  1990:21. 

Diagnosis. — Sculpture  with  rough  as- 
pect; frontal  lobes  each  with  a  median 
emargination;  eyes  protuberant  and  subgl- 
obular;  propodeal  spiracles  mounted  at 
the  apex  of  protuberances. 

Ecology. — Hypogeic  leaf  litter  (including 


bamboo)  dweller  of  cloud  forests  of  the 
Venezuelan  Cordillera  de  la  Costa. 

Comments. — The  series  from  near  Tejer- 
ias  differs  from  the  type  series  in  having 
the  notch  on  the  frontal  lobes  shallower, 
the  propodeal  spiracles  on  higher  turrets, 
finer  costulae  on  the  anterior  postpetiolar 
face  and  metacoxal  spines  more  acute. 

Specimens  Examined. — VENEZUELA, 
Aragua:  Pq.  Nac.  Henri  Pittier,  vie.  Ran- 
cho  Grande,  1500m;  17  km  S  Las  Tejerias, 
1300  m  (MIZA). 

Gnamptogenys  pleurodon  (Emery) 

Ectatomma  (Holcoponera)  pleurodon  Emery  1896: 

47. 
Holcoponera  emery i  Santschi  1929:  463  (nee  Forel 

1901). 
Holcoponera    vidua    Santschi    1929:467;    Brown 

1958:229. 
Gnamptogenys  pleurodon  (Emery);  Brown  1958: 

320;    Kempf    1961:390;     Kempf    1970:325; 

Kempf  1972:114;  Lattke  1990:22. 

Diagnosis. — Pronotum  with  concentric 
costulae;  mesonotum  with  longitudinal 
costulae,  sometimes  surrounded  by  con- 
centric costulae;  anterior  postpetiolar  face 
with  3-4  transverse  costulae  and  dorsal 
face  with  longitudinal  costulae;  metacoxal 
tooth  acicular.  Body  dark  brown,  legs 
brown. 

Ecologxj. — Kempf  (1970)  notes  a  nest 
preference  for  pre-existing  cavities  in 
plants.  It  is  also  a  frequent  forager  on  trees 
and  shrubs.  Usually  found  in  lowland  to 
premontane  humid  forests. 

Comments. — An  occasional  specimens 
may  have  vertical  costulation  on  the  de- 
clivitous propodeal  face,  as  in  striatula,  but 
the  backwards  tilt  of  the  petiolar  node  and 
its  brief  anterior  peduncle  will  identify 
pleurodon. 

Specimens  Examined. — BOLIVIA,  Tumu- 
pasa.  BRAZIL,  Amazonas:  Benjamin  Con- 
stant, Fazenda  Esteio,  80  km  NNE  Ma- 
naus,  80  m;  Iriboca,  Pirelli  Plantation.  CO- 
LOMBIA, Amazonas:  7  km  N  Leticia;  Pu- 
tumayo:  5  km  S  Mocoa,  610m.  ECUADOR, 
Napo:  Sushijindi;  Tena,  400m.  PERU,  Ma- 


180 


Journal  of  Hymenoptera  Research 


dre  de  Dios:  Estacion  Biologica  Cocha  Ca- 
shu,  400;  Tingo  Maria:  Yurac,  108  km  E 
Tingo  Maria.  SURINAM,  Brownsberg  Na- 
tuur  Park.  VENEZUELA,  Amazonas,  Alto 
Rio  Mavaca,  2°01'N  65°07'W,  200m;  Boli- 
var, Auyan  Tepui,  Cafion  del  Diablo,  Isla 
Raton,  500m. 

Gnamptogenys  porcata  (Emery) 

Holcoponera  porcata  Emery  1896:48. 
Holcoponera  uiaguifica  Santschi   1921:81.   NEW 

SYNONYMY. 
Gnamptogenys  magnifica  (Santschi);  Brown  1959: 

320;  Kempf  1972:113. 
Gnamptogenys  porcata  (Emery);  Brown  1958:320; 

Kempf  1972:114;  Lattke  1990:23. 

Diagnosis. — Costate  species,  median  lon- 
gitudinal costae  on  pronotum  inscribed 
anteriorly  by  transverse  costae;  postero- 
dorsal  mesosomal  sculpture  variable:  con- 
centric ovaloids  with  longitudinal  or 
transverse  orientation,  or  with  longitudi- 
nal costae  inscribed  within  whorls;  pice- 
ous  body  with  ferruginous  legs. 

Ecology. — A  dweller  of  premontane  to 
montane  humid  forests.  J.  Longino  (pers. 
comm.)  found  an  incipent  nest  in  Cecropia 
insignis.  One  nest  midden  contained  most- 
ly the  remains  of  beetles:  scotylids,  bostri- 
chids,  curculionids,  besides  isopods.  Nests 
from  one  population  (Venezuela,  Tachira) 
consistently  had  their  brood  chambers  ir- 
regularly wall-papered  with  the  remains 
of  pupal  cocoons. 

Comments. — Santschi  described  magnifi- 
ca as  close  to  porcata  but  differing  in  its 
larger  size  and  the  transverse  costulae  on 
the  posterior  dorsal  mesosoma.  Brown 
(pers.  comm.)  studied  the  magnifica  type  in 
1963  and  noted  it  was  "very  close"  to  por- 
cata with  the  former  slightly  larger  and 
with  a  higher,  thinner  node  as  seen  later- 
ally. Specimens  in  the  USNM  from  Bolivia 
collected  by  Mann  during  1956  bear  anon- 
ymous Holcoponera  magnifica  determina- 
tions. The  study  of  series  from  Venezuela 
representing  several  colonies  from  differ- 
ent populations  show  great  variability  for 
dorsal  mesosomal  sculpturing.  Size  differ- 


ences between  the  two  forms  are  no  more 
than  infraspecific.  One  Honduran  worker 
has  a  wide,  lobe-like  subpetiolar  process. 
Specimens  Examined. — BOLIVIA  (no  oth- 
er data);  COLOMBIA,  Antioquia:  Rio  For- 
ce, 1020m;  Cundinamarca:  above  Tena, 
1300-1600  m;  Valle:  3,2  km  above  Rio 
Agua  CLara,  old  Cali-Buenaventura  rd; 
Rio  San  Juan  (tributary  of  Rio  Digua),  vie. 
Queremal,  1300m;  Municipio  Buenaven- 
tura, 650  m.  COSTA  RICA,  Rio  Toro  Am- 
arillo,  vie.  Guapiles,  <  40  m;  Heredia: 
10°19'N  84°43'W,  800m.  ECUADOR,  Pi- 
chincha:  Tinalandia,  16  km  SE  Sto.  Domin- 
go de  los  Colorados,  680m.  HONDURAS, 
14  km  S  La  Ceiba;  Lombardia.  VENEZUE- 
LA, Tachira:  vie.  Siberia,  39,7  km  WNW 
San  Cristobal,  1200m;  Trujillo:  15  ESE  Bo- 
cono,  1160m. 

Gnamptogenys  rastrata  (Mayr) 

Ectatomma  (Parectatomma)  rastrata  Mayr  1866:89. 

Gnamptogenys  trigona  Emery  1905:114;  Brown 
1958:321;  Kempf  1972:116.  NEW  SYNONY- 
MY. 

Gnamptogenys  rastrata  (Mayr);  Brown  1958:322; 
Kempf  1972:114;  Kempf  1976:52. 

Diagnosis. — Antennal  scapes  surpass 
vertexal  margin  by  no  more  than  their  api- 
cal width;  triangular  and  edentate  man- 
dibles with  a  varying  degree  of  costulae 
and  rugae;  small  tubercle-like  propodeal 
teeth  next  to  conspicuous  spiracles  which 
are  slightly  elevated  above  rest  of  sur- 
rounding cuticle;  postpetiolar  sternum 
with  median  longitudinal  smooth  and 
shining  areas. 

Ecology. — Captured  in  wet  forests,  in- 
cluding montane  areas.  A  millipede  feed- 
er. 

Comments. — G.  trigona  was  separated 
from  rastrata  on  account  of  differences  in 
the  gauge  and  form  of  the  costulation,  as 
well  as  size.  Variability  of  mandibular 
sculpture  on  basal  flange  and  dorsum  is 
nothing  beyond  infraspecific.  The  sculp- 
ture on  the  node  as  seen  dorsally  can  vary 
from  concentric  costulae,  with  transverse 
or  longitudinal  costulae  in  the  middle,  or 


Volume  4,  1995 


181 


completely  transverse.  Costulae  transverse 
on  anterior  petiolar  node  face.  Brown 
(1958:321)  discusses  the  differences  in  size, 
costulae  counts,  length  vs.  width  of  node, 
subpetiolar  process  shape  and  variability 
of  sculpture  on  node  between  the  speci- 
mens of  G.  rastrata  and  G.  trigona  he  ex- 
amined. As  more  specimens  became  avail- 
able the  gaps  were  bridged.  The  petiolar 
node  can  be  wider  tha  long  or  the  oppo- 
site. Size  range  is  HL  0.74-0.90;  WL  1.10- 
1.28  mm. 

Specimens  Examined. — Brazil,  Rio  de  Ja- 
neiro: Jussaral;  Sao  Paulo:  Sao  Bernardo 
do  Campo;  Nova  Teutonia,  27°11'S 
52°23'W,  300-500m  (MCZC). 

Gnamptogenys  regularis  (Mayr) 

Gnamptogenys  regularis  Mayr  1870:965;   Mann 

1922:3;  Brown  1959:229,  319;  Kempf  1972:114; 

Lattke  1990:23. 
Gnamptogenys  rimulosa  var.  splendida  Pergande 

1896:871;  Brown  1959:229. 
Gnamptogenys  fiebrigi    Forel    1909:253;    Brown 

1958:229.^ 

Diagnosis. — Declivitous  propodeal  face 
abruptly  separated  from  dorsal  face,  su- 
perolaterally  with  small  angulate  lobes; 
metacoxal  tooth  apically  rounded  and  not 
triangular;  dark  brown  body,  testaceous 
legs. 

Ecology. — J.  Longino  (pers.  comm.)  re- 
ports observing  a  short  linear  column  of 
workers  walk  to  an  apparent  nest  entrance 
on  the  ground.  He  excavated  it  before  any 
activity  had  begun  and  found  a  small  col- 
ony of  Pseudomyrmex  boopns.  The  related  G. 
horjti  has  a  dietary  preference  for  ants  and 
beetles  (Lattke,  1990).  One  specimen  was 
removed  from  the  stomach  of  a  Tamandua 
tetrndactylus. 

Comments. — The  degree  of  transverse 
costulation  on  the  anterior  nodal  face  has 
traditionally  been  used  to  separate  this 
species  from  horni,  but  as  additional  ma- 
terial accumulated  it  became  impossible  to 
rely  on  the  character.  Transverse  costula- 
tion can  be  totally  absent  or  occupy  all  of 
the  anterior  nodal  face.  Females  tend  to 


have  the  anterior  nodal  face  totally  trans- 
versely costulate.  This  was  noted  for  one 
specimen  by  Mayr  (1871:965)  who  con- 
cluded it  was  not  normal  and  that  a  fe- 
male with  longitudinal  costulae  would 
eventually  be  found.  The  longitudinal  cos- 
tulae on  the  sides  of  the  mesosoma  do  not 
curve  onto  the  declivitous  propodeal  face. 
The  type  of  splendida  was  examined  in  the 
CASC. 

Specimens  Examined. — ARGENTINA, 
Tintina.  BRAZIL,  Amazonas:  Reserva 
Ducke,  vie.  Manaus;  Maturaca.  COLOM- 
BIA, Magdalena:  4  km  N  San  Pedro, 
10°57'N  74°03'W,  550  m.  COSTA  RICA, 
Alajuela:  vie.  Volcan  Arenal,  10°29'N 
84°44'W,  550  m.  Heredia:  10°20'N 
84°04'W,  500  m;  Peninsula  de  Osa:  Sirena, 
8°28'N  83°35'W,  50m;  Puntarenas:  Res. 
Biol.  Carara,  9°47'N  84°36'W,  500m;  vie. 
Guapiles,  Rio  Toro  Amarillo.  BRAZIL, 
Goias:  Anapolis;  Sao  Paulo:  Holambra; 
Agudos;  Distrito  Federal:  Deodoro;  Ama- 
zonas: Maturaca;  Minas  Gerais:  Carmo  da 
Cachoeira;  Bahia:  Salvador.  ECUADOR, 
Pichincha:  Rio  Palenque,  Centro  Cientifi- 
co;  Santo  Domingo  de  los  Colorados;  Pi- 
chilingue:  Los  Rios.  GUIANA,  (NY  Quar- 
antine Stn.).  MEXICO,  Tepic;  Veracruz:  10 
km  NNW  Sontecomapan,  200m.  PERU, 
Tingo  Maria:  Valle  del  Monson;  Junin:  Co- 
lonia  Perene,  Rio  Perene,  18  km  NE  La 
Merced;  Puerto  Maldonado,  260  m.  VEN- 
EZUELA, Sucre:  El  Rincon,  10°36'N 
63°12'W,  100  m;  Guarico:  Hato  Masaguar- 
al. 

Gnamptogenys  reichenspergeri 

(Santschi) 

Acanthoponera  (Anacanthoponera)  reichenspergeri 

Santschi  1929:274. 
Gnamptogenys  reichenspergeri  (Santschi);  Brown 

1958:324;  Kempf  1972:114. 

Diagnosis. — Head  in  frontal  view  with 
concave  vertexal  margin,  vertex  smooth 
and  shining;  anterior  face  of  postpetiole 
and  median  disc  of  gastric  tergum  II 
smooth  and  shining;  metacoxal  spine  ab- 


182 


Journal  of  Hymenoptera  Research 


sent  and  declivitous  face  of  propodeum 
with  small  superolateral  lobes,  spiracles 
lateral  and  not  elevated;  metanotal  suture 
absent. 

Ecology. — From  leaf-litter  samples  of  hu- 
mid forests.  . 

Comments. — A  rarely  collected  ant,  its 
nearest  relative  is  apparently  relicta. 

Specimens  Examined. — BRAZIL,  Jussaral: 
Angra  dos  Rios  (MCZC).  VENEZUELA, 
Amazonas:  Cerro  Ya-pakana  (LACM). 

Gnamptogenys  relicta  (Mann) 

Rhopalopone  relicta  Mann  1916:403. 
Holcoponera  relicta  (Mann);  Brown  1957:491. 
Gnamptogenys  relicta  (Mann):  Brown  1958:229; 

Kempf  1972:114;  Wheeler  &  Wheeler  1975: 

119. 

Diagnosis. — Mandibles,  vertex  and  an- 
terior postpetiolar  face  smooth  and  shin- 
ing, occiput  sometimes  with  very  faint 
transverse  costulae;  last  3  antennal  seg- 
ments form  vague  club;  declivitous  pro- 
podeal  face  with  transverse  costulae,  pro- 
podeal  spiracles  elevated  above  rest  of  cu- 
ticle; pronotal  suture  present  but  fine,  me- 
tanotal suture  well  impressed,  breaking 
sculpture;  no  anterolateral  lobes  on 
declivitous  propodeal  face;  metacoxal 
spine  present. 

Ecology. — Sifted  from  leaf  litter  in  rain 
forests. 

Comments. — The  degree  of  effacement  of 
sculpture,  and  size  can  vary  considerably. 
S.  Cover  reports  a  specimen  from  the  N. 
Range  of  Trinidad. 

Specimens  Examined. — BRAZIL,  Amazo- 
nas: Rio  Taruma,  High  Falls;  Benjamin 
Constant;  Ponta  Negra,  N  of  Manaus;  66 
km  N  Manaus  on  Carcari  road.  COLOM- 
BIA, Amazonas:  7  km  N.  Leticia.  SURI- 
NAM, Raleigh  Vallen-Voltzberg  Res., 
Voltzberg  Camp,  90m.  VENEZUELA,  Bo- 
livar: 49  km  ESE  Tumeremo,  7°28'N 
61°06'W,  200  m. 

Gnamptogenys  rimulosa  (Roger) 

Ponera  rimulosa  Roger  1861:18. 
Gnamptogenys  rimulosa  (Roger);  Brown  1958:324; 
Kempf  1972:114. 


Diagnosis. — Mandibles  with  small  basal 
patch  of  costulae,  rest  smooth  and  shining; 
metacoxa  with  broad,  basal  lobe;  petiolar 
node  longitudinally  costulate  throughout; 
postpetiolar  sternum  smooth  and  shining. 

Ecology. — Unknown. 

Comments. — This  species  seems  to  be 
particularly  close  to  G.  regularis.  Even 
though  regidaris  workers  lack  the  small  la- 
terobasal  costulate  area  on  the  mandibles, 
females  have  it;  there  is  a  slight  depression 
of  the  posterior  mesosomal  dorsum,  ves- 
tigial propodeal  lobes  and  totally  longi- 
tudinal costulae  on  the  node. 

Specimens  Examined. — BRAZIL,  Santa 
Catarina:  Nova  Teutonia:  27°11'S  52°23'W, 
300-500m. 

Gnamptogenys  schmitti  (Forel) 

Emeryella  schmitti  Forel   1901:334;  Wheeler  & 

Wheeler  1952:127. 
Emeryella   schmitti   minor   Wheeler    1936:195; 

Brown  1958:229. 
Gnamptogenys  schmitti  (Forel);  Brown  1958:330; 

Kempf  1972:114. 

Diagnosis. — Very  broad  head,  in  frontal 
view,  and  striking  elongate  mandibles 
without  denticles;  striae  on  anterior  nodal 
face  transverse,  slightly  effaced  laterally; 
striae  on  metanotum  and  propodeum 
transverse;  gastric  sternum  II  smooth  and 
shining. 

Ecology. — A  predator  of  millipedes 
whose  range  has  shrunken  considerably 
this  century  due  to  massive  deforestation 
on  Hispaniola. 

Comments. — This  species  represents  an 
extreme  in  the  development  of  falcate 
mandibles  within  the  genus  (Brown  1958: 
215),  but  probably  represents  a  develop- 
ment independent  of  the  banksi  subgroup 
species.  The  type  series  in  the  MCZC  was 
examined. 

Specimens  Examined. — HAITI,  Diquini. 

Gnamptogenys  sentiferox  Brown 

Gnamptogenys  semiferox  Brown  1958:324;  Kempf 
1972:114. 


Volume  4,  1995  183 

Diagnosis. — Body  with  abundant  coarse  eral  view  subquadrate;  subpetiolar  pro- 
punctation,  especially  on  head;  mandibles  cess  subquadrate,  with  angulate  ventral 
with  about  six  denticles  and  projecting  corners;  postpetiolar  sternum  medially 
basal  lobe;  anterior  clypeal  margin  con-  smooth  and  shining  and  with  lateral  Ion- 
vex;  declivitous  propodeal  face  longitudi-  gitudinal  costulae;  tibiae  and  femora 
nally  costulate.  smooth  and  shining;  metacoxae  with  basal 

Ecology. — A  millipede  hunter  known  triangular  lobe,  dorsally  transversely  cos- 
from  rain  forests  between  900-1200  m  on  tulate  and  laterally  mostly  smooth  and 
Hispaniola.  Also  suffering  from  habitat  shining;  abundant  long  suberect  and  sub- 
shrinkage  due  to  deforestation.  decumbent  hairs  on  body.  Color  ferrugi- 

Comments. — Even  though  it  shares  the  nous, 
lack  of  metacoxal  spines  and  elongate  Queen,  Male. — Unknown, 
mandibles  with  schmitti,  it  does  not  seem  Discussion. — This  species  is  quite  near  to 
to  be  closely  related  to  it.  Its  mandibles  are  lucaris  and  the  differences  between  the 
different,  including  in  cross  section:  shin-  two  species  may  be  infraspecific:  in  lucaris 
ing  and  longitudinally  rugulose,  as  op-  the  petiolar  node  dorsum  is  evenly  con- 
posed  to  opaque  and  striate  in  schmitti.  vex,  the  metacoxal  tooth  is  broadly  trian- 

Specimens    Examined. — DOMINICAN  gular    and    the    subpetiolar   process    has 

REPUBLIC,  Mt.  Diego  de  Ocampo,  3^000  acutely  pointed  angles.  Due  to  the  great 

ft  (Paratype).  differences  that  separate  the  range  of  the 

two  species,  the  apparent  endemicity  of 

Gnamptogenys  siapensis  Lattke,  g^^^^    Gnamptogem/s   to    southern   Brazil, 

new  species  ^^^  ^^xe  scarcity  of  material,   it  seemed 

(Figs.  62,63)  prudent  to  separate  the  forms  until  more 

Type  Material. — Holotype  worker.  VEN-  specimens  are  available. 

EZUELA,   Amazonas:   Alto   Rio  Siapa,  Etymology. — The  species  name  alludes 

1°40'N  64°35'W,  600  m,  4-II-89,  J.  Lattke,  to  the  type  locality. 

leg.  Sifted  leaf  litter  sample.  The  specimen 

1  .  .                            •       11     fi   ^^  Gnamptogenys  stellae  Lattke, 

was  caught  in  an  area  occasionally  flood-  ^         f    &     y 

ed  during  the  rainy  season.  Deposited  in  ^^  ^P     \ 

MIZA.  (P^§^-  ^^'  ^^^ 

Worker.— UL  1.13;  ML  0.44;  HW  1.00;  Type  Material— Holotype  worker.  COS- 
ED 0.25;  SL  0.97;  WL  1.60  mm;  CI  0.89;  SI  TA  RICA,  Estrella  Valley,  April,  1924, 
0.97;  Ol  0.25.  W.M.  Mann,  leg.  Deposited  in  USNM. 

Cephalic  dorsum  with  fine  longitudinal  Worker. — Holotype  measurements:  HL 
costulae;  eyes  placed  behind  midlength;  1.18;  ML  0.57;  HW  1.06;  SL  0.65;  ED  0.13; 
clypeal  lamella  anteriorly  straight  and  lat-  WL  1.55  mm;  CI  0.90;  SI  0.61;  OI  0.12. 
erally  bluntly  angulose;  mandibles  smooth  Head  elongate,  in  frontal  view  with  con- 
and  shining,  subtriangular  with  apical  and  cave  vertexal  margin,  more  or  less  parallel 
basal  margins  joined  through  broad  con-  sided  with  eyes  at  about  midlength;  clyp- 
vexity;  scapes  smooth  and  shining,  sur-  eal  lamella  gradually  projects  forward 
passing  vertexal  margin,  antennal  bullae  from  sides  and  is  medially  weakly  emar- 
partially  visible  beneath  frontal  lobes;  cos-  ginate;  mandibles  almost  linear  and  with 
tulae  longitudinal  throughout  mesosomal  a  double  row  of  low  tubercles  on  apical 
dorsum,  propodeal  declivity  and  petiolar  border,  dorsally  and  laterally  smooth  and 
node;  promesonotal  and  mesometanotal  shining  with  sparse  punctae,  basally  with 
sutures  weakly  impressed;  mesometa-  small  area  of  longitudinal  costulae;  scapes 
pleural  suture  well  impressed;  petiolar  smooth  and  shining,  dorso-ventrally  corn- 
node  more  or  less  as  long  as  wide,  in  lat-  pressed,    bent    at    basal    one-fourth    and 


184 


Journal  of  Hymenoptera  Research 


Figs.  60-61.     Micrographs  of  G.  stellae.  60,  head,  scale  bar  =  0.5  mm;  61,  body,  scale  bar  =  1.0  mm. 


Volume  4,  1995 


185 


Figs.  62-65.     Frontal  view  of  head  and  lateral  view  of  mesosoma  and  petiole.  62-63,  G.  siapensis;  64-65,  G. 
volcano. 


thickest  just  apicad  of  midlength;  cepahlic 
dorsum  with  longitudinal  striae,  diverg- 
ing posterad  at  vertex,  gula  mostly  longi- 
tudinally striate,  weakly  diverging  poster- 
ad; occipital  margin  glabrous. 

Anterior  pronotal  face  and  collar  with 
transverse  striae;  longitudinal  striae  on 
mesosomal  dorsum  and  propodeal  decliv- 
ity, slightly  shallower  on  metanotum  and 
dorsal  propodeal  face;  promesonotal  su- 
ture very  lightly  impressed,  metanotal 
groove  markedly  impressed  and  inter- 
rupting longitudinal  striae.  Mesosomal 
sides  longitudinally  striate,  except  for 
oblique  striae  on  anepisternum;  propodeal 
spiracle  round  and  slightly  raised  above 
rest  of  integument;  procoxae  anteriorly 
and  anterolaterally  smooth  and  shining, 
pc)Sterolaterally  striate;  mesocoxae  with 
dorsal  transverse  striae  that  are  effaced 
posteriorly;  metacoxae  with  transverse 
striae  and  low  dorsobasal  swelling;  tibia 
and  femora  smooth  and  shining;  anterior 


petiolar  node  face  smooth  and  shining, 
laterally  with  weakly  effaced  longitudinal 
striae,  dorsally  diverging  posterad;  node 
wider  posterad  than  anterad;  petiole  in 
lateral  view  slightly  pedunculate,  anteri- 
orly slightly  concave,  dorsally  slightly 
convex,  posterior  face  dropping  sharply; 
subpetiolar  process  anteriorly  projecting 
with  convex  anterior  border  and  posteri- 
orly sinuous;  gastric  terga  I  and  II  with 
longitudinal  striae,  effaced  towards  pos- 
terior margin  of  tergum  II.  Gastric  sterna 
I  and  II  smooth  and  shining;  pilosity 
sparse,  a  row  of  stout  hairs  present  along 
apical  mandibluar  order,  few  standing 
hairs  on  posterior  cephalic  dorsum,  pro- 
notum  and  gaster.  Body  reddish  brown; 
mesosomal  dorsum  and  head  darker 
brown,  leg  and  antennae  ferruginous. 

Queen,  Male. — Unknown. 

Discussion. — The  median  anterior  pro- 
jection of  the  clypeal  lamella  is  unique 
among  the  extant  species  of  the  mordax 


186 


Journal,  of  Hymenoptera  Research 


group.  This  species  seems  to  occupy  an  in- 
termediate position  between  G.  continua 
and  G.  horni.  The  combination  of  clypeal 
configuration,  small  eyes,  subfalcate  man- 
dibles, glabrous  occiput,  mesometanotal 
suture  and  very  brief  petiolar  peduncle 
point  to  this  placement. 

Etymology. — The  species  epithet  is  de- 
rived from  the  Latin  word  for  star,  stella, 
and  alludes  to  the  type  locality. 

Gnamptogenys  striatula  Mayr 

Gnamptogenys    striatula    Mayr    1883:32;    Mann 

1916:404;  Brown  1959:327;  Kempf  1970:325; 

Kempf  1972:115;  Kempf  1976:52;  Lattke  1990: 

23. 
Holcoponera  curtulum  Emery  1896:47;  Forel  1899: 

7.  NEW  SYNONYMY. 
Holcoponera  obscurum  Emery  1896:48;  Lueder- 

wait  1926:238;  Santschi  1929:442. 
Holcoponera    curtula    var.    stolli    Forel    1899:7; 

Brown  1958:228. 
Holcoponera  brasiliensis  Emery,  1902:181;  Brown, 

1958:229. 
Holcoponera  striatula  obscura  var.  angustiloba  For- 
el   1908:341;    Luederwalt    1926:237;    Brown 

1958:229.  Unavailable. 
Holcoponera   striatula    obscura    var.    simplicoides 

Forel  1908:341;  Luederwalt  1926:328.  NEW 

SYNONYMY.  Unavailable. 
Holcoponera  striatula  obscura  var.  angustipleura 

Forel  1908:342;  Brown  1958:229.  Unavailable. 
Holcoponera  curtula  var.  paulina  Forel  1908:342; 

Brown  1958:229. 
Holcoponera  curtula  var.  vollemoeideri  Forel  1912: 

33;  Santschi  1929:453;  Brown  1958:229. 
Holcoponera  striatula  (Mayr);  Luederwalt  1926: 

237;  Santschi  1929:442;  Wheeler  and  Wheeler 

1952:123. 
Holcoponera  striatula  var.  antillana  Santschi  1929: 

444;  Brown  1957:490. 
Holcoponera    rustica    Santschi    1929:446.    NEW 

SYNONYMY. 
Holcoponera   wheeleri  Santschi   1929:448.   NEW 

SYNONYMY. 
Holcopwnera  brasiliensis  var.  pernambucana  Sants- 
chi 1929:452;  Brown  1958:229. 
Holcoponera   brasiliensis  var.   calcarata   Santschi 

1929:452;  Brown  1958:229. 
Holcoponera  brasiliensis  var.  mayri  Santschi  1929: 

453;  Brown  1958:230. 


Holcoponera   brasiliensis   simplicoides   Santschi 

1929:45. 
Holcoponera  brasiliensis  simplicoides  var.  hybrida 

Santschi  1929:455;  Brown  1958:229.  Unavail- 
able. 
Holcoponera  emeryi  var.  recta  Santschi,  1929:465; 

Brown,  1958:229.  NEW  SYNONYMY. 
Holcoponera  regularis  arcuata  Santschi  1929:457; 

Brown  1958:227;  Kempf  1972:112.  NEW  SYN- 
ONYMY. 
Holcoponera    regularis    Santschi    1929:457    (nee 

Mayr  1970);  Brown  1958:227. 
Holcoponera  wasmanni  Santschi  1929:466.  NEW 

SYNONYMY. 
Holcoponera    wasmanni    var.    isthmica    Santschi 

1929:467. 
Gnamptogenys  arcuata  (Santschi);  Brown  1958: 

227,  237;  kempf  1972:112. 
Gnamptogenys  curtula  (Emery);  Brown  1958:227, 

327;  Kempf  1972:112. 
Gnamptogenys    simplicoides    (Santschi);    Brown, 

1958:229,327;  Kempf,  1972:114. 
Gnamptogenys  rustica  (Santschi);  Brown,  1958: 

229,  327;  kempf,  1972:114. 
Gnamptogenys  wasmanni  (Santschi);  Brown  1958: 

229,  238;  Kempf  1972:116. 
Gnamptogenys  wheeleri  (Santschi);  Brown  1958: 

230,  238;  Kempf  1972:116. 

Diagnosis. — Pronotal  costulae  semicir- 
cular; dorsoposterior  mesosomal  costulae 
longitudinal  and  slightly  diverging  cau- 
dad;  mesometanotum  sometimes  with 
semiciruclar  costulae;  triangular  metacox- 
al  teeth;  body  dark  brown. 

Ecology. — A  generalist  epigaeic  forager 
of  humid  forests.  See  Lattke  (1990).  A  mite 
was  found  on  the  anterolateral  petiolar 
side  of  one  specimen  from  Beni,  Bolivia. 

Comments. — This  widespread,  common, 
and  fairly  variable  ant  has  been  the  object 
of  a  lengthy  synonymic  list.  Most  of  the 
names  were  based  upon  differences  in 
size,  gauge  of  costulation,  and  the  shape 
of  the  subpetiolar  process  and  the  meso- 
pisternal  lobe.  These  size  differences  are 
all  normal  for  the  species.  One  can  recog- 
nize coarsely  costulate  and  finely  costulate 
specimens,  as  well  as  intermediate  forms. 
The  finely  costulate  ants  can  be  found  in 
several  places:  s.  Brazil,  Argentina,  Guia- 


Volume  4,  1995  187 

na,  Costa  Rica,  Jamaica  and  Hispaniola.  Cundinamarca:  81  km  de  Bogota;  Fusa- 
Occasional  series  can  be  found  that  have  gasuga;  Valle:  Sevilla;  Cali,  90m.  COSTA 
transverse  costulae  on  part  or  all  of  the  RICA,  Hacienda  La  Pacifica,  50m;  Limon: 
propodeal  declivity;  this  condition  is  not  Finca  La  Lola,  Siquerres;  Heredia:  Finca 
frequent  on  the  finely  costulate  forms.  La  Selva;  Puntarenas:  6  km  SW  Monte- 
Several  series  from  Bolivia  have  a  long  an-  verde,  10°16'N  84°50'W,  900m.  DOMINI- 
terior  slope  of  the  node,  which  approaches  CAN  REPUBLIC,  Clarke  Hall.  ECUA- 
the  condition  in  pleurodon.  The  use  of  the  DOR,  Napo:  Limoncocha,  250m;  Sushijin- 
subpetiolar  process  in  the  separation  of  di:  Sucua:  1500m.  JAMAICA,  St.  Ann,  Ce- 
species  is  of  limited  value  as  a  variation  dar  V.,  760m;  Westmor.  bog,  460m.  EL 
from  subquadrate  to  anteriorly  projecting  SALVADOR,  6,4  Km  N  Quetaltepec. 
lobe  can  be  found  in  series  from  the  same  MARTINIQUE,  (no  other  data).  MEXICO, 
population  or  even  nest.  Brown  (1957:489)  Maria  Madre;  Michoacan:  8  km  SW  Tiqui- 
already  discussed  the  futility  of  using  the  cheo,  430m;  Consequina  slope;  Sinaloa: 
mesepisternal  lobe.  Mazatlan;  Veracruz:  Lago  Catemaco, 
Specimens  determined  as  G.  curtula  by  250m;  5  km  S  Cuernavaca;  Chiapas:  8  km 
Brown  for  the  1958  revision  are  G.  stria-  NE  Huixtla,  225m.  PERU,  Panguana, 
tula,  and  so  are  several  specimens  deter-  9°37'S  74°56'W,  220  m;  Puerto  Maldonado, 
mined  as  H.  wasmanni  var.  isthmica.  Spec-  260m.  PUERTO  RICO,  49  Carr.  186,  km 
imens  labelled  as  type  series  of  H.  emenji  13.2,  Rio  Grande,  Humacao.  VENEZUE- 
recta  were  studied  in  the  USNM  and  LA,  Amazonas:  Alto  Mavaca,  2°01'N 
found  to  be  conspecific  with  striatula  and  65°07'W,  200m;  Tachira:  Las  Cuevas, 
not  with  G.  pleurodon.  I  was  able  to  ex-  7°48'N  71°46'W,  200  m. 
amine  two  G.  striatula  syntypes  in  the 
BMNH.  Bill  Brown  generously  shared  the  Gnamptgenys  strigata  (Norton) 

notes  he  took  during  his  study  of  the  fol-  ,,  ,                .  •    x   m    .      lorr,  a 

..         ^  .     ^r^^^    TT           I     •  Holcoponera  stn^ata  Norton  1871A. 

lowmg  Santschi  types  m  1963:  H.  regularis,  ^,i,,^,,,,,,  conceutnca  Mayr  1870:964;  Emery 

H.  regularis  v.  arcuata,  H.  rustica,  and  H.  1891-167 

wheeleri.  He  concluded  that  they  are  all  the  Holcoponera  simplex  Emery  1896:46;  Kempf  and 

same  species.  He  also  wrote  that  these  Brown  1968:90. 

specimens  plus  additional  types  and  spec-  Holcoponera  satzgeri  Forel  1908:39;  Brown  1957: 

imens  he  studied  (H.  striatula  v.  antillana,  490. 

v.  angustipleura,  H.  wasmanni,  H.  ivasmanni  Holcoponera  simplex  spuria  Forel  1908:39;  Brown 

V.   isthmica.  H.   brasiliensis  v.  calcarata,  v.  1957:490. 

mayri,  H.  curtula  v.  paulina  (det.  Santschi))  Holcoponera    simplex  foreli    Santschi    1929:460; 

perhaps  could  be  separated  into  two  spe-  ^  Brown  1957:490. 

'^ .            •        ,1           1      \-    1                         r>    1.  Gnampto?enus  sniiplex  CEmery);  Brown  195S:229. 

cies  usmg  the  subpetiolar  process.  But  as  „         ,              x       x     /m    /    ^    u            mco 

,        ,     ,        ,                     .       ^,     ,  .  Gnampto^enys  stri'^ata   (Norton);   Brown   1958: 

already  has  been  menhoned,  this  process  339.   ^^^^^   1958:115;    Kempf  and    Brown 

has  proven  to  be  an  unreliable  character  i968:90;  Kempf  1972:115;  Billen  1986:168. 
for  species  determination  in  this  group. 

Material  Examined. — ARGENTINA,  Sal-  Diagnosis. — Roughly  costate;  anterior 

ta:  El  Rey;  Misiones:  Puerto  Iguazu.  BO-  postpetiolar  surface  fairly  flat  and  with  3- 

LIVIA,    Beni:    Blancaflor.    BRAZIL:    Para:  4  transverse  costae;  abundant  long,  sub- 

Tucurul;  Sao  Paulo:   Reserva  Caraguata-  decumbent  and  suberect  hairs  on  body, 

tuba,  40-80  m;  Rio  Grande  do  Sul:  Caixa  Ecology. — A    dweller    of   wet    montane 

do  Sul,  Nossa  Senhora  da  Saude;  Ceara:  transandean  forests,  commonly  taken  in 

Maranganape  Mts.  COLOMBIA,  Meta:  Re-  leaf  litter  samples. 

serva  La  Macarena,  Rio  Guayabero,  270m;  Comments. — Aside  from  aspects  covered 

Quebrada  Chirijara,  Villavicencio,  1400m;  by  Kempf  &  Brown  (1968),  other  charac- 


188 


Journal  of  Hymenoptera  Research 


ters  that  a  vary  are  the  size  of  the  propo- 
deal  spiracle,  costulation  of  the  propodeal 
declivity  and  the  shape  of  the  node. 

Specimens  Examined. — COLOMBIA,  Val- 
le:  Pance,  1700m;  Reserva  Forestal  de  Yo- 
toco.  COSTA  RICA,  Monteverde:  1520m; 
Puntarenas:  1300-1580m;  Alajeula:  Rio 
Penas  Blancas,  800-880m;  Heredia:  17-12 
km  N  Volcan  Barba,  880-1420m;  San  Jose: 
2  km  E  San  Gerardo,  9°27'N  83°43'W, 
1440m.  GUATEMALA,  Sepacuite.  MEXI- 
CO, Chiapas:  Ocosinga;  19  km  NW  Oco- 
zocoautla,  975m;  Veracruz:  5  km  N  Cuil- 
huac. 

Gnamptogenys  sulcata  (Smith) 

Edatomma  sulcatum  F.  Smith,  1858:99. 
Ponera  tomata  Roger  1861:15.  NEW  SYNONY- 
MY. 
Gnamptogenys    lineata    Mayr    1870:965;    Brown 

1958:229.' 
Gnamptogenys  sulcata  var.  cearensis  Forel  1912: 

33;  Brown  1958:229 
Gnamptogenys  sulcata  var.  nitens  Mann  1916:407; 

Brown  1958:229. 
Gnamptogenys  ypirangensis  Borgmeier  1928:229. 
Gnamptogenys  sulcata  bufonum  Weber  1938:208; 

Brown  1958:229. 
Gnamptogenys  sulcata  (Smith);  Brown  1958:329; 

Kempf  192:115;  Lattke  1990:24. 
Gnamptogenys    tomata    (Roger);    Mann    1922:3; 

Brown  1958:319,  329;  Kempf  1968:378;  Kempf 

1972:116;  Lattke  1990:25. 

Diagnosis. — Mandibles  subtriangular; 
dorsal  mesosomal  costulae  convergent 
caudad  and  transverse  on  declivitous  pro- 
podeal face;  metacoxal  tooth  absent,  at 
most  present  as  very  small  tubercle  or 
short  lobe;  very  variable  color:  mesosoma 
black  to  light  brown,  frequently  head,  gas- 
tric apex  and  posterior  margin  of  gastric 
tergum  II  darker  than  rest  of  body.  Legs 
antennae,  and  mandibles  light  to  dark 
brown. 

Ecology. — Used  as  prey  by  Dendrohates 
histrionicus.  See  Lattke  (1990). 

Comments. — Color  is  of  no  help  in  sep- 
arating the  aforementioned  forms,  except 
at  an  occasional  local  level.  Brown  (1958) 
recognized  these  difficulties  and  made  it 


clear  the  situation  would  need  further 
study.  Variation  has  been  observed  in  size 
and  shape  of  the  subpetiolar  process, 
mandibular  width,  and  relative  size  of  the 
eyes.  The  node  can  project  posteriorly  and 
approach  the  condition  in  acuminata  but 
does  not  become  acutely  pointed.  Even 
though  the  propodeal  declivity  is  com- 
monly transversely  costulate,  it  can  some- 
times have  oblique  or  even  longitudinal 
costulae. 

I  examined  the  sulcata  type  in  the 
BMNH  found  it  to  have  transverse  costu- 
lae on  the  posterior  nodal  face.  Roger's  de- 
scription includes  a  key  character  that 
points  to  tornata's  synonymy  with  sulcata: 
"The  petiole,  as  in  rimulosa,  is  concentri- 
cally costulate,  with  the  smallest  circle  or 
oval  in  the  middle  of  the  dorsum,  gradu- 
ally becoming  larger."  This  I  have  inter- 
preted as  meaning  that  the  costulation  on 
the  posterior  face  of  the  petiole  is  trans- 
verse. My  examination  of  Mann's  nitens 
type  reveals  tansverse  costulation  on  the 
posterior  petiolar  face,  longitudinal  cos- 
tulae on  the  declivitous  propodeal  face 
and  very  low  triangular  metacoxal  teeth. 
Borgmeier's  ypirangensis  is  described  with 
transverse  costulae  on  the  posterior  nodal 
face.  Dr.  Ivan  Lobl  of  the  MHNG  kindly 
examined  the  cearensis  type  and  reported 
transverse  costulae  on  the  posterior  nodal 
face.  Dr.  Max  Fischer  of  the  NHMV  gra- 
ciously examined  the  types  of  lineata  and 
reported  transverse  costulation.  What 
could  constitute  a  separate  species  are 
specimens  from  the  Amazonas  Basin 
(MUSP)  that  are  black  with  yellow  man- 
dibles and  are  considerably  larger  than  the 
average  sulcata.  But  in  the  midst  of  so 
much  variability  it  is  prudent  to  await  the 
accumulation  of  additional  evidence  be- 
fore coining  a  name  for  a  few  specimens. 

Specimens  Examined. — BELIZE,  Caves 
Branch.  BOLIVIA,  Rosario.  BRAZIL,  Ama- 
zonas: 24  km  NE  Manaus,  Ig.  Marianil, 
Rio  Branco  Rd.;  Independencia:  Parahyba; 
Para:  Rio  Xingu  (Cachoiera  do  Espelho); 
10  km  N  Tucurul;  Porto  Velho:  Rio  Ma- 


Volume  4,  1995 


189 


67 

Figs.  66-67.     G.  transversa.  66,  frontal  view  of  head;  67,  lateral  view  of  mesosoma  and  petiole. 


deira.  COLOMBIA,  Casanare:  (no  other 
data);  Magdalena:  12  km  ESE  Minca, 
11°08'N  74°06'W,  780m;  Parque  Tayrona, 
Pueblito,  300m;  Meta:  Vista  Hermosa;  S 
Villavicencio,  Cano  El  Buque,  480m;  Na- 
rino:  La  Guayacana;  Sucre,  Est.  Primates 
Inderena;  Valle:  64  km  E  Buanventura, 
570m.  COSTA  RICA,  Sirena:  Peninsula  de 
Osa,  8°28'N  83°35'W,  50m;  Pq.  Nac.  Brau- 
lio  Carrillo,  500m;  Peninsula  de  Osa,  Pq. 
Nac.  Corcovado,  San  Pedrillo,  0-lOOm; 
Puntarenas:  5  km  N  Ciudad  Neily,  780m; 
Heredia,  10°20'N  84°04'W,  500m.  ECUA- 
DOR, Estacion  Rio  Palenque;  Pichincha: 
Tinalandia,  16  km  SE  Santo  Domingo  de 
los  Colorados.  GUATEMALA,  Escuintla. 
HONDURAS,  Ola,  18  km  NE  Catacamas, 
370m.  MEXICO,  Veracruz:  Lago  Catema- 
co,  350m;  Los  Tuxtlas,  10  m  NNW  Sonte- 
comapan,  200m;  trail  above  Presidio,  305- 
490m;  Cordoba.  PANAMA,  Barro  Colora- 
do. PERU,  Tingo  Maria:  Yurac,  107  km  E 
Tingo  Maria;  Madre  de  Dios:  Est.  Biol.  Co- 
cha  Cashu,  400m;  Rio  Tambopata  Res., 
20°50'S  69°20'W,  290m;  Huanuco:  Cueva 
de  la  Lechuzas,  6  km  W  Tingo  Maria  Mar- 
ia; Cueva  de  la  Boca  del  Lobo,  710m,  69 
km  W  Tingo  Maria.  VENEZUELA,  Ama- 
zonas:  Alto  Mavaca,  2°02'N  65°06'W,  250; 
Rio  Baria,  0°50'N  66°10'W,  140m. 

Gnamptogenys  tortuolosa  (Smith) 

Poncra  tortuolosa  F.  Smith  1858:99. 
Gnamptogeui/s   tortuolosa   var.   quitensis   Forel 
1921:133;  Brown  1958:230. 


Ectatoinma  {Gnamptogeriys)  tortuoloswn  (Smith); 

Emery  1896:51;  Mann  1916:406;  Wheeler  and 

Wheeler  1952:134. 
Gnamptogenys  tortuolosa  (Smith);  Brown  1958: 

230;     Kempf    1961:492;     Kempf    1970:325; 

Kempf  1972:116;  Lattke  1990:25. 

Diagnosis. — Mandibles  subtriangular; 
clypeal  lamella  laterally  angular  and  me- 
dianly  straight  to  softly  concave;  propo- 
deum  with  transverse  costulae;  coxal 
tooth  small  sometimes  shaped  as  a  short 
lobe;  tibiae,  femora  and  scapes  smooth 
and  shining.  Piceous  body;  legs  and  an- 
tennae ferruginous. 

Ecology. — Found  nesting  in  rotten  wood 
in  humid  forests  and  foraging  in  grassy  or 
weedy  areas. 

Comments. — Very  constant  in  sculpture 
and  color. 

Specimens  Examined. — BRAZIL,  Rorai- 
ma:  64  km  S  Boa  Vista.  COLOMBIA,  Ama- 
zonas:  Trocha  Buenos  Aires;  Meta:  8  km 
w  Villavicencio;  Mesetas,  La  Uribe,  720m; 
Serrania  La  Macarena.  ECUADOR,  Sucua. 
GUIANA,  Kartabo;  Bartica;  Rupupuni: 
Apoteri,  4°05'N  58°35'W,  100m.  VENE- 
ZUELA, Amazonas:  Alto  Rio  Siapa, 
1°05'N  58°35'W,  600m;  Alto  Rio  Mavaca, 
2°01'N  65°07'W,  200. 

Gnamptogenys  transversa  Lattke, 

new  species 
(Figs.  66,  67) 

Type  Material. — Holotype  worker:  PAN- 
AMA,  Bocas  del  Toro,   Fortuna-Chirigui 


190 


Journal  of  Hymenoptera  Research 


Grande  rd.,  8°47'N  82°12'W,  12/14-VII-78, 
1050m,  D.M.  Olson  (523),  leg.  Premontane 
rainforest  sifted  leaflitter.  Deposited  in 
MCZC. 

Worker. — Holotype  measurements:  HL 
1.30;  HW  1.10;  ML  0.70;  ED  0.28;  SL  1.04; 
WL  1.92  mm;  CI  1.18;  SI  0.95;  OI  0.25. 

Head  in  frontal  view  with  rectangular, 
elongate  head:  sides  broadly  convex,  pos- 
terior margin  slightly  concave;  clypeal  la- 
mella relatively  long,  medianly  slightly 
concave  and  laterally  rounded;  eyes  mod- 
erately prominent.  Costulae  on  head  prin- 
cipally longitudinal,  slightly  convergent 
anterad,  costulae  between  eyes  and  frontal 
lobes  curving  into  depression  laterad  of 
antennal  sclerite.  Mandibles  triangular 
with  blunt  denticles  and  relatively  long 
basal  broader,  rugulose  with  piligerous 
punctae.  Scapes  shining  and  slightly  ru- 
gulose, barely  passing  posterior  edge.  Me- 
sosoma  dorsally  with  longitudinal  costu- 
lae from  pronotum  to  dorsum  of  propo- 
deum,  declivity  with  transverse  costulae; 
promesonotal  suture  visible  as  a  brief 
transverse  depression.  Laterally  with  lon- 
gitudinal costulae  throughout.  Anepister- 
nal  flange  well-developed  along  anterior 
third;  katepisternum  well-defined,  anepis- 
ternum  not  as  well-defined.  Petiole  later- 
ally subcylindrical,  costulae  longitudinal/ 
oblique,  ventral  process  is  an  anteriorly 
projecting  lobe.  Petiole  in  dorsal  view 
wider  posteriorly  than  anteriorly,  costulae 
form  concentric  arches:  transverse  in 
oblique-anterior  view  and  longitudinal  in 
posterior  view.  Gastric  dorsum  with  lon- 
gitudinal costulation,  postpetiolar  sternal 
disc  smooth  and  shining.  Transverse  cos- 
tulae on  forecoxa,  metacoxa  with  a  paral- 
lel-sided denticle  with  a  rounded  apex. 

Antennae,  tibiae,  femora  with  no  pilos- 
ity,  only  decumbent  to  suberect  hairs  and 
sparse  punctulae.  Body  brown,  legs  fer- 
ruginous brown. 

Female,  Male. — Unknown. 

Discussion. — This  species  is  very  close  to 
hartmani  but  hartmani  has  longitudinal  cos- 
tulae on  the  propodeal  declivity;  mostly 


smooth  and  shining  mandibles;  dorsum  of 
petiole  with  mostly  longitudinal  striae; 
longitudinal  striae  on  the  postpetiolar 
sternum;  eyes  not  as  prominent,  and 
scapes  with  more  defined  striae  and  ru- 
gulae. 

Etxjmology. — The  name  alludes  to  the 
transverse  costulae  on  the  propodeal  de- 
clivity. 

Gnamptogenys  triangularis  (Mayr) 

Ectatomma  [Gnamptogenys)   triangularis  Mayr 

1887:544. 
Gnamptogenys  triangularis  (Mayr);  Emery  1905: 

113;  Kusnezov  1954:34;  Brown  1958:323;  Kus- 

nezov  1962:236;  Brown  1958:230  321. 
Ectatomma    (Parectatomma)    triangularis    richteri 

Forel  1913:203;  Luederwalt  1926:236;  Brown 

1958:230  321. 
Ectatomma  (Parectatomma)  aculeaticoxae  Santschi 

1921:82;   Wheeler   and   Wheeler   1952:133. 

NEW  SYNONYMY. 
Gnamptogenys   aculeaticoxae    (Santschi);    Brown 

1958:227,^330;  Kempf  1960:390;  Kempf  1961: 

491;  Kempf  1972:116;  Deyrup,  et.  al.  1989:93; 

Lattke  1990:8. 

Diagnosis. — Promesonotal  suture  weak- 
ly impressed;  node  dorsum  with  trans- 
verse costulae  and  subquadrate  costulae 
and  subquadrate  subpetiolar  process;  first 
gastric  sternum  with  transverse  costulae; 
metacoxal  tooth  long  and  thin.  Piceous 
body. 

Ecology. — Millipede  feeder  found  in  hu- 
mid forests.  The  USA  record  (Deyrup,  et. 
al.  1989:93)  is  undoubtably  a  recent  intro- 
duction and  they  have  apparently  found 
prey  in  the  local  species  of  millipedes,  as 
they  are  well  established. 

Comments. — The  number  of  transverse 
costulae  on  the  petiolar  dorsum  can  vary 
from  8  to  14,  and  those  on  the  pronotum 
from  13  to  23.  Specimens  from  Argentina 
tend  to  have  a  higher  count  but  there  is 
no  gap  separating  the  values.  The  length 
of  the  coxal  teeth  is  variable  and  bears  no 
relation  to  the  number  of  petiolar  costulae. 
Propodeal  teeth  also  show  variation  form 
a   low   mound   to   the  usual   low,   sharp 


Volume  4,  1995 


191 


teeth.  Occasional  specimens  can  have  up 
to  4  transverse  costulae  on  the  anterior 
pronotal  face,  and  rarely  longitudinal  cos- 
tulae on  the  petiolar  node.  Other  traits 
used  by  Santschi  to  separate  aciilenticoxae, 
such  as  degree  of  impression  of  the  pro- 
mesonotal  suture,  gastric  constriction  and 
gauge  of  hairs,  length  vs.  width  of  petiolar 
node  and  the  mandibular  costulation  all 
show  continuous  variation  that  is  best  de- 
scribed as  infraspecific. 

Specimens  Examined. — ARGENTINA,  Tu- 
cuman:  vie.  Horco  Molle,  750-900m;  Tafe- 
cillo;  La  Cavera;  48  km  S  Jujuy.  BOLIVIA, 
Huachi:  Rio  Beni.  BRAZIL,  Espiritu  Santo: 
4  km  W  Santa  Tereza,  Res.  Nova  Lombar- 
dia,  900m;  Parana:  Foz  do  Iguaqu;  Sao  Pau- 
lo: Agudos.  COLOMBIA,  Magdalena:  3  km 
SE  Minca,  ir08  74°06'W,  1050m;  Pq.  Tay- 
rona,  Pueblito,  360m.  COSTA  RICA,  He- 
redia:  La  Selva;  Peninsula  de  Osa,  Pq.  Nac. 
Corcovado,  8°28'N  83°35'W,  0-lOOm.  EC- 
UADOR, Pichincha:  47  km  S  Santo  Domin- 
go, Rio  Palenque,  215m.  GUIANA,  Kamak- 
usa.  PANAMA,  Barro  Colorado.  PERU, 
Tingo  Maria:  Yurac,  108  km  E  Tingo  Maria; 
Puerto  Maldonado,  260  m;  Panguana, 
9°37'S  74°56'W,  220m.  USA,  Florida:  S.  Mi- 
ami. VENEZUELA,  Amazonas:  Alto  Rio 
Mavaca,  2°01'N  65°07'W,  200m. 

Gnamptogenys  volcano  Lattke, 
new  species 
(Fig.  64,  65) 

Type  Material. — Holotype  worker.  COS- 
TA RICA,  Heredia:  18  km  N  Volcan  Barba, 
10°17'N  84°05'W,  800m,  4/14-VII-1986,  J. 
Longino  no.  1383-5.  Wet  forest  litter  sam- 
ple. Deposited  in  LACM.  One  antenna 
lacking. 

Worker. — Holotype  measurements:  HL 
1.38;  ML  0.55;  HW  1.18;  ED  0.25;  SL  1.18; 
WL  1.93  mm;  CI  0.86;  SI  1.00;  OI  0.21. 

Head  in  frontal  view  elongate,  vertexal 
margin  concave;  eyes  prominent  and  situ- 
ated at  mid-length;  clypeal  lamella  relative- 
ly long,  laterally  rounded  and  medianly 
concave;  mandibles  smooth  and  shining, 
with  prominent  piligerous  fossae;  subtrian- 


gular  basal  angle  preceded  by  a  tooth  and 
concavity  (or  notch);  scape  when  laid  back 
barely  surpasses  vertexal  margin;  mesoso- 
mal  dorsum  with  longitudinal  costulae  and 
propodeal  declivity  with  transverse  costu- 
lae, in  dorsal  view  the  costulae  arch  pos- 
terad  and  become  longitudinal;  gaster  with 
longitudinal  costulae,  including  postpetio- 
lar  sternum,  weakly  effaced  anteromedi- 
anly;  metacoxae  with  dorsobasal  tooth, 
dorsally  with  transverse  costulae  and  lat- 
erally mostly  smooth  and  shining;  subpe- 
tiolar  process  with  no  posterior  heel;  tibiae 
and  femora  smooth  and  shining.  Body 
with  numerous  standing  golden  hairs; 
body  brown,  legs  and  antennae  reddish 
brown. 

Queen,  Male. — Unknown. 

Discussion. — Gnamptogenys  volcano  is 
close  to  sulcata  but  the  latter  can  be  recog- 
nized by  the  longitudinal  costulae  on  the 
propdeal  declivity,  broader  head,  rounded 
mandibular  basal  angle  with  no  tooth  and 
the  straight  clypeal  lamella  with  acutely 
angulate  sides.  The  structure  of  the  clypeal 
lamella,  mesepisternum,  and  mandible 
make  this  a  striking  species  within  the  sul- 
cata group. 

Etymologi/. — The  name  of  the  species 
comes  from  the  latin  word  for  volcano,  as 
the  type  locality  is  near  the  Barbas  volcano. 

ACKNOWLEDGEMENTS 

I  gratefully  acknowledge  the  following  indi\'iduals 
for  providing  specimens,  logistical  and  moral  support 
in  field,  lab,  and  museum:  M.L.  Baena,  B.  Bolton,  C. 
Bordon,  C.R.  Brandao,  W.L.  Brown,  F.  Castano  (R.I.P.), 
S.  Cover,  D.  Davis,  E.  de  Bellard,  F.  Fernandez,  M. 
Fischer,  A.  Gentry,  M.  Harrison,  K.  Jaffe,  D.  Kistner, 
Lynda  Lattke,  I.  Lobl,  J.  Longino,  W.P.  MacKay,  A. 
Nunes,  S.  Peck,  R.  Robbins,  E.  Ross,  D.  Smith,  R.  Snell- 
ing,  P.  Ulloa-Chacon,  G.  Umphrey,  and  P.S.  Ward. 

The  following  institutions  are  acknowledged  for 
providing  funds  and  logistical  support:  British  Coun- 
cil, California  Academy  of  Sciences,  Fundacion  para  el 
Desarrollo  de  las  Ciencias  Fisicas,  Matematicas  y  Na- 
turales,  Fundacion  Polar,  Fimdacion  Terramar,  Insti- 
tuto  de  Zoologia  Agricola  (Universidad  Central  de 
Venezuela),  Laboratorio  de  Comportamiento  {Univer- 
sidad Simon  Bolivar)  and  (especially)  the  Smithsonian 
Institution. 


192 


Journal  of  Hymenoptera  Research 


I 


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Arkiv  for  Zoologie  15(7):l-6. 

Wheeler,  W.M.  1930.  A  new  Emeryella  from  Panama. 
Proceedings  of  the  New  England  Zoological  Club  12: 
9-13. 

Wheeler,  W.M.  1936.  Ants  from  Hispaniola  and 
Mona  Island.  Bulletin  of  the  Muscu})i  of  Compara- 
tive Zoology  80:1 93-21 L 

Wheeler,  W.M.'  &  W.M.  Mann.  1914.  The  ants  of  Hai- 
ti. Bulletin  of  the  American  Museum  of  Natural  His- 
tory 33:1-61. 


J.  HYM.  RES. 
Vol.  4,  1995,  pp.  194-203 

Latitudinal  Gradients  in  North  American  Braconid  Wasp  Species 

Richness  and  Biology 

Donald  L.  J.  Quicke  and  Rachel  A.  Kruft 
Department  of  Biology,  Imperial  College  at  Silwood  Park,  Ascot,  Berkshire  SL5  7PY,  U.K. 


Abstract. — Latitudinal  trends  in  the  numbers  of  species  and  genera  of  all  subfamilies  of  Bracon- 
idae  in  America  north  of  Mexico  were  examined  by  compiling  lists  of  published  records  for  states 
(USA)  and  provinces  (Canada)  and  grouping  these  into  approximate  latitudinal  belts.  The  data 
obtained  agree  well  with  past  studies  on  the  species  richness  of  Ichneumonidae  in  North  America 
(Janzen  1981)  with  peaks  in  numbers  of  taxa  occurring  in  the  belt  lying  approximately  between 
37°  and  42°  North.  The  ratio  of  idiobiont  to  koinobiont  genera  and  species  increased  monotonically 
with  decreasing  latitude. 


Most  insect  groups  increase  in  species  using  the  catalogues  of  Townes.  These  cat- 
richness  with  decreasing  latitude  (see  Ste-  alogues  were  based  on  an  enormous 
phens  1989).  However,  there  is  a  body  of  amount  of  work  and  were  therefore  taken 
evidence  that  the  species  richness  and  di-  to  be  relatively  geographically  unbiased, 
versity  of  some  groups  of  insect  parasit-  though  the  possibility  does  exist  that  since 
oids,  whilst  increasing  towards  the  trop-  Townes  himself  did  much  of  his  collecting 
ics,  does  so  at  a  lesser  rate  than  does  the  around  Michigan,  the  taxonomic  coverage 
species  richness  and  diversity  of  herbivo-  oi  that  part  of  the  United  States  may  have 
rous  insects  (Owen  &  Owen  1974;  Gauld  been  relatively  more  complete.  However, 
1986;  Noyes  1989).  Whilst  there  is  good  Janzen's  approach  does  overcome  some  of 
evidence  that  the  species  richness  and  per-  the  difficulties  encountered  when  working 
haps  also  diversity  of  some  groups  of  par-  ^^  tropical  versus  temperate  comparisons 
asitic  Hymenoptera,  such  as  the  Chalci-  ^^  ^^at  sampling  and  taxonomic  effort 
doidea,  increases  towards  the  tropics  (e.g.  ^^*^^^"  *^^  temperate  region  has  been 
Hespenheide  1979),  for  others  it  has  been  comparatively  evenly  distributed  and 
suggested  that  species  richness  may  even  "^°^^  thorough  because  of  the  long  history 

,  ...  ..     1.U        •     ..       •     1  of  collectme  and   revisionary  work  that 

be  greater  m  temperate  than  m  tropical  ,        ,  ^   .    ,  ,  ^^         .  .      , 

.  ,.       J      r        J  .^  1  has  been  carried  out  there,  burprismely, 

regions,  a  trend  referred  to  as  anomalous  ,  r         ,     i  i  i     •      x  t      / 

Janzen    found    that    the    peak    in    North 


diversity  (Owen  &  Owen  1974;  Janzen  & 
Pond  1975;  Janzen  et  al.  1976).  However, 


American  ichneumonid   species   richness 

,                ,     .        1         ,           ,    ,,          ,  occurs  between  38°  and  42°  North,  i.e.  spe- 

these  conclusions  have  been  challenged  on  •         •  ,                          u  c        ^   u     ■ 

^  cies  richness  was  not  found  to  mcrease 

the    grounds   of   sampling    methodology  consistently  with  decreasing  latitude.  The 

(Morrison  et  al.  1978;  Hespenheide  1979),  question  remained,  however,  whether  this 

and  also  because  they  mostly  deal  with  ^^^^^^  ^^^  ^^^^  generally  applicable  to 

just  one  family,  the  Ichneumonidae,  and  ^t^er  groups  of  parasitic  Hymenoptera. 

the  generality  of  these  findings  has  yet  to  indeed  there  is  a  growing  body  of  evi- 

be  demonstrated.  dence  to  suggest  that  different  groups  of 

Janzen  (1981)  examined  the  latitudinal  parasitoids  show  quite  different  trends  in 

distributions  of  members  of  eight  subfam-  their  species  richness  with  respect  to  lati- 

ilies  of  Ichneumonidae  in  North  America  tude  (Gauld  1986;  Askew  1990). 


Volume  4,  1995 


195 


The  present  work  attempts  to  carry  out 
for  the  Braconidae,  a  similar  exercise  to 
Janzen's  (1981)  ichneumonid  study.  The 
Braconidae  are  generally  accepted  as  be- 
ing the  extant  sister  group  of  the  Ichneu- 
monidae  (Sharkey  &  Wahl  1992),  and  col- 
lectively, the  two  families  constitute  the 
superfamily  Ichneumonoidea.  Juillet 
(1964)  obtained  data  from  samples  of  ich- 
neumonoids  caught  in  a  rotary  trap  at  a 
single  site  in  Canada,  which  suggested 
that  members  of  the  Ichneumonidae  'pre- 
fer', and  hence  are  more  diverse  in,  cooler 
and  more  humid  habitats  than  members 
of  the  Braconidae.  Thus,  it  might  be  ex- 
pected that  braconids  will,  as  a  whole,  be 
more  species  rich  at  lower  latitudes  than 
ichneumonids,  and  therefore  might  not 
show  the  same  peak  in  species  richness 
that  Janzen  found  for  Nearctic  Ichneu- 
monidae. Thus,  as  pointed  out  by  Whar- 
ton (1993a),  the  distribution  of  species 
richness  in  the  Braconidae  ought  to  make 
an  interesting  comparison  with  the  Ich- 
neumonidae. 

Catalogues  and  descriptions  of  Nearctic 
Braconidae  usually  provide  data  on  the 
state  or  province  from  which  material  has 
been  recorded  or  described.  We  therefore 
divided  the  states  of  the  USA  and  the  Ca- 
nadian provinces  collectively  into  5 
groups  such  that  each  group  occupied  a 
reasonably  constant  latitudinal  zone  with 
minimal  latitudinal  overlap  between 
groups  (Fig.  1).  The  areas  of  the  five  zones 
are  given  in  Table  2.  Whilst  it  was  not  pos- 
sible to  completely  avoid  overlap,  the  ef- 
fect of  such  overlaps  will  be  to  reduce 
rather  than  enhance  the  likelihood  of  de- 
tecting latitudinal  gradients,  and  conse- 
quently our  results  will  be  conservative. 
The  resolution  of  latitudinal  trends  in  the 
present  study  is  necessarily  somewhat 
coarser  than  that  obtained  by  Janzen 
(1981)  as  information  on  distributions  is 
only  readily  available  in  the  form  of  state 
or  province  records. 

The  primary  sources  of  distributional 
data    were    the    catalogues    of    Shenefelt 


(1969,  1970a,  b,  1972,  1973a,  b,  1974  1975, 
1978),  Shenefelt  &  Marsh  (1976)  and 
Marsh  (1979).  We  then  attempted  to  take 
into  account  many  c^f  the  subsequently 
published  taxonomic  changes  so  as  to  al- 
low for  new  synonymies,  new  generic 
placements,  newly  described  taxa  and 
new  distribution  records  (van  Achterberg 
1977,  1979,  1983,  1985,  1986,  1987,  1988a, 
b;  Capek  &  Achterberg  1992;  Deyrup  1981; 
Haeselbarth  &  Loan  1985;  Huddleston 
1976;  Johnson  1987;  Loan  1979;  Loan  & 
HoUiday  1979;  Marsh  1988,  1989,  1991, 
1993;  Mason  1975,  1976a,  b,  1978,  1979, 
1981,  1987,  1991;  Quicke  &  Kruft  1995; 
Quicke  &  Sharkey  1985;  Riegel  1987;  Ries- 
ke  et  al.  1989;  Saffer  1982;  Sharkey  1985, 
1988;  Sharkey  &  Mason  1986;  Sharkey  & 
Wharton  1985,  1994;  Shaw  1983,  1985, 
1992,  1993;  Stary  &  Marsh  1982;  Wharton 
1977a,  b,  1980,  1983,  1986,  1988,  1993b, 
1994;  Wharton  &  Quicke  1988;  Wharton  et 
al.  1989;  Wheeler  &  Loan  1984;  Whitfield 
1985,  1988a,  b;  Whitfield  &  van  Achter- 
berg 1987;  Whitfield  &  Mason  1994;  Wil- 
liams 1985,  1988).  In  addition,  a  few  un- 
published data  were  included  for  the  fol- 
lowing taxa:  Vipio  (Braconinae)  (Inayatul- 
lah  1992);  Pholetesor  (Microgastrinae)  (J. 
Whitfield,  in  press);  undescribed  genus  of 
Euphorinae  (S.  R.  Shaw,  pers.  comm.).  The 
genus  Celerion  is  excluded  as  its  subfamil- 
ial  status  is  questionable  (Marsh,  pers. 
comm.).  No  data  are  included  for  intro- 
duced species. 

The  treatment  of  subfamilies  largely  fol- 
lows that  of  Quicke  &  Achterberg  (1990) 
except  that  the  Hormiini  and  Meteorini 
are  treated  here  as  separate  subfamilies. 

No  pretence  is  made  that  this  taxonomic 
treatment  is  complete,  but  it  is  hoped  that 
it  is  unbiased  and,  therefore,  that  any  er- 
rors that  may  have  been  included  and  any 
omissions  there  may  be  will  not  affect  the 
results  substantially. 

RESULTS  AND  DISCUSSION 

The  numbers  of  genera  and  species  of 
each  subfamily  recorded  as  occurring  in 


196 


Journal  of  Hymenoptera  Research 


Fig.  1 .     Map  of  North  America  showing  boudaries  of  latihidinal  zones  employed  in  the  present  study. 


each  of  the  five  latitudinal  zones  are  pre- 
sented in  Tables  1  and  2.  The  subfamilies 
are  arranged  in  two  groups,  the  first  being 
idiobionts  (Table  1)  and  the  second  koino- 
bionts  (Table  2).  As  expected,  different 
subfamilies  showed  markedly  different 
latitudinal  trends,  though  nearly  all 
showed  maximum  generic  and  specific 
representation  in  our  latitudinal  zone  IV 


which  corresponds  approximately  to  the 
latitudinal  range  from  37°  and  42°  North. 
None  of  the  larger  subfamilies  was  found 
to  be  most  speciose  in  zone  V.  Four  of 
these,  the  Braconinae,  Microgastrinae,  Ma- 
crocentrinae  and  Opiinae  were  most  di- 
verse in  zone  III.  Our  data  for  the  Alysi- 
inae,  usually  considered  a  more  temperate 
and  northern  group,  suggest  that  within 


Volume  4,  1995 


197 


Table  1.    Distributions  of  genera  and  species  of  braconid  subfamilies  with  respect  to  latitudinal  zones  (see 
Fig.  1).  Idiobiont  taxa 


Sublamilies 

Histcro- 

Rhyssalinao 

iTiLTinao 

Doryctin.ie 

Braconinae 

l-ADthcLinaL' 

Hormimae 

Total 

Genera 

Zone  I 

0 

0 

2 

3 

2 

1 

8 

Zone  II 

2 

1 

8 

9 

3 

2 

25 

Zone  III 

1 

1 

16 

8 

4 

4 

34 

Zone  IV 

2 

1 

20 

7 

3 

6 

39 

Zone  V 

1 

1 

28 

10 

2 

8 

50 

Species 

Zone  I 

0 

0 

2 

4 

3 

1 

10 

Zone  II 

2 

1 

35 

67 

4 

3 

112 

Zone  III 

5 

1 

72 

108 

5 

5 

196 

Zone  IV 

4 

1 

111 

104 

3 

12 

235 

Zone  V 

1 

1 

105 

86 

2 

11 

206 

North  America,  it  is  at  its  richest  in  the 
middle  latitudes  of  the  U.S.A.  (zone  IV)  as 
with  most  other  subfamilies. 

Table  3  provides  a  comparison  of  the 
relative  latitudinal  distributions  of  braco- 
nid genera  and  species  according  to  zone 
and  in  terms  of  taxa  per  unit  area.  Addi- 
tionally, it  shows  the  total  numbers  of 
idiobiont  and  koinobiont  taxa  and  the  ra- 
tios of  numbers  of  koinobiont  to  idiobiont 
taxa  in  each  latitudinal  zone.  The  latter 
show  consistent  trends  for  both  genera 
and  species,  with  the  ratio  of  koinobionts 
to  idiobionts  decreasing  with  decreasing 
latitude. 

It  has  long  been  recognised  that  some 
subfamilies  of  Braconidae  are  most  species 
rich  in  the  temperate  region  whilst  others 
are  tropicocentric.  For  example,  the  Alysi- 
inae,  which  are  entirely  koinobiont  endo- 
parasitoids  of  Diptera,  are  a  principally 
northern  group,  and  whilst  there  are  many 
undescribed  tropical  members  of  the  genus 
Asobam,  a  group  that  attacks  Drosophili- 
dae,  more  detailed  knowledge  of  these  is 
not  likely  to  change  our  concept  of  the  sub- 
families overall  distribution  to  a  major  de- 
gree. In  contrast,  the  Agathidinae  and  Car- 
diochilinae,  two  groups  of  koinobiont  en- 
doparasitoids  of  Lepidoptera  larvae,  are 
both  most  speciose  in  the  tropical  region 


(Sharkey  1992;  Huddleston  &  Walker 
1988),  and  the  latter  in  particular  is  rather 
species  poor  in  temperate  latitudes.  Some 
groups  are  restricted  in  latitudinal  range  by 
the  distributions  of  their  hosts.  For  exam- 
ple, members  of  the  ichneumonid  subfam- 
ily Ctenopelmatinae,  most  tryphonine  ich- 
neumonids,  and  the  braconid  tribes  Ich- 
neutini  and  Proteropini  (both  usually  treat- 
ed as  belonging  to  the  Ichneutinae;  see 
Sharkey  &  Wharton  1994)  are  all  endopar- 
asitoids  of  sawflies,  and  since  sawflies  are 
most  abundant  in  temperate  regions  they 
are  likely  to  support  a  higher  species  rich- 
ness of  parasitoids  there. 

In  the  case  of  the  Braconidae,  life  history 
strategy,  idiobioncy  and  koinobioncy 
(Askew  &  Shaw  1986),  is  very  tightly 
linked  with  whether  the  wasps  develop  as 
ecto-  or  endoparasites  respectively  (Quicke 
&  Achterberg  1990;  Shaw  &  Huddleston 
1991).  It  is  not  possible  therefore,  to  dis- 
cern which  of  these  life  history  features  is 
most  responsible  for  the  observed  trends. 
Further,  as  biology  is  usually  consistent 
within  subfamilies  within  the  Braconidae 
(Shaw  &  Huddleston  1991),  any  latitudi- 
nal trends  in  the  ratios  of  idiobiont  to  ko- 
inobiont taxa  will  necessarily  involve  the 
differential  occurrence  or  representation 
of  subfamilies. 


198 


Journal  of  Hymenoptera  Research 


Table  2.    Distributions  of  genera  and  species  of  braconid  subfamilies  with  respect  to  latitudinal  zones  (see 
Fig.  1).  Koinobiont  subfamilies 


Subfamilies 

<u 

<U 

01 

c 

(C 

« 

.      01 
C 

o 

T3 
O 

s 

C 

01 

c 
'S. 

01 

c 

OJ 

s 

I 

■a 

3 

OJ 

0) 
nj 
C 

■5. 

0 
X, 
I- 

Oi 

"3 
-a 

to 
c 

X. 

T3 

C 
I1 

c 
'G 

c 

u 

0 

c2 

o 

O 

< 

< 

"^ 

D 

< 

U 

5 

2 

u 

Genera 

Zone  I 

2 

1 

2 

13 

6 

1 

0 

0 

3 

0 

0 

0 

Zone  II 

4 

1 

6 

26 

16 

3 

0 

0 

5 

0 

1 

1 

Zone  III 

7 

1 

7 

28 

13 

3 

1 

2 

5 

0 

1 

1 

Zone  IV 

7 

1 

6 

30 

12 

4 

1 

2 

5 

0 

1 

1 

Zone  V 

7 

2 

7 

18 

13 

5 

1 

2 

5 

1 

1 

1 

Species 

Zone  I 

3 

2 

14 

73 

10 

3 

0 

0 

10 

0 

0 

0 

Zone  II 

14 

1 

64 

77 

57 

4 

0 

0 

55 

0 

3 

3 

Zone  III 

33 

2 

147 

131 

54 

6 

1 

2 

77 

0 

4 

10 

Zone  IV 

36 

4 

111 

158 

70 

6 

1 

4 

98 

0 

5 

26 

Zone  V 

31 

5 

122 

55 

34 

7 

1 

2 

50 

1 

5 

24 

Whilst  straightforward  taxonomic  ef- 
fects are  undoubtedly  involved  in  deter- 
mining the  distributions  of  some  groups, 
there  is  a  growing  evidence  that  trends  in 
species  richness  and  diversity  may  be  cor- 
related with  life  history  strategy.  In  partic- 
ular, idiobionts  whose  hosts  by  definition 
do  not  develop  further  following  initial 
parasitization  (Askew  &  Shaw  1986)  ap- 
pear to  be  more  species  rich  at  lower  lat- 
itudes than  parasitoid  taxa  that  display 
the  opposite  strategy,  the  koinobionts. 

Hawkins  et  al.  (1992)  in  an  attempt  to 
understand  better  the  latitudinal  trends  in 
parasitoid  diversity  in  North  America, 
took  a  different  approach  in  that  they  con- 
centrated on  parasitoid  assemblage  size 
per  host  rather  than  on  the  numbers  of 
species  in  random  field  samples.  Their 
data  showed  that  latitudinal  trends  in  spe- 
cies richness  are  influenced  not  only  by 
whether  the  parasitoids  display  idiobiont 
or  koinobiont  life  history  strategies  but 
also  by  host  feeding  niche.  In  particular, 
they  found  that  the  diversity  of  parasit- 
oids of  exophytic  hosts  decreased  with  de- 
creasing latitude  but  that  the  diversity  of 


those  attacking  endophytic  hosts  in- 
creased. 

Several  hypotheses  have  been  proposed 
to  account  for  declines  in  the  species  rich- 
ness of  some  parasitoid  groups  with  de- 
creasing latitude  and  for  trends  in  the  ra- 
tios of  idiobionts  to  koinobionts.  These  in- 
clude Gauld  et  al.'s  (1992)  "nasty"  host 
hypothesis,  resource  fragmentation  hy- 
potheses (Janzen  &  Pond  1975;  Janzen 
1981),  interphyletic  competition  (Eggleton 
&  Gaston  1990),  parasitoid  predation  ef- 
fects (Gauld  1987),  host  predation  effects 
(Rathcke  &  Price  1976),  and  limited  spe- 
ciation  opportunity  as  a  result  of  sup- 
posed low  host  recruitment  potential 
(Shaw  1994).  These  are  not  dealt  with  in 
detail  here  as  they  are  reviewed  in  detail 
by  Hawkins  (1994). 

Our  data  show  that  overall,  the  latitu- 
dinal variation  in  species  richness  in 
North  American  Braconidae  closely  re- 
sembles that  found  by  Janzen  (1981)  for 
Ichneumonidae,  with  a  peak  near  the  cen- 
tral latitudes  of  the  USA  and  with  a  dis- 
tinct decline  in  the  numbers  of  species  and 
genera    occurring    in    the    southernmost 


Volume  4,  1995 


199 


Table  2.    Extended 


Subtamilif 


a. 

3 


< 


'~j 


'J. 
■T. 


11 

6 

2 

0 

2 

1 

1 

0 

1 

1 

1 

2 

0 

0 

58 

22 

14 

2 

2 

6 

1 

1 

1 

2 

2 

12 

2 

0 

1 

131 

23 

17 

2 

2 

5 

1 

1 

1 

2 

2 

13 

2 

1 

1 

143 

21 

16 

2 

2 

7 

2 

1 

1 

1 

2 

15 

1 

0 

1 

143 

10 

15 

2 

2 

8 

2 

1 

1 

2 

2 

10 

2 

1 

1 

123 

18 

6 

7 

0 

2 

7 

1 

0 

1 

2 

2 

2 

6 

0 

0 

171 

122 

61 

25 

3 

25 

31 

1 

4 

4 

2 

25 

25 

17 

0 

1 

634 

182 

41 

30 

9 

53 

49 

1 

5 

4 

1 

36 

57 

14 

1 

1 

961 

169 

43 

32 

8 

61 

55 

1 

8 

3 

2 

32 

70 

13 

0 

1 

1025 

125 

33 

24 

5 

51 

34 

1 

8 

6 

2 

23 

58 

11 

1 

1 

725 

States.  We  have  no  evidence  that  sampling 
in  the  southern  states  has  been  signifi- 
cantly less  intense  than  in  more  northerly 
ones  though  the  possibility  cannot  be  ex- 
cluded. Further,  it  should  be  noted  that  in 
all  five  zones  some  states  and  provinces 
have  been  collected  in  much  more  inten- 
sively than  others.  California,  a  state  that 
has  been  extensively  collected,  was   in- 


cluded in  our  Zone  IV,  though  it  could 
equally  justifiably  have  been  included  in 
Zone  V  as  it  straddles  the  boundary  lati- 
tude more  or  less  evenly.  Reanalysis  of  the 
data  set  with  California  transferred  to 
Zone  V  does  not  reduce  the  peaks  in  spe- 
cies and  generic  richness  in  zone  IV  to  any 
great  extent. 

Extrapolation  of  the  observed  trend  in 


Table  3.    Summary  of  latitudinal  distributions  of  genera  and  species  of  Braconidae  in  North  America  and 
ratios  of  koinobiont  to  idiobiont  taxa 


Total  no. 
of  taxa 

Area 

(10-  km-') 

Total  taxa 
per  10-  km= 

No.  lit 

idiobiont 

taxa 

No.  of 

koinobiont 

taxa 

Koinobiont/ 
Idiobiont 

Genera 

Zone  I 

66 

5.19 

12.3 

8 

58 

7.25 

Zone  II 

156 

5.76 

26.9 

25 

131 

5.20 

Zone  III 

177 

2.62 

66.9 

34 

143 

4.17 

Zone  IV 

182 

2.57 

70.0 

39 

143 

3.64 

Zone  V 

172 

2.63 

63.8 

50 

122 

2.42 

Species 

Zone  I 

181 

5.19 

34.1 

10 

171 

17.10 

Zone  II 

746 

5.76 

127.4 

112 

634 

5.66 

Zone  III 

1157 

2.62 

437.4 

196 

961 

4.90 

Zone  IV 

1260 

2.57 

486.8 

235 

1025 

4.36 

Zone  V 

930 

2.63 

351.3 

206 

724 

3.51 

200 


Journal  of  Hymenoptera  Research 


species  richness  in  North  America  to  more 
general,  temperate /tropical  species  rich- 
ness and  diversity  questions  would  be  un- 
wise. One  reason  for  caution  in  this  re- 
spect is  that  a  substantial  part  of  the 
southern  U.S.A.  and  northern  Mexico  is 
markedly  arid,  and  this  may  restrict  par- 
asitoid  species  richness  as  it  appears  to  do 
in  North  Africa  (unpubl.  data).  Further, 
even  in  areas  of  higher  annual  rainfall, 
ichneumonoids  may  still  be  limited  by 
water  availability  since  it  has  been  pro- 
posed that  many  may  need  to  drink  on  a 
daily  basis  and  rains  are  typically  less  reg- 
ular (Townes  1971).  Townes  suggested 
that  dew  formation  may  be  equally  if  not 
more  important  than  rainfall  per  se  in  reg- 
ulating numbers  and  species  richness  of 
ichneumonoids  in  warmer  climates,  and 
that  it  is  for  this  reason  that  altitude  is  an 
important  factor  in  determining  ichneu- 
monoid  species  richness  in  the  tropics, 
dew  forming  more  regularly  with  increas- 
ing altitude.  In  the  case  of  the  Southern 
U.S.A.,  much  of  the  less  arid  south  is  at 
low  altitude  and  therefore  may  experience 
dew  less  regularly  than  required  by  many 
species  of  parasitoids.  However,  interpre- 
tation of  dew  and  rainfall  data  is  likely  to 
be  far  from  a  simple  matter  of  examining 
the  mean  annual  number  of  dew  days  for 
particular  states,  and  if  water  availability 
is  an  important  factor  in  limiting  ichneu- 
monoid  species  richness  in  the  more 
southern  states,  then  rainfall,  season,  tem- 
perature and  dew  frequency  are  likely  to 
be  involved  together  in  a  complex  inter- 
action. Further,  other  important  sources  of 
water  for  many  insects  of  arid  region  are 
nectar  and  honeydew,  and  availability  of 
these,  and  their  utilization  by  ichneumo- 
noids, may  also  need  to  be  considered. 

ACKNOWLEDGEMENTS 

This  work  was  funded  by  the  NERC  Initiative  in 
Taxonomy.  We  would  like  to  thank  Brad  Hawkins 
(Imperial  College)  for  helpful  discussions  and  com- 
ments on  an  earlier  version  of  the  manuscript. 


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the  generic  relationships  within  the  tribe.  Ento- 
mology  88:  1-112. 

Wharton,  R.A.  1983.  New  species  of  lllidops  and  Bra- 
con  (Hymenoptera:  Braconidae)  of  potential  use 
in  biological  control.  Canadian  Entomologist  115: 
667-672. 

Wharton,  R.A.  1986.  The  braconid  genus  Alysia  (Hy- 
menoptera): a  description  of  the  subgenera  and 
a  revision  of  the  subgenus  Alysia.  Systematic  En- 
tomology 11:  453-504. 

Wharton,  R.A.  1988.  The  braconid  genus  Alysia  (Hy- 
menoptera): A  revision  of  the  subgenus  Anarcha. 
Contributions  of  the  American  E)itomological  Insti- 
tute 25:  1-69. 

Wharton,  R.  A.  1993a.  Bionomics  of  the  Braconidae. 
Annual  Revieio  of  Entomology  38:  121-143. 

Wharton,  R.  A.  1993b.  Review  of  the  Hormiini  (Hy- 


menoptera: Braconidae)  with  a  description  of 
new  taxa.  Journal  of  Natural  History  27:  107-171. 

Wharton,  R.  A.  1994.  New  genera,  species  and  re- 
cords of  New  World  Alysiinae  (Hymenoptera: 
Braconidae).  Proceedings  of  the  Entomological  So- 
ciety of  Washington  96:  630-664. 

Wharton,  R.A.,  and  D.L.J.  Quicke.  1988.  A  new  spe- 
cies of  Bracon  (Hymenoptera:  Braconidae)  para- 
sitic on  Eoreuma  loftini  (Dyar)  (Lepidoptera:  Py- 
ralidae).  Proceedings  of  the  Entomological  Society  of 
Washington  90:  288-293. 

Wharton,  R.A.,  J.  W.  Jr.  Smith,  D.L.J.  Quicke  and  H. 
W.  Browning.  1989.  Two  new  species  of  Digon- 
ogastra  Viereck  (Hymenoptera:  Braconidae)  par- 
asitic on  Neotropical  pyralid  borers  (Lepidop- 
tera) in  maize,  sorghum  and  sugarcane.  Bulletin 
of  Entomological  Research  79:  401-410. 

Wheeler,  A.  G.'jr,  and  C.  C.  Loan,  1984.  Peristenus 
henryi  (Hymenoptera:  Braconidae,  Euphorinae)  a 
new  species  parasitic  on  the  honeylocust  plant 
bug,  Diaphnocoris  chlorionis  (Hemiptera:  Miridae). 
Proceedings  of  the  Entomological  Society  of  Washing- 
ton 86:  669-672. 

Whitfield,  J.B.  1985.  The  Nearctic  species  of  Deuterix- 
ys  Mason  (Hymenoptera:  Braconidae).  Pan-Pacific 
Entomologist  61:  60-67. 

Whitfield,  J.B.  1988a.  Revision  of  the  Neactic  species 
of  the  genus  Stiropius  Cameron  {  =  Bucculatriplex 
Auct.)  with  the  description  of  a  new  related  ge- 
nus (Hymenoptera:  Braconidae).  Systematic  En- 
tomology 13:  373-385. 

Whitfield,  J.B.  1988b.  Two  new  species  of  Paradelius 
(Hymenoptera:  Braconidae)  from  North  America 
with  biological  notes.  Pan-Pacific  Entomologist  64: 
313-319. 

Whitfield,  J.B.  and  C.  van  Achterberg.  1987.  Clarifi- 
cation of  the  taxonomic  status  of  the  genera  Can- 
tJiaroctonus  Viereck,  Noserus  Foerster  and  Pseu- 
davga  Tobias  (Hymenoptera:  Braconidae).  Sys- 
tematic Entomology  12:  509-518. 

Whitfield,  J.  B.  and  W.  R.  M.  Mason.  1994.  Mende- 
sellinae,  a  new  subfamily  of  braconid  wasps  (Hy- 
menoptera, Braconidae)  with  a  review  of  rela- 
tionships within  the  microgastroid  assemblage. 
Systematic  Etitomology  19:  61-76. 

Williams,  D.  J.  M.  1985.'  The  New  World  genus  Lath- 
rapanteles  n.  gen.:  Phylogeny  and  placement  in 
Microgastrinae  (Hymenoptera:  Braconinae:  Co- 
tesiini).  Canadian  Journal  of  Zoology  63:  1962-1981. 

Williams,  D.  J.  M.  1988.  Classification,  phylogeny 
and  zoogeograhic  studies  of  species  of  Sathon 
Mason  (Hymenoptera:  Braconidae).  Quaestiones 
Entomologicae  24:  529-639. 

Williamson,  M.  1988.  Relationship  of  species  number 
to  area,  distance  and  other  variables.  Pages  91- 
115.  in  A. A.  Myers  and  P.S.  Giller  eds  Analytical 
Biogeography.  Chapman  and  Hall,  London. 


J.  HYM.  RES. 
Vol.  4,  1995,  pp.  204-284 

Revision  of  the  World  Genera  of  the  Subtribe  Stigmina 
(Hymenoptera:  Apoidea:  Sphecidae:  Pemphredoninae),  Part  1 

Albert  T.  Finnamore 

Curator,  Invertebrate  Zoology,  Provincial  Museum  of  Alberta,  12845 — 102  Avenue, 

Edmonton,  Alberta  CANADA  T5N  0M6 


Abstract. — The  following  new  genera  are  described:  Araucastigmus,  Aykhustigmus,  Incastigmus, 
Llaqhastigmus,  and  Tziistigmus.  The  following  new  species  are  described:  Araucastigmus  masneri, 
yanillus;  Aykhustigmus  fritzi,  patanaivi,  phasti,  luarawa;  Incastigmus  inti;  Llaqhastigmus  ambiguus,  aus- 
tralis,  chutiyana,  colomhianus,  ecuatorialis,  jatunkirus,  llutanis,  mantanti,  muthus,  nigricollaris,  santan- 
deranus,  sapanis,  shachus,  sharkeyi;  Parastigmus  huecuvus,  nina;  Tzustigmus  khmer,  syam,  veda,  and 
wuming.  New  synonymy:  Stigmus  smithii  Ashmead  is  placed  under  Incastigmus  tlioracicus  (Ash- 
mead).  A  key  to  world  genera  is  presented,  and  history,  biology,  generic  relationships,  and  bio- 
geography  are  discussed.  Species  of  Araucastigmus,  Aykhustigmus,  Llaqhastigmus  and  Parastigmus 
are  keyed  and  described,  and  their  distributions  are  presented. 


INTRODUCTION 

This  contribution  is  intended  as  the  first 
in  a  series  of  papers  dealing  with  the 
world  species  of  the  Stigmina.  Members  of 
the  Stigmina  are  small,  generally  3  to  5mm 
in  length,  and  black  or  red  and  black  in 
coloration.  The  group  contains  an  estimat- 
ed 150  species  distributed  in  all  areas  ex- 
cept the  polar  regions  but  for  the  most 
part  is  found  throughout  the  southern 
hemisphere  tropics.  This  first  paper  de- 
scribes five  new  genera,  Tzustigmus,  Ar- 
aucastigmus, Aykhustigmus,  Incastigmus 
and  Llaqhastigmus,  treats  the  species  of  the 
smaller  genera,  reviews  biology,  taxonom- 
ic  history,  and  presents  an  analysis  of  ge- 
neric phylogeny  before  considering  the 
biogeography  of  the  Stigmina.  Future  pa- 
pers will  deal  with  the  species  of  Incastig- 
mus, Stigmus,  and  Carinostigmus. 

Morphological  Terms  And  Symbols. — Ter- 
minology generally  follows  Bohart  and 
Menke  (1976)  but  in  some  cases  terms 
need  clarification  and  some  new  terms  are 
introduced.  They  are  listed  below: 

Appressed  setae:  setae  forming  an  angle 
close  to  0°  with  the  body  surface. 


Carinate:  a  series  of  relatively  large  ridges 
(see  microcarinate  below)  that  do  not 
impart  a  dull  appearance  to  the  body. 

Lateral  sphere  of  propodeum:  area  of  pro- 
podeum  curving  between  propodeal  en- 
closure and  side. 

LCD:  maximum  diameter  of  lateral  ocellus. 

Mesosoma:  the  thorax  plus  the  propo- 
deum. 

Metasoma:  the  apparent  abdomen  con- 
sisting of  the  abdomen  excluding  the 
first  segment  or  propodeum. 

Micropore  field  (Fig.  32):  a  grouping  of 
very  small  pores  usually  visible  only  by 
scanning  electron  microscope,  but  by 
stereomicroscope  apparent  as  a  discrete 
microsculpture  patch. 

Microcarinate:  a  series  of  parallel  ridges 
so  fine  as  to  impart  a  dull  appearance 
to  the  body. 

Microsculpture:  minute  sculpture  impart- 
ing a  dull  appearance  to  the  body. 

GOD:  least  distance  between  lateral  ocel- 
lus and  eye. 

Preomaular  area:  =  preomaulal  area  of 
Bohart  and  Menke  (1976). 

Sternum:  metasomal  sternum. 

Tergum:  metasomal  tergum. 


Volume  4,  1995 


205 


CARN 


CAS 


FSCA 


IIES 


Transscutellar  sulcus:  the  anterior  trans-      BMNH 
verse  sulcus  of  the  scutellum,  immedi- 
ately posterior  to  the  transscutal  articu- 
lation. BRD 

Transverse  sulcus:  on  pronotal  dorsum, 
the  transverse  sulcus  immediately  pos- 
terior to  the  transverse  carina. 

*:  holotype  examined. 

Species  Treatments. — Descriptions  of  all 
included  species  are  provided.  Descrip- 
tions of  new  species  are  based  on  all  the 
material  examined.  In  species  demonstrat- 
ing variability  descriptions  are  based  on 
representatives  of  the  most  prevalent  phe- 
notype  with  variation  noted  throughout 
the  description.  Collection  data  for  the  pri- 
mary type  material  are  presented  exactly 
as  they  appear  on  the  label,  thus  several 
spellings  for  the  same  locality  and  collec- 
tor, and  several  formats  for  date  of  collec- 
tion will  be  encountered.  Collection  data 
for  secondary  type  specimens  are  present- 
ed in  a  standardized  format  organized  by 
country  and  state  or  province.  In  previ- 
ously described  species  the  Material  Ex- 
amined section  lists  only  localities  of  col- 
lection. 

Sources  Of  Material. — Over  10,000  speci- 
mens from  47  institutions  were  amassed 
for  this  study.  The  following  institutions 
have  provided  material  used  in  this  paper 
or  loaned  types.  The  abbreviation  preced- 
ing the  institution  is  that  used  in  the  text 
to  designate  type  repositories. 

AEI     American    Entomological    Insti- 
tute, 3005  sw  56th  Ave.  Gaines- 
ville, FLORIDA  32608  USA.  V.K. 
Gupta,  the  late  H.K.  Townes,  D.      MCSN 
Wahl. 

ANIC     Australian  National  Insect  Col- 
lection, CSIRO,  Division  of  En- 
tomology, GPO  Box  1700  Can-         MCZ 
berra,  ACT  2601,  AUSTRALIA. 
I.D.  Naumann. 

BISH     Bishop    Museum,    1525    Bernice 
Street,  P.O.  Box  19000-A  Hono- 
lulu, HAWAII  96819  USA.  G.M.       MPEG 
Nishida. 


LILLO 


MACN 


The  Natural  History  Museum, 
Cromwell  Road,  London,  SW7 
5BD  ENGLAND.  C.R.  Vardy. 
Biosystematics  Research  Divi- 
sion, CLBRR,  Agriculture  Cana- 
da, Research  Branch,  K.W.  Neat- 
by  Building,  Ottawa,  Ontario, 
CANADA  KIA  0C6.  L.  Masner. 
Carnegie  Museum,  4400  Forbes 
Avenue,  Pittsburgh,  PENNSYL- 
VANIA 15213  USA.  J.E.  Raw- 
lings. 

California  Academy  of  Sciences, 
Golden  Gate  Park,  San  Francis- 
co, CALIFORNIA  94118  USA. 
W.J.  Pulawski. 

Florida  State  Collection  of  Ar- 
thropods, Division  of  Plant  In- 
dustry, Department  of  Agricul- 
ture, Gainesville,  FLORIDA 
32601  USA.  L.A.  Stange. 
Consejo  Nacional  de  Investiga- 
ciones  Cientificas  y  Tecnicas,  In- 
stitute de  Investigaciones  Ento- 
mologicas  Salta  (INESALT),  9  de 
Julio  14-Casilla  de  Correo  3,  4405 
Rosario  de  Lerma,  Salta,  AR- 
GENTINA. M.A.  Fritz. 
Fundacion  Miguel  Lillo,  Institu- 
te de  Zoologia  Miguel  Lillo  251, 
4000  San  Miguel  de  Tucuman, 
ARGENTINA.  A.  Willink. 
Museo  Argentine  de  Ciencias 
Naturales,  Seccion  Entomologia, 
Angel  Gallardo  470,  Casilla  de 
Correo  220,  Sucursal  5,  1405 
Buenos  Aires,  ARGENTINA.  A. 
Roig  Alsina. 

Museo  Civico  Di  Storia  Naturale 
"Giacomo  Doria"  1-16121  Gene- 
va, via  Brigata  Liguria,  N.  9,  IT- 
ALY. V.  Raineri. 
Museum  of  Comparative  Zoolo- 
gy, The  Agassiz  Museum,  Har- 
vard University,  Cambridge, 
MASSACHUSETTS  02138  USA. 
C.  Vogt. 

Museu  Paraense  Emilio  Goeldi, 
Av.  Magalhaes  Barata,  376 — C.P. 


206 


Journal  of  Hymenoptera  Research 


399,  CEP  66000-Belem,  Para, 
BRASIL.  W.L.  Overal. 
NMW  Naturhistorisches  Museum  Wien, 
2.  Zoologische  Abteilung  (Insek- 
ten),  A-1014  Wien,  Burgring  7, 
oSTERREICH  (Austria).  M.  Fi- 
scher. 

OSU  Oregon  State  University,  Depart- 
ment of  Entomology,  Corvallis, 
OREGON  97331  USA.  G.R.  Fer- 
guson. 

PMA  Provincial  Museum  of  Alberta, 
12845-102  Avenue,  Edmonton, 
Alberta,  CANADA  T5N  0M6. 

RNH  Rijksmuseum  van  Natuurlijke 
Historie,  Raamsteeg  2,  Leiden, 
NETHERLANDS.  K.  van  Ach- 
terberg. 

SEM  Snow  Entomological  Museum, 
University  of  Kansas,  Lawrence, 
KANSAS  66045  USA.  R.W. 
Brooks. 

TMA  Termeszettudomanyi  Muzeum 
Allattara,  H-1088  Budapest,  Bar- 
oss  u.  13  HUNGARY.  J.  Papp. 

UCV  Instituto  de  Zoologia  Agricola, 
Universidad  Central  de  Vene- 
zuela, Maracay  2101-A,  VENE- 
ZUELA. J.  Luis  Garcia. 
USNM  Systematic  Entomology  Labora- 
tory, USDA  %  U.S.  National 
Museum,  NHB  168,  Washing- 
ton, D.C  20560  U.S.A.  A.S.  Men- 
ke. 
ZIN  Insect  Systematics  Laboratory, 
Zoological  Institute,  Russian 
Academy  of  Sciences,  Sankt-Pe- 
terburg  199034  RUSSIAN  FED- 
ERATION. 

ZMD  Zoologisk  Museum,  Department 
of  Entomology,  Universitetspar- 
ken  15,  DK  2100  Kobenhavn, 
DANMARK  (Denmark).  O. 
Lomholdt. 
ZMMU  Zoological  Museum  of  the  Mos- 
cow State  University,  Herzen 
Street  6,  Moscow  103009  RUS- 
SIAN FEDERATION.  A.V.  An- 
tropov. 


HISTORY 

G.W.F.  Panzer  (1804)  described  the  ge- 
nus Stigmus  for  a  single  species,  Stigmus 
pendiilus  from  Europe.  Species  description 
in  the  Stigmina  accumulated  more  or  less 
ad  hoc  until  Kohl  (1890)  produced  the  first 
world  synthesis  of  Stigmus  and  related 
genera.  His  revision  of  the  Pempliredon  ge- 
nus group  provided  a  sound  generic 
grouping  that  is  still,  a  century  later,  the 
basis  of  modern  classification.  Over  the 
period  of  1890-1911,  24  species  were  de- 
scribed by  such  notable  authorities  as 
Kohl  (1892,  1905),  Cameron  (1891),  Fox 
(1892,  1897),  Ashmead  (1900),  Mantero 
(1901),  Rohwer  (1911)  and  Strand  (1911). 
During  this  period  Turner  (1907)  de- 
scribed the  genus  Paracrabro  from  Austra- 
lia. 

By  1920  most  of  the  descriptive  work 
had  shifted  to  fauna  of  the  southern  hemi- 
sphere culminating  with  Tsuneki's  (1954) 
description  of  a  new  subgenus,  Carinostig- 
miis,  accompanying  a  revision  of  the  Asian 
and  European  Stigmus.  In  subsequent  pa- 
pers Tsuneki  (1963,  1966,  1971)  went  on  to 
describe  numerous  species  and  subspecies 
in  the  Oriental  Region  while  Leclercq 
(1959)  described  species  from  the  Afro- 
tropical  Region.  The  Nearctic  fauna  was 
revised  by  Krombein  (1973)  at  which  time 
he  described  a  new  subgenus  Atopostig- 
mus.  Work  on  all  groups  of  sphecoid 
wasps  reached  a  high  point  when  Bohart 
and  Menke  (1976)  published  a  world  ge- 
neric revision,  a  landmark  synthesis  un- 
precedented in  the  history  of  the  group. 
That  work  delineated  the  subtribe  Stig- 
mina to  which  the  following  7  genera 
were  assigned:  Arpnctophilus,  Paracrabro, 
Stigmus,  Carinostigmus,  Spilomena,  Micro- 
stigmus,  and  Xysma.  Subsequently  Krom- 
bein (1984)  described  4  new  species  of  Car- 
inostigmus and  a  new  subgenus  Perissostig- 
mus  from  Sri  Lanka.  Budrys  (1987)  de- 
scribed new  species  from  the  former 
Soviet  far  east.  Antropov  (1992)  described 


Volume  4,  1995  207 

the  genus  Parastigmus  from  the  Neotropi-  pith.  Cells  are  mass  provisioned  or  possi- 

cal  Region.  bly  progressively  provisioned  with  12-30 

Among  the  most  important  publications  aphids  depending  on  prey  size  and  spe- 

influencing  the  classification  of  the  Stig-  cies  of  wasp.  Progressive  provisioning  cre- 

mina  was  Menke's  (1989)  paper  on  new  ates  a  subsocial  situation  where  mothers 

species  of  Arpnctophilus  from  New  Guinea,  and  daughters  are  present  in  the  same 

The  author  indicated   that  the  sub  tribal  nest.   Iwata   (1964)   observed  progressive 

limits  proposed  by  Bohart  and  Menke  in  provisioning  in  nests  of  Cnriiiostignius  ium- 

the  1976  world  revision  should  be  recon-  tai  (Tsuneki)  and  found  larvae  and  adults 

sidered.  He  established  a  new  subtribe,  of  different  ages  in  nests  of  C.  monstrosus 

Spilomenina,  containing  ArpactopJiilus,  (Tsuneki).  The  only  reported  cleptoparas- 

Spilornena,  Microstigmus  and  Xi/sma.  Fur-  ites  are  chrysidid  wasps  of  the  genus  Otn- 

thermore  Menke  was  able  to  unite  these  alus. 

genera  using  a  newly  discovered  apotypic  Unpublished  or  anecdotal  records  indi- 

character  state,  that  of  reduced  papal  seg-  cate  some  species  of  Stigmina  are  sand 

mentation.  Menke's  work  first  of  all  re-  nesting,  a  presumed  plesiotypic  character 

duced  the  Stigmina  to  3  genera:  Pnracrabro  state.  Krombein  (1973,  1984)  in  his  revi- 

(1  Australian  species),  Stigmus  (30  species,  sions  of  Nearctic  Stigmiis  and  Sri  Lankan 

Holarctic,  Neotropical  and  Oriental),  and  Cnrinostignnis,    lent    some    credibility    to 

Carinostigmiis  (25  species,  Afrotropical  and  these  observations  when  he  noted  nesting 

Oriental).  Secondly,  no  synapotypic  char-  differences  in  females  with  2  or  3  mandib- 

acter  states  supporting  the  3  genera  re-  ular   teeth.    Females   with   3   mandibular 

maining   in    the   Stigmina    were   known,  teeth  excavated  nests  in  pithy  twigs  or 

which  meant  that  the  Stigmina  could  be  a  stems  while  those  with  2  teeth  used  pre- 

plesiotypic  assemblage  representing  an  ar-  existing  cavities  in  the  ground.  The  impli- 

tificial   rather   than   a   natural   grouping,  cation  is  that  those  species  constructing 

This  paper  provides  ample  evidence  of  the  nests  evolve  more  complex  mandibular 

monophyly  of  the  Stigmina  and  indicates  dentition   presumably   to    perform   more 

that  9  genera  and  about  150  species  are  specialized  tasks.  Microstigmus  (Spilomen- 

included.  ina)  have  a  bidentate  mandible  that  is  not 

used  in  construction  of  a  complex  nest 
suspended  beneath   a   leaf  and   housing 

Published  records  (Arnold  1924;  Eick-  multiple  generations.  The  foregoing  indi- 
wort  1967;  Iwata  1964;  Janvier  1962;  cates  that  mandibular  dentition  likely 
Krombein  1956,  1958a,  1958b,  1961,  1963,  bears  a  straightforward  relationship  to 
1984;  Rau  1928;  Richardson  1915;  Smith  nest  construction  when  mandibles  are  em- 
1923;  Tsuneki  1970;  Wasbauer  and  Si-  ployed  for  that  purpose, 
monds  1964;  Yasumatsu  and  Watanabe  More  reliable  evidence  of  sand  nesting 
1964)  indicate  that  Stigmus  and  Carinostig-  can  be  inferred  from  the  presence  of  a  fore 
mus  nest  in  twigs  or  pre-existing  cavities  tarsal  rake  and  a  broad  pygidial  plate  in 
and  prey  on  aphids  (Homoptera:  Aphi-  the  female  or  a  remnant  of  that  plate  in 
doidea).  Twig  nests  are  usually  construct-  the  male.  The  fore  tarsal  rake  is  a  series  of 
ed  by  excavating  pithy  twigs  like  Ruhrus,  elongate  setae  which  are  used  to  move 
Sambucus,  or  Sassafras  among  others.  Pre-  sand.  Members  of  Stigmina  lack  a  fore  tar- 
existing  cavities  include  straws  of  thatch,  sal  rake  but  have  spatulate  setae  on  the 
holes  or  borings  in  timber  and  galls.  Nests  rake  area,  possibly  a  secondary  adaptation 
vary  in  length  from  8cm  to  half  a  metre,  to  manipulation  c^f  nest  material.  The  py- 
the  cells  are  constructed  in  linear  series  gidial  plate  is  a  flat  specialized  area  de- 
separated  by  masticated  wood  or  plugs  of  fined  by  a  carina  or  groove  on  the  6th 


BIOLOGY 


208 


Journal  of  Hymenoptera  Research 


metasomal  tergum  in  the  female  and  the 
7th  in  the  male.  In  sand  nesting  species 
this  plate  occupies  a  much  greater  pro- 
portion of  the  tergum  than  in  twig  or  cav- 
ity nesting  species.  Almost  all  Stigmina 
have  a  narrow  pygidial  plate  occupying  a 
small  fraction  of  the  tergum  indicating 
twig  or  cavity  nesting  behaviour.  A  few 
species  have  an  intermediate  pygidial 
plate  and  several  Argentine  species  have 
a  large  pygidial  plate  with  a  remnant  of 
this  plate  in  the  male.  Manfredo  Fritz  (per- 
sonal communication)  has  observed  sand 
nesting  Stigmus  in  Argentina,  he  was  un- 
able to  capture  any  specimens.  Specimens 
from  his  collection,  used  in  this  study, 
contained  females  with  relatively  large 
pygidial  plates  and  with  remnants  in  sev- 
eral males,  possibly  representing  the  only 
true  sand  nesting  Stigmina. 

In  summary,  biological  observations,  al- 
though detailed  in  several  instances,  are 
available  for  few  species  only.  More  ob- 
servations are  necessary  to  document  nest 
construction  and  the  existence  of  social  be- 
havior in  the  group.  The  Stigmina  appear 
to  be  twig  or  cavity  nesting  and  restrict 
prey  to  aphids.  There  is  some  anecdotal 
evidence  of  sand  nesting  behaviour  in 
southern  Neotropical  species.  A  number 
of  new  genera  have  no  behavioral  data 
whatever.  Observations  on  these  genera 
are  often  impeded  by  the  small  size  of  the 
individuals  but  would  offer  a  fertile  field 
of  study  to  a  dedicated  biologist. 

SUBTRIBE  STIGMINA 

Diagnosis. — The  Stigmina,  as  here  de- 
fined, includes  9  genera  with  a  microsetal 
or  micropore  fields  laterally  on  metaso- 
mal sternum  II  (Figs.  44,  45),  more  or  less 
centrally  on  the  fore  wing  stigma  (Figs. 
46,  47)  and  on  the  vertex  between  the  lat- 
eral ocellus  and  the  compound  eye  (Figs. 
3,  8,  20,  26,  32).  The  latter  character  is  not 
fully  developed  in  several  genera  but 
consists  of  several  pits  in  a  small  depres- 
sion. The  stigmal  pit  field  is  diffuse  in  11 
species  from   Chile  and   Argentina   and 


may  not  be  readily  apparent  in  these  in- 
stances. These  characters  are  found  in  all 
members  of  the  Stigmina  and  nowhere 
else  in  the  Pemphredoninae.  They  are 
thus  autapotypies  that  unambiguously 
delineate  the  Stigmina.  Described  genera 
include  Carinostigmus  Tsuneki,  Paracrabro 
Turner,  Parastigmus  Antropov,  and  Stig- 
mus Panzer.  In  addition  to  Paracrabro  and 
Parastigmus,  five  new  genera  are  pro- 
posed and  33  of  the  58  species  are  treated 
in  the  present  paper:  Tzustigmus  (5  spe- 
cies. Oriental,  far  eastern  Palearctic),  Par- 
astigmus (4  species,  southern  Neotropi- 
cal), Araucastigmus  (3  species,  Chile),  Par- 
acrabro (1  species,  Australian),  Aykhustig- 
mus  (4  species.  Neotropical),  Incastigmus 
(25  species.  Neotropical),  and  Llaqhastig- 
mus  (14  species.  Neotropical). 

Description. — Head.  Palpal  formula  6-4; 
mandibular  socket  closed;  antennal  sock- 
ets placed  low  on  frons  near  or  touching 
clypeus  (Fig.  1);  frons  unmodified  by 
grooves,  except  occasionally  along  inner 
eye  margin;  vertex  with  a  micropore  field 
or  at  least  a  few  associated  pits  between 
lateral  ocellus  and  eye  (Figs.  3,  8,  20,  26, 
32);  genal  setae  usually  normal,  not  elon- 
gate setae  scattered  among  short  setae  (ex- 
ceptions among  some  males  of  Stigmus 
and  Carinostigmus). 

Mesosoma.  Pronotum  with  transverse 
carina  present,  at  least  laterally  (Fig.  2); 
omaulus  present;  episternal  sulcus  unde- 
veloped; hypersternaulus  present,  obli- 
quely oriented  across  mesopleuron  (Fig. 
4);  fore  tarsus  of  female  without  rake;  mid 
coxa  with  oblique  dorsal  carina;  inner 
hind  coxal  carina  absent;  hind  coxa  with- 
out elongate  bristle;  metasternum  with 
shallow  posterior  emargination;  propo- 
deal  enclosure  present  (Fig.  10);  stigma  of 
fore  wing  enlarged,  lenticular  with  dis- 
crete micropore  field  (Figs.  46,  47);  mar- 
ginal cell  normal  size,  subequal  to  or  larg- 
er than  stigma,  closed  and  bordering  an- 
terior wing  margin;  2  submarginal  cells; 
submarginal  cell  II  quadrate;  submarginal 
cell  I  out  of  line  with  respect  to  marginal 


Volume  4,  1995 


209 


Figs.  1-5.     Carinostigmiis  sp.  9.  1,  Head,  frontal  view.  2,  head  and  prothorax,  dorsal  view.  3,  micropore  field 
between  lateral  ocellus  and  compound  eye.  4,  mesosoma,  lateral  view.  5,  mesosoma,  dorsal  view. 


and  discoidal  cells;  submarginal  cell  I 
without  spur,  closed;  recurrent  vein  re- 
ceived by  submarginal  cell  I;  2  discoidal 
cells;  hind  wing  cells  closed. 

Metasoma.  Petiole  present,  longer  than 
wide,  setae  short,  scattered  over  surface; 
tergum  I  without  lateral  carina;  male  ster- 


na without  elongate  specializeci  setae; 
sterna  II  and  often  III  with  lateral  micro- 
setal  patches  (Figs.  44,  45);  sternum  VI 
without  apical  pegs;  pygidial  plate  present 
in  female,  almost  always  absent  in  male; 
sternum  VIII  of  male  elongate,  volsella  di- 
vided into  digitus  and  cuspis. 


210  Journal  of  Hymenoptera  Research 


KEY  TO  GENERA  OF  STIGMINA 

1.  Hind  wing  submedian  cell  reduced,  cu-a  positioned  about  halfway  from  wing  base  to 
origin  of  media  (Fig.  50);  Old  World,  except  Australian  Region     ....  Carinostigmus  Tsuneki 

1'.  Hind  wing  submedian  cell  normal  size,  cu-a  positioned  next  to  origin  of  media  (Fig.  49) 

2 

2.  Hind  wing  media  diverging  beyond  cu-a  (Fig.  49)  or  mesopleuron  coarsely  areolate  to 
sternopleural  region  (Figs.  22,  28)     3 

2'.  Hind  wing  media  diverging  before  cu-a  (Fig.  48);  mesopleuron  not  entirely  coarsely  sculp- 
tured (Fig.  34)    5 

3.  Mesopleuron  coarsely  sculptured  to  sternopleural  region  (Figs.  22,  28);  male  with  prom- 
inent genal  carina  separated  from  mid  outer  orbit  by  a  distance  subequal  to  basal  width 

of  mandible  (Fig.  21);  Neotropical    Aykhustigmus  Finnamore,  new  genus 

3'.  Mesopleuron  mostly  unsculptured,  shiny;  genal  carina,  if  present,  close  to  eye  margin 
(Fig.  9)    4 

4.  Interantennal  tubercle  absent  (Fig.  40);  eyes  not  margined  by  a  carina:  petiole  carinate; 
Japan    Stigtnus  Panzer,  in  part 

4'.  Interantennal  tubercle  present  (Fig.  7);  eyes  margined  by  a  carina  (Fig.  9);  petiole  smooth, 

round,  at  most  microcarinate;  Oriental,  far  eastern  Palearctic     

Tztistigmus  Finnamore,  new  genus 

5.  Acetabular  carina  absent  (Fig.  11);  southern  Neotropical    6 

5'.  Acetabular  carina  present  (Fig.  43);  widespread    7 

6.  Pygidial  plate  small,  narrow,  present  in  female  only;  mandible  in  male  bidentate  and  in 
female  tridentate  at  apex;  body  shiny,  without  microsculpture  (Figs.  14,  15);  Chile   .... 

Araucastigmus  Finnamore,  new  genus 

6'.  Pygidial  plate  broad,  present  in  both  sexes,  evanescent  in  male;  mandible  of  both  sexes 
bidentate  at  apex;  microsculpture  present  on  head  and  often  most  of  body;  Argentina, 
Chile     Parastigmus  Antropov 

7.  Vertex  micropore  field  absent;  Australia    Paracrabro  Turner 

7'.  Vertex  micropore  field  present  (Figs.  20,  26,  32)    8 

8.  Mandibles  in  male  bidentate,  female  uni-,  bi-,  or  usually  tridentate;  male  clypeal  apex 
usually  with  lateral  bevel  (Fig.  42);  scutum  without  trace  of  median  groove  or  postero- 
median pit  but  may  be  ridged  (Fig.  41);  apicoventral  mandibular  tooth  in  female,  acute 
(Fig.  39);  widespread,  except  Afrotropical  and  Australian  Regions    .  .  Stigtnus  Panzer,  in  part 

8'.  Mandibles  tridentate  in  both  sexes;  male  clypeal  apex  without  bevel;  scutum  usually  with 
median  groove  or  posteromedian  pit  (Figs.  33,  35);  females  lacking  median  scutal  groove 
have  apicoventral  mandibular  tooth  enlarged  and  broadly  truncate  (Fig.  37);  Neotropical 
9 

9.  Scutum  with  median  groove  or  posteromedian  pit  (Figs.  33,  35);  apicoventral  mandibular 
tooth  in  female,  acute  (Fig.  39);  labrum  quadrilobed;  throughout  Neotropical  Region 

Incastigvtus  Finnamore,  new  genus 

9'.  Scutum  without  trace  of  median  groove  or  posteromedian  pit  (Fig.  38);  female  with  api- 
coventral mandibular  tooth  enlarged,  broadly  truncate  (Fig.  37);  labrum  usually  bilobed, 
rarely  4-lobed;  Neotropical:  restricted  to  continental  South  America   

Llaqhastigmus  Finnamore,  new  genus 


Volume  4,  1995 


211 


Tzustigmus  Finnamore,  new  genus 
(Figs.  6-11,  44,  45,  47) 

Derivation  of  Name. — Tzu,  a  Chinese 
term  meaning  master  or  teacher,  in  this 
case  referring  to  the  relatively  basal  posi- 
tion of  the  genus  within  the  Stigmina  and 
its  subsequent  value  in  the  inference  of 
polarization  of  character  states  in  the  sub- 
tribe. 

Diagnosis. — The  combination  of  an  in- 
terantennal  tubercle,  the  hind  wing  me- 
dia diverging  just  beyond  cu-a,  and  a 
mostly  unsculptured  mesopleuron  will 
separate  this  genus  from  all  others  in  the 
Stigmina. 

Description. — Head.  Labrum  quadri- 
lobed  with  median  emargination  narrow 
and  deep;  mandibular  apex  bidentate  in 
male,  tridentate  in  female;  apicoventral 
mandibular  tooth  of  female  acute;  man- 
dible without  inner  basal  tooth;  male 
clypeal  apex  unmodified,  not  bevelled;  3 
clypeal  teeth  in  female;  interantennal  tu- 
bercle developed;  frontal  carina  absent; 
vertex  with  micropore  field  undeveloped 
consisting  of  only  a  few  associated  pits  in 
a  puncture-like  depression;  inner  orbits 
converging  below;  eyes  margined  by  a 


carina;  occipital  carina  present,  simple  in 
female,  foveolate  in  male;  occipital  carina 
complete,  not  intersecting  hypostomal  ca- 
rina. 

Mesosoma.  Scutum  without  median 
groove;  notaular  grooves  normal,  not 
elongate.  Acetabular  carina  absent,  omau- 
lus  curving  forward  toward  prothorax. 
Scrobal  sulcus  absent  to  weakly  present. 
Hypoepimeral  area  without  coarse  sculp- 
ture. Hind  tibia  with  2  or  3  spines  on  pos- 
terior margin.  Mid  basitarsus  of  male  as 
long  or  longer  than  next  3  tarsomeres 
combined.  Fore  wing  setae  absent  in  cel- 
lular area.  Hind  wing  media  diverging 
just  after  cu-a;  submedian  cell  of  normal 
size,  not  reduced. 

Metasoma.  Petiole  smooth  and  round, 
occasionally  finely  microcarinate.  Pygidial 
plate  narrow,  present  in  female  only.  Dig- 
itus elongate,  toothed;  cuspis  not  lobed 
but  multitoothed. 

Type  Species. — Tzustigmus  syam  new 
species. 

Distribution. — Oriental,  far  eastern  Pale- 
arctic. 

Species  Transferred. — Carinostigmus  rhin- 
ocerus  Budrys. 


KEY  TO  SPECIES  OF  TZUSTIGMUS 
(Males  of  T.  rhinoccnis  and  T.  veda  are  unknown) 

1.  Hypoepimeral  area  with  a  single  ventral  carina  forming  upper  margin  of  scrobal  sulcus 
(Fig.  9);  scrobal  sulcus  foveolate;  male  fore  basitarsus  unmodified;  female  with  area  ad- 
jacent to  propodeal  enclosure  unsculpted  at  least  posterolaterally;  south  India  to  Vietnam 

3.  syam  Finnamore,  new  species 

1'.  Hypoepimeral  area  carinate  on  ventral  half;  scrobal  sulcus  not  evident;  male  fore  basitar- 
sus angulate  medially;  female  with  area  adjacent  to  propodeal  enclosure  finely  carinate   ...    2 

2.  At  least  dorsal  surface  of  petiole  microcarinate;  Thailand  to  Vietnam    

1.  khmer  Finnamore,  new  species 

2'.  Dorsal  surface  of  petiole  unsculptured,  shiny,  dorsolateral  sulci  may  be  present 3 

3.  Pronotal  transverse  sulcus  unsculptured;  south  India     4.  veda  Finnamore,  new  species 

3'.  Pronotum  carinate  on  transverse  sulcus     4 

4.  Stemopleural  region  closely  and   finely  striatopunctate;  Russian   Federation:   far  east 

2.  rhinocerus  (Budrys) 

4'.  Stemopleural  region  shiny,  punctate  only;  Taiwan   5.  iviiming  Finnamore,  new  species 


212 


Journal  of  Hymenoptera  Research 


1.  Tzustigmus  khmer  Finnamore, 
new  species 

Derivation  of  Name. — The  species  epithet, 
khmer,  refers  to  the  11th  century  invaders 
of  the  Dvaravati  Kingdom  then  occupying 
present  day  Thailand  which  is  within  the 
distribution  range  of  this  species. 

Diagnosis. — This  species  is  readily  dis- 
tinguished from  its  relatives  by  the  micro- 
carinate  petiole  (at  least  dorsally). 

Description. — Male.  Length  4.5  mm. 

Head.  Flagellomeres  without  visible 
tyli.  Head  shiny,  without  microsculpture. 
Clypeus  obscured  by  dense  appressed  se- 
tae that  extend  along  inner  orbits  to  half 
height  of  scape.  Inner  and  outer  orbit  mar- 
gined by  coarsely  foveolate  sulcus  that  is 
absent  ventrally  and  anterodorsally.  Frons 
lower  two  thirds  with  irregular,  coarse, 
transverse  carinae.  Vertex  with  sparse,  ev- 
anescent punctures.  Ocelli  closer  to  each 
other  than  to  eye,  OOD  1.4  X  LOD.  Gena 
with  sparse  punctures  and  sparse  setae 
that  increase  in  density  ventrally.  Occipi- 
tal carina  foveolate,  more  coarsely  so  ven- 
trally. 

Mesosoma.  Pronotum  with  transverse 
sulcus  longitudinally  carinate;  side  longi- 
tudinally carinate.  Scutum  shiny,  sparsely 
punctate,  with  anterior  transverse  micro- 
carinae  becoming  longitudinal  adjacent  to 
admedian  lines,  posterior  edge  margined 
by  a  transverse  row  of  short  carinae.  Hy- 
poepimeral  area  microcarinate  on  ventral 
half.  Scrobal  sulcus  not  or  at  most  only 
slightly  evident.  Sternopleural  region 
shiny,  without  microsculpture,  sparsely 
punctate.  Omaulus  and  hypersternaulus 
coarsely  foveolate.  Scutellum  entirely  or 
mostly  shiny,  without  microsculpture  an- 
teriorly, but  sometimes  present  posterior- 
ly. Metapleuron  carinate.  Propodeum 
with  side  entirely  finely  longitudinally 
carinate,  dorsolateral  carina  evanescent, 
lateral  spheres  shiny  with  microcarinae, 
enclosure  with  irregularly  areolate  sculp- 
ture. Fore  basitarsus  angulate  at  basal 
third. 


Metasoma.  Petiole  round,  longitudinal- 
ly microcarinate  dorsally  and  laterally, 
ventrolateral  carinae  faintly  indicated  pos- 
teriorly. Terga  II-V  usually  with  large 
sparse  punctures  in  a  more  or  less  trans- 
verse median  band.  Sterna  IV-VI  with  api- 
cal band  of  dense  punctures  up  to  one 
puncture  diameter  apart,  apical  margins 
impunctate.  Metasoma  with  microsculp- 
ture so  fine  as  to  impart  an  oily  sheen  that 
is  most  evident  on  the  terga. 

Color.  Ground  color  black.  White:  pron- 
otal  lobe.  Brown  to  yellow-brown:  man- 
dible, palpi,  antenna,  legs,  except  coxae, 
tegula,  stigma. 

Female.  Length  5  mm.  Similar  to  male 
except  as  follows:  clypeus  sparsely  setose, 
underlying  sculpture  visible,  punctures 
sparse  and  up  to  several  diameters  apart 
on  median  area  which  is  slightly  bulging. 
OOD  2.3  X  LOD.  Body  sculpture  stronger 
than  in  male,  punctation  closer  than  in 
male  particularly  on  sternopleural  region 
where  fine  punctures  are  evenly  distrib- 
uted up  to  2  diameters  apart.  Mesopleu- 
ron  longitudinally  carinate  from  dorsal 
hypoepimeral  area  to  hypersternaulus. 
Fore  basitarsus  straight,  not  angled  at  bas- 
al third.  Petiole  round,  entirely  microcar- 
inate. Abdominal  tergum  VI  with  narrow, 
shiny,  impunctate  pygidial  plate. 

Material  Examined  {26,  29). — Holotype: 
6,  VIET  NAM:  15  km  nw.  of  Dalat  [Da 
Lat],  1850  m,  5-V-1960.  L.W.  Quate  (BISH). 
Paratypes:  THAILAND:  Chiangmai:  Doi 
pui,  2-IV-1958.  T.C.  Maa  (d:  1  BISH).  VI- 
ETNAM: Fyan,  1200  m.  11-VII-9-VIII- 
1961.  N.R.  Spencer  (9:  1  BISH).  VIET- 
NAM: Mt.  Lang  Bian,  1500-2000  m.  19- V- 
8-VI-1961.  N.R.  Spencer  (9:1  BISH). 

Distribution  (Map  1). — Thailand  to  Vi- 
etnam. 

2.  Tzustigmus  rhinocerus  (Budrys), 
new  combination 

Carinostigmus  rhinocerus  Budrys  1987:54,  9,  c?. 
Holotype:  9  (not  examined),  Russian  Feder- 
ation: Khabarovsk,  Khekhtsir  Range,  Levaya 
River  (ZIN). 


Volume  4,  1995  213 

Diagnosis. — The  combination  of  a   hy-  late.     Fore     basitarsus     straight,     sub- 

poepimeral  area  carinate  on  ventral  half,  cylindrical,  not  bent  or  otherwise  modi- 

sternopleural    region   closely    and    finely  fied. 

striatopunctate,  propodeum  with  carinate  Metasoma.  Petiole  round,  shiny  with 
lateral  spheres  and  petiole  shiny  without  dorsolateral  carina  partially  developed  an- 
microcarinae  (dorsolateral  carinae  are  teriorly.  Terga  II-V  with  medium  size, 
faintly  indicated)  separate  this  species  widely  scattered  sparse  punctures;  micro- 
from  others  in  the  genus.  sculpture  on  anterior  terga  imparts  an  oily 
Description. — Male.  Not  seen.  sheen  to  metasoma.  Sterna  IV  and  V  with 
Female.  Length  6.5  mm.  apical  bands  of  punctures,  punctures  at 
Head.  Clypeus  with  relatively  long  de-  least  2  diameters  apart;  sternum  VI  with 
curved  setae,  not  obscuring  underlying  close  nearly  contiguous  punctures, 
sculpture,  punctures  coarse  and  contigu-  Color.  Ground  color  black.  White:  pron- 
ous  on  median  area  which  is  strongly  otal  lobe.  Brown  to  yellow-brown:  man- 
bulging.  Inner  and  outer  orbit  margined  dibles  except  apex,  palpi,  scape,  underside 
by  a  coarsely  foveolate  sulcus  which  is  ab-  of  flagellum,  legs  except  coxae  and  hind 
sent  ventrally  and  anterodorsally.  Frons  femur,  tegula,  pygidial  plate, 
irregularly,  transversely  carinate  adjacent  Material  Examined  (19  paratype). — 
to  lower  inner  orbit  becoming  striatopunc-  RUSSIAN  FEDERATION:  Khabarovsk: 
tate    toward    vertex.    Scapal    basin    dull,  Khekhtsir,  18th  km. 

densely  microsculptured.  Vertex  shiny  an-  Distribution  (Map  2). — Russian  Federa- 

teriorly,  obscurely  striatopunctate,  punc-  tion  far  east, 
tures  sparse  with  most  over  3  diameters 

apart;  posterior  vertex  with  transverse  mi-  3.  Tzustigmus  syam  Fmnamore, 

crosculpture.  Lateral  ocelli  closer  to  each  ^^^  species 

other  than  to  eyes,  OOD  2.6  X  LOD.  Gena  Derivation  of  Name. — The  species  epithet, 

striatopunctate,  more  densely  so  ventrally.  syam,  is  a  Sanskrit  term  for  dark  used  by 

Occipital  carina  simple,  not  foveolate.  12th  century  Khmers  to  refer  to  the  dark 

Mesosoma.   Pronotum  with  transverse  skin  of  the  people  migrating  south  from 

sulcus  longitudinally  carinate;  side  longi-  southern  China  at  that  time.  It  is  used  here 

tudinally  carinate.  Scutum  shiny,  sparsely  to  refer  to  the  generally  dark  appearance 

punctate,  with  anterior  transverse  micro-  of  this  species. 

carinae  that  become  longitudinal  adjacent  Diagnosis. — The    unsculpted    (non-cari- 

to  admedian  lines,  posterior  edge  mar-  nate)  hypoepimeral  area  will  distinguish 

gined  by  transverse  row  of  short  carinae.  this  species  from  others  in  the  genus. 

Mesopleuron  longitudinally  carinate  from  Description. — Male.  Length  4  mm. 

dorsal  part  of  hypoepimeral  area  to  hy-  Head.  Flagellomeres  without  tyli.  Head 

persternaulus  (which  is  coarsely  foveola-  shiny  with  microsculpture  on  frons,  gena 

te).  Sternopleural  region  shiny  with  dense,  and  posterior  vertex.  Clypeus  obscured  by 

fine  punctures  (that  are  less  then  1  diam-  dense  appressed  setae  which  extend  along 

eter  apart),  somewhat  finely  striatopunc-  inner  orbits  to  a  point  Va  height  of  scape, 

tate  toward  posterior.  Omaulus  foveolate.  Inner    and    outer    orbit    margined    by    a 

Scutellum  shiny  anteriorly,  sparsely  punc-  coarsely  foveolate  sulcus  except  ventrally 

tured,  microsculptured  on  posterior  third,  and    dorsally.    Frons    microsculptured, 

Metapleuron  longitudinally  carinate.  Pro-  without  carinae  except  along  inner  orbital 

podeal     side     longitudinally     carinate  margin  where  they  are  evanescent.  Vertex 

throughout;  dorsolateral  carina  present,  ir-  sparsely  punctate,  with  minute  punctures, 

regular;  lateral  spheres  carinate  to  propo-  appearing    impunctate.    Ocelli    closer    to 

deal  enclosure  which  is  irregularly  areo-  each  other  then  to  eyes;  OOD  1.4  X  LOD. 


214 


Journal  of  Hymenoptera  Research 


Figs.  6-11.  Tziistigmus  syam  9.  6,  head,  frontal  view.  7,  head  and  prothorax,  dorsal  view.  8,  micropore  field 
between  lateral  ocellus  and  compound  eye.  9,  mesosoma,  lateral  view.  10,  mesosoma,  dorsal  view.  11,  me- 
sosoma,  ventrolateral  view. 


Gena  almost  impunctate,  with  microsculp- 
ture  present  to  absent.  Occipital  carina 
foveolate,  areolae  becoming  evanescent 
dorsally. 

Mesosoma.  Pronotum  with  transverse 
sulcus  longitudinally  carinate;  side  longi- 
tudinally carinate.  Scutum  shiny,  sparsely 
punctate,  without  anterior  microcarinae; 
with  transverse  row  of  short  longitudinal 


carinae  next  to  posterior  margin.  Hypoe- 
pimeral  area  microsculptured,  not  cari- 
nate; scrobal  sulcus  distinct,  carinate  and 
foveolate.  Sternopleural  region  shiny, 
without  microsculpture,  sparsely  micro- 
punctate.  Omaulus  and  hypersternaulus 
coarsely  foveolate.  Scutellum  shiny,  with 
several  punctures  mediolaterally.  Meta- 
pleuron  not  carinate.  Propodeal  side  with 


Volume  4,  1995 


215 


evanescent  carinae,  less  so  in  the  discal 
area;  dorsolateral  area  a  series  of  areolae, 
lateral  spheres  shiny,  weakly  carinate  ba- 
sally  becoming  evanescent  and  absent  to- 
ward midline;  propodeal  enclosure  areo- 
late.  Fore  basitarsus  straight,  cylindrical. 

Metasoma.  Petiole  round,  shiny,  with- 
out microsculpture  or  dorsolateral  carina. 
Terga  impunctate  (this  character  is  vari- 
able in  other  species  and  as  more  speci- 
mens are  encountered  may  prove  to  be 
punctate  in  some  specimens).  Sterna  IV  to 
VI  with  apical  bands  of  small  punctures 
less  dense  than  in  previous  species.  Terga 
with  fine  microsculpture  so  as  to  impart 
an  oily  sheen. 

Color.  Ground  color  black.  White:  pron- 
otal  lobe.  Brown  to  yellow-brown:  man- 
dibles except  apically,  palpi,  antenna,  teg- 
ula,  legs. 

Female.  Length  4.5  mm.  Similar  to  male 
except  as  follows:  clypeus  with  3  median 
teeth,  sparsely  setose,  underlying  sculp- 
ture visible  medially;  median  area  bulging 
but  not  strongly  so  and  shiny  with  punc- 
tures 1.5  to  3  diameters  apart.  OOD  2.6  X 
LOD.  Occipital  carina  not  as  strongly  are- 
olate.  Gena  striatopunctate  ventrally.  Pro- 
podeal side  more  strongly  carinate  than  in 
male.  Tergum  VI  with  a  narrow  shiny  py- 
gidial  plate. 

Materinl  Exatnined  {26 ,  89). — Holotype: 
9,  THAILAND:  Chiengmai  Prov.,  10  km 
w.  Fang,  V-5-69,  G.R.  Ballmer,  on  tree 
trunk  (PMA).  Paratypes:  INDIA:  Karna- 
taka:  Bangalore,  916  m,  9-XII-1973,  Ghor- 
pade  collection,  Bangalore  (9:  1  ZMD). 
Nilgiri  Hills,  Naduvatam,  6000  ft.  May 
1958,  P.S.  Nathan.  George  R.  Ferguson 
Collection,  1972  {6:1  OSU).  THAILAND: 
Chiengmai:  10  km  w.  Fang,  5-V-69,  G.R. 
Ballmer,  on  tree  trunk  (9:2  PMA).  Doi 
Suthep-Pui  natn.  Park,  Konthathan  water- 
fall area,  600  m,  20-27-X-1979.  Zool.  Mus. 
Copenhagen  Exped.  (9:1  ZMD).  VIET- 
NAM: Dalat  [Da  Lat]  6  km  s.,  1400—1500 
m,  9-VI-7-VII-1961,  N.R.  Spencer,  9:  1 
BISH.  Fyan,  900  -1000  m.  ll-VII-9-VIII-'61, 
N.R.  Spencer  (9:1  BISH).  Fyan,  1200  m. 


ll-VII-9-VIII-'61,  N.R.  Spencer  (9:1  BISH, 
6:  1  BISH).  The  male  was  collected  at  a 
light  trap. 

Distribution  (Map  3). — India  to  Vietnam. 

4.  Tzustigmus  veda  Finnamore, 
new  species 

Derivation  of  Name. — The  species  epithet 
was  arbitrarily  selected  for  the  species. 
Veda,  in  Hindu  perspective  is  the  eternal 
wisdom  realized  by  the  ancient  seers  or 
sages  preserved  over  thousands  of  years 
by  generations  of  professional  reciters  in 
oral  transmission. 

Diagnosis. — The  non-carinate,  smooth, 
transverse  pronotal  sulcus  is  diagnostic 
for  this  species. 

Description. — Male  unknown. 

Female.  Length  4.5  mm. 

Head.  Head  shiny,  faint  microsculpture 
on  frons  and  posterior  vertex.  Clypeus 
with  3  median  teeth,  sparsely  setose,  un- 
derlying sculpture  visible  medially,  shiny, 
closely  punctate,  punctures  1-2  diameters 
apart.  Inner  and  outer  orbit  margined  by 
a  coarsely  foveolate  sulcus  except  ventral- 
ly and  dorsally.  Frons  with  fine  transverse 
microcarinae  and  some  coarse  transverse 
carinae  near  inner  orbital  sulcus.  Vertex 
shiny  with  fine  microsculpture  posterior- 
ly, punctures  fine,  sparse.  Lateral  ocelli 
closer  to  each  other  than  to  eyes,  OOD  1.9 
X  LOD.  Gena  shiny,  sparsely  punctate 
dorsally,  closely  punctate  to  striatopunc- 
tate ventrally.  Occipital  carina  simple,  ob- 
scurely foveolate  ventrally. 

Mesosoma.  Pronotum  with  transverse 
sulcus  unsculptured  although  longitudi- 
nal carinae  are  present  at  extreme  dorso- 
lateral end,  side  longitudinally  carinate. 
Scutum  shiny,  sparsely  punctate,  with  fine 
anterior  transverse  microcarinae  that  turn 
longitudinally  adjacent  to  admedian  lines; 
with  transverse  row  of  short  carinae  next 
to  posterior  margin.  Hypoepimeral  area 
carinate  over  most  of  its  area.  Scrobal  sul- 
cus evanescent.  Omaulus  and  hyperster- 
naulus  foveolate.  Sternopleural  region 
shiny,  without  microsculpture,  punctures 


216 


Journal  of  Hymenoptera  Research 


fine  1-2  diameters  apart.  Scutellum  shiny 
anteriorly,  posterior  Vz  microsculptured 
and  sparsely  punctate.  Metapleuron  cari- 
nate.  Propodeum  carinate  laterally,  dor- 
solateral carina  present,  lateral  spheres 
finely  carinate  to  propodeal  enclosure 
which  is  areolate. 

Metasoma.  Petiole  round,  shiny,  with- 
out microsculpture,  dorsolateral  carina 
faintly  indicated  anteriorly.  Terga  with 
few  fine  evanescent  punctures  and  with 
microsculpture  imparting  oily  sheen.  Ster- 
na III-V  with  sparse  bands  of  punctures, 
VI  closely  punctate,  punctures  less  than  1 
diameter  apart  near  apex. 

Color.  Ground  color  black.  White:  pro- 
notal  lobe.  Brown  to  yellow-brown:  man- 
dibles except  apex,  palpi,  scape,  tegula, 
legs. 

Material  Examined  (1$). — Holotype:  9, 
S.  INDIA:  Cinchona,  Anamalai  Hills  [An- 
aimalai],  3500  ft.  May  1959,  P.S.  Nathan. 
George  R.  Ferguson  Collection  (OSU). 

Distribution  (Map  4). — Southern  India. 

5.  Tzustigmus  wuming  Finnamore, 
new  species 

Derivation  of  Name. — The  species  epithet, 
wuming,  is  a  Chinese  term  derived  from 
Taoist  philosophy  referring  to  that  which 
is  unnameable. 

Diagnosis. — Resembling  T.  rhinocerus  in 
having  the  hypoepimeral  area  carinate  on 
ventral  half,  the  propodeum  with  closely 
carinate  lateral  spheres,  and  the  petiole 
shiny  without  microcarinae.  Unlike  rhino- 
cerus, wuming  has  a  smooth,  shiny,  sparse- 
ly punctate  sternopleural  region.  Addi- 
tionally the  pleuron  of  wuming  is  unsculp- 
tured  immediately  above  hypersternaulus, 
while  in  rhinocerus  it  is  sculptured  from 
the  mid  hypoepimeral  area  to  hyperster- 
naulus and  sternopleural  region. 

Description. — Male.  Length  4  mm. 

Head.  Flagellomeres  without  tyli.  Head 
shiny  with  microsculpture  on  frons,  vertex 
and  occasionally  gena.  Clypeus  obscured 
by  dense,  appressed  setae  which  extend 
along  inner  orbits  to  about  half  height  of 


scape.  Inner  and  outer  orbits  margined  by 
a  coarsely  foveolate  sulcus  except  ventral- 
ly  and  dorsally.  Frons  microsculptured  on 
scapal  basin,  elsewhere  with  relatively 
coarse,  irregularly  transverse  carinae 
which  meet  dorsally.  Vertex  sparsely 
punctate  with  minute  punctures.  Ocelli 
closer  to  each  other  than  to  eyes;  OOD  1.8 
X  LOD.  Gena  sparsely  punctate  dorsally, 
more  densely  punctate  ventrally;  micro- 
sculpture  present  to  absent.  Occipital  ca- 
rina foveolate. 

Mesosoma.  Pronotum  with  transverse 
sulcus  longitudinally  carinate;  side  longi- 
tudinally carinate.  Scutum  shiny,  sparsely 
punctate;  with  transverse  anterior  micro- 
carinae that  turn  longitudinally  adjacent 
to  admedian  lines;  posterior  edge  mar- 
gined by  a  transverse  row  of  short  carinae. 
Hypoepimeral  area  carinate.  Scrobal  sul- 
cus indistinct,  carinae  not  extending  ven- 
trally to  hypersternaulus.  Sternopleural 
region  shiny,  without  microsculpture, 
sparsely  and  obscurely  punctate.  Omaulus 
and  hypersternaulus  coarsely  foveolate. 
Scutellum  shiny,  without  microsculpture 
except  faintly  microsculptured  on  poste- 
rior third;  sparsely  punctate.  Metapleuron 
carinate  on  ventral  half.  Propodeal  side 
carinate,  dorsolateral  area  with  slightly 
more  coarse  irregular  sculpture,  lateral 
spheres  carinate;  propodeal  enclosure  are- 
olate. Fore  basitarsus  angulate  at  basal 
third. 

Metasoma.  Petiole  round,  shiny,  with- 
out microsculpture  or  dorsolateral  carina 
except  remnant  anteriorly.  Terga  sparsely, 
obscurely  punctate.  Sterna  III-VI  with 
broad  bands  of  punctures  most  of  which 
are  less  than  1  diameter  apart.  Terga  with 
fine  microsculpture  imparting  oily  sheen. 

Color.  Ground  color  black.  White:  pro- 
notal  lobe.  Brown  to  yellow-brown:  palpi, 
antenna,  legs  except  coxae,  tegula,  stigma. 

Female.  Length  5-6  mm.  Similar  to  male 
except  as  follows:  clypeus  sparsely  setose, 
underlying  sculpture  visible  on  median 
area,  punctures  sparse  (1-2  diameters 
apart)  on  median  area  which  is  slightly 


Volume  4,  1995  217 

bulging,  with  3  weakly  developed  apical  in  both  sexes;  apicoventral  tooth  of  female 
teeth.  Frons  with  irregular  transverse  ca-  acute;  inner  basal  tooth  in  at  least  females 
rinae  not  as  strong  as  in  male  and  some-  of  some  species.  Clypeal  apex  in  male 
times  evanescent  dorsally  on  midline,  without  bevelled  modifications;  4  clypeal 
OOD  2.2-2.3  X  LOD.  Sternopleural  region  teeth  present  in  female,  although  median 
slightly  more  densely  punctate  than  in  teeth  are  often  fully  reduced  imparting  a 
male,  punctures  pin  prick  fine.  Fore  basi-  deep,  broad,  median  emargination.  Inter- 
tarsus  straight,  not  angulate.  Abdominal  antennal  tubercle  absent.  Frontal  carina 
tergum  VI  with  narrow  shiny  pygidial  absent  in  male,  present  but  simple  in  fe- 
plate.  Mandibles  yellow-brown.  male.  Vertex  without  micropore  field.  In- 

Material  Examined  26,  4  9). — Holotype:  ner  orbits  parallel  in  female,  converging 

9,  TAIWAN:  Wushe,  1150  m,  III-16-'83,  below  in  male.  Eye  not  margined  by  a  ca- 

H.  &  M.  Townes  (AEI).  Paratypes:  TAI-  rina.   Occipital   carina    simple,   complete, 

WAN:  Wushe,  1150  m,  III-23-'83,  H.  &  M.  not  intersecting  hypostomal  carina. 

Townes.  1150  m  (d:  1  AEI).  Wushe,  IV-2-  Mesosoma.  Scutum  without  median 

83,  Henry  Townes  (9:1  AEI).  Wushe,  1150  groove;  notaular  groove  normal,  short,  not 

m,  IV-19-83,  Henry  Townes  (9:  1  AEI).  elongate.  Acetabular  carina  absent;  omau- 

Wushe,  1150  m,  IV-26-83,  Henry  Townes  lus   curving   forward    toward   prothorax. 

{6:  1  AEI).  Wushe,  1150  m,  V-3-83,  Henry  Scrobal  sulcus  absent  in  male,  evanescent 

Townes  (9:  1  AEI).  in    female.    Hypoepimeral    area    without 

Distribution. — (Map  5).  Taiwan.  coarse  sculpture.  Mid  basitarsus  in  male 

n  A  r>  A  cTir-ii/fi  tc  a    >.  Unmodified,  as  long  as  or  longer  than  next 

PARASTIGMUS  Antropov  _  ^                           u-      j   u-    j  i.u-        -..u  ^ 

^  3  tarsomeres  combmed.  Hmd  tibia  with  2 

Parnsti;^miis   Antropov    1992:97.   Type   species  or  3  spines  on  posterior  margin.  Fore  wing 

Parastigmus  argentinus  Antropov,   1992,   by  without  setae  in  cellular  area.  Hind  wing 

original  designation  and  monotypy.  ^^^^^  diverging  before  cu-a;  submedian 

Diagnosis. — The  combination  of  a   mi-  cell  of  normal  size,  not  reduced. 

crosculptured  body,  the  hind  wing  media  Metasoma.  Petiole  carinate.  Pygidial 

diverging  just  before  cu-a,  a  broad  pygid-  plate  broad,  present  in  both  sexes.  Digitus 

ial  plate,  and  the  lack  of  an  acetabular  ca-  elongate,    toothed;    cuspis    not    lobed, 

rina  will  separate  this  genus  from  others  toothed. 

in  the  Stigmina.  Distribution  (Map  34). — Southern  Neo- 

Description. — Head.   Labrum   quadri-  tropical. 

lobed,  with  median  emargination  narrow  Species  Transferred. — Stigwus  patagonicus 

and  deep.  Mandible  with  apex  bidentate  Mantero. 


KEY  TO  SPECIES  OF  PARASTIGMUS 

1  Males     2 

1'  Females   5 

2  Mesosoma  entirely  dull,  microsculptured;  Chile     2.  hiiectanis  Finnamore,  new  species 

2'  Scutellum  or  sternopleural  region  shiny,  without  microsculpture;  Argentina     3 

3  Sternum  II  and  apices  of  terga  I  and  II,  red   1.  argcntinus  Antropov 

3'  Metasoma  all  black     4 

4  Pedicel  subequal  to  length  of  first  flagellomere  (1:1)    4.  patagonicus  (Mantero) 

4'  Pedicel  longer  than  first  flagellomere  (1:0.7) 3.  nina  Finnamore,  new  species 

5  Apex  of  clypeus  transverse  with  4  small  equally  developed  teeth;  labrum  not  exposed; 
Chile    2.  htieciivus  Finnamore,  new  species 


218  Journal  of  Hymenoptera  Research 

5'  Apex  of  clypeus  with  a  broad,  deep  median  emargination  or  biemargination  exposing 
labrum;  lateral  teeth  prominent;  median  lobe  present  or  absent;  Argentina     6 

6  Pedicel  longer  than  first  flagellomere  (1:0.6-0.7);  mesosoma  red  or  black     

3.  nina  Finnamore,  new  species 

6'  Pedicel  subequal  to  length  of  first  flagellomere  (1:0.9-1.1);  mesosoma  black    7 

7  Clypeus  biemarginate,  median  lobe  developed;  clypeus  and  inner  orbits  bare,  without 
appressed  setae     1.  argentinus  Antropov 

7'  Clypeus  with  single  broad  median  emargination,  median  lobe  evanescent  or  absent;  clyp- 
eus and  inner  orbits  with  appressed  setae  that  partially  obscure  surface 

4.  patagonicus  (Mantero) 


1.  Parastigmus  argentinus  Antropov  somewhat  shiny,  weakly  microsculptured. 

Scutum  without  appressed  setae;  punc- 

Parastigmus  argentinus  Antropov  1992:99  $ .  Ho-  .                                             i.        i.      i 

lotype:  9  (not  examined),  ARGENTINA:  Ne-  "'"^  f?"''^"'  ^^^^^^^^^'^  ^^'^^^^^^  g^^^^es 

uquen,  Collon  Cura,  19-1-1980.  WiUink,  Fi-  fo^^olate.    Scutellum    sparsely    punctate, 

dalgo,  Dominguez,  Claps  (LILLO).  punctures  evanescent.  Scrobal  sulcus  ab- 
sent. Mesopleuron  impunctate.  Hyperster- 

Diagnosis. — The  male  is  recognized  by  naulus   and    omaulus   weakly    foveolate. 

its  red  metasomal  sternum  II,  longitudi-  Metapleuron   shiny,   weakly   microsculp- 

nally  microsculptured  scutum,  and  length  tured.  Propodeum  with  areolate  sculpture 

of  flagellomere  I  which  is  0.75  X  length  of  over  most  of  surface,  sculpture  finer  to- 

pedicel.  Males  of  other  species  have  a  first  ward  metapleuron;  dorsal  enclosure  not 

flagellomere  length  subequal   to   pedicel  defined  by  a  carina. 

and /or  microsculpture  of  scutum   dem-  Metasoma.    Microsculptured,    punctate 

onstrating  no  particular  orientation.  The  at  apices  of  terga  II-VI,  more  coarsely  so 

metasoma  of  other  species  is  black  and  on  posterior  terga;  terga  V  and  VI  sparsely 

may  also  prove  to  be  black  in  other  spec-  punctate  over  entire  dorsal  area.  Tergum 

imens  of  this  species.  The  female  is  rec-  VII  with  ill-defined  pygidial  plate,  punc- 

ognized  by  its  biemarginate  clypeal  edge  tured,   shiny,   impunctate   apically.   Setal 

with  median  lobe  developed.  density  abruptly  increasing  on  tergum  IV 

Description. — Male.  Length  3.5  mm.  to  posterior  metasoma. 

Head.  Flagellomeres  without  tyli.  Ratio  Color.  Ground  color  black.  White:  palpi, 
of  pedicel  to  flagellomere  I  is  1:0.75.  Head  labrum,  mandible  except  apex,  scape  ven- 
coarsely  microsculptured,  without  shiny  trally,  pronotal  lobe,  anterior  spot  on  teg- 
areas.  Clypeus  and  subantennal  area  ob-  ula.  Brown  to  yellow-brown:  antenna  ex- 
scured  by  dense  appressed  setae  which  cept  ventral  scape,  fore  leg  beyond  coxa, 
extend  broadly  up  the  inner  orbits  to  a  mid  leg  beyond  coxa,  hind  leg  except  coxa 
point  1.6  X  scapal  length.  Frons  and  ver-  and  femur,  tegula  except  anterior  spot, 
tex  with  sparse  obscure  punctures,  ap-  stigma.  Red:  apices  of  metasomal  terga  I 
pearing  impunctate  at  low  magnification,  and  II,  sterna  II,  base  of  III  and  VII. 
Ocelli  closer  to  each  other  than  to  eyes;  Female.  Length  4  mm.  Similar  to  male 
OOD  1.6  X  LOD.  Genal  punctures  eva-  except  as  follows:  ratio  of  pedicel  to  fla- 
nescent.  gellomere  I  is  1:1.1.  Clypeus  almost  de- 

Mesosoma.   Pronotum  with  transverse  void  of  setae  and  punctures,  shiny,  with- 

sulcus  longitudinally  carinate.  Scutum  out    microsculpture;    apex    biemarginate 

longitudinally  microsculptured;  scutellum  with  lateral  teeth  prominent  and  median 

and  mesopleuron  above  hypers ternaulus  lobe  developed.  Inner  orbits  with  several 

microsculptured;   sternopleural   region  setae    that    do    not    obscure    underlying 


Volume  4,  1995  219 

sculpture.  Subantennal  area  without  setae.  Punctures  of  frons  and  vertex  sparse,  ev- 

Frons  somewhat  shiny,  with  weak  micro-  anescent.  Ocelli  closer  to  each  other  than 

sculpture.  Vertex  posterior  to  ocelli,  trans-  to  eyes;  OOD  1.4-1.6  X  LOD.  Genal  punc- 

versely  microsculptured.  OOD  2.2  X  LOD.  tures  evanescent. 

Upper  frons  and  vertex  sparsely  punctate.  Mesosoma.  Entirely  microsculptured. 

Lower  gena  more  densely  punctate  than  Pronotum  with  transverse  sulcus  longitu- 

upper  gena,  punctures   1-2   diameters  dinally  carinate;  pronotal  side  longitudi- 

apart   near   mandibular   base.    Scutum  nally  carinate.  Scutum  and   scutellum 

sparsely  punctate.  Mesopleuron  shiny,  sparsely  punctate,  with  many  large  im- 

without    microsculpture;    scrobal    sulcus  punctate  areas;  microsculpture  simple,  not 

slightly   impressed;   sternopleural   region  oriented    in    any    direction.    Notaular 

closely  punctate,  punctures  minute,   1-3  grooves  deeply  impressed,  coarsely  fov- 

diameters    apart.    Pygidial    plate    broad,  eolate.  Scrobal  sulcus  absent.  Mesopleu- 

shiny,  punctate  on  midline,  more  broadly  ron   impunctate,   without  carinae   except 

punctate  basally.  Colored  as  in  male  but  occasionally   at   lower   posterior   margin, 

without  red.  Omaulus  and  hypersternaulus  foveolate. 

Material  Examined  {16,  19). — ARGEN-  Metapleuron  microsculptured,  without  ca- 

TINA:  Neuquen:  Collon  Cura,  19-1-1980.  rinae,  impunctate.  Propodeal  sculpture  ir- 

Willink,  Fidalgo,  Dominguez,  Claps.  regularly  areolate  on  dorsal  and  posterior 

Distribution  (Map  6). — Argentina.  surface.  Propodeal  enclosure  not  evident. 

Propodeal  side  with  sculpture  evanescent 

2.  Parastigmus  huecuvus  Finnamore,  toward  metapleuron. 

new  species  Metasoma.  Microsculptured  except  ter- 

Derivation  of  Name. — Huecuvue,  from  gum  VII.  Punctures  of  terga  and  sterna 
Araucanian  mythology  (Chile),  are  spirits  small,  sparse,  evanescent.  Pygidial  plate 
able  to  change  into  any  form  they  wished  shiny  medially,  without  microsculpture,  2 
for  the  purpose  of  doing  evil,  in  this  case  or  3  punctures  laterally.  Setal  density  in- 
creating  confusion  among  taxonomists  creasing  toward  posterior  metasoma. 
due  to  aberrant  characters.  Color.  Ground  color  black.  White:  Man- 

Diagnosis. — Males   are   distinguished  dible  except  apex,  scape  ventrally,  prono- 

from  other  Parastigmus  by  the  entirely  mi-  tal  lobe,  anterior  spot  on  tegula.  Brown  to 

crosculptured  mesosoma  and  black  abdo-  yellow-brown:  palpi,  antenna  except  ven- 

men.  Other  species  have  a  shiny  sterno-  tral  scape,  stigma,  legs  but  coxae  and  fem- 

pleural  region,  and  the  metasoma  is  in  ora  tend  to  be  darker  brown  and  fore  leg 

part    red    in   patagonicus.    The    female    is  tends  to  be  lighter  yellow-brown, 

unique   in   the   genus   in   having  4   sub-  Female.  Length  4  mm.  Similar  to  male 

equally  developed  clypeal  teeth  and  with-  except  as  follows:  clypeus  entirely,  evenly 

out  a  broad   median  emargination.   The  setose  although  setae  not  obscuring  un- 

clypeus  in  females  of  other  species  has  a  derlying    surface,    shiny,    microsculpture 

broad  median  emargination  in  which  the  absent,  with  several  relatively  large  punc- 

median  teeth  or  lobe  is  recessed  or  absent  tures  subapically  along  apex.  Clypeal  apex 

resulting  in  prominent  lateral  teeth.  with  4  subequally  developed  teeth,  with- 

Description. — Male.  Length  3.5  mm.  out  a  broad  median  emargination.  Suban- 

Head.  Flagellomeres  without  tyli.  Ratio  tennal  area  and  inner  orbits  without  setae, 

of  pedicel  to  flagellomere  I  is  1:0.9.  Head  OOD  2  X  LOD.  Pygidial  plate  shiny,  im- 

entirely  microsculptured.  Clypeus  and  punctate  over  lateral  third;  median  third 

subantennal  area  obscured  by  dense  ap-  slightly  raised,  shiny,  without  microsculp- 

pressed  setae  which  extend  along  inner  ture,  irregularly  punctate  and  irregularly, 

orbits  to  a  point  1.5  times  length  of  scape,  finely  rugose. 


220 


Journal  of  Hymenoptera  Research 


Material  Examined  (36,  109). — Holo- 
type:  9 ,  CHILE:  Santiago — Cordillera,  Re- 
serva  Rio  Clarillos  1-20-II-1989.  R.  Miller 

6  L.  Stange.  Malaise  Trap  (FSCA).  Para- 
types:  CHILE:  Concepcion:  Concepc.  19/ 
12  1907.  P.  Herbst  (9:1  NMW).  The  19/ 
12  and  the  7  in  1907  are  inscribed  in  red 
ink  on  the  specimen  label,  in  the  latter 
case  the  7  is  written  over  the  4  in  1904. 
Specimen  bears  a  second  label  with  the 
numbers  17  and  84.  Concepc.  27/12  1907. 
P.  Herbst  (6:2  NMW).  The  27/12  and  the 

7  are  inscribed  in  red  ink  on  the  specimen 
label  in  the  latter  case  the  7  in  1907  is  writ- 
ten over  the  3  in  1903  or  the  4  in  1904. 
Concepc.  3-1908.  P.  Herbst  (6:  1  NMW). 
The  3  and  8  are  inscribed  in  red  ink,  the 
latter  over  the  4  in  1904.  Santiago:  Cordil- 
lera, Reserva  Rio  Clarillos  1-20-II-1989.  R. 
Miller  &  L.  Stange.  Malaise  Trap  (9:  5 
FSCA,  2  PMA).  Cordillera,  Reserva  Rio 
Clarillos  21-28-11-1989.  R.  Miller  &  L. 
Stange.  Malaise  Trap  (9:1  FSCA). 

Distribution. — (Map  7).  Chile. 

3.  Parastigmus  nina  Finnamore, 
new  species 

Derivation  of  Name. — Nina  is  a  Quechua 
term  meaning  fire,  referring  to  the  red  me- 
sosoma  of  the  holotype. 

Diagnosis. — The  short  flagellomere  I 
(0.6-0.7  X  length  of  pedicel)  distinguishes 
this  species  from  others  in  the  genus  ex- 
cept males  of  argentinus.  In  all  other  spe- 
cies the  length  of  flagellomere  I  equals 
0.9-1.1  of  the  pedicel  length,  except  male 
of  argentinus  which  differs  from  nina  in  its 
red  second  metasomal  sternum.  Parastig- 
mus nina  has  a  pedicekflagellomere  I  ratio 
of  1:0.6-0.7,  black  metasoma,  and  the  me- 
sosoma  may  be  red  or  black,  that  of  the 
female  holotype  is  extensively  red. 

Description. — Male.  Length  4  mm. 

Head.  Flagellomeres  without  tyli.  Ratio 
of  pedicel  to  flagellomere  I  is  1:0.7.  Head 
microsculptured  except  near  frontal  carina 
where  microsculpture  is  evanescent.  Clyp- 
eus  and  subantennal  area  obscured  by 
dense    appressed    setae    which   extend 


broadly  up  the  inner  orbits  to  a  point  al- 
most twice  the  length  of  the  scape.  Frons 
and  vertex  irregularly  punctate  (1-5  di- 
ameters apart).  Ocelli  slightly  closer  to 
each  other  than  to  eyes;  OOD  1.5  X  LOD. 
Genal  punctures  evanescent. 

Mesosoma.  Scutum,  scutellum,  meso- 
pleuron,  and  upper  metapleuron  exten- 
sively shiny,  microsculptured  in  part.  Pro- 
notum  with  transverse  sulcus  longitudi- 
nally carinate;  side  weakly  carinate.  An- 
terior quarter  of  scutum  partially  ob- 
scured by  dense  appressed  setae,  micro- 
sculptured;  median  and  posterior  areas  of 
scutum  shiny  with  evanescent  microsculp- 
ture, punctures  sparse,  notaular  grooves 
foveolate.  Scutellum  shiny,  sparsely  punc- 
tate anteriorly,  microsculptured  posteri- 
orly. Scrobal  sulcus  absent.  Mesopleuron 
immediately  ventral  to  hypersternaulus, 
minutely  punctured;  elsewhere  punctures 
sparse,  evanescent.  Hypersternaulus  and 
omaulus  weakly  foveolate.  Mesopleuron 
microsculptured  posteriorly,  mostly  shiny 
anteriorly.  Metapleuron  microsculptured 
over  most  of  surface  except  dorsal  shiny 
patch.  Propodeum  coarsely  microsculptu- 
red on  side,  finely  areolate  dorsally,  pos- 
teriorly, and  on  lateral  sphere.  Propodeal 
enclosure  not  defined  by  a  carina. 

Metasoma.  Microsculptured,  impuncta- 
te  except  tergum  VII.  Sterna  II-IV  minute- 
ly punctate  posteriorly.  Pygidial  plate  ill- 
defined,  shiny  medially,  without  micro- 
sculpture,  several  punctures  present  lat- 
erally. Setal  density  gradually  increasing 
toward  posterior  metasoma. 

Color.  Ground  color  black.  White:  man- 
dible except  apex,  labrum,  scape  ventrally, 
pronotal  lobe,  anterior  spot  on  tegula. 
Brown  to  yellow-brown:  palpi,  antenna 
except  ventral  scape,  stigma,  legs  except 
coxae,  hind  leg  is  darker  than  other  legs. 

Female.  Length  4.0-4.5  mm.  Similar  to 
male  except  as  follows:  ratio  of  pedicel  to 
flagellomere  I  is  1:0.6-0.7.  Clypeus  sparse- 
ly setose,  shiny,  sparsely  punctate  with 
median  bulge;  apex  broadly  and  deeply 
emarginate  over  median  0.45;  lateral  teeth 


Volume  4,  1995  221 

acute,  prominent;  median  teeth  absent.  In-  the  pedicel  subequal  in  length  to  flagello- 

ner  orbits  and  subantennal  area  not  setose,  mere  I,  and  the  broadly  emarginate  clyp- 

OOD  2.3  X  LOD.  Mesosoma  red  or  black,  eus  with  prominent  lateral  teeth  but  with- 

the  holotype  has  a  red  mesosoma  with  out  a  median  lobe. 

black  in  the  following  areas:  propodeal  en-  Description. — Male.  Length  4.5-5.0  mm. 

closure,  mid  ventral  line,  coxae,  mid  tro-  Head.  Flagellomeres  without  tyli.  Ratio 

chanter,  mid  femur,  hind  leg;  otherwise  of  pedicel  to  flagellomere  I  is  1:1.  Head 

colored  as  in  the  male.  The  9  paratype  has  coarsely  microsculptured,  without  shiny 

the  same  color  pattern  as  the  male.  Pygid-  areas.  Clypeus  and  subantennal  area  ob- 

ial  plate  broad,  shiny,  sparsely  punctate  scured  by  dense  appressed  setae  which 

medially.  extend  broadly  up  inner  orbit  to  a  point 

Material  Examined  {16,  2$). — Holotype:  1.5   X   length  of  scape.  Upper  frons  and 

9,  ARGENTINA:  Neuquen,  Collon  Cura  lower  gena  sparsely  punctate.  Ocelli  closer 

19-1-1980.    Willink,   Fidalgo,    Dominguez,  to  each  other  than  to  eye.  OOD  1.6-1.7  X 

Claps  (LILLO).  Paratypes:  ARGENTINA:  LOD. 

Neuquen:  Collon  Cura  19-1-1980.  Willink,  Mesosoma.   Pronotum  with  transverse 

Fidalgo,  Dominguez,  Claps  {6:  1  LILLO).  sulcus  longitudinally  carinate.  Scutum 

Santa  Cruz:  3  km  s.  Rio  Chico  at  Hwy.  3;  and  scutellum  microsculptured,  without 

XII-1 1-1966,  90m,  E.I.  Schlinger  &  M.E.  Ir-  appressed  setae,  sparsely  punctate;  notau- 

win  (9:1  CAS).  lar  grooves  foveolate.  Mesopleuron  shiny. 

Distribution  (Map  8). — Argentina.  without  microsculpture  ventral  to  hypo- 

epimeral  area  which  has  evanescent  mi- 

4.  Parastigmus  patagonicus  (Mantero)  crosculpture  (may  be  longitudinally  ori- 

new  combination  ented).  Punctures  on  sternopleural  region 

^,.             ,        .       »,     ,       ir>ni  mr,   ^   TT  1  minute,  3-4  diameters  apart.  Scrobal  sul- 

Sti';^miis  imtamnicus  Maniero  1901:199  d.  Holo-  .                ,        ,      , 

lype-  6,  Rio  Sta.  Cruz,  Silvestri  (MCSN).  ^^^  absent,  mdicated  only  by  change  m 

microsculpture.    Hypersternaulus    and 

Diagnosis. — The    male    can    be    distin-  omaulus   weakly    foveolate.    Metapleural 

guished  from  all  other  species  in  the  genus  microsculpture  evanescent.  Propodeum 

by  the  combination  of  a  pedicel  subequal  microsculptured,   areolate   over   most   of 

to  length  of  flagellomere  I,  and  a  shiny  surface  except  near  metapleuron  where  it 

mesopleuron  without  microsculpture  ven-  is  carinate;  propodeal  enclosure  not  indi- 

tral  to  hypoepimeral  area.  The  male  of  this  cated  by  a  carina. 

species  is  superficially  similar  to  huecuvus  Metasoma.  Metasoma  microsculptured 
with  which  it  shares  a  long  first  flagello-  except  tergum  VII,  punctures  evanescent 
mere.  In  addition  to  the  mesopleural  mi-  but  more  evident  on  apices  of  apical  terga. 
crosculpture  the  length  of  the  petiole  may  Pygidial  plate  ill-defined,  shiny,  sparsely 
be  used  to  separate  the  two  species;  the  punctate.  Setal  density  gradually  increas- 
petiole  slightly  shorter  than  tergum  I  in  ing  toward  posterior  of  metasoma. 
huecuvus  and  slightly  longer  than  tergum  Color.  Ground  color  black.  White:  palpi, 
I  in  patagonicus.  The  petiole  length  should  labrum,  mandible  except  apex,  scape  ven- 
be  used  with  caution  since  it  is  subject  to  trally,  pronotal  k^be,  anterior  spot  on  teg- 
variation  in  closely  related  genera  and  has  ula.  Brown  to  yellow-brown:  antenna  ex- 
been  excluded  from  descriptions  for  that  cept  scape  ventrally,  fore  leg,  mid  leg, 
reason.  The  number  of  specimens  avail-  hind  leg  except  coxa  and  femur,  tegula  ex- 
able  for  this  genus  is  insufficient  to  deter-  cept  anterior  spot, 
mine  variation  in  petiole  length.  Female.  Length  6.0-6.5  mm.  Similar  to 

Females  can  be  separated   form  other  male  except  as   follows:   clypeus  evenly 

species  in  the  genus  by  the  combination  of  covered  with  dense  setae  that  partially  ob- 


222 


Journal  of  Hymenoptera  Research 


scure  sculpture  beneath  and  extend  up  in- 
ner orbit  to  a  point  subequal  to  length  of 
scape.  Clypeus  without  microsculpture, 
shiny,  punctures  minute  and  sparse.  Clyp- 
eal  apex  broadly  emarginate,  lateral  teeth 
prominent,  median  lobe  (teeth)  reduced  to 
a  small  mid  point.  Subantennal  area  with- 
out setae.  OOD  1.8-1.9  X  LOD.  Mesosoma 
with  scrobal  sulcus  slightly  more  im- 
pressed than  in  male.  Propodeum  more 
coarsely  areolate  than  male.  Pygidial  plate 
broad,  shiny  with  submedian  puncture 
row  that  converges  apically  with  the  me- 
dian. 

Material  Examined  {56,  2  9).— ARGEN- 
TINA: Catamarca:  Andalhuallas  2000m. 
Santa  Cruz:  Rio  Santa  Cruz  (Type  locali- 
ty). Tucuman:  Amaicha.  Amaicha  del  Val- 
le  1900m. 

Distribution  (Map  9). — Argentina. 

ARAUCASTIGMUS  Finnamore, 

new  genus 

(Figs.  12-15) 

Derivation  of  Name. — The  genus  is 
named  after  the  Araucanian  people  of 
Chile  the  country  of  origin  of  most  known 
specimens. 

Diagnosis. — The  combination  of  an  ace- 
tabular carina  present,  hind  wing  media 
diverging  just  before  cu-a  and  the  meso- 
soma (except  propodeum)  without  mi- 
crosculpture will  separate  this  genus  from 
all  others  in  the  Stigmina. 

Description. — Head.  Labrum  quadri- 


lobed  with  a  narrow  deep  median  emar- 
gination.  Mandible  with  apex  bidentate  in 
male,  tridentate  in  female;  apicoventral 
tooth  acute  in  female;  inner  basal  tooth  ab- 
sent. Clypeal  apex  in  male  without  bev- 
elled modification,  in  female  with  4  teeth. 
Interantennal  tubercle  absent.  Frontal  ca- 
rina present,  simple.  Vertex  with  micro- 
pore field  present,  diffuse.  Inner  orbits 
converging  below.  Eye  not  margined  by  a 
carina.  Occipital  carina  present,  simple, 
complete,  and  separated  from  hypostomal 
carina. 

Mesosoma.  Scutum  without  median 
groove,  notaular  grooves  normal,  short, 
not  elongate.  Acetabular  carina  absent, 
omaulus  curving  forward  toward  protho- 
rax.  Scrobal  sulcus  absent.  Hypoepimeral 
area  without  coarse  sculpture.  Mid  basi- 
tarsus  of  male  unmodified,  as  long  or  lon- 
ger than  next  3  tarsomeres  combined. 
Hind  tibia  with  2  or  3  spines  on  posterior 
margin.  Fore  wing  with  setae  in  cellular 
areas.  Hind  wing  media  diverging  before 
cu-a;  submedian  cell  of  normal  size,  not 
reduced. 

Metasoma.  Petiole  carinate.  Pygidial 
plate  narrow,  absent  in  male.  Digitus 
somewhat  elongate;  cuspis  with  a  large 
lobe  bearing  several  teeth. 

Type  Species. — Araiicastigmiis  glabratus 
(Kohl),  new  combination. 

Distribution  (Map  35). — Southern  Neo- 
tropical: Chile. 

Species  Transferred. — Stigmus  glabratus 
Kohl. 


KEY  TO  SPECIES  OF  ARAUCASTIGMUS 

1  Propodeum  sculptured  throughout,  area  adjacent  to  metapleuron  microsculptured  (Figs. 

14,  15)    2.  masneri  Finnamore,  new  species 

1'  Propodeal  side  with  basal  shiny  area  adjacent  to  metapleuron     2 

2  Propodeum  microsculptured,  shiny  without  microsculpture  along  mid  dorsal  line  (outside 
enclosure)  in  male  and  also  on  mid  posterior  line  in  female     

3.  yanillus  Finnamore,  new  species 

2'  Propodeum  with  sculpture  in  addition  to  microsculpture  at  least  on  dorsolateral  areas  in 
male;  female  without  shiny  areas  on  propodeum  other  than  adjacent  to  metapleuron   .  . 
1.  glabratus  (Kohl) 


Volume  4,  1995 


223 


Figs.  12-17.  Araucastigmus  and  Paracrabro.  12,  Amucastigmus  masneri  3,  head,  frontal  view.  13,  A.  masneri  6, 
head,  dorsal  view.  14,  A.  masneri  6,  mesosoma,  lateral  view.  15,  A.  masneri  6 ,  mesosoma,  dorsal  view.  16, 
Paracrabro  froggatti  9,  mandible,  outer  view.  17,  Paracrabro  froggatti  9,  mandible,  inner  view. 


1.  Araucastigmus  glabratus  (Kohl), 
new  combination 

Stigmus  glabratus  Kohl  1905:358  9.  Holotype*: 
9,  CHILE:  Corral  [Coual?],  3-1904,  P.  Herbst 

(NMW). 

Ding)iosis. — Propodeum  with  shiny  area 
adjacent  to  metapleuron  and  with  both 
microsculpture  and  sculpture  at  least  on 
dorsolateral  areas.  Other  species  are  either 


microsculptured  throughout  or  macro- 
sculpture  is  absent  or  shiny  areas  are 
much  more  extensive. 

Description. — Male.  Length  3.2-5  mm. 

Head.  Flagellomeres  without  tyli.  Head 
shiny,  mostly  without  microsculpture, 
sparsely  punctate.  Clypeus,  subantennal 
area  and  inner  orbit  obscured  by  dense 
appressed  setae  which  extend  broadly  up 
inner  orbit  to  a  point  1.5  X  length  of  scape. 


224 


Journal  of  Hymenoptera  Research 


Frons  impunctate,  weakly  microsculp- 
tured.  Vertex  transversely  microsculp- 
tured.  Ocelli  closer  to  each  other  than  to 
eyes.  OOD  1.2-1.3  X  LOD. 

Mesosoma.  Pronotal  transverse  carina 
foveolate  laterally;  transverse  sulcus  un- 
sculptured;  side  irregularly  carinate.  Scu- 
tum and  scutellum  microsculptured.  Scu- 
tum usually  with  a  transverse  row  of  mi- 
crocarinae  next  to  posterior  margin;  punc- 
tation  variable,  ranging  from  small  sparse 
pinpricks  to  larger  more  dense  (3^  diam- 
eters apart)  punctures  more  or  less  concen- 
trated in  the  notaular  region;  anterior  scu- 
tal  margin  nearly  contiguously  punctate 
between  notaular  grooves.  Scutellum 
sparsely  punctate  on  disc.  Mesopleuron 
shiny,  microsculpture  at  most  weakly  evi- 
dent, nearly  impunctate;  scrobal  sulcus  ab- 
sent; hyperstemaulus  and  omaulus  weakly 
foveolate.  Metapleuron  impunctate,  weak- 
ly microsculptured.  Propodeal  side  shiny 
adjacent  to  metapleuron,  otherwise  micros- 
culptured and  finely  areolate.  Propodeal 
enclosure  weakly  defined,  not  carinate. 

Metasoma.  Weakly  microsculptured. 
Punctures  evanescent  on  terga  I-III;  poste- 
rior band  of  punctures  on  terga  IV- VII  that 
increases  in  width  and  puncture  density  to- 
wards posterior  metasoma.  Sterna  minute- 
ly punctured. 

Color.  Ground  color  black.  White  to 
cream:  palpi,  mandibles  except  apically, 
antenna  ventrally  except  apical  flagello- 
meres,  pronotal  lobe,  tegula,  fore  leg,  mid 
leg  except  base  of  coxa  and  median  area 
of  femur,  hind  leg  except  base  of  coxa  and 
basal  fj  of  femur.  Yellow-brown:  apical 
flagellomeres  of  antenna,  base  of  mid  and 
hind  coxae,  median  area  of  mid  femur, 
basal  %  of  hind  femur.  Orange:  apex  of  ter- 
gum  VII  of  metasoma. 

Female.  Length  4-6  mm.  Similar  to  male 
except  as  follows:  clypeus  without  micro- 
sculpture,  evenly  punctured  and  setose,  se- 
tae not  obscuring  underlying  sculpture; 
punctures  small  about  1-2  diameters  apart; 
apex  with  2  median  teeth  separated  by  a 
shallow  emargination;  setae  extend  from 


clypeus  over  subantennal  area  and  up  in- 
ner orbit  to  a  point  slightly  greater  than 
that  of  scapal  apex.  OOD  1.8-2.0  X  LOD. 
Pygidial  plate  present,  microsculptured, 
with  several  punctures.  Antenna  beyond 
pedicel  brown;  coxae,  femora,  and  hind  tib- 
ia suffused  with  brown;  remaining  area  of 
legs  more  yellowed  than  in  male.  Apex  of 
metasomal  tergum  VI  orange. 

Material  Examined  {266, 43  9 ).— ARGEN- 
TINA: Neuquen:  Pucara,  Lago  Lacar.  Pu- 
cara,  P.  Nac.  Lanin.  San  Martin  Andes. 
CHILE:  Arauco:  Contulmo,  Palo  Botado. 
Aysen:  Lago  Frio.  Puerto  Cisnes.  Cautin: 
Afunalhue  Villarrica.  Chiloe:  Chiloe  Is- 
land, Dalcahue.  Curico:  Cord.  Curico,  El 
Coigo.  Cubillo,  C.  Curico.  Estero  La  Juala, 
Los  Queues,  Cord.  Curico.  Los  Lagos:  Cor- 
ral (type  locality).  Malleco:  P.  N.  Contul- 
mo. Victoria,  4km  s.,  300m.  Nuble:  Las 
Trancas,  Cord.  Nuble  (Chilian).  O'Higgins: 
Graneros.  Santiago:  El  Canelo.  Lo  Canas. 
Talca:  Altos  de  Vilches,  1200m.  Valparaiso: 
Los  Andes.  Vina  del  Mar.  Valparaiso. 

Distribution  (Map  10). — Argentina,  Chile. 

2.  Araucastigmus  masneri  Finnamore, 
new  species 

Derivation  of  Name. — The  species  is 
named  in  honour  of  Lubomir  Masner,  col- 
lector of  half  the  known  specimens. 

Diagnosis. — Propodeum  without  shiny 
areas.  Other  species  in  the  genus  have  a 
shiny  area  on  the  propodeum  adjacent  to 
the  metapleuron. 

Description. — Male.  Length  4.0-5.5  mm. 

Head.  Flagellomeres  without  tyli.  Head 
sparsely  punctate,  shiny,  usually  without 
microsculpture  except  on  frons  but  larger 
specimens  have  evanescent  microsculpture 
on  vertex  and  gena.  Clypeus  and  subanten- 
nal area  obscured  by  dense  appressed  setae 
which  extend  up  inner  orbit  to  a  point 
slightly  greater  than  height  of  scape.  Frons 
impunctate,  microsculptured.  Vertex  mi- 
nutely and  sparsely  punctate.  Ocelli  closer 
to  each  other  than  to  eyes.  OOD  1.8-1.9  X 
LOD. 

Mesosoma.    Pronotal   transverse   carina 


Volume  4,  1995  225 

foveolate  laterally,  transverse  sulcus  un-  type:  (5,  CHILE:  Cautin,  Conguillio  Nation- 
sculpted;  pronotal  side  with  a  few  short  ca-  al  Park,  1150m  4-II-1988  L.  Masner,  Aurau- 
rinae.  Scutum  with  contiguous  punctures  carin  [Araucarial ,  Araucariaceae],  NotJiofa- 
between  notaular  grooves  near  anterior  gus  [Fagaceae]  (BRD).  Paratypes:  ARGEN- 
margin,  otherwise  impunctate  or  nearly  so;  TINA:  Neuquen:  Lanin,  Pucara  15-III-1971 
shiny,  without  microsculpture;  posterior  Schajovskoy  (9:  4  IIES).  CHILE:  Arauco: 
margin  weakly  longitudinally  microcari-  Pichinahuel,  Cord.  Nahuelbuta  20-28-1- 
nate;  notaular  groove  deeply  impressed,  1959  L.  Pefia  {6:1  BRD).  Aysen:  Lago  Frio 
foveolate.  Scutellum  shiny  on  anterior  half,  20/23-1-61  L.  Peiia  {6:  1  MCZ).  Cautin: 
microsculptured  on  posterior  half;  with  same  data  as  holotype  ( c5 :  2  BRD,  2  PM A; 
several  scattered  punctures  on  transverse  9:  4  BRD,  5  PMA).  Los  Lagos:  Osomo, 
midline.  Mesopleuron  shiny,  without  mi-  Puyehue  Nat.  Pk.,  Antillanca,  1200m  16-11- 
crosculpture,  punctures  sparse,  obscure;  1988  L.  Masner,  NotJwfngus  [Fagaceae]  near 
scrobal  sulcus  absent.  Hyperstemaulus  and  treeline  {6:  1  BRD).  Magallanes:  Laguna 
omaulus  foveolate.  Metapleuron  weakly  Amarga,  12-60  Pena  (9:  1  IIES).  Laguna 
microsculptured,  impunctate.  Propodeum  Amarga,  Natales  XII-14-21-60  Luis  Pena 
without  shiny  areas,  entirely  microsculp-  (9:1  OSU).  Malleco:  Cord,  de  las  Raices, 
tured;  evanescent,  irregular,  areolae  on  lat-  40km  e.  Curacautin  6-7-II-1979,  1200m  D. 
eral  spheres;  propodeal  enclosure  not  de-  &  M.  Davis  &  B.  Akerbergs  (9:1  USNM). 
fined  by  a  carina,  with  irregular  carinae  Curacautin,  R.  Blanco  11-64  Luis  E.  Pefia  (d: 
medially  and  on  anterior  margin.  1  AEI).  Icalma,  31-XII-1958  Luis  E.  Pena  (9: 

Metasoma.  Terga  dull,  microsculptured,  1  BRD).  Liucura  1800m,  Jan.  1959  L.E.  Pefia 

punctures  evanescent;  posterior  metasoma  (9:1  OSU).  Nuble:  Las  Trancas  1600m  19- 

more  shiny  with  punctures  forming  more  22-1979  (9:1  PMA).  Las  Trancas,  Chilian 

or  less  discrete  apical  bands  on  terga.  Ster-  1-19-22-1979,  1600m  L.  Pena  (9:  1  AEI). 

na  microsculptured,  impunctate.  O'Higgins:  Graneros  1100m  4-III-62  Pena 

Color.  Ground  color  black.  White:  palpi,  (9:1  MCZ).  Valdivia:  30km  w.  La  Union, 

mandible  except  apex,  pronotal  lobe,  ante-  Las  Traucas  500m  8-11-II-1988  PT,  L.  Mas- 

rior    spot    on    tegula.    Brown    to    yellow-  ner  {6:1  BRD). 

brown:  scape,  pedicel,  at  least  flagellomer-         Distribution  (Map  11). — Argentina:  Neu- 

es  I-V  and  occasionally  I-XI,  tegula  except  quen.  Chile, 
anterior  spot,  trochanters,  fore  and  mid  tib- 

iae  and  tarsi,  occasionally  bases  of  sterna         3.  Araucastigmus  yanillus  Finnamore, 
IV,  V  and  apices  of  VI  and  VII.  ^^^  species 

Female.  Length  4.5-5.5  mm.  Similar  to  Derivation  of  Name. — The  name  yanillus 
male  except  as  follows:  clypeus  shiny,  is  derived  from  two  Quechua  terms,  yana 
evenly  punctured  and  setose,  setae  not  ob-  meaning  black  and  illay  meaning  bright  or 
scuring  underlying  sculpture;  punctures  shiny,  in  reference  to  the  shiny  black  ap- 
minute,  2-3  diameters  apart.  Clypeal  apex  pearance  of  this  species, 
with  2  teeth  separated  by  shallow  median  Diagnosis. — The  extensive  shiny  areas  in 
emargination.  Subantennal  area  less  dense-  the  female  propodeum  and  in  the  male 
ly  setose  than  clypeus.  OOD  2.0-2.1  X  propodeum,  the  lack  of  sculpture  overlying 
LOD.  Scutum  and  mesopleurc^n  more  microsculpture,  and  the  lateral  shiny  area 
coarsely  punctate  than  in  male.  Pygidial  next  to  the  metapleuron  will  separate  this 
plate  weakly  microsculptured,  with  several  species  from  all  others  in  the  genus.  Other 
punctures.  Flagellum  of  antenna  brown  to  species  have  the  propodeum  entirely  mi- 
black,  mid  tibia  occasionally  black,  sterna  crosculptured  or  they  have  sculpture  on  at 
black.  least  lateral  spheres. 

Material    Examined    {106,    209). — Holo-  Description. — Male.  Length  4  mm. 


226  Journal  of  Hymenoptera  Research 

Head.  Flagellomeres  without  tyli.  Head  anterior  spot  on  tegula,  trochanters,  fore 
sparsely  punctate,  shiny,  without  micro-  and  mid  tarsi,  coxae  ventrally.  Yellow- 
sculpture  except  sometimes  evanescent  mi-  brown:  scape,  pedicel,  flagellomeres  I-VII 
crosculpture  on  gena.  Clypeus  and  suban-  ventrally,  coxae  dorsally,  femora,  tibiae, 
tennal  area  obscured  by  dense  appressed  hind  tarsus,  and  sterna  beyond  II  more  or 
setae  which  extend  up  inner  orbit  to  a  less  suffused. 

point  1.1  X  length  of  scape.  Lower  frons  Female.  Length  4.5-5.5  mm.  Similar  to 

impunctate,  shiny;  upper  frons  with  sev-  male  except  as  follows:  clypeus  sparsely 

eral  scattered  punctures.  Vertex  sparsely  and  evenly  setose,  more  sparsely  setose 

punctate.  Ocelli  closer  to  each  other  than  to  than  in  other  species,  punctures  minute 

eyes.  OOD  1.5-1.6  X  LOD.  with   several   larger   punctures   medially. 

Mesosoma.  Pronotal  transverse  carina  Clypeal  apex  with  2  teeth  separated  by 
foveolate  laterally,  several  longitudinal  ca-  shallow  median  emargination.  Subanten- 
rinae  sometimes  present  near  median  re-  nal  area  without  setae.  OOD  2.1  X  LOD. 
gion  of  transverse  sulcus.  Scutum  with  Propodeum  with  sculpture  on  lateral 
variable,  non-contiguous  punctures  and  a  spheres  variable  from  nearly  completely 
few  fine,  transverse  microcarinae  on  ante-  shiny  (most  specimens)  to  almost  entirely 
rior  margin  between  notaular  grooves;  mi-  microsculptured,  without  sculpture  over- 
crosculpture  present  between  admedian  lying  microsculpture;  area  adjacent  to  me- 
lines.  Scutum  otherwise  shiny,  without  mi-  tapleuron  shiny.  Pygidial  plate  narrow 
crosculpture,  mostly  impunctate,  few  with  median  area  minutely  punctate.  Mid 
punctures  clustered  posterior  to  notaular  and  hind  legs  brown  except  trochanters, 
groove;  notaular  groove  deeply  impressed.  Material  Examined  (8(?,  109). — Holotype: 
foveolate;  posterior  margin  of  scutum  9,  CHILE:  Cautin:  Conguillio  National 
smooth,  without  transverse  row  of  longi-  Park  1150m  4-II-1988  L.  Masner,  A?/rni/cann 
tudinal  carinae.  Scutellum  shiny  on  ante-  [Araiicaria? ,  Araucariaceae],  Nothofagiis  [Fa- 
rior  half,  microsculptured  on  posterior  half;  gaceae]  (BRD).  Para  types:  ARGENTINA: 
a  few  punctures  along  transverse  median  Neuquen:  Pucara  (S.M.  Andes)  XII-6-16- 
discal  area.  Mesopleuron  shiny,  impuncta-  1968  C.C.  Porter  {6:1  MCZ).  CHILE:  Ar- 
te above  stemopleural  region;  scrobal  sul-  auco:  Pichinahuel,  Cord.  Nahuelbuta  10- 
cus  absent.  Hypersternaulus  foveolate;  20-1-1959  L.  Pena  (9:1  BRD).  Cautin:  same 
omaulus  not  foveolate,  usually  evanescent  data  as  holotype  (6:5  BRD,  2  PMA;  9:  1 
to  absent  toward  hypersternaulus  but  con-  BRD,  5  PMA).  Curico:  El  Coigo  Jan-1961  J. 
tinned  ventrally.  Metapleuron  shiny,  with-  Foerster  (9:1  BRD).  Nuble:  Las  Trancas, 
out  microsculpture.  Propodeum  shiny  lat-  1400m  Chilian  XII-19-21-1983  Luis  Pena 
erally  adjacent  to  metapleuron,  otherwise  (9:1  AEI). 

microsculptured    without    raised    lines  Distribution  (Map  12). — Argentina:  Ne- 

(sculpture)  except  on  median  area  of  pro-  uquen.  Chile, 
podeal  enclosure  where  several  more  or 


PARACRABRO  Turner 
(Figs.  16,  17) 


less  transverse  carinae  are  usually  evident. 
Propodeal  enclosure  not  defined  by  a  ca- 
rina. Paracrabro  Turner  1907:274.  Type  species:  Para-' 

Metasoma.   Weakly   microsculptured,  cratro /ro^^am' Turner,  1907  by  original  des- 

somewhat  shiny  compared  to  other  spe-  ignation  and  monotypy. 

cies;  punctures  minute,  sparse,  evanescent.  Diagnosis. — The  presence  of  an  acetab- 

not  forming  apical  bands  on  posterior  ter-  ular  carina,  hind  wing  media  diverging 

ga.  Sterna  microsculptured,  impunctate.  just  before  cu-a,  and  the  lack  of  a  discrete 

Color.  Ground  color  black.  White:  palpi,  micropore  field  on  the  vertex  will  separate 

mandible  except  apically,  pronotal   lobe,  this  genus  from  all  others  in  the  Stigmina. 


Volume  4,  1995 


227 


Description. — Head.  Labrum  weakly 
quadrilobed  (appears  bilobed),  median 
lobes  evanescent.  Mandible  with  apex  bi- 
dentate  in  male,  pentadentate  in  female; 
apicoventral  tooth  in  female  acute;  inner 
basal  tooth  absent.  Clypeal  apex  in  male 
without  bevelled  modifications;  female 
clypeus  with  4  apical  teeth  although  me- 
dian teeth  are  greatly  reduced  imparting  a 
bidentate  appearance.  Interantennal  tuber- 
cle present,  protruding.  Frontal  carina 
present  as  a  simple  raised  line  immediate- 
ly above  interantennal  tubercle.  Vertex 
with  loose  association  of  pits  rather  than 
micropore  field;  pits  shifted  anteriorly  in 
female.  Inner  orbits  parallel  in  female, 
converging  below  in  male.  Eye  margined 
by  a  carina  on  inner  orbit  in  female,  en- 
tirely margined  in  male.  Occipital  carina 
foveolate,  more  or  less  raised  ventrally, 
not  intersecting  hypostomal  carina. 

Mesosoma.  Scutum  without  median 
groove,  notaular  groove  not  elongate.  Ac- 
etabular carina  present,  omaulus  intersect- 
ing acetabular  carina,  subomaulus  absent. 
Scrobal  sulcus  absent  in  female,  evanes- 
cent in  male.  Hypoepimeral  area  without 
coarse  sculpture  except  in  area  of  scrobal 
sulcus  in  male,  microsculptured  only.  Mid 
basitarsus  of  male  flattened,  curved  with 
posterior  subapical  expansion.  Hind  tibia 
excluding  apex,  with  4  posterior  spines  in 
female,  2  in  male.  Fore  wing  cellular  area 
without  setae  in  male,  setae  present  in  fe- 
male. Hind  wing  media  diverging  at  cu-a; 
hind  wing  submedian  cell  not  reduced,  of 
normal  size. 

Metasoma.  Petiole  rounded,  carinae 
present  apically  and  basally.  Pygidial 
plate  narrow,  trough-like,  absent  in  male. 
Male  genitalia  unusually  elongate  relative 
to  other  genera  in  the  subtribe,  digitus 
elongate  relative  to  cuspis. 

Distribution  (Map  13). — Australia. 

Paracrabro  froggatti  Turner 

Paracrabro  froggatti  Turner  1907:275    9.   Holo- 
type  9  (not  examined),  Australia:  Victoria. 


Description. — See  generic  description 
above. 

Material  Examined  {IS,  1  9).— AUSTRA- 
LIA: New  South  Wales:  Elizabeth  Beach, 
21km  s.  Forster.  Western  Australia:  Gas- 
coyne  Exp.  Sta.,  Carnarvon. 

Distribution  (Map  13). — Australia. 

AYKHUSTIGMUS  Finnamore, 

new  genus 

(Figs.  18-29) 

Derivation  of  Name. — Aykhu  is  a 
Quechua  term  meaning  extravagant  or 
fancy  in  reference  to  the  unusually  com- 
plex sculpturing  found  in  members  of 
this  genus. 

Diagnosis. — The  coarsely  sculptured 
mesopleuron  will  separate  this  genus 
from  all  others  in  the  Stigmina. 

Description. — Head.  Labrum  quadri- 
lobed, with  a  shallow  broad  median  emar- 
gination.  Mandible  with  apex  in  male  bi- 
dentate, in  female  tridentate;  apicoventral 
tooth  in  female  acute;  inner  basal  tooth  ab- 
sent. Clypeal  apex  in  male  without  bev- 
elled modification;  female  clypeal  apex 
with  4  teeth.  Interantennal  tubercle  absent. 
Frontal  carina  absent.  Vertex  with  micro- 
pore field  between  lateral  ocellus  and  eye 
margin.  Inner  orbits  converging  below. 
Eye  margined  by  a  strong  carina.  Occipital 
carina  present,  complete,  not  joining  hy- 
postomal carina,  simple  in  female,  raised 
and  foveolate  in  male. 

Mesosoma.  Scutum  with  median  groove 
usually  absent  but  occasionally  evanescent 
posteriorly;  notaular  groove  normal,  not 
elongate.  Acetabular  carina  present,  inter- 
cepted by  omaulus.  Scrobal  sulcus  pres- 
ent, evanescent.  Hypoepimeral  area 
coarsely  sculptured.  Mid  basitarsus  in 
male  unmodified.  Posterior  margin  of 
hind  tibia  with  2  or  3  spines.  Fore  wing 
with  setae  absent  from  cellular  area.  Hind 
wing  with  media  diverging  before  or  after 
cu-a;  submedian  cell  of  normal  size,  not 
reduced. 

Metasoma.  Petiole  carinate.  Pygidial 


228 


Journal  of  Hymenoptera  Research 


plate  narrow,  absent  in  male.   Genitalia  Distribution  (Map  36). — Neotropical, 

with    digitus    elongate,    acute,    cuspis  south  of  the  Brazilian  Highlands  although 

(lobed?)  with  tooth.  a  single  record  exists  for  Venezuela,  pos- 

Type  Species. — Ai/khiistigmus  patanawi  sibly  an  introduced  species  or  mislabelled 

new  species.  specimen. 


KEY  TO  SPECIES  OF  AYKHUSTIGMUS 
(Male  of  A.  phasti  is  unknown) 

1  Males     2 

r  Females   4 

2  Hind  wing  media  diverging  before  cu-a  (Fig.  48);  vertex  and  gena  striatopunctate  (Figs. 
19,  21);  frons  irregularly  carinate;  scutum  closely  and  coarsely  punctate  to  striatopunctate 
posteriorly  (Fig.  23)    2.  patanaivi  Finnamore,  new  species 

2'  Hind  wing  media  diverging  after  cu-a  (Fig,  49);  head  at  most  with  microsculpture;  scutum 
with  smaller  more  widely  separated  punctures    3 

3  Frons  with  pair  of  small  swellings  anterior  to  mid  ocellus  on  each  side  of  frontal  carina 

1.  fritzi  Finnamore,  new  species 

3'  Frons  smoothly  rounded,  without  swellings     4.  waraiva  Finnamore,  new  species 

4  Hind  wing  media  diverging  before  cu-a  (Fig.  48);  vertex  transversely  striatopunctate;  upper 
frons  striatopunctate     2.  patanaivi  Finnamore,  new  species 

4'  Hind  wing  media  diverging  after  cu-a  (Fig.  49);  frons  and  vertex  at  most  with  microsculp- 
ture         5 

5  Frons  with  a  pair  of  broad  swellings  anterior  to  mid  ocellus  that  are  visible  in  lateral  profile 

1.  fritzi  Finnamore,  new  species 

5'  Frons  evenly  rounded,  without  swellings 6 

6  Clypeus  with  4  apical  teeth  (Fig.  24);  scutum  microsculptured  on  anterior  Va  to  Vi,  shiny 
and  punctate  to  striatopunctate  on  posterior  half  (Fig.  29)    

4.  ivarawa  Finnamore,  new  species 

6'  Clypeus  with  2  apical  teeth;  scutum  shiny,  without  microsculpture,  carinate  on  median 
third    3.  phasti  Finnamore,  new  species 


1.  Aykhustigmus  fritzi  Finnamore, 
new  species 

Derivation  of  Name. — The  species  is 
named  in  honour  of  Manfredo  A.  Fritz, 
collector  of  most  of  the  specimens  I  have 
seen  of  the  genus. 

Diagnosis. — The  hind  wing  media  di- 
verging after  cu-a,  swellings  on  the  upper 
frons,  and  microsculpture  on  the  anterior 
scutum  will  distinguish  this  species  from 
all  others  in  the  genus. 

Description. — Male.  Length  4  mm. 

Head.  Flagellomeres  without  tyli.  Clyp- 
eus and  subantennal  area  obscured  by 
dense  appressed  setae  that  extend  up  in- 
ner orbits  to  a  point  about  %  length  of 
scape.  Eye  margined  by  a  coarse  carina 


that  is  interrupted  ventrally  near  base  of 
mandible.  Carina  foveolate,  except  on  ver- 
tex and  separated  from  outer  orbit  by  a 
distance  subequal  to  basal  width  of  man- 
dible. Frons  microsculptured  and  impunc- 
tate  on  lower  face;  shiny,  with  evanescent 
microsculpture  on  upper  surface  to  lateral 
ocellus  where  it  is  sparsely  punctate  with 
punctures  at  least  3  diameters  apart.  Up- 
per frons  with  a  swelling  on  each  side  of 
frontal  line  that  is  just  visible  in  lateral 
profile.  Vertex  microsculptured,  with  a 
circular  depression  on  midline  posterior  to 
lateral  ocelli.  Posterior  vertex  with  an  ev- 
anescent transverse  ridge  (posterior  to  cir- 
cular depression)  connecting  outer  orbital 
carinae.  Ocelli  closer  to  each  other  than  to 


Volume  4,  1995 


229 


eyes.  OOD  1.8  x  LOD.  Gena  microsculp- 
tured,  punctures  evanescent;  irregular  ev- 
anescent carinae  radiate  from  occipital  ca- 
rina on  lower  genal  area.  Occipital  carina 
considerably  raised,  coarsely  foveolate  to 
mid  ventral  line. 

Mesosoma.  Pronotum  with  transverse 
carina  greatly  raised,  toothed  at  lateral 
end;  carina  continued  on  side  to  antero- 
ventral  corner  which  is  produced  anteri- 
orly as  a  tooth  that  surpasses  dorsolateral 
tooth  in  size.  Pronotum  with  transverse 
sulcus  longitudinally  carinate;  setae 
sparse,  not  obscuring  sculpture.  Pronotal 
side  longitudinally  carinate.  Scutum  mi- 
crosculptured  anteriorly,  shiny  evanescent 
microsculpture  posteriorly.  Scutum  be- 
tween notaular  grooves,  striatopunctate, 
elsewhere  sparsely  punctate.  Scutellum 
shiny  anteriorly,  microsculptured  posteri- 
orly, with  a  foveolate  transverse  sulcus  on 
lateral  disc.  Mesopleuron  coarsely  areolate 
from  subalar  fossa  to  sternopleural  region. 
Scrobal  sulcus  indicated  by  slight  depres- 
sion. Hypersternaulus  and  omaulus 
coarsely  foveolate.  Mesosternum  micro- 
sculptured,  punctate  next  to  areas  of  are- 
olate sculpture  on  sternopleural  region. 
Metapleuron  microsculpture  evanescent 
with  a  few  irregular  longitudinal  carinae. 
Hind  wing  media  diverging  after  cu-a. 
Propodeum  coarsely  areolate,  except  area 
next  to  metapleuron  which  is  weakly  car- 
inate. 

Metasoma.  Terga  shiny,  without  mi- 
crosculpture, impunctate.  Sterna  shiny, 
without  microsculpture,  with  several 
punctures  on  lateral  margins. 

Color.  Ground  color  black.  White:  pro- 
notal lobe.  Yellow-brown:  palpi,  mandible 
except  apex,  antenna,  tegula,  wing  veins, 
stigma,  legs. 

Female.  Length  3.5  mm.  Similar  to  male 
except  as  follows:  clypeus  with  2  median 
teeth  separated  by  a  deep  semicircular 
emargination;  setose,  although  less  so 
than  male;  setae  partially  obscuring 
densely  punctate  surface.  Subantennal 
area  and  inner  orbits  sparsely  setose.  Inner 


orbits  margined  by  an  evanescent,  non- 
foveolate  carina.  Eye  elsewhere  not  mar- 
gined by  carina.  Frons  between  swellings 
and  lateral  ocelli,  shiny,  weakly  micro- 
sculptured  with  coarse  sparse  punctures 
(3  or  more  diameters  apart).  Vertex  with- 
out circular  depression  or  transverse  ridge 
but  with  frontal  line  continued  as  a  sulcus 
on  midline  posterior  to  ocelli.  OOD  1.7  X 
LOD.  Gena  with  coarse  sparse  punctures. 
Occipital  carina  simple,  not  raised  or  fo- 
veolate. Pronotum  with  teeth  less  promi- 
nent than  in  male.  Metasomal  terga  and 
sterna  shiny  with  sparse  punctures;  ster- 
num VI  densely  punctate.  Pygidial  plate 
narrow,  shiny,  bisected  by  a  longitudinal 
ridge,  with  several  coarse  punctures. 

Material  Examined  {16,  2  9). — Holotype: 
6,  PARAGUAY:  S.  Bernardino,  Mitte 
Decbr.,  Fiebrig  (NMW).  Paratypes:  BO- 
LIVIA: Km  335  de  Cochabamba  a  Santa 
Cruz,  Fritz  -Martinez  Col.  (9:  1  IIES). 
BRAZIL:  Estado  do  Ceara:  Ceara,  Baturite 
17-6-1908,  Ducke  (9:1  MPEG). 

Distribution  (Map  14). — Brazil:  Ceara. 
Bolivia,  Paraguay. 

2.  Aykhustigmus  patanawi  Finnamore, 
new  species 

Derivation  of  Name. — The  species  epithet 
is  derived  from  two  Quechua  terms, 
"pata"  meaning  margin  and  "nawi" 
meaning  eye,  in  reference  to  the  margined 
eye  of  this  species. 

Diagnosis. — Hind  wing  media  diverging 
before  cu-a  and  vertex  with  transverse  car- 
inate sculpture. 

Description. — Male.  Length  4.2  mm. 

Head.  Flagellomeres  without  tyli.  Clyp- 
eus and  subantennal  area  obscured  by 
dense  appressed  setae  that  extend  narrow- 
ly up  inner  orbit  to  a  point  about  IV3 
height  of  scape.  Eye  margined  by  a  coarse 
foveolate  carina  that  is  interrupted  ven- 
trally  at  base  of  mandible.  Carina  separat- 
ed from  mid  point  of  outer  orbit  by  a  dis- 
tance subequal  to  basal  width  of  mandi- 
ble. Frons  with  frontal  surface  impunctate, 
microsculptured;  irregular  transverse  ca- 


230 


Journal  of  Hymenoptera  Research 


Figs.  18-23.  Aykhustigmus  patanawi  6.  18,  head,  frontal  view.  19,  head  and  prothorax,  dorsal  view.  20,  mi- 
cropore field  between  lateral  ocellus  and  compound  eye.  21,  head,  lateral  view.  22,  mesosoma,  lateral  view. 
23,  mesosoma,  dorsal  view. 


rinae  laterally  next  to  inner  orbital  carina. 
Upper  frons  with  coarse  more  or  less  ir- 
regular striatopunctate  sculpture  radiating 
from  mid  ocellus,  punctures  less  than  1  di- 
ameter apart;  small  swelling  on  either  side 
of  mid  line  just  anterior  to  mid  ocellus. 
Vertex  densely  microsculptured  with 
large,  median,  circular  depression  poste- 


rior to  lateral  ocelli;  several  short  carinae 
radiating  from  lateral  ocellus.  Posterior 
vertex  with  a  series  of  3  or  4  transverse, 
striatopunctate  ridges  extending  between 
outer  orbits  posterior  to  orbital  carinae. 
OOD  0.9-1.3  X  LOD.  Gena  entirely  cari- 
nate  or  lower  area  areolate,  sculpture  in 
either  case  continuous  from  occipital  ca- 


Volume  4,  1995  231 

rina  to  hypostomal  carina  and  outer  orbit-  notch;  setae  dense,  obscuring  underlying 

al  margin.  Occipital  carina  considerably  sculpture.  Setae  less  dense  on  subantennal 

raised,  lamella-like,  coarsely  foveolate  to  area  and  inner  orbit.  Inner  orbit  margined 

mid  ventral  line.  by  a  non-foveolate  carina  that  extends  to 

Mesosoma.   Pronotum  with  transverse  vertex.  Outer  orbit  without  carina.  Frons 

carina  considerably  raised,  toothed  at  lat-  with  lower  area  impunctate,  densely  mi- 

eral  end;  carina  continued  ventrally  on  crosculptured;  upper  area  with  evanescent 

propodeal    side    to    anteroventral    corner  microsculpture,  shiny,  obscurely  striato- 

which  is  produced  anteriorly  as  tooth  sub-  punctate;    carinae,    if    present,    radiating 

equal  in  size  to  dorsolateral  tooth.  Prono-  from  mid  ocellus  anteriorly.  Frons  with 

turn  with  transverse  sulcus  longitudinally  small  swelling  on  either  side  of  mid  line, 

carinate;  setae  sparse,  not  obscuring  sculp-  Vertex  with  reduced  circular  depression 

ture.  Pronotal  side  with  irregular  longitu-  posterior  to  ocelli  (not  evident  in  1  speci- 

dinal  carinae.  Scutum  microsculptured  an-  men).    Vertex    densely    microsculptured, 

teriorly  and  in  depressions  medially  and  with  a  series  of  irregular,  transverse  fine 

posteriorly;  striatopunctate,  punctures  carinae  between  and  posterior  to  outer  or- 

coarse,    variable    from    relatively    sparse,  bits.    Gena    carinate    and    punctate    near 

about  1  diameter  apart  on  disc  to  nearly  mandible,  elsewhere  microsculptured. 

contiguous;  notaular  groove  deeply  im-  OOD  1.9-2.1  X  LOD.  Occipital  carina  sim- 

pressed,    foveolate,   continued   over   half  pie,    complete,    not    raised    or    foveolate. 

length  of  scutum.  Scutellum  microsculp-  Punctation  of  metasoma  as  in  male  except 

tured,  with  foveolate  transverse  sulcus  on  sternum    VI    densely    punctate.    Pygidial 

mid    lateral    disc.    Mesopleuron   coarsely  plate  narrow,  shiny,  bearing  a  lateral  row 

areolate  from  subalar  fossa  to  sternopleu-  of  punctures. 

ral   region.   Scrobal   sulcus   indicated   by  Material  Examined  {16,  3  9). — Holotype: 

slight    depression.    Hypersternaulus    and  6,  ARGENTINA:  B.  Aires,  La  Plata  (Fac. 

omaulus  coarsely  foveolate.  Mesosternum  Agronomia)    X-XI-68    C.    Porter    (MCZ). 

microsculptured,  sparsely  punctate.  Hind  Paratypes:  ARGENTINA  :  Buenos  Aires: 

wing  media  diverging  before  cu-a.  Meta-  Moreno,  Fritz  (9:1  IIES).  Moreno,  12-73 

pleuron  microsculptured,  few  short  cari-  Fritz  (9:1  IIES).  Tigre  XI-39  Viana  Leg. 

nae  on  posterior  margin.  Propodeum  mi-  (9:1  IIES). 

crosculptured  and  coarsely  areolate,  ex-  Distribution   Map   15. — Argentina:  Bue- 
cept  for  side  next  to  metapleuron  which  is  nos  Aires, 
without  areolae  and  partially  without  mi- 
crosculpture. ^-  Aykhustigmus  phasti  Finnamore, 

Metasoma.  Terga  shiny  with  slight  oily  ^^^  species 

sheen,  appearing  impunctate  on  anterior  Derivation  of  Name. — Phasti  is  a  Quech- 

terga  (punctures  minute,  sparse,  evanes-  ua  term  meaning  naked  or  bare  in  refer- 

cent)  but  punctures  increasing  in  size  and  ence  to  the  lack  of  microsculpture  on  the 

density  on  posterior  terga.  Sterna  shiny  scutum  and  the  resulting  naked  appear- 

with  slight  oily  sheen,  punctures  sparse  ance. 

throughout.  Diagnosis. — Male  unknown.  Female:  the 

Color.  Ground  color  back.  White:  pro-  absence  of  microsculpture  on  the  scutum 

notal  lobe.  Yellow-brown  to  brown:  palpi,  will  distinguish  this  species  from  others  in 

mandible  except  apex,  antenna,  legs,  teg-  the  genus.  In  addition  the  hind  wing  me- 

ula,  wing  veins,  stigma.  dia  diverges  after  cu-a,  the  vertex  is  with- 

Female.  Length  4.5  mm.  Similar  to  male  out  a  transverse  carinae,  and  the  clypeal 

except  as  follows:  clypeus  with  2  median  apex  has  2  teeth, 

teeth   separated   by   a   deep   semicircular  Descripition. — Female.  Length  4  mm. 


232  Journal  of  Hymenoptera  Research 

Head.  Clypeus  with  2  teeth  on  apical  sparse,  evanescent  on  anterior  terga,  more 

margin   separated  by   a   deep   U-shaped  apparent  on  posterior  terga.  Pygidial  plate 

notch;  surface  partially  obscured  by  dense  with   slight   raised   median   basal    ridge, 

appressed  setae  which  extend  up  suban-  Sterna    shiny,    sparsely    punctate    except 

tennal  area  and  narrowly  up  inner  orbit  to  sternum  VI  which  is  densely  punctate, 

about  %  height  of  scape.  Inner  orbit  with  Color.  Ground  color  black.  White:  man- 

non-foveolate  carina  that  is  evanescent  to-  dible  on  basal  third,  pronotal  lobe.  Yel- 

ward  vertex.  Outer  orbit  not  margined,  low-brown  to  brown:  palpi,  mandible  ex- 

Frons   microsculptured  on   scapal  basin,  cept  apex  and  basal  third,  antenna,  legs, 

impunctate;    upper    frons    more    or    less  tegula,  wing  veins,  stigma, 

shiny  with  evanescent  microsculpture.  Material  Examined  (39). — Holotype:    9, 

punctures  sparse.  Vertex  sometimes  with  ARGENTINA:    San   Isidro   1-1976   B.A. 

poorly  defined  depression  posterior  to  lat-  (PMA).  Paratypes:  ARGENTINA  :  Entre 

eral  ocelli  on  mid  line  or  depression  ab-  Rios:  Palmar  Colon  1-74  Fritz  (9:1  IIES). 

sent;  densely  microsculptured  in  contrast  VENEZUELA  :  Zulia:  6km  w.  La  concep- 

to  upper  frons.  OOD  1.8-1.9  X  LOD.  Gena  cion,  18-VI-1976  A.S.  Menke  &  D.  Vincent 

microsculptured,  punctate   toward   man-  (9:1  USNM). 

dibular  base.  Occipital  carina  simple,  com-  Distribution  (Map  16). — Argentina,  Ven- 

plete,  not  raised  or  foveolate.  ezuela.  The  Venezuelan  record  represents 

Mesosoma.   Pronotum  with  transverse  the  only  collection  locality  north  of  the 

carina    considerably    raised,    lateral    end  Brazilian   Highlands   for   the   genus   and 

forming  right  angle  that  continues  ven-  could   represent  an  introduction  of  this 

trally  on  carinate  pronotal  side  to  antero-  species  to  that  area, 
ventral  corner  which  is  produced   as   a 

small    tooth.    Pronotum   with    transverse  4.  Aykhustigmus  warawa  Finnamore, 
sulcus  longitudinally  carinate.  Scutum  ^^^  species 
shiny,   without   microsculpture   except  Derivation  of  Name. — The  species  epithet, 
weakly   at   extreme   anterolateral  corner,  warawa,    is    a    Quechua    term    meaning 
median  third  (between  notaular  grooves)  adorned    in    reference    to    the    elaborate 
striatopunctate;  lateral  area  punctate,  usu-  sculpture  of  this  species, 
ally  more  densely  punctate  next  to  notau-  Diagnosis. — The  combination  of  the  hind 
lar  groove.  Scutellum  shiny  or  weakly  mi-  wing    media    diverging    after    cu-a;    and 
crosculptured  anteriorly,  microsculptured  frons  without  swellings,  smoothly  round- 
posteriorly;  a  foveolate,  transverse  sulcus  ed  in  both  sexes;  and  the  female  clypeal 
on  mid  lateral  disc;  punctures  few,  present  edge  with  4  teeth,  and  scutum  microsculp- 
on  anterolateral  disc.  Mesopleuron  coarse-  tured  on  anterior  V4  or  more  will  separate 
ly  areolate  from  subalar  fossa  to  sterno-  this  species  from  all  others  in  the  genus, 
pleural  region.  Scrobal  sulcus  indicated  by  Description. — Male.  Length  3.0-4.5  mm. 
a  slight  depression.  Hypersternaulus  and  Head.  Flagellomeres  without  tyli.  Clyp- 
omaulus  coarsely  foveolate.  Mesosternum  eus   and   subantennal   area   obscured  by 
microsculptured,  sparsely  punctate.  Meta-  dense  appressed  setae  that  extend  up  in- 
pleuron  weakly  microsculptured,  a   few  ner  orbit  about  ll^  length  of  scape.  Inner 
short  carinae  on  posterior  margin.  Hind  orbit  margined  by   non-foveolate  carina 
wing  media  diverging  after  cu-a.  Propo-  extending  %  up  frons.  Outer  orbit  mar- 
deum  with  microsculpture  evanescent  or  gined  from  base  of  mandible  to  vertex  by 
absent  over  most  of  its  surface;  coarsely  foveolate  carina  or  evanescent  non-foveo- 
areolate,   except   next   to   metapleuron  late  ridge  in  smaller  specimens;  at  mid 
which  is  irregularly  carinate.  point  the  carina  is  separated  from  orbit  by 

Metasoma.    Terga    shiny;    punctures  distance  subequal  to  basal  width  of  man- 


Volume  4,  1995  233 

dible.  Lower  frons  microsculptured;  upper         Color.  Ground  color  black.  White:  pron- 

frons   shiny,   weakly   microsculptured,  otal  lobe.  Yellow-brown  to  brown:  palpi, 

punctate.    Vertex    microsculptured    with  mandibles  except  apically,  antenna,  legs, 

puncture-like  depression  on  mid  line  pos-  tegula,  wing  veins,  stigma, 
terior  to  lateral  ocelli;  transverse  evanes-         Female.  Length  4.0-4.5  mm.  Similar  to 

cent  ridge  connects  laterally  with  outer  or-  male  except  as  follows:  clypeus  with  4 

bital  carina.  Lateral  ocelli  closer  to  each  teeth  on  apical  margin,  lateral  teeth  locat- 

other  than  to  eye.  OOD  1.3-1.4  X  LOD.  ed  below  inner  antennal  socket  margin 

Gena  microsculptured,  punctate  near  base  and  often  obscured  by  overhanging  clyp- 

of  mandible.   Occipital   carina   complete,  eal  setae.  Surface  of  clypeus  and  subanten- 

foveolate,  not  raised.  nal  area  obscured  by  dense  appressed  se- 

Mesosoma.   Pronotum  with  transverse  tae  which  extend  up  inner  orbit  3/4  length 

carina  raised,  lateral  end  with  small  tooth;  of  scape.  Outer  orbit  unmargined.  Lower 

carina  continued  on  pronotal  side  to  an-  ggna  striatopunctate.  OOD  1.4-1.6  X  LOD. 

teroventral  corner  which  is  produced  as  Occipital    carina    simple,    complete,    not 

tooth  surpassing  in  size  that  of  dorsolat-  raised  or  foveolate.  Pygidial  plate  narrow, 

eral  tooth.  Pronotum  with  transverse  sul-  shiny,   sparsely  punctate.   Sternum   VI 

cus  longitudinally  carinate;  setae  sparse,  densely  punctate. 

not    obscuring    sculpture.    Pronotal    side         Material   Examined   (13(?,   13  9).— Holo- 

with  irregular,  longitudinal  carinae.  Scu-  ^yp^.  ^^  ^rq   [ARGENTINA]:  Salta:  Ro- 

tum   with   microsculpture  variable   from  ^^^^^  ^^^j  Lerma  10-84  Fritz  (IIES).  Para- 

densely   microsculptured   to   microsculp-  ^^^^^.  ARGENTINA:  Buenos  Aires:  48- 

tured  on  anterior  margin  only;  punctures  889  (9:  1  MACN).  Burzaco  Sea  L.  12-13- 

present  on  median  third  (between  notau-  iji.2974  c  R.  Vardy  B.M.  1974-204  (9:1 

lar  grooves  which  extend  posteriorly  as  pMNH).  Moreno,  M.A.  Fritz  (d:  1  IIES). 

somewhat  polished  low  ridges);  lateral  ar-  Moreno  1-73,  Fritz  (9:2  IIES).  La  Plata 

eas  impunctate  to  sparsely  punctate;  pos-  ^^^^  Agronomia)  X-XI-68,  C.  Porter  {6:1 

terior  margin  and  often  median  area  stria-  j^^^).  Jujuy:  Palpala  11-1968,  M.  Aczel 

topunctate.  Scutellum  shmy,  weakly  mi-  ^^^  ^  ^^^^  ^^^^^,  p^^.^^^  ^2-56,  Fritz  {6: 

crosculptured  anteriorly,  punctured;  pos-  ^  ^^^^^.^  ^^^^  P^^^^  ^^_^^^  P^.^^ 

terior     microsculptured,     jn^P^^nctate;  (^,  4  HES).  Urundel  26-1-950,  R.  Golbach 
without  foveolate  sulcus  on  disc  laterally.  p^pp^    ^^  ^^^  ^^^^^_  ^^^^  ^_ 

Mesopleuron  coarsely  areolate  from  sub-  1^    ^,      .  .n     1   ?■,T-c-^    t  t 

,      /        ,      ,  1         1        •        c       u  1  74,  Martinez  (9:1  IIES).  Tucuman:  Las 

alar  fossa  to  sternopleural  region.  Scrobal  '  v.t  c^  r^  1  ur     /  j^    1  ttttcx   c 

sulcus    indicated    by    slight    depression.  ^^S^^^^^/J'^^l.^f^f  ^"  ^f /i  "^^  \^ 

Hypersternaulus   and   omaulus   coarsely  Janvier  21-X-1950,  M.  Aczel  (d:  1  LILLO). 

foveolate.  Mesosternum  microsculptured,  ^an   Miguel,  Tucuman   18-XI-1979,   L.A. 

sparsely    punctate.    Metapleuron    shiny,  Stange  (d:  1  FSCA).  San  Pedro  de  Cola- 

without  or  with  evanescent  microsculp-  1^«'  Foerster  (9:1  IIES).  Trancas,  Fritz  {6: 

ture,  several  short  carinae  on  posterior  1  "ES,  9:  1  IIES).  BOLIVIA:  Beni:  Ichilo 

margin.  Hind  wing  media  diverging  after  Yapacani  2-71,  Fritz  ((5:1  IIES,  9:  1  IIES). 

cu-a.      Propodeum      microsculptured,  BRAZIL:  Santa  Catarina:  Nova  Teutonia 

coarsely  areolate,  except  for  side  next  to  Feb.  1957,  F.  Plaumann  (9:2  OSU).  Sao 

metapleuron    which    is    shiny,    unsculp-  Paulo:   Campinas   Mar.    1924,   F.X.    Wil- 

tuj-ed.  Hams  {6:1  BISH).  Unplaced:  Minas,  Ja- 

Metasoma.  Terga  shiny  with  slight  oily  care  Dec.  1-1907  Carn.  Mus.  Ace.  3533  ( 9 : 

sheen,  punctures  sparse  and  evanescent.  1  CARN). 

Sterna  similar  to  terga  except  punctures         Distribution  (Map  17).— Argentina,  Bo- 
more  dense.  livia,  southern  Brazil. 


234 


Journal  of  Hymenoptera  Research 


Figs.  24-29.  Aykhustigmus  warawa  9 .  24,  head,  frontal  view.  25,  head  and  prothorax,  dorsal  view.  26,  micro- 
pore field  between  lateral  ocellus  and  compound  eye.  27,  head,  lateral  view.  28,  mesosoma,  lateral  view.  29, 
mesosoma,  dorsal  view. 


INCASTIGMUS  Finnamore,  new  genus 
(Figs.  30-35) 

Derivation  of  Name. — Incastigmus  is 
named  after  the  indigenous  Inca  people  of 
South  America. 

Diagnosis. — The  presence  of  a  median 
groove  or  posteromedian  pit  on  the  scu- 
tum will  separate  this  genus  from  all  oth- 
ers in  the  Stigmina. 


Description. — Head.  Labrum  quadri- 
lobed  with  narrow,  deep  median  emargi- 
nation.  Mandible  with  apex  in  both  sexes 
tridentate;  apicoventral  tooth  in  female 
acute;  inner  basal  tooth  absent.  Clypeal 
apex  in  male  without  bevelled  modifica- 
tions; with  4  teeth  in  female.  Interantennal 
tubercle  absent.  Frontal  line  absent.  Micro- 
pore field  present.  Inner  orbits  converging 


I 


Volume  4,  1995 


235 


below.  Eye  at  most  partially  margined  by 
a  carina.  Occipital  carina  complete,  not 
joining  hypostomal  carina,  simple  in  fe- 
male; raised,  foveolate  in  male. 

Mesosoma.  Scutum  with  a  median 
groove  or  at  least  a  posteromedian  pit;  no- 
taular  groove  usually  elongate.  Acetabular 
carina  intercepting  omaulus.  Scrobal  sul- 
cus present.  Hypoepimeral  area  without 
coarse  sculpture.  Mid  basitarsus  of  male 
elongate,  as  long  as  next  3  tarsomeres 
combined.  Posterior  margin  of  hind  tibia 
with  2  or  3  spines.  Fore  wing  without  se- 
tae in  cellular  area.  Hind  wing  media  di- 
verging before  cu-a;  submedian  cell  of 
normal  size,  not  reduced. 

Metasoma.  Petiole  carinate.  Pygidial 
plate  narrow,  absent  in  male.  Digitus  elon- 
gate relative  to  cuspis,  clubbed. 

Type  Species. — Incastigmus  inti  new  spe- 
cies. 

Distribution. — Neotropical. 

Species  Transferred. — Stigmus  hexagonalis 
Fox,  S.  neotropiciis  Kohl,  S.  smithii  Ash- 
mead,  and  S.  thorncicus  Ashmead.  The  25 
species  of  Incastigmus  are  the  subject  of  the 
forthcoming  Part  2  of  this  revision. 

Incastigmus  inti  Finnamore,  new  species 

Derivation  of  Name. — Inti  is  a  Quechua 
term  meaning  sun  in  reference  to  the  cen- 
tral position  of  this  species  as  type  for  the 
genus. 

Diagnosis. — The  combination  of  the  fol- 
lowing characters  will  separate  this  spe- 
cies from  all  others  in  the  genus.  Micro- 
pore field  on  vertex  oval.  Scutum  with 
median  and  notaular  grooves  complete 
from  anterior  to  posterior  margin,  without 
regular  longitudinal  carinae  between 
grooves.  Pronotal  lobe  white,  rounded 
conical,  not  flattened  or  toothed  or  dark. 
Male  antenna  with  tylus  on  apical  flagel- 
lomere.  Female  clypeus  with  2  elongate 
setae  issuing  from  2  narrowly  separated 
pits  on  median  lobe.  Female  frons  shiny, 
without  microsculpture  anterolaterally  to 
mid  ocellus. 

Description. — Male.  Length  2.5-4.0  mm. 


Head.  Flagellomeres  without  special- 
ized setae;  II-XI  with  tyli,  that  on  flagel- 
lomere  XI  imparting  asymmetrical  shape; 
basal  flagelk^mere  length  2.2  X  apical 
width;  penultimate  flagellomere  length  1.4 
X  apical  width.  Head  microsculptured  on 
lower  frons,  mid  frons  and  vertex  but 
shiny  on  upper  lateral  frons,  punctures 
sparse.  Clypeus  obscured  by  dense  ap- 
pressed  setae  which  extend  up  inner  orbits 
little  more  than  height  of  antennal  socket. 
Micropore  field  present  as  discrete  patch. 
Ocelli  closer  to  each  tether  than  to  com- 
pound eyes.  OOD  1.7  X  LOD.  Gena  mi- 
crosculptured, sparsely  punctate,  non-car- 
inate,  without  ventral  tooth  or  swelling. 
Occipital  carina  foveolate. 

Mesosoma.  Transverse  carina  of  prono- 
tum  toothed  laterally,  continued  ventrally 
to  pronotal  side  and  terminating  in  a  ven- 
trolateral tooth;  transverse  sulcus  longitu- 
dinally carinate.  Pronotal  side  carinate, 
lobe  conical  with  rounded  apex.  Scutum 
entirely  microsculptured  to  weakly  mi- 
crosculptured with  shiny  patches;  notau- 
lar groove  complete  to  posterior  margin; 
median  groove  complete  to  anterior  mar- 
gin; punctures  sparse.  Scutellum  micro- 
sculptured.  Mesopleuron  variable  from 
entirely  microsculptured  to  weakly  shiny 
with  reduced  microsculpture,  sparsely 
punctate  to  impunctate.  Preomaular  area 
with  sparse  setae  that  do  not  obscure  un- 
derlying sculpture.  Hypersternaulus, 
omaulus  and  scrobal  sulcus  foveolate.  Me- 
tapleuron  usually  microsculptured.  Pro- 
podeum  shiny,  generally  without  micro- 
sculpture,  uniformly  coarsely  areolate  ex- 
cept anterolaterally  without  sculpture. 

Metasoma.  Terga  shiny,  microsculpture 
and  punctures  evanescent  (appearing  im- 
punctate). Sterna  weakly  microsculptured, 
punctures  sparse. 

Color.  Ground  color  black.  White:  man- 
dible medially,  pronotal  lobe.  Yellow- 
brown:  palpi,  mandible  subapically,  an- 
tenna, fore  leg  except  coxa,  mid  leg  except 
coxae,  hind  trochanter  and  tarsus,  tegula, 
metasomal  sternum  VIII. 


236 


Journal  of  Hymenoptera  Research 


Figs.  30-35.  Incastigmiis  inti.  30,  9  head,  frontal  view.  31,  S  head  and  prothorax,  dorsal  view.  32,  6  micropore 
field  between  lateral  ocellus  and  compound  eye.  33,  i  mesosoma,  dorsal  view.  34,  9  mesosoma,  lateral  view. 
35,  9  mesosoma,  dorsal  view. 


Female.  Length  3.0-4.25  mm.  Similar  to 
male  except  as  follows:  antenna  without 
tyli,  penultimate  flagellomere  length  1.25 
X  apical  width.  Clypeus  shiny,  usually 
sparsely  punctate,  occasionally  moderate- 
ly dense  punctation  grouped  in  median 
area;  median  lobe  with  2  elongate  setae  is- 
suing from  2  narrowly  separated  pits; 
apex  with  truncate  median  lobe.  Scutum 
with  several  irregular  carinae  between  no- 


taular  and  median  grooves.  Color  as 
above  but  variable  to  dark  antennal  fla- 
gellum  and  dark  femora. 

Material  Examined  (258c5  109  9).— Holo- 
type:  S,  ECUADOR:  Napo  Prov.  Tena, 
sweep  15-11-1986  A.T.  Finnamore  (PMA). 
Paratypes:  BOLIVIA:  La  Paz:  Chulumani 
1,700m  25-III-1979  M.  Cooper  B.M.  1979- 
216  (9:  2  BMNH).  Chulumani  1,700m  30- 
III-1979  M.  Cooper  B.M.  1979-216  (9:1 


Volume  4,  1995 


237 


BMNH).  Chulumani  1,700m  31-III-1979  M. 
Cooper  B.M.  1979-216  {6:  1  BMNH).  Chu- 
lumani 1,700m  3-IV-1979  M.  Cooper  B.M. 
1979-216  {6:1  BMNH.  9:  2  BMNH).  Co- 
roico-Chulomani  III-7-1968  Garcia  &  Por- 
ter (9:  1  MCZ).  Coroico,  Sta.  Barbara 
1100m  4-1-1975  (9:  1  PMA).  Tumupasa 
Dec.  W.M.  Mann,  Mulford  Biol.  Expd. 
1921-1922  {6:1  USNM).  Yungas,  13km  s. 
Caranavi  850m  27-III-1973  J.  Helava  (9:2 
PMA).  Unplaced:  Las  Juntas  Dec.  1913, 
Steinbach  (9:  1  CARN).  BRAZIL:  Mato 
Grosso:  Itaum  111-1974  M.  Alvarenga  (9: 
3  AEl).  Para:  Belem  IPEAN  Xll-1-4-1969 
J.M.  &  B.A.  Campbell  (9:  1  BRD).  CO- 
LOMBIA: Caqueta:  Florencia  480m  31-X- 
1971  M.  Cooper  B.M.  1972-275  (9:  1 
BMNH).  Yuruyaco,  73k  sw.  Florencia  3-II- 
1979  M.  Cooper  B.M.  1979-106  (9:  1 
BMNH).  Yuruyaco,  73k  sw.  Florencia  12- 
11-1979  M.  Cooper  B.M.  1979-106  {6:  I 
BMNH).  Yuruyaco,  73k  sw.  Florencia  13- 
11-1979  M.  Cooper  B.M.  1979-106  {6:  1 
BMNH).  Putumayo:  Mocoa  13-IV-1974  M. 
Cooper  B.M.  1974-327  {6:1  BMNH).  Mo- 
coa 16-1V-1974  M.  Cooper  B.M.  1974-327 
(9:5  BMNH).  Mocoa  18-IV-1974  M.  Coo- 
per B.M.  1974-327  {6:1  BMNH).  Mocoa 
19-rV-1974  M.  Cooper  B.M.  1974-327  {6:1 
BMNH).  Mocoa  14-VI-1974  M.  Cooper 
B.M.  1974-548  {6:1  BMNH).  Mocoa  24- 
Vl-1974  M.  Cooper  B.M.  1974-548  {6:  2 
BMNH).  Mocoa  26-V1-1974  M.  Cooper 
B.M.  1974-548  (9:2  BMNH).  Mocoa  16- 
VII-1978  M.  Cooper  B.M.  1978-431  (9:1 
BMNH).  Mocoa  7-IX-1974  M.  Cooper  B.M. 
1975-33  {6:  1  BMNH).  Mocoa  27-IX-1974 
M.  Cooper  B.M.  1975-33  {6:  1  BMNH). 
Mocoa  600m  26-111-1976  M.  Cooper  B.M. 
1976-290  {6:1  BMNH).  Mocoa  18-111-1978 
M.  Cooper  B.M.  1978^31  {6:  1  BMNH). 
Villa  Garzon,  8  mi  s.  Mocoa  3-V1II-1978  M. 
Cooper  B.M.  1978^31  {6:1  BMNH).  Vau- 
pes:  Mitu  ll-V-1974  M.  Cooper  B.M. 
1974-327  {6:1  BMNH).  ECUADOR:  Car- 
chi:  Chical  1250m  0-56N  78-1 IW,  1-VlII- 
1983  J.  Rawlings,  M.  Smyers  (9:1  CARN). 
Napo:  Coca,  May  1965  Luis  Pefia  {6:  1 
AEl).   Coca   &   Napo   Rivers,   V-1-12-65, 


Luis  Pefia  {6:1  AEl).  Limoncocha  250m, 
15-28- VI-1976  S.  &  J.  Peck  {6:  2  BRD,  9: 
2  BRD).  Misahualli,  down  Rio  Napo  19-11- 
1983  L.  Huggert  {6:15  PMA,  9:  2  PMA). 
Muyuna,  5km  w.  Tena  29-IX-1978  M.  Coo- 
per B.M.  1979-20  (9:  1  BMNH).  Puerto 
Misahualli  350m  11-1983  Sharkey  {6:  2 
PMA,  9:  3  PMA).  Puerto  Misahualli, 
30km  e.  350m  11-1983  Sharkey  {6:  9  PMA). 
Santa  Cecilia  III-25-31-1969  P.  &  P.  Span- 
gler  ((5:1  USNM).  Tena  9-14-II-1971  M. 
Cooper  B.M.  1972-275  {6:1  BMNH).  Tena 
8-VII-1976  S.  &  J.  Peck  (9:3  PMA).  Tena 
400m  11-1983  M.  Sharkey  {6:6  PMA,  9:  1 
PMA).  Tena,  sweep  15-11-1986  A.T.  Fin- 
namore  {6:  12  PMA,  9:  3  PMA).  Tena, 
sweep  18-11-1986  A.T.  Finnamore  (9:  1 
PMA).  Tena,  12km  sw.  500m  8-11-VII- 
1976  S.  &  J.  Peck  (9:1  BRD).  Tena-Puyo 
Hwy.  5km  n.  Santa  Clara  21-11-1986 
sweep,  T.  Thormin  &  J  Wojcicki  {6:  1 
PMA).  Pastaza:  Puyo  960m  1-8-X-1970  ]. 
&  M.  Sedlacek  {6:  8  BISH).  Puyo  900- 
960m  1-8-X-1970  J.  &  M.  Sedlacek  {6:7 
BISH).  Puyo  22km  sw.  900m  14-16-VII- 
1976  S.  &  J.  Peck,  forest  {6:  1  BRD,  9:  2 
BRD).  Puyo  23km  se  19-V-1977  P.J.  Span- 
gler  &  D.R.  Givens  #58  (9:  1  USNM). 
Puyo  44km  s.  21  May  1977  DL  &  SS  Vin- 
cent {6:  5  USNM,  9:  2  USNM).  Puyo 
18km  n.  1100m  14-8-1982  R.  Hensen  &  \. 
Aptroot  {6:1  RNH).  Pichincha:  Tinalan- 
dia  800m  111-1983  L.  Masner  &  M.  Sharkey 
{6:2  PMA).  Zamora:  Rio  Jumboe  1-IV- 
1965  Pefia  {6:5  MCZ,  9:  2  MCZ).  Zamora 
lV-4-1965  Pena  {6:  1  AEl).  Unplaced: 
Cumbaratza  (E)  Xl-21-1970  Luis  E.  Peha 
{6:  1  AEl).  Mera  26-1-1923  F.X.  Williams 
{6:1  BISH).  PARAGUAY:  Guaira:  w.  Vil- 
larica,  Caballero,  1-72  Pena  {6:1  IlES).  Un- 
placed: Piareta  12-71  Pena  {6:  1  IIES). 
PERU:  Cuzco:  Agua  Calliente  21- 
28-X1I-1983  L.  Huggert  {6:  8  PMA,  9: 
IPMA).  Machu  Picchu  l-XII-1965  H.  &  M. 
Townes  (9:1  AEl).  Qunicemil,  750m  nr. 
Marcapata  September  1962  Luis  Pena  {6: 
8  AEl,  9:  3  AEl).  Huanuco:  Cayumba, 
35km  s.  Tingo  Maria  800m  l-XI-1973  J.M. 
Schunke  B.M.  1974-37  (9:1  BMNH).  Las 


238 


Journal  of  Hymenoptera  Research 


Palmas  5km  sw.  1000m  X-16-1954  E.I. 
Schlinger  &  E.S.  Ross  {6:2  CAS).  Las  Pal- 
mas lOmi  sw.  1000m  IX-26-1954  E.I. 
Schlinger  &  E.S.  Ross  (9:4  CAS).  Monson 
Valley,  Tingo  Maria  IX-21-1954  E.I. 
Schlinger  &  E.S.  Ross  ((5:1  CAS).  Monson 
Valley,  Tingo  Maria  23-IX-1954  E.I.  Schlin- 
ger &  E.S.  Ross  (c?:  6  CAS,  9:  3  CAS). 
Monson  Valley,  Tingo  Maria  X-10-1954 
E.I.  Schlinger  &  E.S.  Ross  {6:4:  CAS,  9:  3 
CAS).  Monson  Valley,  Tingo  Maria  X-19- 
1954  E.I.  Schlinger  &  E.S.  Ross  (9:1  CAS). 
Monson  Valley,  Tingo  Maria  X-21-1954 
E.I.  Schlinger  &  E.S.  Ross  {6:1  CAS,  9:  1 
CAS).  Monson  Valley,  Tingo  Maria  X-26- 
1954  E.I.  Schlinger  &  E.S.  Ross  (9:  1  CAS). 
Monson  Valley,  Tingo  Maria  XI-21-1954 
E.I.  Schlinger  &  E.S.  Ross  {6:  1  CAS). 
Monson  Valley,  Tingo  Maria  XI-29-1954 
E.I.  Schlinger  &  E.S.  Ross  {6:1  CAS,  9:  1 
CAS).  Monson  Valley,  Tingo  Maria  XII-2- 
1954  E.I.  Schlinger  &  E.S.  Ross  (9:1  CAS). 
Tingo  Maria  26-29-1-1984  L.  Huggert  {6: 
14  PMA,  9:  3  PMA).  Tingo  Maria  30-1- 
1984  L.  Huggert  {6:  6  PMA,  9:  1  PMA). 
Tingo  Maria,  1km  e.  15-VIII-1971  P.S.  & 
H.L.  Broomfield.  MT  dense  woodland 
B.M.  1971-486  (9:1  BMNH).  Tingo  Maria, 
26  mi.  e.  XII-10-54  1100m  E.I.  Schlinger  & 
E.S.  Ross  (9:1  CAS).  Tingo  Maria,  67  mi. 
e.  X-4-54  350m  E.I.  Schlinger  &  E.S.  Ross 
(9:1  CAS).  Tocache  2-II-1984  L.  Huggert 
{6:  4  PMA,  9:  1  PMA).  Junin:  Colonia 
Perene,  Rio  Perene  18  mi  ne  La  Merced  I- 
3-55  E.I.  Schlinger  &  E.S.  Ross  {6:2  CAS, 
9:  1  CAS).  Paratuchali  22-1-1984  L.  Hug- 
gert {6:10  PMA,  9:  4  PMA).  Satipo  18-1- 
1984  L.  Huggert  {6:10  PMA,  9:  1  PMA). 
Satipo  19-24-1-1984  L.  Huggert  {6:  11 
PMA,  9:  6  PMA).  Lima:  Magdalena  [del 
Mar?]  [Lima?]  Mar  28-10  CHT  Townsend 
(9:1  USNM).  Loreto:  Iquitos,  ne.  Rio  Na- 
nay  6-n-1984  L.  Huggert  {6:  10  PMA). 
Iquitos,  Gransa  UNAP  9-II-1984  L.  Hug- 
gert {6:2  PMA).  Iquitos,  Quisto  Cocha  5- 
11-1984  L.  Huggert  {6:  1  PMA).  Iquitos, 
Barilla  10-11-1984  L.  Huggert  {6:1  PMA). 
Madre  de  Dios:  Laberinto,  70km  w.  Pto. 
Maldonado  on  Rio  Madre  de  Dios  1-2-1- 


1984  A.T.  Finnamore  (9:  1  PMA).  Pto. 
Maldonado  1-11-1-1984  L.  Huggert  {6:50 
PMA,  9:  14  PMA).  Tambopata  Reserve, 
50km  s.  Pto.  Maldonado  on  Rio  Tambo- 
pata 3-8-1-1984  A.T.  Finnamore  {6:  1 
PMA,  9:  1  PMA).  Ucayali:  Tacshitea, 
88km  n.  Pucallpa  jet.  Rio  Callaria  &  Rio 
Ucayali  22-25-1-1984  A.T.  Finnamore  {6:1 
PMA).  VENEZUELA:  Zulia:  El  Tucuco, 
45km  sw.  Machiques,  5-6-VI-1976  A.S. 
Menke  &  D.  Vincent  (9:1  USNM). 

Distribution  (Map  18). — Bolivia,  Brazil, 
Ecuador,  Paraguay,  Peru,  Venezuela. 

Incastigmus  thoracicus  (Ashmead), 
new  combination 

Stigmus  thoracicus  Ashmead  1900:223  9.  Holo- 
type*:  9,  St.  Vincent  W.I.,  H.  Smith  238/ W. 
Indies  99-331  (BMNH  type  Hym  21.885). 

Stigmus  smithii  Ashmead  1900:223  2 .  New  syn- 
onymy. Holotype*:  S,  W.  Indies  (BMNH 
type  Hym  21.886).  99-331.  Ashmead  incor- 
rectly sexed  the  type  specimen  when  he  de- 
scribed sniitliii  based  on  a  single  female.  The 
holotype  is  a  male. 

Diagnosis. — The  red  petiole  is  sufficient 
to  distinguish  this  species  in  lighter  col- 
ored specimens.  Darker  (and  lighter)  spec- 
imens are  distinguished  by  the  extreme  re- 
duction of  the  median  scutal  groove,  shiny 
hypoepimeral  area,  and  rounded  corners 
of  the  transverse  pronotal  carina. 

Description. — Male.  Length  3.0-3.8  mm. 

Head.  Flagellomeres  without  tyli  or  spe- 
cialized setae;  basal  flagellomere  length 
2.0  X  apical  width;  penultimate  flagello- 
mere length  1.1  X  apical  width.  Head  al- 
most uniformly  microsculptured;  upper 
frons  and  vertex  with  slightly  less  micro- 
sculpture  and  with  sparse  obscure  punc- 
tures. Clypeus  obscured  by  dense  ap- 
pressed  setae  which  extend  broadly  up  in- 
ner orbits  to  height  of  scape.  Micropore 
field  present  as  discrete  patch.  Ocelli  clos- 
er to  each  other  than  to  eyes.  OOD  2.0  X 
LOD.  Gena  microsculptured,  sparsely 
punctate,  non-carinate,  without  ventral 
swelling  or  tooth.  Occipital  carina  foveo- 
late. 


Volume  4,  1995  239 

Mesosoma.   Pronotum  with  transverse  Female.  Length  3.9  mm.  Similar  to  male 

carina  rounded  laterally,  not  toothed  or  except   as    follows:   basal    flagellomere 

produced;  transverse  sulcus  with  evanes-  length  3.0  X  apical  width.  Clypeus  shiny, 

cent  longitudinal  carinae;  pronotal   lobe  with  several  punctures,  setae  sparse;  clyp- 

normal,  not  toothed;  side  with  several  ev-  eal  apex  with  median  teeth  separated  by 

anescent  carinae.  Scutum  shiny,  weakly  shallow  emargination;  median  lobe  with  2 

microsculptured;  notaular  groove  present  elongate  setae  issuing  from  2  narrowly 

anteriorly,  not  elongate;  median  groove  separated  pits.   Inner  orbits  sparsely  se- 

absent,  pit  evident  posteriorly  as  one  of  tose,  not  obscuring  sculpture.  OOD  2.5  X 

many  evanescent  fovea  in  transverse  row  LOD.  Color,  as  above  for  light  form  except 

next  to  posterior  margin;  punctures  min-  white  on  scape  and  orange  on  clypeus  and 

ute,  sparse.  Scutellum  and  mesopleuron  propodeal  dorsum. 

shiny,   without   microsculpture;   impunc-  Material  Examined  {66,  59). — DOMIN- 

tate  except  sternopleural  region  with  min-  ICA:  Pont  Casse;  Springfield;  Springfield 

ute    sparse    punctures;   preomaular    area  Plantation.  GRENADA:  2500  feet.  Baltha- 

with  sparse  setae;  hypersternaulus  with-  zar,  windward  side;  Botanical  Garden.  ST. 

out  foveae;  scrobal  sulcus  and  omaulus  VINCENT:  (no  other  data), 

weakly    foveolate.    Metapleuron    micros-  Distribution  (Map  19). — Dominica,  Gre- 

culptured  on  ventral  half,  otherwise  shiny,  nada,  St.  Vincent, 
impunctate.    Propodeum    shiny,    without 

microsculpture  over  most  of  basolateral  LLAQHASTIGMUS  Finnamore, 

area  and  dorsolateral  spheres;  basolateral  new  genus 

area    adjacent    to    metapleuron    without  ^  ^§^'      ~     -' 

sculpture;  dorsal  enclosure  areolae  eva-  Derivation  of  Generic  Name. — Llaqha  is  a 

nescent,  areolae  small  relative  to  larger  ev-  Quechua  term  meaning  dark,  in  reference 

anescent  areolae  of  dorsolateral  spheres,  to  the  generally  black  coloration  of  species 

the  2  groups  of  areolae  separated  by  a  in  this  genus, 

smooth  unsculptured  area.  Diagnosis. — Females  of  this  genus  can  be 

Metasoma.    Terga    shiny,   without   mi-  separated  from  all  others  in  the  Stigmina 

crosculpture;    punctures    sparse,    evanes-  by    the   enlarged,    truncate,    apicoventral 

cent.    Sterna   weakly   microsculptured,  mandibular  tooth.  Females  in  other  genera 

punctures  sparse  but  increasing  in  density  have  an  acute  apicoventral  tooth.  Males 

on  posterior  metasoma.  can  be  separated  with  a  combination  of  a 

Color.  Light  form:  ground  color  black,  tridentate  mandible  and  a  scutum  without 

White:  palpi,  mandible  except  apex,  spot  a  trace  of  a  posteromedian  pit  or  groove, 

on  pronotal  lobe,  fore  tibia  and   tarsus.  Males  in  other  genera  have  a  bidentate 

mid  tibia  and  tarsus,  hind  tibia  on  basal  mandible  or  at  least  a  posteromedian  pit 

half  or  more  and  tarsus.  Yellow-orange:  on  the  scutum. 

antenna,  mesosoma  except  propodeal  dor-  Description. — Head.  Labrum  bilobed 
sum,  fore  leg  basal  to  tibia,  mid  leg  basal  with  a  small  median  notch  (labrum  quad- 
to  tibia,  hind  leg  basal  to  tibia  and  apex  of  rilobed  in  2  species  both  of  which  lack  a 
tibia,  petiole  except  ventral  apex,  apex  of  subomaulus).  Mandible  with  apex  triden- 
1  or  more  sterna.  Dark  form:  ground  color  tate;  apicoventral  tooth  in  female  en- 
black.  White:  palpi,  mandible  except  apex,  larged,  apex  truncate;  without  inner  basal 
spot  on  pronotal  lobe,  basal  ring  of  hind  tooth.  Clypeal  apex  unmodified  in  male, 
tibia.  Yellow-brown:  antenna,  pronotum  without  bevelled  edge;  in  female  with  4 
except  dorsally  and  spot  on  pronotal  lobe,  teeth.  Interantennal  tubercle  absent.  Fron- 
mesopleuron  anteriorly,  fore  leg,  mid  leg,  tal  carina  absent.  Vertex  with  micropore 
hind  trochanter  and  tarsus,  tegula.  field   between    lateral   ocellus    and    com- 


240 


Journal  of  Hymenoptera  Research 


pound  eye.  Inner  orbits  converging  below. 
Eye  not  margined  by  carina.  Occipital  ca- 
rina complete,  not  joining  hypostomal  ca- 
rina, simple  in  female,  foveolate  in  male, 
otherwise  unmodified. 

Mesosoma.  Scutum  without  trace  of  me- 
dian groove  or  posteromedian  pit;  notau- 
lar  groove  normal,  not  elongate.  Acetab- 
ular carina  intercepting  omaulus.  Scrobal 
sulcus  often  evanescent.  Hypoepimeral 
area  without  coarse  sculpture.  Mid  basi- 
tarsus  of  male  elongate,  as  long  or  longer 
than  next  3  tarsomeres  combined.  Poste- 


rior margin  of  hind  tibia  with  2  or  3 
spines.  Fore  wing  without  setae  in  cellular 
area.  Hind  wing  media  diverging  before 
cu-a;  submedian  cell  of  normal  size,  not 
reduced. 

Metasoma.  Petiole  carinate.  Pygidial 
plate  narrow,  absent  in  male.  Digitus  elon- 
gate. 

Ti/pe  Species. — Llaqhastig?nus  jatunkirus 
Finnamore,  new  species. 

Distribution  (Map  37). — Neotropical,  re- 
stricted to  South  America  (subtropical 
highland  or  montane  forest?). 


KEY  TO  SPECIES  OF  LLAQHASTIGMUS 

1  Male     2 

1'  Female     15 

2  Gena  ventrolaterally  and  sternopleural  region  with  relatively  dense,  elongate  setae;  Peru 

13.  shachus  Finnamore,  new  species 

2'  Gena  and  sternopleural  region  sparsely  pubescent  with  short  inconspicuous  setae     ....       3 

3  Preomaular  area  densely  setose,  obscuring  sculpture;  mesosoma  often  extensively  red;  se. 
Brazil 8.  mantanti  Finnamore,  new  species 

3'  Preomaular  area  sparsely  setose;  mesosoma  black  except  pronotal  lobe  which  is  usually 
white    4 

4  Scape,  fore  femur,  and  pronotal  lobe,  black;  mesosoma  with  scrobal  sulcus  foveolate; 
clypeus  sparsely  setose,  underlying  punctures  visible;  Ecuador,  Colombia    

3.  chiitiyana  Finnamore,  new  species 

4'  One  or  more  of  scape,  fore  femur,  or  pronotal  lobe,  yellow-brown  or  white;  mesosoma 
with  scrobal  sulcus  usually  smooth,  rarely  foveolate;  clypeus  usually  obscured  by  dense 
appressed  setae     5 

5  Subomaulus  absent     6 

5'  Subomaulus  present   7 

6  Petiole  with  3  or  4  coarse  lateral  carinae;  Bolivia   2.  australis  Finnamore,  new  species 

6'  Petiole  microcarinate  laterally;  Ecuador,  Colombia     

11.  santanderamis  Finnamore,  new  species 

7  Scape  brown  with  yellow-brown  at  base  and  apex  only;  flagellum  and  femora  black; 
propodeum  with  area  adjacent  to  enclosure  finely  carinate,  lateral  spheres  without  large 
areolae;  transscutellar  sulcus  with  foveae  small  or  absent;  scrobal  sulcus  not  foveolate, 
only  slightly  impressed;  Ecuador     5.  ecuatorialis  Finnamore,  new  species 

7'  Scape  yellow-brown  at  least  ventrally;  one  or  more  femora  usually  yellow  to  yellow- 
brown;  propodeum  with  area  adjacent  to  enclosure  irregularly  sculptured,  lateral  spheres 
coarsely  areolate;  transscutellar  sulcus  usually  with  prominent  foveae;  scrobal  sulcus  of- 
ten foveolate   8 

8  Head  and  mesosoma  coarsely,  uniformly  microsculptured,  dull;  microsculpture  of  frons 
not  distinct  from  that  of  vertex;  scutum  with  small  evanescent  punctures;  Peru,  Ecuador, 
Venezuela    9.  tmithiis  Finnamore,  new  species 

8'  Vertex,  anterior  to  mid  ocellus,  more  shiny  than  frons  (less  microsculpture)  and/or  me- 
sosoma with  mid  and  lower  mesopleuron  shiny,  without  or  with  little  microsculpture; 
punctation  of  scutum  distinct  or  absent    9 

9  Scutum  shiny,  without  microsculpture  on  posterior  %;  scutellum  punctures  sparse,  min- 


Volume  4,  1995  241 

ute,  appearing  impunctate;  scrobal  sulcus  coarsely  foveolate;  Ecuador,  Colombia,  Vene- 
zuela     14.  sharkeyi  Finnamore,  new  species 

9'  Posterior  half  of  scutum  microsculptured  and /or  punctate  or  scrobal  sulcus  smooth,  with- 
out foveae   10 

10  Pronotal  lobe  brown  to  black;  posterior  half  of  scutum  shiny,  punctures  sparse;  scrobal 
sulcus  only  slightly  evident,  not  foveolate;  Ecuador,  Colombia     

10.  nigricollaris  Finnamore,  new  species 

10'  Pronotal  lobe  white;  posterior  half  of  scutum  punctate  and /or  microsculptured;  scrobal 
sulcus  usually  foveolate   11 

11  Transverse  sulcus  of  pronotum  without  longitudinal  carinae,  sulcus  smooth,  usually  with- 
out sculpture     12 

ir  Transverse  sulcus  of  pronotum  with  longitudinal  carinae   13 

12  Propodeum  with  few  carinae  on  lateral  sphere,  shiny  toward  metapleuron;  petiole  (mea- 
sured dorsally)  longer  than  first  tergum;  Peru    7.  Uiitanis  Finnamore,  new  species 

12'  Propodeum  with  dense  irregular  carinae  on  lateral  sphere  and  with  microsculpture  ad- 
jacent to  metapleuron;  petiole  (measured  dorsally)  shorter  than  first  tergum;  Bolivia,  Peru 
12.  sapanis  Finnamore,  new  species 

13  Occipital  carina  simple,  not  foveolate  or  raised  ventrally;  tarsi  white;  Ecuador 

1.  atnbigutis  Finnamore,  new  species 

13'  Occipital  carina  at  least  finely  foveolate  and  usually  slightly  raised  ventrally;  tarsi  white 
to  yellow  brown   14 

14  Scrobal  sulcus  coarsely  foveolate;  scutum  coarsely  but  sparsely  punctured  in  single 
known  specimen;  Colombia   4.  colombiamis  Finnamore,  new  species 

14'  Scrobal  sulcus  finely,  if  at  all,  foveolate;  scutum  finely  to  coarsely  punctured;  Venezuela 
to  Bolivia     6.  jatunkirus  Finnamore,  new  species 

15  Gena  ventrally  and  sternopleural  region  with  relatively  dense,  elongate  setae;  Peru 

13.  shachus  Finnamore,  new  species 

15'  Gena  ventrally  and  sternopleural  region  sparsely  pubescent  with  short  inconspicuous 
setae     16 

16  Fore  coxa  and  usually  some  of  prothorax  red;  se.  Brazil    

8.  mantanti  Finnamore,  new  species 

16'  Prothorax  and  fore  coxa  black   17 

17  Microsculpture  of  head  and  scutum  uniform  throughout;  scutum  with  small  obscure 
punctures;  Peru,  Ecuador,  Venezuela    9.  muthiis  Finnamore,  new  species 

17'  Microsculpture  of  vertex  not  as  dense  as  frons  (vertex  more  shiny);  scutum  often  shiny 

or  with  large  punctures    18 

18  Pronotal  lobe  dark,  brown  to  black 19 

18'  Pronotal  lobe  white     20 

19  Scrobal  sulcus  coarsely  foveolate;  dorsal  mandibular  tooth  acute;  scape  black;  Ecuador, 
Colombia     3.  chutiyana  Finnamore,  new  species 

19'  Scrobal  sulcus  finely  foveolate;  dorsal  mandibular  tooth  broad;  scape  yellow;  Ecuador, 
Colombia     10.  nigricollaris  Finnamore,  new  species 

20  Propodeum  closely  and  finely  carinate  on  area  adjacent  to  enclosure;  Ecuador     

5.  ecuatorialis  Finnamore,  new  species 

20'  Propodeum  coarsely  areolate  on  area  adjacent  to  enclosure    21 

21  Transverse  sulcus  of  pronotum  without  longitudinal  carinae,  smooth,  usually  without 
sculpture   22 

21 '  Transverse  sulcus  of  pronotum  with  at  least  a  few  longitudinal  carinae    23 

22  Clypeus  with  elongate  specialized  setae  on  apical  edge  except  for  narrow  median  emar- 
gination;  Peru    7.  llutanis  Finnamore,  new  species 

22'  Clypeus  with  elongate  specialized  setae  on  median  and  lateral  teeth  only;  Peru,  Bolivia 

12.  sapanis  Finnamore,  new  species 

23  Scutum  shiny,  without  microsculpture  on  posterior  %;  punctures  minute,  sparse,  appear- 


242  Journal  of  Hymenoptera  Research 

ing  impunctate  posteriorly;  scrobal  sulcus  foveolate;  Ecuador,  Venezuela    

14.  sharkeyi  Finnamore,  new  species 

23'  Scutum  with  microsculpture  and/or  punctures  posteriorly;  scrobal  sulcus  foveolate  or 

not   24 

24  Scrobal  sulcus  entirely  foveolate;  clypeal  punctures  minute,  relatively  dense  and  more  or 
less  equally  distributed  on  raised  area  above  median  teeth;  Colombia 

4.  colotnbiamis  Finnamore,  new  species 

24'  Scrobal  sulcus  usually  foveolate  anteriorly  or  not  at  all,  occasionally  entirely  foveolate; 
clypeal  punctures  sparse  with  relatively  large  more  or  less  impunctate  area  above  median 
teeth     25 

25  Clypeal  teeth  black;  Venezuela  to  Bolivia   6.  jatiinkiriis  Finnamore,  new  species 

25'  Clypeal  teeth  yellow-brown;  Ecuador   1.  ambiguus  Finnamore,  new  species 


1.  Llaqhastigmus  ambiguus  Finnamore,  ocellus.  Occipital  carina  simple,  not  raised 

new^  species  ventrally  or  foveolate. 

Derivation  ofName.-The  species  epithet,  M^sosonia.  Transverse  sulcus  of  pro- 
ambiguus,  is  from  the  Latin  "ambiguus"  "°^^"^  ^/^^  longitudinal  carinae.  Scutum 
meaning  of  doubtful  nature,  in  reference  nucrosculptured,  with  fine  sparse  punc- 
to  the  single  known  locality  and  lack  of  ^"^^'  "^^'"^  *^^^  ^  diameters  apart  (some- 
material  to  adequately  judge  variation.  ^'"^^^  mterspersed  with  more  coarse,  al- 

Diagnosis.-The  simple  occipital  carina  ^^^^S^    ^^^^^^y    ^P^^^^    punctures)    and 

that  is  not  raised  ventrally  or  foveolate  concentrated  on  centre  of  disc.  Transscu- 

will  distinguish  males  of  this  species  from  *^"^^  ^^^^"^  foveolate.   Preomaular  area 

all  others  and  females  from  most  other  sparsely  setose,  setae  not  obscuring  un- 

species  in  the  genus.  Males  of  all  other  derlying  sculpture.  Subomaulus  present, 

species  and  females  of  most  species  have  Scrobal  sulcus  usually  with  irregular  fo- 

an  occipital  carina  at  least  slightly  raised  ^^ae  on  at  least  anterior  third;  foveae  vari- 

ventrally  bearing  at  least  several  foveae.  ^^^^  f^om  almost  absent  to  present  on  en- 

Additionally,  female  fl/7?&/g» us  are  without  ti^e  scrobal  sulcus.  Sternopleural  region 

red  coloration,  have  yellow-brown  clypeal  microsculptured  with  sparse  (apparently 

teeth,  normal  genal  setae,  differing  mi-  absent)  setae.  Propodeum  shiny,  area  ad- 

crosculpture  density  between  frons  and  jacent  to  enclosure  and  lateral  sphere  with 

vertex,  and  scutum  with  microsculpture  several  large  areolae  composed  of  relative- 

and  sparse  punctures.  Females  likely  to  be  ly  low  carinae. 

confused  with  this  species  have  one  or  Metasoma.  Petiole  with  3  or  4  coarse  ca- 

more  of  the  following:  clypeal  teeth  black,  rinae  laterally. 

mesosoma  red  in  part,  setae  of  lower  gena  Color.  Ground  color  black.  White:  man- 
elongate  and  relatively  dense,  frons  and  dible  except  apically,  palpi,  pronotal  lobe, 
vertex  with  uniform  microsculpture  den-  tarsi.  Yellow  to  yellow-brown:  antenna 
sity,  and /or  scutum  shiny  apparently  (generally  yellow  ventrally  to  brown  dor- 
without  punctures.  sally  on  apical  flagellomeres),  tegula,  fore 

Description. — Male.  Length  3.4^.6  mm.  and  mid  legs  except  tarsi,  wing  veins,  api- 

Head.  Sculpture  of  clypeus  obscured  by  cal  metasomal  sterna.  Brown:  mandibular 

dense   appressed   setae.    Gena    ventrally,  apex,  stigma  of  fore  wing,  hind  leg  except 

with  sparse  straight  setae  that  are  sub-  tarsus. 

equal  to  or  less  than  greatest  width  of  fore  Female.  Length  3.9  mm.  Similar  to  male 

basitarsus.  Microsculpture  of  frons  more  except  as  follows:  apicoventral  mandibu- 

dense  than  that  of  vertex  adjacent  to  mid  lar  tooth  enlarged,  truncate,  much  larger 


Volume  4,  1995 


243 


Figs.  36-41.  Llaqhastigmus  and  Stigmus.  36,  Llaqhastigmiis  jatunkirus  9 ,  head,  frontal  view.  37,  L.  jatiinkinis  9, 
mandible,  outer  view.  38,  L.  jatunkirus  9,  mesosoma,  dorsal  view.  39,  Stigmus  sp.  9,  mandible,  outer  view. 
40,  Stigmus  sp.  9,  head,  frontal  view.  41,  Stigmus  sp.  9,  mesosoma,  dorsal  view. 


than  other  teeth.  Clypeal  apex  quadriden- 
tate  as  follows:  a  lateral  lobe  flanking  lat- 
eral edge  of  labrum,  a  pair  of  smaller  me- 
dian teeth  separated  from  lateral  lobes  by 
broad  emarginations  and  separated  from 
each  other  by  a  narrow  U-shaped  emar- 
gination.  Clypeal  apex  with  specialized 
flattened,  elongate,  setae  on  teeth  but  ab- 
sent from  emarginate  interspaces;  2  setae 


per  median  tooth  and  6-7  on  lateral  lobe. 
Clypeus  shiny,  microsculpture  absent, 
punctures  3^  diameters  apart  on  disc. 
Apical  margin  of  clypeus  yellow-brown. 

Material  Examined  {27 S,  19). — Holo- 
type:  9,  ECUADOR:  Pich.  Prov.  Guay- 
llabamba,  10km  n.  on  Rio  Pisque,  2500m 
11-1983.  M.  Sharkey,  L.  Masner  (BRD). 
Paratypes:  ECUADOR:  Pichincha:  Guay- 


244 


Journal  of  Hymenoptera  Research 


llabamba,  10km  n.  on  Rio  Pisque,  2500m 
11-1983.  L.  Masner,  M.  Sharkey  {6:2  BRD, 
2  PMA).  10km  n.  Guayllabamba  26-11- 
1983.  Masner  &  Sharkey  (cJ:  4  BRD,  19 
PMA). 

Distribution. — (Map  20).  Ecuador. 

2.  Llaqhastigmus  australis  Finnamore, 
new  species 

Derivation  of  Name. — The  species  epithet 
is  derived  from  the  Latin  "austral"  mean- 
ing southern  in  reference  to  the  southern 
locality  of  the  only  known  specimen. 

Diagnosis. — Female  unknown.  Male:  la- 
brum  quadrilobed  with  a  deep  median 
notch;  subomaulus  absent,  petiole  coarse- 
ly carinate  (carinae  normal)  laterally.  L. 
australis  and  santanderanus  are  the  only 
species  in  the  genus  lacking  a  subomaulus 
and  having  a  quadrilobed  labrum  (at  least 
in  males,  females  for  both  species  are  un- 
known). In  all  other  species  in  the  genus 
the  subomaulus  is  present  and  the  labrum 
is  bilobed  with  a  slight  median  emargi- 
nation. 

Description. — Male.  Length  5.0  mm. 

Head.  Sculpture  of  clypeus  obscured  by 
dense  appressed  setae.  Labrum  quadri- 
lobed with  a  deep  median  emargination. 
Gena  ventrally,  with  sparse  straight  setae 
that  are  subequal  to  or  less  than  greatest 
width  of  fore  basitarsus.  Microsculpture  of 
frons  more  dense  than  that  of  vertex.  Oc- 
cipital carina  raised  ventrally  and  finely 
foveolate  at  least  ventrally. 

Mesosoma.  Transverse  sulcus  of  pro- 
notum  with  longitudinal  carinae.  Scutum 
microsculptured,  punctures  sparse  more 
or  less  evenly  distributed  and  separated 
by  5-10  puncture  diameters.  Transscutel- 
lar  sulcus  not  foveolate.  Preomaular  area 
sparsely  setose,  setae  not  obscuring  un- 
derlying sculpture.  Subomaulus  absent, 
represented  as  an  evanescent  ridge  near 
posteroventral  apex  of  pronotum,  not  car- 
inate or  intersecting  omaulus.  Scrobal  sul- 
cus weakly  impressed  anteriorly,  attenu- 
ated posteriorly,  not  foveolate.  Sterno- 
pleural  region  shiny  without  microsculp- 


ture, sparsely  punctate,  sparsely  setose. 
Propodeum  shiny,  area  adjacent  to  enclo- 
sure without  microsculpture,  with  several 
large  areolae  composed  of  low  carinae;  lat- 
eral sphere  shiny,  with  evanescent  micro- 
sculpture,  areolae  smaller. 

Metasoma.  Petiole  with  3  coarse  lateral 
carinae. 

Color.  Ground  color  black.  White:  man- 
dible except  apically,  palpi,  pronotal  lobe. 
Yellow-brown:  scape,  pedicel,  fore  and 
mid  legs  except  coxae,  hind  tibia  and  tar- 
sus, tegula,  apical  metasomal  sterna. 

Female.  Unknown. 

Material  Examined  (IS). — Holotype:  6, 
BOLIVIA:  Cochabamba,  Prov.  Carrasco, 
Empalme  3000m  11-1971.  Fritz  y  Martinez 
(IIES). 

Distribution  (Map  21). — Bolivia. 

3.  Llaqhastigmus  chutiyana  Finnamore, 
new  species 

Derivation  of  Name. — The  species  epithet 
is  derived  from  the  Quechua  terms 
"ch'uti",  meaning  naked  and  "yana", 
meaning  black,  in  reference  to  the  rela- 
tively hairless  condition  of  this  black  spe- 
cies. 

Diagnosis. — The  combination  of  the 
black  scape  and  pronotal  lobe,  scrobal  sul- 
cus foveolate,  and  the  apicodorsal  man- 
dibular tooth  of  female  acute  (normal)  will 
separate  this  species  from  all  others  in  the 
genus.  All  other  species  except  nigricollaris 
have  a  white  pronotal  lobe.  L.  chutiyana  is 
distinguished  from  nigricollaris  by  the 
black  scape,  foveolate  scrobal  sulcus  and 
acute  apicodorsal  mandibular  tooth  in  the 
female;  nigricollaris  has  a  yellow-brown 
scape,  partially  foveolate  scrobal  sulcus 
and  enlarged  apicodorsal  mandibular 
tooth  in  the  female. 

Description. — Male.  Length  3.9-5.0  mm. 

Head.  Sculpture  of  clypeus  not  ob- 
scured by  setae.  Clypeus  shiny  without 
microsculpture,  setae  and  punctures 
sparse  and  at  least  1  diameter  apart  on 
disc.  Gena  ventrally  with  evanescent  mi- 
crosculpture, punctures  and  setae  sparse, 


Volume  4,  1995 


245 


setae  less  than  greatest  width  of  fore  ba- 
sitarsus.  Microsculpture  of  frons  more 
dense  than  that  of  vertex.  Occipital  carina 
raised  ventrally  and  foveolate  on  at  least 
that  area. 

Mesosoma.  Transverse  sulcus  of  pro- 
notum  with  longitudinal  carinae.  Scutum 
microsculptured,  punctures  small  and 
sparse,  punctures  more  or  less  evenly  dis- 
tributed although  somewhat  less  dense  on 
posterior  disc.  Transscutellar  sulcus  fo- 
veolate. Preomaular  area  asetose.  Subo- 
maulus  present.  Scrobal  sulcus  coarsely 
foveolate.  Sternopleural  region  shiny  often 
with  evanescent  microsculpture,  punc- 
tures sparse  and  minute.  Propodeum 
shiny  without  microsculpture,  area  adja- 
cent to  enclosure  and  lateral  sphere  with 
large  areolae  composed  of  relatively  high 
carinae. 

Metasoma.  Petiole  with  2  or  3  lateral  ca- 
rinae. 

Color.  Ground  color  black.  Yellow- 
brown  to  brown:  mandibles  except  apical- 
ly,  palpi,  fore  and  mid  tibiae  and  tarsi, 
hind  tibia  on  inner  side  and  tarsus,  tegula, 
apical  metasomal  sterna. 

Female.  Length  4.1-4.4  mm.  Similar  to 
male  except  as  follows:  apicoventral  man- 
dibular tooth  enlarged,  truncate  and  much 
larger  than  other  teeth.  Clypeal  apex 
quadridentate  as  follows:  lateral  lobe 
flanking  edge  of  labrum,  pair  of  smaller 
median  teeth  separated  from  lateral  lobe 
by  deep  emargination  and  from  each  other 
by  U-shaped  notch.  Clypeal  apex  with  4 
specialized,  flattened,  elongate  setae  on 
lateral  lobe  and  2  per  median  tooth,  setae 
absent  from  emarginate  interspaces.  Clyp- 
eus  shiny,  microsculpture  absent;  punc- 
tures sparse,  3  or  more  diameters  apart. 
Apical  margin  of  clypeus  black. 

Material  Examined  (8(5,  3  9). — Holotype: 
(5,  ECUADOR:  Tungurahua,  Banos 
2,000m.  21-X-1974.  M.  Cooper  B.M.1975- 
33  (BMNH).  Paratypes:  COLOMBIA:  Val- 
le:  Penas  Blancas  22-Xi-1974-L  R.  Wilk- 
erson.  Malaise  Trap  (9:1  FSCA).  ECUA- 
DOR: Tungurahua:  Banos  2,000m.  17-X- 


1974.  M.  Cooper  B.M. 1975-33  {6:  1 
BMNH).  Banos  2,000m.  21-X-1974.  M. 
Cooper  B.M.1975-33  {6:  2  BMNH,  1 
PMA).  Banos.  c.2,000m.  7-X-1978.  M.  Coo- 
per B.M.  1979-20  (9:  2  BMNH).  Banos. 
c.2000m  19-X1-1978  M.  Cooper  B.M.  1979- 
20  {6:  2  BMNH).  Banos;-Tungurahua 
Norths.-:2300m;  Leg.  R.  Hensen  et  A.  Ap- 
troot;  11-8-1982  {S:\  RNH). 

Distribution. — (Map  22).  Colombia,  Ec- 
uador. 

4.  Llaqhastigmus  colombianus 

Finnamore,  new  species 

Derivation  of  Name. — The  species  epithet, 
colombianus,  is  derived  from  the  country 
Colombia,  where  all  specimens  of  this  spe- 
cies have  been  collected. 

Diagnosis. — The  following  combination 
will  separate  this  species  from  others  in 
the  genus.  Occipital  carina  foveolate  ven- 
trally, pronotum  black  except  white  pro- 
notal  lobe,  coarsely  foveolate  scrobal  sul- 
cus usually  with  3  or  4  large  foveae  and 
sulcus  usually  diminishing  in  width  pos- 
teriorly, and  female  clypeus  with  fine 
punctures  about  2  diameters  apart  and 
equally  distributed  on  median  third.  This 
species  is  similar  to  jatiinkirus  and  sharkeyi 
from  which  it  differs  with  the  coarsely 
foveolate  scrobal  sulcus  and  in  the  female 
with  the  homogeneously  punctate  median 
clypeal  region. 

Description. — Male.  Length  3.9  mm. 

Head.  Sculpture  of  clypeus  obscured  by 
dense  appressed  setae.  Gena  ventrally 
with  sparse  straight  setae  that  are  sub- 
equal  to  or  less  than  greatest  width  of  fore 
basitarsus.  Microsculpture  of  frons  more 
dense  than  that  of  vertex.  Occipital  carina 
raised  ventrally  and  foveolate. 

Mesosoma.  Transverse  sulcus  of  pro- 
notum with  longitudinal  carinae.  Scutum 
microsculptured,  punctures  sparse  and 
relatively  coarse.  Transscutellar  sulcus 
foveolate.  Preomaular  area  sparsely  se- 
tose, setae  not  obscuring  underlying 
sculpture.  Subomaulus  present.  Scrobal 
sulcus  with  3  or  4  coarse  foveae,  sulcus 


246 


Journal  of  Hymenoptera  Research 


Figs.  42^7.  Stigmus,  Tzustigmus  and  Incastigmiis  42,  Stigmus  sp.  <?,  clypeus,  showing  lateral  bevels.  43,  Stig- 
nnis  sp.  6 ,  mesosoma,  ventral  view.  44,  Tzustigmus  syam  9,  metasomal  sternum  2.  45,  T.  syam  9,  micropore 
field  on  metasomal  sternum  2.  46,  Incastigmus  inti  6,  micropore  field  on  fore  wing  stigma.  47,  T.  syam  ?, 
micropore  field  on  fore  wing  stigma. 


entirely  foveolate,  somewhat  narrower 
posteriorly.  Sternopleural  region  weakly 
microsculptured,  punctures  and  setae 
sparse  almost  absent  from  lower  area.  Pro- 
podeum  shiny,  area  adjacent  to  enclosure 
and  lateral  sphere  with  several  large  are- 
olae composed  of  relatively  low  carinae. 

Metasoma.  Petiole  with  several  carinae 
laterally. 


Color.  Ground  color  black.  White:  pro- 
notal  lobe.  Yellow-brown:  palpi,  mandible 
except  apically,  antenna,  tegula,  stigma, 
fore  and  mid  legs  except  coxae,  hind  tar- 
sus, apical  metasomal  tergum.  Brown: 
hind  leg  except  tarsus. 

Female.  Length  3.8-4.7  mm.  Similar  to 
male  except  as  follows:  apicoventral  man- 
dibular  tooth   enlarged,   truncate,   much 


Volume  4,  1995 


247 


larger  than  other  teeth.  Clypeal  apex 
quadridentate  as  follows:  lateral  lobe 
flanking  lateral  edge  of  labrum,  pair  of 
smaller  median  teeth  separated  from  lat- 
eral lobes  by  broad  emarginations  and 
separated  from  each  other  by  broad  U- 
shaped  notch.  Clypeal  apex  with  special- 
ized flattened,  elongate,  setae  on  teeth  and 
most  of  emarginate  interspaces;  with  2  se- 
tae per  median  tooth,  3  per  lobe  and  3  on 
lateral  emarginations.  Clypeus  shiny, 
without  microsculpture,  punctures  sparse 
laterally  but  raised  median  area  evenly 
and  finely  punctate  with  punctures  up  to 
3  diameters  apart,  no  part  of  median 
raised  area  without  punctures.  Apical 
clypeal  margin  black.  Scutum  sometimes 
without  microsculpture  on  disc.  Antenna 
and  legs  more  brown  than  male. 

Material  Examined  {IS,  89). — Holotype 
9,  COLOMBIA:  Dept.  Valle,  Penas  Blan- 
cas  1750m  10km  w.  Call,  very  wet  pre- 
montane  forest,  R.C.  Wilkerson  29-1-1975 
Malaise  Trap  (FSCA).  Paratypes:  CO- 
LOMBIA: Antioquia:  Antioquia  1800m 
13-22-IV-1973  (9:1  PMA).  Caqueta:  Flo- 
rencia  480m  31-X-5-XI-1971  M.  Cooper 
B.M.  1972-275  (9:1  BMNH).  Valle:  Pance 
19-XI-1974-1  R.  Wilkerson  Malaise  Trap 
(9:1  FSCA).  Pance  CVC  (1700m)  15km  w. 
Call,  very  wet  premontane  forest,  R.C. 
Wilkerson  23-XII  1974  {6:  I  FSCA,  9:  3 
FSCA).  Pance  CVC  (1700m)  15km  w.  Cali, 
very  wet  premontane  forest,  R.C.  Wilker- 
son 28-X-1974  Malaise  Trap  (9:1  FSCA). 
Penas  Blancas  21-XI-1974-3  R.  Wilkerson 
Malaise  Trap  (9:  1  FSCA). 

Distribution  (Map  23). — Colombia. 

5.  Llaqhastigmus  ecuatorialis 
Finnamore,  new  species 

Derivation  of  Name. — The  species  epithet 
is  derived  from  a  combination  of  the  coun- 
try of  origin  of  the  specimens,  Ecuador, 
and  the  English  word  equator  in  reference 
to  the  distribution  of  the  species. 

Diagnosis. — The  combination  of  a  brown 
scape  that  is  yellow-brown  at  base  and 
apex,  and  a  scrobal  sulcus  that  is  absent 


or  weakly  impressed  and  without  foveae 
will  separate  this  species  from  all  others 
in  the  genus.  All  other  species,  except  chit- 
tiyann  and  santanderatius  have  the  scape 
yellow  ventrally  and  often  a  foveolate 
scrobal  sulcus.  Llaqhastigmus  chutiyana  has 
a  black  pronotal  lobe  while  santanderanus 
lacks  a  subomaulus;  ecuatorialis  has  a 
white  pronotal  lobe  and  a  subomaulus  is 
present.  This  species  also  has  finely  cari- 
nate  sculpture  on  the  propodeum  adjacent 
to  enclosure  further  distinguishing  it  from 
other  species. 

Description. — Male.  Length  3.9-4.3  mm. 

Head.  Clypeal  sculpture  obscured  by 
dense  appressed  setae.  Gena  ventrally, 
with  sparse  straight  setae  that  are  sub- 
equal  to  or  less  than  greatest  width  of  fore 
basitarsus.  Microsculpture  of  frons  more 
dense  than  that  of  vertex.  Vertex  micro- 
sculpture  in  some  specimens  is  unusually 
dense  but  never  as  dense  as  that  of  frons. 
Occipital  carina  ventrally  raised  and  fo- 
veolate. 

Mesosoma.  Transverse  sulcus  of  pro- 
notum  with  longitudinal  carinae.  Scutum 
microsculptured,  with  sparse  scattered 
punctures.  Transscutellar  sulcus  finely 
foveolate  or  foveae  absent.  Preomaular 
area  sparsely  setose,  setae  not  obscuring 
underlying  sculpture.  Subomaulus  pres- 
ent. Omaulus  present  but  in  one  specimen 
attenuated  ventrally  and  not  reaching  ac- 
etabular carina.  Scrobal  sulcus  evanescent, 
slightly  impressed,  not  foveolate.  Hypoe- 
pimeral  area  microsculptured.  Sterno- 
pleural  region  shiny  without  microsculp- 
ture, punctures  and  setae  sparse.  Propo- 
deum with  microsculpture  especially  out- 
side enclosure,  area  adjacent  to  enclosure 
and  propodeal  side  with  fine  irregular  ca- 
rinae which  meet  dorsolaterally  to  form 
small  irregular  areolae  composed  of  low 
carinae. 

Metasoma.  Petiole  with  3  or  4  carinae 
laterally. 

Color.  Ground  color  black.  White:  man- 
dible basally,  pronotal  lobe.  Yellow  to  yel- 
low-brown: palpi,  scape  on  basal  and  api- 


248 


Journal  of  Hymenoptera  Research 


48 


49 


Figs.  48-50.     Stigmina,  hind  wing.  48,  Llaqhastigmus.  49,  Tzustigmus.  50,  Carinostigmus. 


cal  ring,  sometimes  pedicel,  tegula,  legs 
except  coxae  and  femora.  Brown:  scape 
except  basal  and  apical  ring,  femora. 

Female.  Length  4.5  mm.  Similar  to  male 
except  as  follows:  apicoventral  mandibu- 
lar tooth  enlarged,  much  larger  than  other 
teeth.  Clypeal  apex  quadridentate  as  fol- 
lows: acutely  triangular  lateral  tooth 
flanking  lateral  edge  of  labrum  and  sur- 
passing median  teeth;  pair  of  smaller  me- 
dian teeth  separated  from  lateral  teeth  by 
broad  U-shaped  emarginations  and  sepa- 
rated from  each  other  by  narrow  U- 
shaped  notch.  Clypeal  apex  with  special- 
ized, flattened,  elongate  setae  present  on 
teeth  but  absent  from  most  of  emarginate 
interspace  with  the  exception  of  single 
seta  on  emargination  near  lateral  tooth. 
Clypeus  shiny,  with  evanescent  micro- 
sculpture   on   upper   median   area    only. 


punctures  sparse  and  present  only  on  me- 
dian area.  Lateral  clypeal  tooth  red  at 
apex.  Both  known  females  have  omaulus 
which  is  attenuated  ventrally  and  widely 
separated  from  acetabular  carina.  This 
character  is  variable  in  the  male  (omaulus 
usually  intersecting  acetabular  carina)  and 
may  also  be  so  in  the  female. 

Material  Examined  (llcJ,  2  9). — Holo- 
type:  6,  ECUADOR:  Los  Duendes,  S.  Bo- 
livar 10-VH965  Pena  (MCZ).  Paratypes: 
ECUADOR:  Imbabura:  Otavalo;  -Qui- 
chinche-2700m  Leg.  R.  Hensen  et  A.  Ap- 
troot  8-7-1982  (6:  1  RNH).  S.  Otavalo 
3300m  L8-9-1971  Luis  E.  Pena  ($:  AEI). 
Pichincha:  San  Rafael  ca.  Quito  2500m  21- 
29-VI-1975  Col.  C.  Porter  (9:1  LILLO).  S. 
Bolivar:  same  data  as  holotype  (6:6  MCZ, 
1  PMA).  Tungurahua:  Banos,  Tungurahua 
Norths.  2300m  Leg.  R.  Hensen  et  A.  Ap- 


Volume  4,  1995  249 

troot  11-8-1982  (c?:  1  RNH).  Rio  Chota  VI-  ae.   Sternopleural   region   shiny,   with   at 

10-1965  1800m  Luis  Pena.  ((5:1  AEI).  most    evanescent    microsculpture,    punc- 

Distribution  (Map  24). — Ecuador.  tures  small  and  sparse,  setae  sparse.  Pro- 

podeum  shiny,  area  adjacent  to  enclosure 

6.  Llaqhastigmus  jatunkirus  Finnamore,  ^^^^    i^^^^.^!    ^^^^^^    ^-^^    g^^^j.^!    ^arge, 

new  species  somewhat  irregular  areolae  composed  of 

Derivation  of  Name. — The  species  epithet  low  lying  carinae;  side  without  sculpture 

is  derived  from  two  Quechua  words;  "ja-  adjacent  to  enclosure, 

tun"  meaning  large  and  "kiru"  meaning  Metasoma.  Petiole  with  3  or  4  coarse  lat- 

tooth,  referring  to  the  enlarged  apicoven-  eral  carinae. 

tral  mandibular  tooth  in  the  female  of  this  Color.  Ground  color  black.  White:  palpi, 
species.  pronotal  lobe.  Yellow-white:  tarsi.  Yellow 
Diagnosis. — The  following  combination  to  yellow-brown:  mandible  basally,  scape 
of  characters  will  separate  this  species  and  pedicel  usually  entirely  but  at  least 
from  all  others  in  the  genus.  Scape  yellow-  ventrally,  basal  flagellomeres  ventrally, 
brown,  vertex  microsculpture  of  less  den-  fore  and  mid  legs  except  coxae  and  tarsi, 
sity  than  that  of  frons,  lower  gena  without  sometimes  hind  tibia,  tegula,  tergum  VI 
modified  setae,  occipital  carina  raised  ven-  on  apical  half  or  more  and  VII,  sternum 
trally  and  foveolate,  transverse  pronotal  VI  and  VII.  Brown:  usually  hind  leg  except 
sulcus  longitudinally  carinate,  pronotal  tarsi  and  occasionally  tibia, 
lobe  white,  scutum  microsculptured,  su-  Female.  Length  3.8-5.1  mm.  Similar  to 
bomaulus  present,  preomaular  area  male  except  as  follows:  apicoventral  man- 
sparsely  setose,  scrobal  sulcus  impressed  dibular  tooth  enlarged,  truncate,  much 
and  finely  if  at  all  foveolate,  and  propo-  larger  than  other  teeth.  Clypeal  apex 
deum  areolate  on  dorsolateral  area  out-  quadridentate  as  follows:  lateral  lobe 
side  enclosure.  Females  in  addition  to  the  flanking  edge  of  labrum,  pair  of  median 
foregoing,  have  the  median  clypeal  area  teeth  separated  from  lateral  lobe  by  broad 
mostly  impunctate  apically  and  punctate  emargination  and  separated  from  each 
basally.  other  by  U-shaped  notch.  Clypeal  apex 
Description. — Male.  Length  3.7-5.0  mm.  with  specialized,  flattened,  elongate  setae 
Head.  Sculpture  of  clypeus  obscured  by  distributed  on  teeth  and  often  in  emargin- 
dense  appressed  setae.  Gena  ventrally,  ate  interspaces.  Clypeus  shiny,  micro- 
with  sparse  straight  setae  that  are  sub-  sculpture  absent,  both  lateral  and  median 
equal  to  or  less  than  greatest  width  of  fore  areas  sparsely  punctate  and  impunctate 
basitarsus.  Microsculpture  of  frons  more  apicomedially.  Clypeal  teeth  black, 
dense  than  that  of  vertex  adjacent  to  mid  Material  Examined  (38(5  639). — Holo- 
ocellus.  Occipital  carina  ventrally  raised  type:  9,  VENEZUELA:  Merida:  Merida, 
and  foveolate.  Sta.  Rosa  2000m  15-V-15-VI-1981  A.  Bri- 
Mesosoma.  Transverse  pronotal  sulcus  ceno  &  F.  Suarez  (BRD).  Paratypes:  BO- 
with  longitudinal  carinae.  Scutum  micro-  LI VI A:  La  Paz:  Chulumani  1,700m.  25-III- 
sculptured,  punctures  sparse  with  density  1979  M.  Cooper  B.M.  1979-216  (9:  2 
and  size  variable.  Transscutellar  sulcus  BMNH).  Chulumani  1,700m.  30-111-1979 
foveolate.  Preomaular  area  sparsely  se-  M.  Cooper  B.M.  1979-216  (d:  1  BMNH,  9: 
tose,  setae  not  obscuring  underlying  1  BMNH).  Chulumani  1,700m.  2-IV-1979 
sculpture.  Subomaulus  present.  Scrobal  M.  Cooper  B.M.  1979-216  (9:1  BMNH,  1 
sulcus  impressed  with  foveae  variable  NMW).  COLOMBIA:  Cauca:  Popayan, 
from  complete  to  absent,  usually  anterior  1,800m.  lO-X-1974  M.  Cooper  B.M.  1975- 
half  of  sulcus  weakly  foveolate  but  rarely  33  {6:  2  BMNH,  9:  1  BMNH).  Santander 
entirely  foveolate  or  rarely  without  fove-  del  Norte:  Prima  1,700m  29-V-1965  J.  &  B. 


250 


Journal  of  Hymenoptera  Research 


Bechyne  (9:1  UCV).  Valle  Prov.:  summit 
w.  of  Call  6-II-1977  2000m  M.  Breed,  CD. 
Michener  (9:1  SEM).  ECUADOR:  Loja: 
Loja  III-23-26-1965  Luis  Pena  {S:  1  AEI). 
Pastaza:  22km  sw.  Puyo  900m  14-16-VII- 
1976  S.  &  J.  Peck,  forest  (9:2  BRD).  Rio 
Jumboe  (Zamora)  l-IV-1965  Pefia  {6:  1 
MCZ).  PERU:  Cuzco:  Agua  Calliente  21- 
28-XII-1983  L.  Huggert  (9:3  PMA).  Ma- 
chu  Picchu  1900m.  IX-4/ 19-1964  C.C.  Por- 
ter (9:  3  MCZ).  Machu  Picchu  XI-29-1965 
H.  &  M.  Townes  {6:1  AEI,  9 :  2  AEI).  Ma- 
chu Picchu  XII-1-1965  H.  &  M.  Townes 
((?:  4  AEI,  9:  4  AEI).  Machu  Picchu  11-24- 
27-1968  A.  Garcia  &  C.  Porter  ( 9  1  MCZ). 
Machu  Picchu  (ruins)  2400m  20-IV-1983  C. 
&  M.  Vardy  B.M.  1983-217  (9:1  BMNH). 
Huanuco:  Tingo  Maria,  26  mi.  e.  XII-10- 
1954  1100m  E.I.  Schlinger  &  E.S.  Ross  (9: 
1  CAS).  Tingo  Maria,  40  mi.  s.  Carpish 
Mts.  XII-28-1954  E.I.  Schlinger  &  E.S.  Ross 
(9:1  CAS).  Pasco:  Oxapampa  2,200m  6- 
III-1979  M.  Cooper  B.M.  1979-216  (9:1 
BMNH).  Oxapampa  2,200m  8-III-1979  M. 
Cooper  B.M.  1979-216  {6:  1  BMNH). 
VENEZUELA:  Lara:  Yacambu  1200m  V-3- 
1981  H.K.  Townes  (c?:  3  AEI).  Yacambu 
1200m  V-7-1981  H.K.  Townes  {6:  6  AEI, 
9:  11  AEI).  Yacambu  1200m  V-10-1981 
H.K.  Townes  (9:3  AEI).  Yacambu  1200m 
V-13-1981  H.K.  Townes  (9:  5  AEI).  Ya- 
cambu N.P.  1200m  cloud  forest  7-V-1981 
H.  Townes  {6:4:  PMA,  9 :  2  PMA).  Yacam- 
bu N.P.  1200m  cloud  forest  lO-V-1981  H. 
Townes  {6:  2  PMA,  9:  1  PMA,  1  BRD). 
Yacambu  1200m  cloud  forest  13-V-1981  H. 
Townes  {6:  3  PMA).  Merida:  Merida 
1800m  ll-V-1981  L.  Masner  1984.149  {6:2 
PMA,  9:  7  PMA).  Merida,  small  valley 
across  R.  Chama  6-V-1981  L.  Masner 
1984.157  (9:1  PMA).  Merida,  Sta.  Rosa 
2000m  3-13-V-1981  L.  Masner  pan  i{6:l 
PMA).  Merida,  Sta.  Rosa  2000m  15-V-15- 
VI-1981  A.  Briceno  &  F.  Suarez  {6:  1 
PMA).  Merida,  Sta.  Rosa  2000m  15-VI-15- 
VII-1981  A.  Briceno  &  F.  Suarez  {6:2 
PMA).  Merida,  Sta.  Rosa  2000m  15-VII-15- 
VIII-1981  A.  Briceno  &  F.  Suarez  {6:2 
PMA).  Mucuy  nr.  Tobay  2200m  cloud  for- 


est 5-V-1981  L.  Masner  {6:1  PMA).  Tobay 
2,200m  IV-30-1981  H.K.  Townes  {6:  2 
AEI).  Tobay  2,200m  V-1-1981  H.K. 
Townes  (9:1  AEI).  Valle  de  Culata  Ruta 
7,  18-VII-1988  C.  Porter  &  L.  Stange  {6:1 
FSCA). 

Distribution  (Map  25). — Venezuela  to 
Bolivia. 

7.  Llaqhastigmus  llutanis  Finnamore, 
new  species 

Derivation  of  Name. — The  species  epithet 
is  derived  from  the  Quechua  "Uutan" 
meaning  strange,  in  reference  to  the  un- 
usual unsculptured  transverse  pronotal 
sulcus. 

Diagnosis. — This  species  can  be  distin- 
guished by  the  combination  of  the  trans- 
verse sulcus  of  pronotum  without  carinae, 
propodeum  with  lateral  sphere  mostly  un- 
sculptured and  crossed  by  only  1  or  2  low 
carinae,  and  female  clypeal  edge  with 
elongate  specialized  setae  along  most  of 
margin.  This  species  is  most  similar  to  sa- 
panis  both  of  which  differ  from  all  others 
in  the  genus  by  their  smooth  unsculptured 
transverse  pronotal  sulcus.  The  other  di- 
agnostic characters  will  separate  these  2 
closely  related  species. 

Description. — Male.  Length  4.4^.5  mm. 

Head.  Sculpture  of  clypeus  obscured  by 
dense  appressed  setae.  Gena  ventrally, 
with  sparse  straight  setae  subequal  to  or 
less  than  greatest  width  of  fore  basitarsus. 
Microsculpture  of  frons  more  dense  than 
that  of  vertex  adjacent  to  mid  ocellus.  Oc- 
cipital carina  ventrally,  raised  and  foveo- 
late. 

Mesosoma.  Transverse  pronotal  sulcus 
smooth,  without  sculpture.  Scutum  mi- 
crosculptured,  punctures  sparse  but  occa- 
sionally relatively  coarse.  Transscutellar 
sulcus  foveolate.  Preomaular  area  sparsely 
setose,  setae  not  obscuring  underlying 
sculpture.  Subomaulus  present.  Scrobal 
sulcus  weakly  impressed,  smooth,  not  fo- 
veolate. Sternopleural  region  shiny  to 
weakly  microsculptured,  punctures 
sparse.  Propodeum  shiny,  area  adjacent  to 


Volume  4,  1995  251 

enclosure  mostly  unsculptured,  lateral  have  the  sculpture  of  the  preomaular  area 

sphere  crossed  by  at  most  a  few  low  ca-  only  partially  obscured  by  setae,  they  still 

rinae  forming  large  areolae.  have  greater  density  of  setae  on  the  preo- 

Metasoma.  Petiole  with  2  fine  carinae  maular  area  than  other  species  except  for 

defining  lateral  region.  sliaduis  which  has  elongate  setae.  Addi- 

Color.  Ground  color  black.  White:  palpi,  tionally  almost  all  specimens  of  this  spe- 

pronotal   lobe.   Yellow   to   yellow-brown:  cies  have  some  red  coloration  on  the  me- 

mandibles  except  apically,  scape,  pedicel,  sosoma.   The   red   coloration  is   variable, 

at  least  ventrally  on  basal  flagellomeres  rarely  absent  but  almost  always  present 

but  sometimes  entire  undersides,  fore  and  on  fore  coxa  and  lateral  pronotum  and  can 

mid  legs  except  coxae  and  femora,  hind  extend  over  most  of  mesosoma  except  the 

trochanter    and    tarsus.    Brown:    femora,  propodeum.  It  is  the  only  species  in  the 

hind  tibia.  genus  with  red  on  the  mesosoma.  Finally 

Female.  Length  5.0  mm.  Similar  to  male,  the  female  has  an  apicoventral  mandibu- 
except  as  follows:  apicoventral  mandibu-  lar  tooth  only  slightly  enlarged  in  com- 
lar  tooth  enlarged,  truncate,  much  larger  parison  with  the  median  and  dorsal  teeth, 
than  other  teeth.  Clypeal  apex  quadriden-  Females  of  all  other  species  in  the  genus 
tate  as  follows:  lateral  lobe  flanking  lateral  have  a  greatly  enlarged  apicoventral  man- 
edge  of  labrum,  pair  of  smaller  median  dibular  tooth.  As  a  consequence  of  the 
teeth  separated  from  lateral  lobe  by  broad  small  apicoventral  mandibular  tooth,  fe- 
emargination  and  separated  from  each  males  of  this  species  can  be  confused  with 
other  by  narrow  U-shaped  notch.  Clypeal  those  of  Incastigmiis.  The  bilobed  labrum 
apex  with  specialized,  elongate,  flattened  with  a  small  median  notch  indicate  this 
setae  distributed  over  all  but  median  species  is  a  Llaqhastigmus  rather  than  In- 
notch.  Clypeus  shiny,  microsculpture  ab-  castigmus  which  has  a  quadrilobed  la- 
sent,  punctures  dense  laterally  and  sparse  brum. 

medially  with  impunctate  area  above  me-  Description. — Male.  Length  4.6-4.9  mm. 

dian  teeth.  Apical  clypeal  margin  red.  Head.  Sculpture  of  clypeus  obscured  by 

Material  Examined  (?>S ,  19). — Holotype:  dense   appressed   setae.    Gena   ventrally, 

9,  PERU:  Machu  Picchu  XI-30-1965  H.  &  with  sparse  straight  setae  that  are  sub- 

M.  Townes  (AEl).  Paratypes:  PERU:  Cuz-  equal  to  or  less  than  greatest  width  of  fore 

co:   Machu   Picchu   XI-29-1965   H.   &   M.  basitarsus.  Microsculpture  of  frons  more 

Townes  {6:1  AEI).  Machu  Picchu  Xl-30-  dense  than  that  of  vertex  adjacent  to  mi- 

1965  H.  &  M.  Townes  {6:1  AEI,  1  PMA).  docellus.  Occipital  carina  ventrally,  raised 

Distribution  (Map  26). — Peru.  and  foveolate. 

Mesosoma.   Transverse  sulcus  of  pro- 

8.  Llaqhastigmus  mantanti  Finnamore,  ^^^^^  carinate.  Scutum  microsculptured, 

new  species  sometimes   shiny   medially,   punctures 

Derivation  of  Name. — The  species  epithet  sparse,  posterior  margin  with  transverse 

mantanti  is   derived   from   two   Quechua  row  of  short  carinae.  Transscutellar  sulcus 

words,   "manta"  a   preposition  meaning  foveolate.  Sculpture  of  preomaular  area 

"from"  and  "anti"  meaning  "east".  The  obscured  by  dense  appressed  setae  that 

name  refers  to  the  distribution  of  this  spe-  are  usually  continued   along  midventral 

cies  in  eastern  Brazil.  line  to  mid  coxae.  Subomaulus  present. 

Diagnosis. — The  preomaular  area  sculp-  Scrobal  sulcus  impressed,  foveolate.  Ster- 
ture  that  is  obscured  by  setae  in  male  and  nopleural  region  shiny,  weakly  micro- 
partially  obscured  by  setae  in  female  will  sculptured,  sparsely  punctate,  with  in- 
separate  this  species  from  all  others  in  the  creasing  setal  density  ventrally.  Propo- 
genus.  Although  females  of  this  species  deum  usually  microsculptured  and  usu- 


252  Journal  of  Hymenoptera  Research 

ally  with  relatively  small  areoleae  adjacent  eira  (9:1  PMA).  Parana:  Campina  Grande 

to  enclosure  that  are  composed  of  relative-  nr.  Curitiba  11-23-1966  H.  &  M.  Townes 

ly  coarse  carinae.  Some  specimens  have  (6:1  AEI).  Pernambuco:  Caruaru  IV-1972 

larger  more  typical  areolae  adjacent  to  en-  M.  Alvarenga  (9:  2  PMA).  Caruaru  V- 

closure.  1972.  J.  Lima  (9:1  PMA).  Curuaru  [sic] 

Metasoma.  Petiole,  laterally  smooth  or  900m  IV-1972  (9:1  BRD).  Rio  de  Janeiro: 

with  several  fine  carinae.  III-5-1966  H.  &  M.  Townes  (9:   1  AEI). 

Color.  Ground  color  black.  White:  palpi,  Guanabara  11-1972.   M.   Alvarenga  ( 9 :   1 

base  of  mandible,  pronotal  lobe,  at  least  PMA).  Mangaratiba,  Muriqui  VII-1969  M. 

fore  tarsus,  sometimes  all  tarsi.  Yellow  to  Alvarenga  (9:1  AEI).  Teresopolis  III-12- 

yellow-brown:   antenna,  mandible  medi-  1966  H.  &  M.  Townes  (d:  1  AEI).  Santa 

ally,  tegula,  legs  except  at  least  fore  tarsus  Catarina:  Nova  Teutonia  27'^'11'B.52"23'L. 

and  sometimes  hind  femur  and  tibia.  Red:  8-X-1937  Plaumann.  B.M.1938-312  {6:   1 

at  least  pronotal  side  to  entire  mesosoma  BMNH).  Sao  Paulo:  Sao  Paulo  XII-31-1968 

except  legs,  pronotal  lobe,  tegula  and  pro-  V.N.  Alin  {6:1  USNM).  San  Paulo  20-1- 

podeum;  all  males  have  some  mesosomal  1978  [label  in  Russian]  {6:2  ZMMU). 

red  coloration.  Brown:  occasionally  hind  Distribution   (Map  27). — Southeastern 

femur  and  tibia.  Brazil. 

Female.  Length  4.8-5.0  mm.  Similar  to 

male  except  as  follows:  apicoventral  man-  ^-  Llaqhastigmus  muthus  Finnamore, 

dibular  tooth  enlarged,  but  only  slightly  ^^^  species 

larger    than    other    teeth.    Clypeal    apex  Derivation  of  Name. — The  species  epithet 

quadridentate  with  lateral  lobe  flanking  is   derived   from   the  Quechua   "muthu" 

lateral  edge  of  labrum,  pair  of  small  me-  which  means  dull,  in  reference  to  the  de- 

dian  teeth  separated  from  lateral  lobe  by  gree  of  microsculpture  found  on  members 

broad  emargination  and  separated  from  of  this  species. 

each   other   by    U-shaped   notch.    Apical  Diagnosis. — The  dense  uniform  micro- 

clypeal  edge  with  specialized,  elongate,  sculpture  on  the  head   (except  clypeus) 

flattened  setae  on  teeth  but  not  on  emar-  and  mesosoma  (except  sometimes  sterno- 

ginate   interspaces   (2   setae   per   median  pleural  region  and  propodeum)  will  sep- 

tooth   and   4   per   lateral   lobe).   Clypeus  arate  this  species  from  all  others  in  the  ge- 

shiny,  microsculpture  absent,  punctures  nus.  In  all  other  species  the  microsculp- 

variable  from  almost  absent  to  sparsely  ture  of  the  frons  is  of  greater  density  than 

distributed  over  entire  surface.  that  of  the  vertex. 

Material    Examined    {66,    19  9). — Holo-  Description. — Male.  Length  3.5^.0  mm. 

type:    9 ,  BRAZIL:  Caruaru,  Pernambuco  Head.  Sculpture  of  clypeus  obscured  by 

900m  IV-1972  M.  Alvarenga  (BRD).  Para-  dense   appressed   setae.    Gena    ventrally, 

types:  BRAZIL:  Bahia:  7-VIII-1983  Cepec  with  sparse  straight  setae  that  are  sub- 

Itabuna  P.P.  Benton  (9:  1  BMNH).  Gua-  equal  to  or  less  than  greatest  width  of  fore 

nabara:  Floresta  da  Tijuca  IV-1966  Alvar-  basitarsus.   Microsculpture  of  frons   and 

enga  &  Seabra  (9:  1  AEI).  Represa  Rio  vertex  of  uniform  (subequal)  density.  Oc- 

Grande,  Guanabara  XII-1967  F.H.  Oliveira  cipital  carina  ventrally,  slightly  raised  and 

(9:1  BRD).  Represa  Rio  Grande  1-1968  M.  finely  foveolate. 

Alvarenga  (9:  2  PMA).  Represa  Rio  Gran-  Mesosoma.   Transverse  sulcus  of  pro- 

de  11-1972  P.M.  Oliveira  (9:2  PMA).  Re-  notum  with  longitudinal  carinae.  Scutum 

presa  Rio  Grande  VII-1972  M.  Alvarenga  densely   microsculptured,   impunctate. 

(9:2  BRD).  Minas  Gerais:  Serra  do  Car-  Transscutellar  sulcus  foveolate.  Preomau- 

aca,  Sta.  Barbara  1-1970  (9:1  PMA).  Serra  lar  area  sparsely  setose,  setae  not  obscur- 

do  Caraca,  S.  Barbara  III-1971  P.M.  Oliv-  ing  underlying  sculpture.  Subomaulus 


Volume  4,  1995  253 

present.  Scrobal  sulcus  weakly  impressed  FSCA).  Lima:  Chancay  shrubs  nr.  river  40 

with  evanescent  usually  irregular  foveae  miles  n.  of  Lima  29-VII-1971  P.S.  &  H.L. 

best  developed  anteriorly.   Sternopleural  Broomfield  B.M. 1971^86  fertile  irrigated 

region  densely  microsculptured,  impunc-  region  in  arid  coastal  desert  {S :  I  BMNH). 

tate.  Propodeum  weakly  microsculptured  Chosica   XI-1961    N.L.H.    Krauss    {9    1 

to  shiny;  area  adjacent  to  enclosure  and  USNM).  Cupiche,  10km  n.  Chosica  25-VI- 

lateral  area  carinate;  carinae  converging  2-VII-1974  C.  Porter  &  L.  Stange  (J:  1  LIL- 

dorsolaterally  to  form  small  irregular  are-  LO,   9 :  2  LILLO).  Cupiche  26-VI-1976  C. 

olae.  Porter,  C.  Calmbacher  {S:  I  FSCA).  Lima 

Metasoma.  Petiole  with  several  coarse  13-XI-1950   E.S.   Ross   (9:   2  CAS).   Lima 

carinae  laterally.  City,  S.  Marcos  Univ.  campus  26-27-IV- 

Color.  Ground  color  black.  White:  palpi,  1983  Mai.  trap  C.&  M.  Vardy  B.M.1983- 

mandible  except  apically,  pronotal  lobe.  217  (9:4  BMNH,  1  PMA).  San  Geronimo 

Yellow   to   yellow-brown:   scape,   pedicel  ca.    Chosica    1-5-VII-1976    C.    Porter,    C. 

and  basal  flagellomeres  ventrally,  tegula,  Calmbacher  (9:3  FSCA).  San  Geronimo 

fore  and  mid  legs  except  coxae,  hind  tro-  30-VII-1982  R.B.  Miller,  L.A.  Stange  (9:2 

chanter,  hind  tarsus.  Brown:  flagellomeres  FSCA).    VENEZUELA:    Distrito   Federal: 

except  as  indicated  above,  hind  leg  except  nr.  Caracas  28-VIII-1943  D.G.  Hall  (9:  1 

trochanter  and  tarsus.  USNM). 

Female.  Length  3.7-4.6  mm.  Similar  to  Distribution  (Map  28). — Venezuela,  Fe- 
male except  as  follows:  apicoventral  man-  uador,  Peru, 
dibular    tooth   enlarged,    truncate,    much 

larger    than    other    teeth.    Clypeal    apex  ^^-  Llaqhastigmus  nigricollaris 

quadridentate  as  follows:   lateral  lobe  Finnamore,  new  species 

flanking  lateral  edge  of  labrum,  pair  of  Derivation  of  Name. — The  species  epithet 

smaller  median  teeth  separated  from  lat-  is  derived  from  two  latin  words,  niger  and 

eral  lobe  by  broad  emargination  and  from  collaris,  in  reference  to  the  dark  pronotal 

each  other  by  narrow  U-shaped   notch,  collar  characteristic  of  this  species. 

Clypeal  apex  with  specialized,  elongate.  Diagnosis. — The  combination  of  a  brown 

flattened  setae  present  on  median  teeth  (2  or  black  pronotal  lobe,  antennal  scape  yel- 

per  tooth)  and  on  lateral  lobe  (6-7  per  low-brown  and  in  the  female  the  apico- 

lobe)    with    occasionally    single    seta    on  dorsal  mandibular  tooth  (in  addition  to 

emarginate  interspace.  Clypeus  shiny,  apicoventral  tooth)  enlarged  and  truncate, 

sparsely  punctate,  area  immediately  above  will  separate  this  species  from  all  others 

median   teeth   impunctate   and    area   be-  in  the  genus. 

tween   antennal   sockets   most   densely  Description. — Male.  Length  3.7-3.9  mm. 
punctate  with  punctures  1  diameter  or  less  Head.  Sculpture  of  clypeus  obscured  by 
apart  (elsewhere  punctures  3  or  more  di-  dense   appressed    setae.    Gena    ventrally, 
ameters  apart).  Lateral  lobe  yellow-brown  with  sparse  straight  setae  that  are  sub- 
to  red-brown.  equal  to  or  less  than  greatest  width  of  fore 

Material    Examined    {86,    16  9). — Holo-  basitarsus.  Microsculpture  of  frons  more 

type:  6 ,  PERU:  Samne,  ca.  40km  ne.  Tru-  dense  than  that  of  vertex  adjacent  to  mid 

jillo,  Prov.  La  Libertad  7°59's  78°4rw  elev.  ocellus.  Occipital  carina  ventrally,  raised 

ca.    1500m    12-17-VII-1975   C.    Porter,    L.  and  foveolate. 

Stange    (LILLO).    Paratypes:    PERU:    San  Mesosoma.   Transverse  sulcus  of  pro- 

Bartolome   VII-1913   C.T.    Brues    {6:    1  notum  with  longitudinal  carinae.  Scutum 

MCZ).  La  Libertad:  same  data  as  holotype  shiny,  weakly  microsculptured  especially 

{6:  1  LILLO,  1  PMA).  15km  e.  Laredo  19-  on  disc,  punctures  sparse.  Transscutellar 

VII-1982  R.B.  Miller,  L.  A.  Stange  {6:  1  sulcus  foveolate.  Preomaular  area  sparsely 


254  Journal  of  Hymenoptera  Research 

setose,    setae    not    obscuring    underlying  XII-1974  Malaise  trap  (?:  1  FSCA).  Penas 

sculpture.    Subomaulus    present.    Scrobal  Blancas  1750m  10km  w.  Call,  very  wet 

sulcus  weakly  impressed,  irregularly  fo-  premontane  forest,  R.C.  Wilkerson  20-22- 

veolate  anteriorly,  occasionally  entirely  fo-  1-1975  Malaise  trap  ( $ :  3  FSCA).  Penas 

veolate.  Sternopleural  region  weakly  mi-  Blancas  1750m  10km  w.  Cali,  very  wet 

crosculptured  with  sparse  punctures  and  premontane  forest,  R.C.   Wilkerson  31-1- 

setae.    Propodeum    shiny,    with    several  1975  Malaise  trap  (9:3  FSCA,  1  PMA). 

coarse  areolae  adjacent  to  enclosure  com-  Penas    Blancas    12-11-1975    R.    Wilkerson 

posed  of  relatively  high  carinae.  Malaise  trap  (9:3  FSCA,  1  PMA).  Penas 

Metasoma.  Petiole  with  2  to  4  coarse  ca-  Blancas  17-19-III-1975  R.  Wilkerson  Mal- 

rinae  laterally.  aise  trap  (9:1  FSCA).  ECUADOR:  Napo: 

Color.  Ground  color  black.  Yellow  to  El  Chaco  2000m  11-1983  M.  Sharkey,  L. 
yellow-brown:  palpi,  mandibles  medially,  Masner  {6:1  BRD).  PERU:  Huanuco:  Tin- 
scape,  pedicel  and  basal  flagellomeres  go  Maria,  40  mi.  s.  Carpish  Mts.  XII-28- 
ventrally,  tegula,  fore  and  mid  legs  except  1954  E.I.  Schlinger  &  E.S.  Ross  (9:1  CAS), 
coxae  and  usually  except  femora,  hind  tro-  Distribution  (Map  29). — Colombia,  Ec- 
chanter  and  tarsus.  Brown:  mandible  ba-  uador,  Peru, 
sally,  antenna  other  than  noted  above, 
pronotal  lobe,  femora,  hind  tibia.  l^-  Llaqhastigmus  santanderanus 

Female.  Length  4.2-5.0  mm.  Similar  to  Finnamore,  new  species 

male,  except  as  follows:  apicoventral  and  Derivation  of  Name. — The  species  epithet 

apicodorsal    mandibular    teeth    enlarged,  is  derived  from  the  Colombian  state  of 

truncate.  Clypeal  apex  quadridentate  as  Santander   Del   Norte,   the   northernmost 

follows:  lateral  lobe  flanking  lateral  edge  record  of  the  species. 

of  labrum,  pair  of  small  median  teeth  sep-  Diagnosis. — The  species  is  distinguished 
arated  from  lateral  lobe  by  deep,  broad,  by  the  lack  of  a  subomaulus,  and  a  petiole 
emargination  and  from  each  other  by  shal-  that  is  finely  carinate  to  microcarinate  lat- 
low  notch.  Clypeal  apex  with  specialized,  erally.  This  species  may  prove  to  be  con- 
flattened,  elongate  setae  on  lateral  lobe  (4  specific  with  australis,  at  present  there  are 
or  5)  and  on  median  teeth  (2  per  tooth)  but  too  few  specimens  to  assess  variation 
generally  absent  on  emarginate  interspac-  within  either  species, 
es.  Clypeus  shiny,  microsculpture  absent.  Description. — Male.  Length  4.8-5.2  mm. 
punctures  sparse  (3  or  more  diameters  Head.  Sculpture  of  clypeus  obscured  by 
apart  laterally,  slightly  more  dense  medi-  dense  appressed  setae.  Gena  ventrally, 
ally),  lateral  lobe  red-brown.  with  sparse  straight  setae  that  are  sub- 

Material    Examined    {26,    16  9 ). — Holo-  equal  to  or  less  than  greatest  width  of  fore 

type:   9,  COLOMBIA:  Valle  Dept.:  Penas  basitarsus.  Microsculpture  of  frons  only 

Blancas  1750m   10km  w.  Cali,  very  wet  slightly  more  dense  than  that  of  vertex, 

premontane  forest,  R.C.  Wilkerson  15-1-  Occipital  carina  ventrally,  raised  and  fov- 

1975  Malaise  trap  (FSCA).  Paratypes:  CO-  eolate. 

LOMBIA:  Cauca:  Popayan  1,800m  11-X-  Mesosoma.   Transverse  sulcus  of  pro- 

1974    M.    Cooper    B.M. 1975-33    {16:    1  notum  with  longitudinal  carinae.  Scutum 

BMNH,  9:  1  BMNH).  Valle:  Cali  900m  17-  microsculptured,    punctures    sparse,    ob- 

1-1972    M.    Cooper    B.M.1972-275    (9:    1  scure.  Transscutellar  sulcus  not,  or  at  most 

BMNH).  Pance  CVC  1700m  15km  w.  Cali,  obscurely   foveolate.    Preomaular   area 

very  wet  premontane  forest,  R.C.  Wilker-  sparsely  setose,  setae  not  obscuring  un- 

son  23-XII-1974  Malaise  trap  (9:  1  FSCA).  derlying   sculpture.   Subomaulus    absent. 

Penas  Blancas  1750m  10km  w.  Cali,  very  Scrobal  sulcus  slightly  impressed,  foveae 

wet  premontane  forest,  R.C.  Wilkerson  23-  absent  or  slightly  evident  anteriorly.  Ster- 


Volume  4,  1995 


255 


nopleural  region  microsculptured,  sparse- 
ly punctate.  Propodeum  microsculptured, 
area  adjacent  to  enclosure  with  fine  irreg- 
ular carinae,  some  small  areolae  dorsola- 
terally  that  are  composed  of  fine,  low  ca- 
rinae. 

Metasoma.  Petiole  microcarinate  later- 
ally or  with  fine  carinae  laterally. 

Color.  Ground  color  black.  White:  man- 
dible except  apically,  pronotal  lobe.  Yel- 
low-brown: palpi,  antenna,  tegula,  fore 
and  mid  legs  except  coxae,  hind  trochan- 
ter and  tarsus.  Brown:  hind  femur  and  tib- 
ia. 

Female.  Unknown. 

Material  Examined  (2c?). — Holotype  6, 
ECUADOR:  Napo:  Pastaza,  Sebundoi, 
2600m  11-15-IX-1977  L.  Peha  B.M.1978- 
293  (BMNH).  Paratype:  COLOMBIA: 
Santander  Del  Norte:  Oroque  lO-Vl-1965 
J.&B.  Bechyne  {6:1  UCV). 

Distribution  (Map  30). — Colombia,  Ec- 
uador. 

12.  Llaqhastigmus  sapanis  Finnamore, 
new  species 

Derivation  of  Name. — The  species  epithet 
is  derived  from  the  Quechua  "sapan" 
meaning  lone,  in  reference  to  the  single 
known  male  specimen. 

Diagnosis. — This  species  can  be  separat- 
ed from  all  others  in  the  genus  with  the 
following  combination  of  characters: 
transverse  sulcus  of  pronotum  smooth 
and  propodeum  areolate  dorsolaterally. 
The  only  other  species  known  to  have  a 
smooth  pronotal  sulcus  is  Ihitanis  which 
differs  from  sapmnis  in  its  non-areolate  pro- 
podeum. 

Description. — Male.  Length  3.8  mm. 

Head.  Sculpture  of  clypeus  obscured  by 
dense  appressed  setae.  Gena  ventrally, 
with  sparse  straight  setae  that  are  sub- 
equal  to  or  less  than  greatest  width  of  fore 
basitarsus.  Microsculpture  of  frons  more 
dense  than  that  of  vertex.  Occipital  carina 
ventrally,  raised  and  foveolate. 

Mesosoma.  Transverse  sulcus  of  pro- 
notum smooth,  without  carinae.  Scutum 


microsculptured,  punctures  relatively 
coarse  and  sparse  (usually  3  or  more  di- 
ameters apart  on  disc).  Transscutellar  sul- 
cus foveolate.  Preomaular  area  sparsely 
setose,  setae  not  obscuring  underlying 
sculpture.  Subomaulus  present.  Scrobal 
sulcus  weakly  impressed,  not  foveolate. 
Sternopleural  region  microsculptured, 
punctures  sparse  and  evanescent.  Propo- 
deum microsculptured,  area  adjacent  to 
enclosure  and  dorsolateral  region  areolate, 
areolae  composed  of  relatively  low  cari- 
nae; side  irregularly  carinate. 

Metasoma.  Petiole  carinate  laterally. 
Color.  Ground  color  black.  White:  man- 
dible except  apically,  pronotal  lobe.  Yel- 
low-brown: palpi,  scape,  ventrally  on  ped- 
icel and  basal  flagellomeres,  tegula,  fore 
and  mid  legs  except  coxae  and  femora, 
hind  tarsus.  Brown:  antenna  except  as  not- 
ed, fore  and  mid  femora,  hind  leg  except 
tarsus  and  coxa. 

Female.  Length  4.0-4.1  mm.  Similar  to 
male  except  as  follows:  apicoventral  man- 
dibular tooth  enlarged,  truncate,  much 
larger  than  other  teeth.  Clypeal  apex 
quadridentate  as  follows:  lateral  tooth 
flanking  lateral  edge  of  labrum,  pair  of 
smaller  median  teeth  separated  from  lat- 
eral tooth  by  broad  emargination  and  sep- 
arated from  each  other  by  narrow  U- 
shaped  notch.  Clypeal  apex  with  special- 
ized, flattened,  elongate  setae  confined  to 
teeth  (2  per  median  tooth  and  3  or  4  per 
lateral  tooth).  Clypeus  shiny,  without  mi- 
crosculpture, punctures  sparse  and  rela- 
tively more  dense  between  antennal  sock- 
ets. Apical  margin  of  clypeus  black. 

Material  Examined  {IS,  49).— Holotype: 
9,  PERU:  Machu  Picchu  XI-28-1965  H.& 
M.  Townes  (AEl).  Paratypes:  BOLIVIA: 
Cochabamba:  Carrasco,  Empalme  3000m 
11-1971  Fritz  y  Martinez  {6:  1  IIES).  PERU: 
Cuzco:  Machu  Picchu  XIl-1-1965  H.  &  M. 
Townes  (9:1  AEI).  Machu  Picchu  11-24- 
27-1968  A.  Garcia  &  C.  Porter  (9 :  2  MCZ). 
Distribution  (Map  31).— Bolivia,  Peru. 


256 


Journal  of  Hymenoptera  Research 


13.  Llaqhastigmus  shachus  Finnamore, 
new  species 

Derivation  of  Name. — The  species  epithet 
is  derived  from  the  Quechua  "sh'achu", 
meaning  hairy  in  reference  to  the  dense 
pilosity  of  this  species. 

Diagnosis. — This  species  can  be  distin- 
guished by  the  presence  of  dense,  elongate 
setae  on  the  lower  gena  and  sternopleural 
region.  In  the  male,  setae  of  lower  gena 
are  bent  apically;  in  the  female  only  a  few 
near  the  hypostomal  carina  are  bent  api- 
cally. In  both  sexes  the  lower  genal  setae 
are  longer  than  the  greatest  width  of  the 
fore  basitarsus.  Females  of  this  species 
have  a  quadridentate  mandible  due  to  a 
deep  U-shaped  notch  in  the  apicodorsal 
mandibular  tooth  (visible  only  by  spread- 
ing the  mandibles).  All  other  species  have 
tridentate  mandibles  and  short  sparse  se- 
tae on  the  lower  genal  region.  Setae  in  oth- 
er species  are  never  longer  than  greatest 
width  of  fore  basitarsus.  Other  species 
have  little  or  no  setae  on  the  sternopleural 
region. 

Description. — Male.  Length  4.4  mm. 

Head.  Sculpture  of  clypeus  obscured  by 
dense  appressed  setae.  Gena  ventrally, 
with  dense  elongate  setae  that  are  about 
twice  the  greatest  width  of  fore  basitarsus 
and  bent  90'  inward  toward  oral  cavity. 
Microsculpture  of  frons  slightly  more 
dense  than  that  of  vertex.  Occipital  carina 
ventrally,  raised  and  foveolate. 

Mesosoma.  Transverse  pronotal  sulcus 
carinate.  Scutum  microsculptured,  punc- 
tures small  and  sparse.  Transscutellar  sul- 
cus foveolate.  Subomaulus  present.  Preo- 
maular  area  densely  setose,  setae  erect  not 
obscuring  underlying  sculpture.  Scrobal 
sulcus  weakly  impressed,  a  few  evanes- 
cent irregular  foveae  anteriorly.  Sterno- 
pleural region  weakly  microsculptured, 
densely  and  minutely  punctate  (punctures 
about  2  diameters  apart  ventrally);  setae 
dense  ventrally  and  elongate,  slightly 
shorter  than  genal  setae  and  with  many 
tips  bent  90°  toward  posterior.  Propodeum 


weakly  microsculptured,  area  adjacent  to 
enclosure  and  dorsolateral  sphere  with 
numerous  areolae  composed  of  relatively 
low  carinae,  side  more  or  less  vertically 
carinate  with  carinae  merging  into  areolae 
of  dorsolateral  sphere. 

Metasoma.  Petiole  laterally,  with  4  lon- 
gitudinal coarse  carinae. 

Color.  Ground  color  black.  White:  man- 
dibles basally,  pronotal  lobe.  Yellow- 
brown:  Palpi,  mandible  medially,  scape 
ventrally,  tegula,  fore  and  mid  legs  except 
coxae  and  femora,  hind  tarsus. 

Female.  Length  4.8-5.4  mm.  Similar  to 
male  except  as  follows:  mandible  quadri- 
dentate, apicodorsal  tooth  with  deep  U- 
shaped  notch  thereby  imparting  quadri- 
dentate appearance  to  the  mandible;  api- 
coventral  mandibular  tooth  enlarged, 
truncate,  much  larger  than  other  teeth. 
Clypeal  apex  quadridentate  as  follows: 
lateral  lobe  flanking  lateral  edge  of  la- 
brum,  pair  of  median  teeth  separated  from 
lateral  lobe  by  broad  emargination  and 
from  each  other  by  U-shaped  notch.  Clyp- 
eal apex  with  specialized,  flattened,  elon- 
gate setae  distributed  over  most  of  apex. 
Clypeus  shiny,  without  microsculpture, 
punctures  sparse  (5-10  diameters  apart). 
Apical  clypeal  margin  black.  Genal  and 
sternopleural  setae  not  as  dense  or  as  long 
as  those  in  male.  Omaulus  ending  ven- 
trally, not  continued  forward  to  acetabular 
carina. 

Material  Examined  (16,  2  9). — Holotype: 
9,  PERU:  Huanuco  IX-16-1954  E.I.  Schlin- 
ger  &  E.S.  Ross  (CAS).  Paratypes:  ECUA- 
DOR: Loja:  14km  w.  Catamayo  1770m  9- 
VII-1989  L.  Stange  &  R.  Miller  (9:  1 
FSCA).  PERU:  Huanuco:  Huanuco  IX:16- 
1954  E.I.  Schlinger  &  E.S.  Ross  {6:  1  CAS). 
Distribution  (Map  32). — Ecuador,  Peru. 

14.  Llaqhastigmus  sharkeyi  Finnamore, 
new  species 

Derivation  of  Name. — This  species  is 
named  in  honour  of  Michael  J.  Sharkey, 
co-collector  of  the  primary  type  material. 

Diagnosis. — The  combination  of  a  white 


Volume  4,  1995  257 

pronotal  lobe;  shiny  scutum  with  punc-  along  edge  except  for  median  teeth  which 
tures  minute  and  sparse,  without  micro-  bear  2  additional  setae  each.  Clypeus 
sculpture  on  posterior  %  except  narrow  shiny,  punctures  minute  but  dense  later- 
posterior  margin;  and  a  coarsely  foveolate  ally  and  larger  and  more  sparse  medially 
scrobal  sulcus  will  separate  this  species  (1  to  2  diameters  apart).  Apical  margin  of 
from  all  others  in  the  genus.  clypeus  black. 

Description. — Male.  Length  3.6-4.7  mm.  Material  Examined  {46 ,  4  9). — Holotype: 

Head.  Sculpture  of  clypeus  obscured  by  S ,  ECUADOR:  Napo  Prov.:  Baeza,  5km  s. 

dense   appressed    setae.    Gena    ventrally,  11-1983    2000m    Sharkey,    Masner    (BRD). 

with  sparse  straight  setae  that  are  sub-  Paratypes:    COLOMBIA:    Santander   del 

equal  to  or  less  than  greatest  width  of  fore  Norte:  Prima  1700m  27-V-1965  J.  &  B.  Be- 

basitarsus.  Microsculpture  of  frons  more  chyne   (9:    1    UCV).    ECUADOR:    Napo: 

dense  than  that  of  vertex.  Occipital  carina  Baeza,    5km    s.    11-1983    2000m    Sharkey, 

ventrally,  raised  and  foveolate.  Masner  (9:1  BRD).  El  Chaco  2000m  II- 

Mesosoma.  Transverse  pronotal  sulcus  1983  M.  Sharkey,  L.  Masner  {6:1  PMA). 

with  longitudinal  carinae.  Scutum  shiny  Pastaza,    Reventador    1750m    3-5-X-1977 

on  most  of  posterior  %,  microsculptured  L.E.   Pena   B.M.1978-293   {6:    1    BMNH). 

anteriorly  and  narrowly  on  posterior  mar-  VENEZUELA:  Aragua:   Hac.   Portapan 

gin;  punctures  minute,  sparse,  10  or  more  1700m  15-VIII-1969  J.  &  B.  Bechyne  (9:1 

diameters  apart.  Transscutellar  sulcus  fo-  UCV).  Rancho  Grande  1100m  16-X1-1967  J. 

veolate.  Preomaular  area  sparsely  setose,  &  b.  Bechyne  (9:  1  UCV).  Rancho  Grande, 

setae  not  obscuring  underlying  sculpture.  Portachuelo  1100m  22-V-1981  J.A.  Clavijo, 

Subomaulus  present.  Scrobal  sulcus  im-  J. L.  Garcia  ((5:  1  UCV).  Falcon:  Curimagua 

pressed,    foveolate.    Sternopleural   region  uoOm  l-XII-1971  J.  &  B.  Bechyne  (9:1 

shiny,  without  microsculpture,  punctures  UCV). 

minute    and    sparse.    Propodeum    shiny,  Distribution    (Map    33).— Ecuador,    Co- 

without  microsculpture,  area  adjacent  to  lombia  Venezuela, 
enclosure  and  lateral  sphere  with  areolae 

composed  of  relatively  high  carinae;  side  GENERIC  RELATIONSHIPS 
unsculptured  toward  metapleuron. 

Metasoma.  Petiole  carinate  laterally.  The  following  hypothesis  of  relation- 
Color.  Ground  color  black.  White:  pro-  ships  of  genera  among  the  Shgmma  is 
notal  lobe.  Yellow  to  yellow-brown:  palpi,  based  on  outgroup  analysis  of  character 
mandible  medially,  scape,  pedicel,  ven-  state  distribution.  The  characters  listed  be- 
trally  on  basal  flagellomeres  or  more,  teg-  ^^w  were  polarized  by  outgroup  analysis 
ula,  fore  and  mid  legs  except  coxae,  hind  with  the  Spilomenina.  Morphology  is  after 
tarsus.  Brown:  antennal  flagellum  except  Bohart  and  Menke  (1976)  unless  otherwise 
as  noted  above,  hind  leg  except  tarsus.  indicated  in  introduction,  only  characters 
Female.  Length  4.2^.6  Similar  to  male  not  sufficiently  treated  by  those  authors 
except  as  follows:  apicoventral  mandibu-  are  explained  in  detail  below.  In  the  fol- 
iar tooth  enlarged,  truncate,  much  larger  lowing  analysis  0  denotes  the  plesiotypic 
than  other  teeth.  Clypeal  apex  quadriden-  state  and  1  to  5  denote  various  apotypic 
tate  as  follows:  lateral  tooth  flanking  lat-  expressions  of  a  character  and  do  not  nec- 
eral  edge  of  labrum,  pair  of  smaller  me-  essarily  represent  transition  series.  Data 
dian  teeth  separated  from  lateral  tooth  by  matrices  for  the  Stigmina,  Pemphredoni- 
broad  emargination  and  separated  from  na,  Spilomenina,  and  Ammoplanina  are 
each  other  by  narrow  U-shaped  notch,  presented  in  Tables  I-IV  respectively.  A 
Clypeal  apex  with  5  or  6  specialized,  elon-  cladogram  illustrating  relationships  is  pre- 
gate,   flattened  setae  equally   distributed  sented  in  Fig.  51.  The  cladogram  is  based 


258 


Journal  of  Hymenoptera  Research 


o 
c 

•§ 


••3      '-a 


3 

2 


o 

JQ 

b 


I  S      I  3 

•I  .1  -I  I 

*•  ^^  ■*"  ••■ 

CA  TSs  CA  ■'irf 


3     ^ 


Fig.  51.     Cladogram  depicting  relationships  among  genera  of  Stigmina. 


on  the  following  characters  and  was  de-     Stigmina  and  the  Pemphredonina,  palpal 

veloped  without  electronic  assistance.  segment  length  generally  exceeds  twice  its 

Character  Polarization  ^^^t^^'  P^^P^  greatly  exceed  hypostomal 


carma. 


1.  Palpal  formula.  1— Palpal  formula  6-A,  palpal  segments 

0 — Palpal  formula  6-4,  palpal  segments     short,  palpi  inconspicuous.  Found  in  the 

long,   palpi   conspicuous.    Found    in    the     Ammoplanina  (genera  examined:  Piilver- 


Volume  4,  1995 


259 


ro,  Ammoplanus,  Ammoplanops,  Parammo- 
planus,  Ammoplanellus,  Timberlakena),  pal- 
pal segment  length  generally  about  twice 
its  width  or  less,  palpi  at  most  slightly  ex- 
ceed hypostomal  carina. 

2 — Palpal  formula  5-4,  palpal  segments 
short,  palpi  inconspicuous.  This  character 
is  synapotopic  for  the  Spilomenina  and 
was  the  justification  used  by  Menke  (1989) 
to  separate  the  group  from  the  Stigmina. 
Palpal  segments  are  short,  particularly 
basal  segments  which  may  be  no  longer 
than  wide,  palpi  at  most  slightly  exceed 
hypostomal  carina. 

2.  Labrum  shape. 

0 — Labrum  truncate  to  broadly  round- 
ed, wider  than  long.  This  character  state  is 
found  in  Arpactophilus,  Microstigmus,  Xys- 
ma,  Pulverro,  Ammoplanops,  and  many  oth- 
er apoid  and  vespoid  wasps.  This  charac- 
ter state  in  the  genera  listed  above  may 
represent  a  reversal  from  2.3  or  2.4.  Only 
a  few  species  and  specimens  were  avail- 
able for  dissection  in  these  groups,  fur- 
thermore the  difference  between  truncate 
and  broadly  rounded  is  sometimes  slight, 
rendering  the  polarity  of  this  character  es- 
pecially difficult  to  establish. 

1 — Labrum  triangular,  longer  than 
wide.  Present  in  Pemphredon,  Polemistus, 
and  Carinostigmus,  as  well  as  most  Am- 
pulicinae  (Ampulex)  and  Ammophilini 
(Sphecinae).  Sides  of  triangle  are  out- 
curved,  basal  and  apical  regions  are  not 
distinguished.  This  state  may  represent 
the  correct  plesiotypic  expression  of  the  la- 
brum in  the  Pemphredonini. 

2 — Labial  apex  distinguished  from  base. 
In  Passnloecus  the  lateral  margins  of  the  la- 
brum are  incurved  resulting  in  a  state  in 
which  the  base  is  abruptly  broadened  and 
thereby  distinguished  from  the  apex.  The 
abrupt  shoulder  is  present,  although  mod- 
ified, in  all  genera  listed  under  2.3,  2.4, 
and  presumably  2.5  although  I  have  not 
seen  specimens  of  the  latter. 

3 — Labrum  emarginate  to  bilobed.  This 
development  ranging  from  a  slight  emar- 


gination  found  in  Paracrabro  to  the  deeply 
bilobed  expression  found  in  Diodontus, 
Llaqhastigmus,  Arpactophilus,  Ammop)lamts, 
Parammop)lanus,  Ammoplanellus,  Timberlak- 
ena and  as  well  is  present  in  those  genera 
listed  under  2.4.  The  labrum  of  Llaqhastig- 
mus is  considered  a  character  reversal 
from  2.4  caused  by  a  reduction  of  the  me- 
dian lobes. 

4 — Labrum  4-lobed.  This  character  state 
results  from  an  apical  shift  of  the  lateral 
shoulders  (see  character  state  2.2  above) 
imparting  a  4-lobed  appearance  to  the  la- 
brum. Found  in  Stigmus,  Tzustigmus,  Ar- 
aucastigmus,  Parastigmus,  Aykhustigmus,  In- 
castigmus,  Arpactophilus,  and  Spilomena. 

5 — Labrum  6-lobed.  I  have  not  seen 
specimens  bearing  this  character.  Menke 
(1989)  noted  undescribed  Arpactophilus 
having  a  "six  toothed  labrum". 

3.  Mandibular  socket. 

0— Open. 
1 — Closed. 

4.  Apical  mandibular  teeth. 

0 — Mandible  with  2  apical  teeth.  This 
character  state  is  found  in  males  of  all  gen- 
era of  Stigmina,  except  Incastigmus  and 
Llaqhastigmus,  it  also  occurs  in  both  sexes 
of  all  genera  in  the  Spilomenina  and  spo- 
radically in  all  genera  of  the  Pemphredon- 
ina. 

1 — Mandible  unidentate,  apically  ter- 
minating in  a  single  point  or  truncate. 

2 — Mandible  tridentate  apically  (Figs. 
37,  39).  Females  of  most  genera  of  Stig- 
mina (except  Parastigmus,  Paracrabro  and 
some  Stigmus)  and  males  of  Incastigmus 
and  Llaqhastigmus.  The  character  state  also 
occurs  in  some  species  of  Passaloecus  and 
Pemphredon. 

3 — Mandible  with  4  apical  teeth. 

4 — Mandible  with  5  apical  teeth. 

5 — Manciible  with  6  apical  teeth. 

5.  Apicoventral  mandibular  tooth  in 

females  having  a  tridentate  mandible 
(character  4.2). 

0 — Apicoventral  tooth  acute,  similar  in 
size  to  other  teeth  (Fig.  39). 


260 


Journal  of  Hymenoptera  Research 


1 — Apicoventral  tooth  enlarged  and 
truncate  (Fig.  37).  This  character  state  oc- 
curs in  females  of  Llaqhastigmus  and  in- 
dependently in  females  of  some  Pemphre- 
don. 

6.  Clypeal  apicolateral  bevel. 

0 — Clypeus  unmodified  apicolaterally. 

1 — Clypeal  apex  bevelled  apicolaterally 
(Fig.  42).  The  lateral  clypeal  bevel  is  an  au- 
tapotypic  character  state  occurring  in  Stig- 
miis.  It  occurs  only  in  males  and  is  most 
developed  in  species  from  the  Neotropical 
Region. 

7.  Clypeus  size. 

0 — Clypeus  not  reduced,  antennal  sock- 
et usually  more  than  its  diameter  from 
nearest  clypeal  edge;  labrum  generally 
flat,  lightly  sclerotized,  mostly  or  entirely 
obscured.  Found  in  males  and  most  fe- 
males. 

1 — Clypeus  reduced,  antennal  socket 
about  its  diameter  from  nearest  clypeal 
edge,  labrum  convex,  heavily  sclerotized, 
entirely  exposed,  or  almost  so.  Some  fe- 
males of  a  few  genera. 

8.  Clypeal  teeth. 

0 — Clypeus  without  teeth  or  lobes  on 
apical  margin. 

1 — Clypeus  with  single  median  tooth  or 
lobe. 

2 — Clypeus  with  2  median  teeth. 

3 — Clypeus  with  3  teeth. 

4 — Clypeus  with  4  teeth. 

5 — Clypeus  with  2  lateral  teeth,  median 
teeth  lost. 

9.  Interantennal  tubercle. 

0 — Reduced  or  absent,  usually  repre- 
sented by  a  small,  raised  conical  point,  sel- 
dom exceeding  half  width  of  antennal 
scape  (Fig.  36). 

1 — Protruberant,  often  exceeding  width 
of  antennal  scape  (Figs.  6,  7). 


10.  Frontal  carina. 

0 — Frontal  carina  absent,  indicated  by  a 
groove,  complete  or  in  part,  from  mido- 
cellus  to  clypeus. 

1 — Frontal  carina  absent,  frons  flat, 
without  indication  of  groove  or  carina. 

2 — Frontal  carina  present  as  a  simple 
raised  line,  complete  or  in  part,  from  mi- 
docellus  to  ocellus. 

3 — Frontal  carina  strongly  raised,  la- 
mellate, usually  between  antennal  sockets. 

11.  Vertex  micropore  field. 

0 — Vertex,  between  lateral  ocellus  and 
eye,  of  similar  microsculpture  and  punc- 
tation  to  adjacent  areas;  without  group  of 
loosely  associated  punctures  or  discrete 
pore  field. 

1 — Vertex,  between  lateral  ocellus  and 
eye,  with  a  loosely  associated  group  of 
punctures  (Figs.  3,  8). 

2 — Vertex,  between  lateral  ocellus  and 
compound  eye,  with  a  micropore  field,  ap- 
pearing as  a  discrete  patch  of  fine  micro- 
sculpture  relative  to  adjacent  microsculp- 
ture (Figs.  19,  20,  25,  26,  31,  32). 

12.  Eye  margins. 

0 — Eye  not  margined  by  a  carina. 
1 — Eye  at  least  partially  margined  by  a 
carina. 

13.  Occipital  Carina. 

0 — Occipital  carina  present,  simple. 
1 — Occipital  carina  raised,  foveolate. 
2 — Occipital  carina  incomplete  dorsally, 
present  ventrolaterally. 
3 — Occipital  carina  absent. 

14.  Occipital-hypostomal  carinae. 

0 — Occipital  carina  intersecting  hypo- 
stomal  carina. 

1 — Occipital  carina  ending  ventrally  not 
intersecting  hypostomal  carina. 

2 — Occipital  carina  forming  a  complete 
circle,  not  intersecting  hypostomal  carina. 


Volume  4,  1995 


261 


15.  Pronotal  collar, 

0 — Pronotal  collar  of  normal  length, 
transverse  sulcus  without  median  longi- 
tudinal raised  area  (Figs.  7,  10). 

1 — pronotal  collar  elongate,  transverse 
sulscu  with  median  longitudinal  area 
raised  (Figs.  2,  5). 

16.  Transverse  pronotal  carina. 

0 — Present,  at  least  laterally. 
1 — Absent. 

17.  Median  scutal  groove. 

0 — Scutum  without  median  groove  or 
pit  (Figs.  38,  41). 

1 — Scutum  with  a  median  groove  or  at 
least  a  median  posterior  pit  (Figs.  33,  35). 

18.  Notaular  grooves. 

0 — Notaular  grot^ves  unmodified,  pres- 
ent anterolaterally  (Figs.  38,  41). 

1 — Notaular  grooves  elongate,  often  to 
posterior  margin  of  scutum  (Figs.  33,  35). 

19.  Acetabular  carina. 

0 — Acetabular  carina  absent  (Fig.  11). 
1 — Acetabular  carina  present  (Fig.  43). 

20.  Omaulus. 

0 — Omaulus  present. 
1 — Omaulus  absent. 

21.  Episternal  sulcus. 

0 — Episternal  sulcus  present. 
1 — Episternal  sulcus  absent. 

22.  Hypersternaulus. 

0 — Hypersternaulus  present. 
1 — Hypersternaulus  absent. 

23.  Orientation  of  hypersternaulus. 

0 — Obliquely  oriented. 
1 — Horizontally  oriented. 

24.  Scrobal  sulcus. 

0— Present  (Fig.  34). 
1— Absent  (Fig.  14). 


25.  6  midbasitarsus. 

0 — Midbasitarsus  elongate,  longer  than 
remaining  tarsomeres  combined. 

1 — Midbasitarsus,  straight,  cylindrical, 
shorter  than  remaining  tarsomeres  com- 
bined. 

2 — Midbasitarsus  short,  modified — 
curved,  excavated,  expanded  or  combina- 
tions thereof. 

26.  Hindtibial  spines. 

0 — One  or  more  rows  of  spines  on  pos- 
terior surface. 

1 — 2-4  widely  separated  spines  on  pos- 
terior surface. 

2 — Without  spines  on  posterior  surface. 

27.  Propodeal  enclosure. 

0 — Present. 
1 — Absent. 

28.  Stigma  size. 

0 — Stigma  of  fore  wing  of  normal  size, 
smaller  than  marginal  cell. 

1 — Stigma  of  fore  wing  enlarged  about 
size  of  marginal  cell  or  larger. 

29.  Stigma  shape. 

0 — Stigma  of  fore  wing  broadly  lentic- 
ular. 

1 — Stigma  of  fore  wing  subglobular. 

30.  Stigma  micropore  field. 

0 — Stigma  of  fore  wing  without  micro- 
pore field,  dorsal  surface  uninterrupted. 

1 — Stigma  of  fore  wing  with  micropore 
field,  appearing  as  a  discrete  circular  or 
elliptical  microsculpture  patch  interrupt- 
ing dorsal  surface  (Figs.  46,  47). 

31.  Marginal  cell  size. 

0 — Marginal  cell  not  reduced,  larger 
than  stigma. 

1 — Marginal  cell  reduced,  smaller  than 
stigma. 

32.  Marginal  cell  veins. 

0 — Marginal  cell  closed. 
1 — Marginal  cell  open. 


262 


Journal  of  Hymenoptera  Research 


33.  Submarginal  cells. 

0 — Two  submarginal  cells. 
1 — One  submarginal  cell. 
2 — Submarginal  cells  absent. 

34.  Submarginal  cell  position. 

0 — Outer  veinlet  of  submarginal  cell  1 
out  of  line  with  outer  veinlet  of  marginal 
and  discoidal  cells. 

1 — Outer  veinlet  of  submarginal  cell  1 
in  line  with  outer  veinlet  of  marginal  and 
discoidal  cells. 

35.  Submarginal  cell  I  veins. 

0 — Submarginal  cell  1  closed. 
1 — Submarginal  cell  1  open. 

36.  Discoidal  cell  number. 

0 — Fore  wing  with  three  discoidal  cells. 
1 — Fore  wing  with  two  discoidal  cells. 
2 — Fore  wing  with  one  discoidal  cell. 

37.  Hind  wing  media  divergence. 

0 — Hind  wing  media  diverging  before 
or  at  cu-a. 

1 — Hind  wing  media  diverging  just  af- 
ter cu-a. 

2 — Hind  wing  media  diverging  far  after 
cu-a. 

38.  Hind  wing  submedian  cell. 

0 — Hind  wing  submedian  cell  about 
half  or  more  length  of  median  cell. 

1 — Hind  wing  submedian  cell  about 
one  third  length  of  median  cell. 

39.  Hind  wing  cells. 

0 — Closed  cells  present  in  hind  wing. 
1 — Hind  wing  without  closed  cells. 

40.  Petiole  length. 

0 — Petiole  undeveloped,  abdomen  ses- 
sile. 

1 — Petiole  developed,  composed  of  ster- 
num only,  wider  than  long.  Some  Micros- 
tigmus  approximate  this  character  state 
but  the  apparent  petiole  is  composed  of 
both  tergum  and  sternum. 


2 — Petiole  developed,  composed  of  ster- 
num only,  longer  than  wide. 

41.  Petiolar  sculpture. 

0 — Petiole  carinate — 2  or  more  longitu- 
dinal carinae. 

1 — Petiole  punctate. 

2 — Petiole  smooth  and  round. 

42.  Sternum  I  carinae. 

0 — Metasomal  sternum  1  without  basal 
carinae. 

1 — Metasomal  sternum  1  with  single 
basal  medial  carina. 

2 — Metasomal  sternum  1  with  two  basal 
median  carinae. 

43.  Sternum  1  basal  groove. 

0 — Metasomal  sternum  1  basal  trans- 
verse groove  undeveloped. 

1 — Metasomal  sternum  1  with  a  trans- 
verse groove  or  constriction  just  posterior 
to  base. 

44.  Tergum  1  lateral  carina. 

0 — Metasomal  tergum  1  with  a  lateral 
carina. 

1 — Metasomal  tergum  1  lateral  carina 
absent. 

45.  Sternum  2  microsetal  patches. 

0 — Metasomal  sternum  2  without  mi- 
crosetal patches. 

1 — Metasomal  sternum  2  with  discrete 
microsetal  patch  laterally  (Figs.  44,  45). 

46.  Pygidial  Plate. 

0 — Pygidial  plate  present,  broad. 
1 — Pygidial    plate    present,    narrow, 
trough-like. 

2 — Pygidial  plate  absent. 

47.  Male  Sternum  8. 

0 — Metasomal  sternum  8  broad,  not 
narrowed  apically. 

1 — Metasomal  sternum  8  triangular, 
gradually  narrowed  apically. 

2 — Metasomal  sternum  8  narrowly  tri- 
angular, base  differentiated  from  apex. 


Volume  4,  1995 


263 


Fig.  52.  World  distribution  and  post  continental  drift  dispersal  routes  for  Carinostigmus  (light  shading)  and 
other  genera  in  the  Stigmina  (dark  shading).  Wide  lines  indicate  generic  level  dispersals,  narrow  lines  indicate 
species  group  dispersal. 


3 — Metasomal  sternum  8  abruptly  nar- 
rowed, apical  portion  forming  a  pseudo- 
sting,  elongate  and  parallel-sided. 

4 — Metasomal  sternum  8  spatulate  api- 
cally. 

48.  Genitalia. 

0 — Digitus  and  cuspis  lobular,  more  or 
less  equal  in  length. 

1 — Digitus,  elongate-triangular,  longer 
than  cuspis. 

2 — Digitus  elongate,  sides  parallel, 
greatly  exceeding  cuspis. 

49.  Microsculpture. 

0 — Present  on  most  of  body,  dull. 
1 — Absent,  body  shiny. 

DISCUSSION 

The  relationship  of  subtribes  within  the 
Pemphredoninae  remains  the  same  as  that 
presented  in  Bohart  and  Menke  (1976). 
The  sister  subtribe  to  the  Stigmina  is  pre- 
sumed to  be  the  Spilomenina,  based  on 
the  shared  presence  of  a  4-lobed  labrum 
(character  2.4  above).  The  Ammoplanina 


(synapotypies:  loss  of  transverse  pronotal 
carina  16.1  and  loss  of  omaulus  20.1)  is 
considered  immediate  outgroup  to  the 
Stigmina  +  Spilomenina  based  on  the  syn- 
apotypic  presence  of  an  enlarged  fore 
wing  stigma.  The  immediate  outgroup  to 
the  Stigmina  +  Spilomenina  +  Ammo- 
planina is  the  Pemphredonina  based  on 
the  synapotypic  loss  of  the  third  submar- 
ginal  cell  and  the  presence  of  specialized 
setae  on  the  apicomedial  clypeal  apex  in 
females.  The  Pemphredoninae  are  in  need 
of  critical  cladistic  analysis. 

In  the  Stigmina  the  analysis  of  charac- 
ters resulted  in  three  equally  parsimoni- 
ous solutions.  Electronic  analysis  of  the 
data  would  have  undoubtedly  resulted  in 
many  more.  The  cladogram  (Figure  51) 
was  selected  on  the  basis  of  the  best  fit 
with  plate  tectonics.  The  recognition  of 
new  genera  was  based  on  the  cladistic 
analysis  coupled  with  biogeography  and 
morphology.  Morphological  distinctions 
for  genera  are  consistent  with  those  for 
genera  in  Bohart  and  Menke  (1976).  A 
dendrogram  placing  Pamcrabro  in  a  posi- 


264 


Journal  of  Hymenoptera  Research 


Fig.  53.     Distribution  of  27  species  of  Stigmina  in  the  Caribbean. 


tion  basal  to  Carinostigmus  and  Stigmus 
was  presented  by  Bohart  and  Menke 
(1976).  The  presence  of  a  number  of  apo- 
typic  character  states  suggest  that  the  ar- 
rangement presented  here  with  Carinostig- 
mus in  the  basal  position,  is  correct.  These 
character  states  include  the  presence  of  an 
acetabular  carina,  a  clypeus  with  four 
teeth  on  the  apical  margin,  two  to  four 
spines  on  the  posterior  surface  of  the  hind 
tibia,  and  an  elongate  digitus  that  greatly 
exceeds  the  length  of  the  cuspis. 

A  composite  distribution  map  of  the 
Stigmina  is  presented  in  Fig.  52,  Carinos- 
tigmus is  plotted  against  all  other  genera 
in  the  Stigmina.  The  presence  of  genera  on 
two  southern  hemisphere  continents  and 
their  absence  on  other  continents  is  indic- 
ative of  a  group  radiating  at  the  time  of 
continental  separation.  Carinostigmus  is  re- 
stricted to  the  Old  World,  generally  south- 
ern hemisphere  (based  on  large  numbers 
of  undescribed  species  from  southern 
regions)  but  is  absent  from  Australia  (an 


introduced  species  is  present  in  New 
Guinea).  The  other  genera  occur  in  the 
western  hemisphere  {Araucastigmus,  Ay- 
khustigmus,  Incastigmus,  Llaqhastigmus,  Par- 
astigmus,  Stigmus),  across  the  Palaearctic 
Region  with  several  outliers  on  the  east 
asian  islands  and  the  orient  (Stigmus, 
Tzustigmus),  and  Australia  (Paracrabro). 
The  only  major  zone  of  contact  between 
Carinostigmus  and  the  other  genera  is  in 
the  Oriental  Region;  a  minor  contact  zone 
occurs  in  southern  Spain.  Elsewhere  Cari- 
nostigmus is  separated  from  the  other  gen- 
era by  the  Himalayas,  the  Gobi  and  Sahara 
Deserts,  and  the  Atlantic  Ocean. 

Outliers  in  Chile,  India,  Thailand,  Japan, 
Taiwan,  Vietnam,  Sumatra  and  Australia 
represent  distributions  of  relict  genera  and 
species.  The  only  anomaly  is  the  presence 
of  Carinostigmus  in  New  Guinea  which 
probably  represents  a  recent  introduction. 
The  Stigmina  are  restricted  in  distribution 
by  2  primary  barriers,  those  of  xeric  con- 
ditions and  water.  The  effect  of  xeric  con- 


Volume  4,  1995 


265 


ditions  is  evident  in  Fig.  52  with  the  ab- 
sence of  genera  from  major  desert  regions 
(some  exceptions  in  the  New  World).  The 
effect  of  water  is  evident  in  the  Caribbean 
fauna  (Fig.  53).  This  composite  distribu- 
tion map  of  27  species  shows  only  7  spe- 
cies on  the  Caribbean  Islands,  of  those  six 
occur  within  the  area  influenced  by  the 
Orinoco  River  current  where  3  are  endem- 
ic (including  Incastigmiis  thoraciciis).  A 
fourth  endemic  species  (an  undescribed 
Stigmus)  occurs  on  Hispaniola.  Many  of 
the  islands  of  the  Lesser  Antilles  are  with- 
in sight  of  neighbouring  islands.  The  fail- 
ure of  the  Stigmina  to  traverse  these  short 
distances  is  indicative  of  the  effectiveness 
of  water  as  a  barrier  to  dispersal  and  also 
to  the  lack  of  long  distance  dispersal 
mechanisms  within  this  group.  This  point 
must  be  born  in  mind  when  attempting  to 
interpret  the  present  distribution  patterns 
of  genera.  If  there  was  not,  at  some  time 
in  the  past,  an  overland  route  between 
points  presently  separated  by  water,  then 
it  is  unlikely  that  species  of  Stigmina 
would  disperse  between  those  points. 

In  South  America  Stigmus  has  a  C- 
shaped  distribution  restricted  to  the  high- 
lands surrounding  the  Amazon  basin  and 
restricted  by  the  lowlands  of  the  Rio  Tir- 
ana in  the  south.  Araucnstigmus  (Map  35) 
is  restricted  to  Chile,  for  the  most  part  to 
the  Pacific  side  of  the  continental  divide 
and  to  the  north  restricted  by  extreme  des- 
ert conditions.  There  are  3  species,  and 
thanks  to  Lubomir  Masner,  an  abundance 
of  specimens.  Parastigmus  (Map  34)  occurs 
on  the  east  side  of  the  divide  although  1 
species,  the  most  plesiotypic  in  the  genus 
occurs  in  Chile,  possibly  indicating  a  com- 
mon regional  origin.  In  Argentina  Paras- 
tigmus appears  to  follow  the  distribution 
of  grassy  steppe  vegetation  surrounding 
the  xeric  vegetation  of  Patagonia.  The  ge- 
nus is  represented  by  4  species  known 
from  16  specimens.  Aykhustigmus  (Map 
36),  represented  by  4  species  known  from 
37  specimens,  occurs  in  the  Brazilian 
Highlands  south  to  the  Rio  Pirana  and  the 


15.5°C  mean  July  isotherm.  The  only  ex- 
ception is  a  single  specimen  recently  col- 
lected from  the  Venezuelan  state  of  Zulia 
believed  to  be  an  introduction.  Llaqhastig- 
mus  (Map  37)  follows  a  distribution  simi- 
lar to  but  not  as  extensive  as  Stigmus.  Thir- 
teen species  are  confined  to  the  Andes 
while  a  single  species  occurs  across  the 
Brazilian  Highlands.  Llaqhastigmus  is  not 
known  from  the  Guyana  Shield  but  could 
well  occur  there.  Incastigmus  with  25  spe- 
cies is  not  only  the  largest  Neotropical  ge- 
nus but  has  the  broadest  distribution.  It  is 
limited  in  the  south  by  the  15.5°C  mean 
July  isotherm,  on  the  west  by  xeric  con- 
ditions and  to  the  north  by  the  10°C  mean 
January  isotherm  which  is  the  approxi- 
mate boundary  of  the  Nearctic  and  Neo- 
tropical Regions.  The  composite  map  (Fig. 
54)  demonstrates  the  limiting  influences  of 
winter  temperatures  and  rainfall  on  the 
distribution  of  Stigmina  in  the  Neotropics. 

About  200  million  years  ago  a  single 
landmass  stretched  from  pole  to  pole, 
Laurasia  to  the  north,  Gondwanaland  to 
the  south.  The  central  part  of  this  great 
landmass  was  dominated  by  xeric  condi- 
tions. Areas  of  rainforest  existed  in  the  ex- 
treme south  (southern  South  America, 
South  Africa,  Antarctica  and  Australia), 
seasonal  or  monsoon  rains  existed  in  the 
mid  latitudes  (southeast  Asia,  Arabian 
Peninsula)  (Lamb  1977).  Superimposed  on 
these  extremes  of  wet-dry  conditions  were 
areas  of  exposed  basement  layer  or  granite 
shield  characterized  by  localized  or 
edaphic  aridity  (Axelrod  1972).  Similar 
conditions  occur  today  in  the  Tipui  system 
of  the  Guiana  Shield.  The  predrift  uplift- 
ing of  the  continents  along  rift  lines  al- 
lowed water  to  flow  east  and  west  from 
the  South  American-African  divide  there- 
by slowly  exposing  the  basement  layer 
through  erosion.  The  exposure  of  the  base- 
ment layer  in  this  region  has  probably  had 
more  consequences  for  floral  and  faunal 
radiation  than  any  other  single  post  drift 
event. 

The  Stigmina  belong  to  the  apoid  lin- 


266 


Journal  of  Hymenoptera  Research 


,.i_--=c:s;^ 


..■^> 


10°C   January  mean 


^^F'^'-^^- 


>••      • 


>•• 


.^, 


%. 


NORTH,  CENTRAL,  AND  SOUTH  AMERICA 

500 


I 

15.5°C   July  mean 
^    10mra   rainfal 1 


MILES 


^^ 


<=^ 


Fig.  54.     Composite  distribution  of  Incastigmus,  Araucastigmus,  Parastigmus  and  Aykhustigmus  in  the  Neotrop- 
ical region. 


eage  (Apoidea)  of  Hymenoptera,  of  which 
bees  form  the  dominant  component.  There 
is  a  well  known  coevolutionary  relation- 
ship between  bees  and  angiosperms.  The 
earliest  flowers  were  probably  beetle  pol- 
linated (Michener  1979).  Bees  were  prob- 
ably an  already  diversified  west  gond- 
wanan  group  of  sandnesting  lineages 
(Michener  1979)  by  the  time  the  first  flow- 
ering plants  (Magnoliaceae)  spread  south- 
ward from  Laurasia  (Raven  and  Axelrod 


1974).  Michener  (1979)  for  bees,  and  Ax- 
elrod (1972)  and  Raven  and  Axelrod 
(1974)  for  angiosperms  hypothesize  a  xeric 
west  Gondwanaland  radiation  of  these 
groups.  The  influences  of  interior  xeric 
conditions,  thousands  of  island-like  patch- 
es of  arid  exposed  basement  layer,  along 
with  a  group  of  wasps  (bees)  preadapted 
for  pollination  contributed  to  an  explosive 
radiation  of  bees  and  angiosperms. 

Conversely  the  southern  rainforests  pre- 


Volume  4,  1995  267 

sented  a  habitat  unsuitable  for  sand  nest-  ney  and  likely  introduced  Gondwanaland 
ing  organisms  perhaps  because  of  high  flora  and  fauna  into  Southeast  Asia.  If  this 
water  tables,  high  humidity  and  the  likely  tectonic  event  occurred,  then  gondwana- 
persistence  of  fungi  in  brood  cells.  I  hy-  land  flora  and  fauna  (Stigmina)  could  be 
pothesize  that  twig-nesting  behaviour  de-  introduced  to  the  Laurasian  continent  at  a 
veloped  early  in  the  apoid  lineages  as  a  very  early  period.  This  would  allow  the 
response  to  increasingly  wet  conditions  to  lineages  of  Stigmina  to  be  in  a  position  to 
the  south.  It  is  also  likely  that  twig-nesting  colonize  the  Nearctic  Region  via  Europe 
developed  more  than  once  since  there  are  and  Beringia  at  65  million  B.P. 
several  apparently  unrelated  types  of  ae-  The  rifting  of  South  America  from  Af- 
rial  nest  architecture.  It  is  the  twig  nesting  rica  commenced  during  the  same  period 
groups  that  reach  the  greatest  diversity  in  (180  million  B.P.)  and  resulted  in  the  sep- 
the  rainforests  today.  Lineages  of  bees,  aration  of  southern  Africa  from  southern 
and  the  Stigmina  likely  radiated  from  one  South  America  by  a  narrow  body  of  wa- 
or  the  other  of  the  two  extreme  gondwan-  ter.  The  inability  of  the  Stigmina  to  cross 
aland  ecosystems.  Modern  distributions  a  water  barrier  is  instrumental  in  my  ex- 
show  presumed  relict  bees  inhabiting  planation  of  the  absence  of  genera  other 
south  temperate  xeric  areas  while  pre-  than  Carinostigmus  from  Africa.  Although 
sumed  relict  Stigmina  inhabit  south  wet  India  likely  once  held  more  genera  of  Stig- 
temperate  or  tropical  rainforest.  Based  on  mina  than  it  does  today,  that  fauna  was 
present-day  distribution  and  on  the  hy-  almost  certainly  lost  when  the  craton  trav- 
pothesized  relationships  between  genera,  elled  over  a  hot  spot  in  the  crust  on  its 
the  Stigmina  probably  existed  prior  to  northward  journey.  Severe  volcanism  on 
continental  breakup,  over  200  million  the  Indian  plate  likely  caused  extinction  of 
years  ago.  Some  support  is  lent  to  this  biota  in  virtually  all  its  ecosystems.  What 
statement  by  the  existence  of  specialized  fauna  the  plate  brought  to  Asia  is  proba- 
pemphredonine  fossils  in  the  Upper  Cre-  bly  the  result  of  a  brief  encounter  with 
taceous,  about  135  million  B.P.  (Evans  north  Africa  before  colliding  with  Asia 
1969,  1973).  Stigmina  at  the  time  of  conti-  about  45  million  years  B.P.  By  the  late  Cre- 
nental  breakup  would  likely  have  oc-  taceous  Period  (135  million  years  B.P.) 
curred  in  the  southern  rainforests  (south-  Stigmus  and  Tzustigmiis  probably  colo- 
ern  South  America,  South  Africa,  Antarc-  nized  Laurasia.  The  Nearctic  fauna  con- 
tica,  Australia)  and  in  the  southern  savan-  sists  of  eastern  and  western  species 
na  surrounding  the  midcontinental  xeric  groups  likely  the  result  of  colonization  by 
area.  Stigmus   via   both    Europe   and    Beringia 

Of  pivotal  importance  in  this  analysis  is  about  65  million  years  B.P. 
a  report  on  the  tectonics  of  the  Indonesian  Based  on  relationships  presented  in  the 
region  (Hamilton  1979).  Hamilton  pre-  cladogram  and  present  distributions  it  is 
sents  evidence  of  Jurassic  (180  million  likely  that  the  Stigmina  originated  in  the 
B.P.)  rifting  of  eastern  Gondwanaland.  southern  hemisphere  continents,  especial- 
Present-day  land  masses  of  Thailand,  Bur-  ly  Antarctica  (Fig.  52)  over  200  million 
ma,  Malaysian  Peninsula,  Sumatra  and  years  B.P.  Australia  was  colonized  once 
Borneo  rifted  from  the  northern  Australia-  (Paracrabro).  South  America  was  colonized 
New  Guinea-Antarctica  continental  mar-  at  a  very  early  stage  (180  million  B.P.)  on 
gin  and  moved  north  to  collide  with  the  at  least  3  separate  occasions,  and  more  re- 
Eurasian  plate.  Furthermore  Audley-  cently  by  Stigmus  from  the  Nearctic  Re- 
Charles  (1987)  found  an  uninterrupted  gion  about  5  million  B.P.  North  America 
pollen  record  indicating  the  craton  was  was  colonized  at  least  twice,  3  times  if  one 
not  submerged  during  its  northward  jour-  includes  the  incursion  of  Neotropical  fau- 


268 


Journal  of  Hymenoptera  Research 


na  (Incastigmus)  into  Central  America 
about  5  million  B.P.  The  Palaearctic  was 
colonized  at  least  twice;  finally  the  Ori- 
ental Region  was  colonized  on  at  least  2 
occasions. 

The  great  diversity  of  the  Neotropical 
fauna  is  due  to  3  major  factors.  The  wet 
adapted  fauna  that  colonized  it,  periodic 
aridity  that  likely  restricted  that  fauna  to 
islands  of  forest,  and  mountains  that  al- 
lowed extremes  in  climatic  conditions  to 
exist  in  close  proximity.  The  xeric  areas  of 
southern  Argentina  afford  the  best  oppor- 
tunity to  collect  relict  stigmine  species  in 
the  New  World. 

By  contrast  the  relatively  depauperate 
nature  of  the  Afrotropical  fauna  may  be 
the  result  of  a  xerically  adapted  original 
fauna  succumbing  to  the  spread  of  mesic 
conditions  over  the  entire  continent,  in- 
cluding the  Sahara  (Axelrod  and  Raven 
1978).  There  was  no,  or  very  little,  of  the 
faunal  island  effect  that  occurred  in  South 
America.  An  increasingly  wet  Africa  prob- 
ably afforded  little  opportunity  for  the 
xerically  adapted  fauna  to  survive.  Africa 
is  again  in  a  dry  phase  with  rainforest  re- 
stricted to  very  limited  areas.  The  more 
equitable  continental  outliers  such  as  the 
mesic  islands  of  the  Comoros  or  Mada- 
gascar probably  still  harbour  remnants  of 
relict  faunas. 

The  interactions  in  the  Oriental  Region 
are  complex.  Most  of  the  relict  taxa  are 
found  there.  Species  of  relict  groups  have 
highly  localized  distributions  thus  ac- 
counting for  their  relative  absence  in  most 
collections.  There  is  a  possibility  that  their 
distributions  are  tied  to  forest  over  lime- 
stone terrain.  Whitmore  (1984)  reported 
that  limestone  terrain  is  not  common  in 
Malaysia,  but  the  flora  of  a  mainland  for- 
est over  limestone  formation  had  23%  of 
its  species  endemic  to  the  country  with 
11%  endemic  to  limestone.  Islands  like 
Borneo  or  Sulawesi  are  melange  forma- 
tions (Hamilton  1979)  resulting  from  near- 
by subduction  zones  indicating  that  lime- 
stone has  likely  been  present  throughout 


the  islands  history.  Although  no  habitat 
information  is  available  for  the  relict  fau- 
na, it  is  likely,  because  of  the  high  degree 
of  endemic  flora  that  limestone  forest  may 
offer  fertile  grounds  for  collecting  speci- 
mens of  these  ancient  apoid  wasp  lin- 
eages. 

Little  mention  has  been  made  of  the  or- 
igins of  Cnrinostigmus.  Two  scenarios  exist 
based  on  the  cladogram,  present  distri- 
bution, and  the  number  of  undescribed 
species.  In  the  first  scenario  I  hypothesize 
an  African  origin  with  the  genus  in  place 
at  the  time  of  rifting  from  South  America 
or  shortly  thereafter.  In  the  second  scenar- 
io I  hypothesize  an  Oriental  origin  with 
more  recent  colonization  of  Africa.  The 
Oriental  Region  is  a  mix  of  land  masses  of 
Laurasian  and  Gondwanaland  origin,  and 
most  species  of  Carinostigmus  occur  there 
(mostly  undescribed).  If  a  cladistic  analy- 
sis of  species  within  Carinostigmus  were  to 
demonstrate  a  Laurasian  origin  then  the 
age  of  lineages  in  the  Stigmina  could  be 
moved  back  to  a  Pangean  origin  about  230 
million  B.P.  It  would  also  imply  a  north- 
south  Pangean  split  of  lineages  within  the 
Stigmina  with  Carinostigmus  of  northern 
(Laurasian)  origin  and  all  other  genera  of 
southern  origin  (Gondwanaland).  Such  a 
scenario  is  compatible  with  the  cladogram 
presented  here  and  with  ecosystem  recon- 
structions of  that  period. 

ACKNOWLEDGEMENTS 

I  wish  to  thank  the  reviewers  of  the  manuscript, 
R.M.  Bohart,  A.S.  Menke,  and  W.J.  Pulawski,  for  their 
time  and  the  numerous  suggestions  for  improve- 
ments to  the  manuscript. 

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Volume  4,  1995 


271 


Table  1.    Data  matrix  for  the  Stigmina 


Carino 

Ti-ustig 

Arjuca 

Parac 

Parastig 

Aykhus 

Stignitis 

Llaqh 

Incast 

1.  Palpal  formula 

2.  Labrum  shape 

3.  Mandibular  socket 

0 

1 

1 

0 
4 
1 

0 
4 
1 

0 
4 
1 

0 
4 

1 

0 

4 

1 

0 
4 
1 

0 
3 
1 

0 
4 
1 

4.  Apical  manibular  teeth 

03 
29 

06 

29 

06 
2? 

06 

49 

06 
09 

06 

29 

06 
0,29 

26 
29 

26 

29 

5.  Apicoventral  tooth 

6.  Clypeal  bevel 

7.  Clypeus  size 

8.  Clypeal  teeth 

9.  Interantennal  tubercle 

0 
0 
0 
2 
1 

0 
0 
0 
3 
1 

0 
0 
0 
4 
0 

na 

0 

1 

5 

1 

na 

0 
1 

4,5 
0 

0 
0 
0 
4 

0 

0 
1 

0 

2,4 

0 

1 
0 

0 
4 
0 

0 
0 

0 
4 
0 

10.  Frontal  carina 

2 

0 

2 

2 

0,2 

0 

0 

0 

0 

11.  Vertex  micropore  field 

12.  Eye  margins 

13.  Occipital  carina 

14.  Occi.-hypo.  carina 

15.  Pronotal  collar 

1 

1 

0,1 

2 

1 

1 

1 

0,1 

2 

0 

1 
0 
0 

2 
0 

1 
1 
1 

2 
0 

1 
0 
0 
2 
0 

2 

1 

0,1 

2 

0 

2 

1 

0,1 

1,2 

0 

2 

1 

0,1 
2 
0 

2 

1 

0,1 

2 

0 

16.  Transverse  pronotal  carina 

0 

0 

0 

0 

0 

0 

0 

0 

0 

17.  Median  scutal  groove 

18.  Notaular  grooves 

19.  Acetabular  carina 

0,1 

0 

0 

0 
0 
0 

0 
0 
0 

0 
0 
1 

0 
0 
0 

0,1 
0 

1 

0 
0 
1 

0 
0 

1 

1 

1 
1 

20.  Omaulus 

0 

0 

0 

0 

0 

0 

0 

0 

0 

21.  Epistemal  sulcus 

22.  Hypersternaulus 

23.  Orientation  of  hypersternaulus 

24.  Scrobal  sulcus 

1 

0 
0 
0,1 

1 
0 

0 
0,1 

1 

0 
0 

1 

1 
0 
0 
0 

1 
0 
0 
0,1 

1 

0 
0 
0 

1 
0 
0 
0 

1 

0 
0 
0 

1 

0 
0 
0 

25.   6  midbasitarsus 

0 

0 

0 

2 

1 

1 

2 

0 

0 

26.  Hindtibial  spines 

27.  Propodeal  enclosure 

28.  Stigma  size 

2 
0 
1 

1 

0 

1 

1 

0 

1 

1 
0 
1 

1 
0 

1 

1 

0 

1 

1 

0 

1 

1 

0 

1 

1 

0 

1 

29.  Stigma  shape 

30.  Stigma  pore  field 

31.  Marginal  cell  size 

32.  Marginal  cell  veins 

0 

1 

0 
0 

0 

1 

0 
0 

0 

1 

0 
0 

0 
1 
0 
0 

0 

1 

0 
0 

0 

1 

0 
0 

0 

1 

0 
0 

0 

1 

0 
0 

0 

1 

0 
0 

33.  Submarginal  cells 

34.  Submarginal  position 

0 
0 

0 
0 

0 
0 

0 
0 

0 
0 

0 
0 

0 
0 

0 
0 

0 
0 

35.  Submarginal  veins 

0 

0 

0 

0 

0 

0 

0 

0 

0 

36.  Discoidal  cell  number 

1 

1 

1 

1 

1 

1 

1 

1 

1 

37.  Media  divergence 

2 

1 

0 

0 

0 

0,1 

0 

0 

0 

38.  Submedian  cell 

1 

0 

0 

0 

0 

0 

0 

0 

0 

39.  Hind  wing  cells 

0 

0 

0 

0 

0 

0 

0 

0 

0 

40.  Petiole  length 

41.  Petiolar  sculpture 

2 
2 

2 
0,2 

2 

0 

2 
0 

2 
0 

2 
0 

2 
0 

2 
0 

2 
0 

42.  Sternum  I  carinae 

0 

0 

0 

0 

0 

0 

0 

0 

0 

43.  SI  basal  groove 

0 

0 

0 

0 

0 

0 

0 

0 

0 

44.  Tl  lateral  carina 

1 

1 

1 

1 

1 

1 

1 

1 

1 

45.  S2  microsetae 

1 

1 

1 

1 

1 

1 

1 

1 

1 

46.  Pygidial  Plate 

47.  Male  Sternum  8 

1 

3 

1 
3 

1 

3 

0 
3 

1 

3 

1 

3 

1 

3 

1 

3 

1 

3 

48.  Genitalia 

0 

2 

0 

2 

2 

2 

2 

2 

2 

49.  Microsculpture 

1 

0,1 

1 

0 

0 

0 

0 

0 

0 

0  =  plesiotypic  character  state,  1-6  =  apotypic  character  states,  na  =  not  applicable,  Carino  =  Carinostigmus, 
Tzustig  =  Tziistigtmis,  Arauca  =  Araucastigiiiiis,  Parac  =  Paracrabro,  Parastig  =  Parastigmus,  Aykhus  =  Ay- 
kliustigmiis,  Llaqh  =  Uaqluistigniiis,  Incast  =  Incastigmns. 


272 


Journal  of  Hymenoptera  Research 


Table  2.    Data  matrix  for  the  Pemphredonina 


Diodontus 

Pemphredon 

Passalcecus 

Polemistus 

1.  Palpal  formula 

0 

0 

0 

0 

2.  Labrum  shape 

3 

1 

2 

1 

3.  Mandibular  socket 

1 

1 

1 

1 

4.  Apical  manibular  teeth 

0,1 

2,  3,  4,  5 

0,2 

0,1 

5.  Apicoventral  tooth 

na 

0,1 

0 

na 

6.  Clypeal  bevel 

0 

0 

0 

0 

7.  Clypeus  size 

0 

0 

0 

0 

8.  Clypeal  teeth 

3 

1,  2,  3 

1,2,3 

1 

9.  Interantennal  tubercle 

0 

0 

0 

1 

10.  Frontal  carina 

2 

2 

0 

2 

11.  Vertex  micropore  field 

0 

0 

0 

0 

12.  Eye  margins 

0 

0 

0 

1 

13.  Occipital  carina 

0 

0 

0 

1 

14.  Occi.-hypo.  carina 

1 

1 

2 

2 

15.  Pronotal  collar 

0 

0 

0 

0 

16.  Transverse  pronotal  carina 

0 

0 

0 

0 

17.  Median  scutal  groove 

0 

0 

0 

0 

18.  Notaular  grooves 

0 

0 

0 

1 

19.  Acetabular  carina 

0 

0 

0 

0 

20.  Omaulus 

0 

0 

1 

0 

21.  Episternal  sulcus 

0 

0 

0 

0 

22.  Hypersternaulus 

0 

0 

0 

0 

23.  Orientation  of  hypersternaulus 

0 

0 

1 

1 

24.  Scrobal  sulcus 

0 

0 

0 

0 

25.   6  midbasitarsus 

1 

1 

1 

1 

26.  Hindtibial  spines 

0 

0 

2 

2 

27.  Propodeal  enclosure 

0 

0 

1 

0 

28.  Stigma  size 

0 

0 

0 

0 

29.  Stigma  shape 

0 

0 

0 

0 

30.  Stigma  pore  field 

0 

0 

0 

0 

31.  Marginal  cell  size 

0 

0 

0 

0 

32.  Marginal  cell  veins 

0 

0 

0 

0 

33.  Submarginal  cells 

0 

0 

0 

0 

34.  Submarginal  position 

0 

0 

0 

0 

35.  Submarginal  veins 

0 

0 

0 

0 

36.  Discoidal  cell  nvimber 

0 

0 

0 

0 

37.  Media  divergence 

0 

0 

0 

0 

38.  Submedian  cell 

0 

0 

0 

0 

39.  Hind  wing  cells 

0 

0 

0 

0 

40.  Petiole  length 

1 

2 

1 

2 

41.  Petiolar  sculpture 

0 

1 

0 

0 

42.  Sternum  I  carinae 

1 

1 

1 

1 

43.  SI  basal  groove 

0 

0 

0 

0 

44.  Tl  lateral  carina 

0 

0,1 

0 

0 

45.  S2  microsetae 

0 

0 

0 

0 

46.  Pygidial  Plate 

0 

1 

2 

2 

47.  Male  Sternum  8 

1 

1 

3 

3 

48.  Genitalia 

0 

0 

0 

0 

49.  Microsculpture 

0 

0,1 

0 

0 

0  =  plesiotypic  character  state,  1-6  = 

=  apotypic  character  states,  na  =  not 

applicable. 

Volume  4,  1995 


273 


Table  3.    Data  matrix  for  the  Spilomenina 


Arpnclop\u\iiy- 

Spilonicna 

Microstigmu^ 

Xxi^-nia 

1.  Palpal  formula 

2 

1 

2 

2 

2.  Labrum  shape 

3,4,5 

4 

0? 

0? 

3.  Mandibular  socket 

1 

1 

1 

1 

4.  Apical  manibular  teeth 

0 

0 

0 

0 

5.  Apicoventral  tooth 

na 

na 

na 

na 

6.  Clypeal  bevel 

0 

0 

0 

0 

7.  Clypeus  size 

1 

1 

1 

1 

8.  Clypeal  teeth 

2 

2 

2 

1 

9.  Interantennal  tubercle 

0 

0 

0 

0 

10.  Frontal  carina 

2,3 

0,1 

2 

2 

11.  Vertex  micropore  field 

0 

0 

0 

0 

12.  Eye  margins 

1 

0 

0 

0 

13.  Occipital  carina 

2 

2,3 

3 

3 

14.  Occi.-hypo.  carina 

2 

na 

na 

na 

15.  Pronotal  collar 

0 

0 

0 

0 

16.  Transverse  pronotal  carina 

0 

0 

0 

0 

17.  Median  scutal  groove 

0 

0 

0 

0 

18.  Notaular  grooves 

0 

0 

0 

0 

19.  Acetabular  carina 

0 

0 

0 

0 

20.  Omaulus 

0 

0 

0 

0 

21.  Episternal  sulcus 

1 

1 

1 

1 

22.  Hypersternaulus 

0 

0,1 

0 

1 

23.  Orientation  of  hypersternaulus 

1 

1 

1 

na 

24.  Scrobal  sulcus 

0,1 

1 

1 

1 

25.   6  midbasitarsus                   -- 

1 

26.  Hindtibial  spines 

2 

2 

2 

2 

27.  Propodeal  enclosure 

1 

0 

1 

1 

28.  Stigma  size 

1 

1 

1 

1 

29.  Stigma  shape 

0 

0 

0 

0 

30.  Stigma  pore  field 

0 

0 

0 

0 

31.  Marginal  cell  size 

0 

0 

0 

0 

32.  Marginal  cell  veins 

0 

0 

0 

1 

33.  Submarginal  cells 

0,1 

0 

1 

2 

34.  Submarginal  position 

0 

0 

0 

0 

35.  Submarginal  veins 

0 

0 

0 

1 

36.  Discoidal  cell  number 

1 

1 

1 

2 

37.  Media  divergence 

0 

0 

0 

na 

38.  Submedian  cell 

0 

0 

0 

na 

39.  Hind  wing  cells 

0 

0 

0 

1 

40.  Petiole  length 

0 

0 

0 

0 

41.  Petiolar  sculpture 

na 

na 

na 

na 

42.  Sternum  I  carinae 

2 

2 

2 

2 

43.  SI  basal  groove 

0 

0 

0 

0 

44.  Tl  lateral  carina 

0 

0 

0 

0 

45.  S2  microsetae 

0 

0 

0 

0 

46.  Pygidial  Plate 

2 

2 

2 

2 

47.  Male  Sternum  8 

2 

48.  Genitalia 

1 

49.  Microsculpture 

0 

0 

0 

0 

0  =  plesiotypic  character  state,  1-6  =  apotypic  character  states,  na   =  not  applicable,  blank  cells  indicate 
specimens  were  unavailable  and  character  state  could  not  be  determined  from  literature. 


274 


Journal  of  Hymenoptera  Research 


Table  4.    Data  matrix  for  the  Ammoplanina 


Pulv 

Ammops 

Ammous 

Paramo 

Ammoplus 

Timber 

1.  Palpal  formula 

1 

1 

1 

1 

1 

1 

2.  Labrum  shape 

0? 

0? 

3 

3 

3 

3 

3.  Mandibular  socket 

1 

1 

1 

1 

1 

1 

4.  Apical  manibular  teeth 

0 

0 

0 

0 

0 

0 

5.  Apicoventral  tooth 

na 

na 

na 

na 

na 

na 

6.  Clypeal  bevel 

0 

0 

0 

0 

0 

0 

7.  Clypeus  size 

0 

0 

1 

1 

1 

1 

8.  Clypeal  teeth 

2 

2 

2 

2 

2 

2 

9.  Interantennal  tubercle 

0 

0 

0 

0 

0 

0 

10.  Frontal  carina 

0,1 

1 

0 

1 

0 

1 

11.  Vertex  micropore  field 

0 

0 

0 

0 

0 

0 

12.  Eye  margins 

0 

0 

0 

0 

0 

0 

13.  Occipital  carina 

2 

2 

2 

2 

0 

3 

14.  Occi.-hypo.  carina 

2 

1 

1 

1 

1 

na 

15.  Pronotal  collar 

0 

0 

0 

0 

0 

0 

16.  Transverse  pronotal  carina 

0 

0,1 

1 

1 

1 

1 

17.  Median  scutal  groove 

0 

0 

0 

0 

0 

0 

18.  Notaular  grooves 

0 

0 

0 

0 

0 

0 

19.  Acetabular  carina 

0 

0 

0 

0 

0 

0 

20.  Omaulus 

1 

1 

1 

1 

1 

1 

21.  Episternal  sulcus 

0 

0 

0 

0 

0 

0 

22.  Hypersternaulus 

1 

1 

1 

1 

1 

1 

23.  Orientation  of  hypersternaulus 

na 

na 

na 

na 

na 

na 

24.  Scrobal  sulcus 

1 

1 

1 

1 

1 

1 

25.   3  midbasitarsus 

1 

1 

1 

0 

0 

26.  Hindtibial  spines 

1 

2 

2 

2 

2 

2 

27.  Propodeal  enclosure 

1 

1 

1 

1 

1 

1 

28.  Stigma  size 

1 

1 

1 

1 

1 

1 

29.  Stigma  shape 

0 

0 

1 

1 

1 

1 

30.  Stigma  pore  field 

0 

0 

0 

0 

0 

0 

31.  Marginal  cell  size 

1 

1 

1 

1 

1 

1 

32.  Marginal  cell  veins 

0 

0 

0 

1 

1 

1 

33.  Submarginal  cells 

1 

1 

1 

1 

1 

0,1,2 

34.  Submarginal  position 

0 

1 

0 

0 

0 

0 

35.  Submarginal  veins 

0 

0 

0 

0 

0 

0,1 

36.  Discoidal  cell  number 

1 

1 

1 

1 

1 

1 

37.  Media  divergence 

0 

0 

2 

2 

2 

na 

38.  Submedian  cell 

0 

0 

0 

0 

0 

na 

39.  Hind  wing  cells 

0 

0 

0 

0 

0 

1 

40.  Petiole  length 

0 

0 

0 

0 

0 

0 

41.  Petiolar  sculpture 

na 

na 

na 

na 

na 

na 

42.  Sternum  I  carinae 

0 

0 

0 

0 

0 

0 

43.  SI  basal  groove 

1 

1 

1 

1 

1 

1 

44.  Tl  lateral  carina 

1 

1 

1 

1 

1 

1 

45.  S2  microsetae 

0 

0 

0 

0 

0 

0 

46.  Pygidial  Plate 

0 

0 

0 

0 

2 

2 

47.  Male  Sternum  8 

4 

4 

2 

2 

2 

2 

48.  Genitalia 

0 

0 

1 

1 

1 

49.  Microsculpture 

0 

0 

0 

0 

0 

1 

0  =  plesiotypic  character  state,  1-6  =  apotypic  character  states,  na  =  not  applicable,  blank  cells  indicate 
specimens  were  unavailable  and  character  state  could  not  be  determined  from  literature.  Pulv  =  Pulverro, 
Ammops  =  Ammoplmiops,  Ammous  =  Ammoplanus,  Paramo  =  Parammoplanus,  Ammoplus  =  Ammoplanellus, 
Timber  =  Tunherlakena. 


Volume  4,  1995 


275 


276 


Journal  of  Hymenoptera  Research 


Map  5. 
Tzustigmus  wuming 

I t 


<l!^^=^=;>^ 


Map  7. 

Parastigmus 

huecuvus 


Map  8, 
Parastigmus 
n  i  na 


Volume  4,  1995 


277 


^2^^=^=D^^ 


Map   9. 
Parast igmus 
patagonicus 


^^2^"=^=^^ 


Map  11. 
Araucas  t  igmus 
masneri 


^^F^=^^ 


Map  10, 
Araucast igmus 
glabratus 


^2^=^=I>^ 


Map  12. 
Araucas  t  i  gmus 
yani  J lus 


278 


Journal  of  Hymenoptera  Research 


Map  13. 

Paracrabro  froggatti 


il?2^'=^^^ 


Map  15. 

Aykhustigmus 

patanawi 


Map  \k. 
Aykhustigmus 
f ri  tzi 


0^"^=^^ 

%«^ 

a 
\ 

ly 

'           P 

Map    16. 

Aykhustigmus 

phasti 

%K   ^ 

•» 

Volume  4,  1995 


279 


/?7=^^^^ 


Map  17. 
Aykhustigmus 
wa  rawa 


^^2^""^=^^^ 


Map. 19. 
Incastigmus 
thoraci  cus 


^F=^^^ 


Map  18. 

I  ncast igmus 

i  nt  i 


^^'^^^^       „. 


Map  20. 
Llaqhastigmus 
ambi guus 


280 


Journal  of  Hymenoptera  Research 


^^^2==^=^^ 


Map  23. 
Llaqhastigmus 
col omb! anus 


^^^'^^^^ 


Map  22. 

Llaqhastigmus 
chut iyana 


ii'^^^^^^ 


Map  Ik. 
Llaqhastigmus 
ecuatori  al is 


Volume  4,  1995 


281 


Z!^^'^'^^^ 


Map  25. 
Ll aqhas t igmus 
jatunki  rus 


^PT^""^^^^ 


Map  27. 
Ll aqhas t igmus 
mantant i 


iJf^'^^i^J^ 


Map  26. 
Llaqhas ti  gmus 
1 1 utani  s 


^^^^^^^^ 


Map  28. 
Llaqhas tigmus 
mu thus 


282 


Journal  of  Hymenoptera  Research 


/:^^'=^^^ 


Map  29. 

LI  aqhast igmus 

nigricollaris 


^  J 


/^^"^^^^ 


Map  31  . 

L  laqhas  t  i  gmus 

sapani  s 


/^2='^=^^ 


Map  30. 

LI aqhast igmus 

santanderana 


^'T^''^^^ 


Map  32. 

LI  aqhast igmus 

shachus 


Volume  4,  1995 


283 


/?2'^^=D^^ 


Map  33. 
Llaqhast igmus 
sharkeyi 


^=^=^^ 


Map  Zk. 
Parast  igmus 


^^^'^^^ 


Map  36. 
Aykhust igmus 


284 


Journal  of  Hymenoptera  Research 


iJ^=?^=^>5<?" 


a 


Map  37. 
Llaqhastigmus 


J.  HYM.  RES. 
Vol.  4,  1995,  pp.  285-293 

The  Family-Group  Names  of  the  Ichneumoninae 
(Hymenoptera:  Ichneumionidae) 

David  B.  Wahl  and  W.R.M.  MASONt 

(DBW)  American  Entomological  Institute,  3005  SW  56th  Ave.,  Gainesville,  FL  32608,  U.S.A.; 
(WRMM)  Biological  Resources  Division,  CLBRR,  Agriculture  Canada,  Ottawa, 

Ontario  KIA  0C6,Canada 


Abstract. — The  family-group  names  in  the  subfamily  Ichneumoninae  are  listed.  Valid  names  are 
in  accordance  with  the  International  Code  of  Zoological  Nomenclature  and  differences  with  the  names 
used  by  Townes  are  discussed.  The  historical  background  to  Townes'  rejection  of  certain  Opinions 
of  the  International  Commission  for  Zoological  Nomenclature  is  examined;  we  conclude  Townes 


was  m  error. 


The  subfamily  Ichneumoninae  consti- 
tutes an  exceedingly  large  and  diverse 
group  of  parasitoid  wasps,  and  is  one  of 
the  most  taxonomically  and  nomenclatur- 
ally  confusing  groups  of  ichneumonids. 
The  genera  w^ere  never  revised  by 
Townes  as  part  of  his  monographs  of  ich- 
neumonid  subfamilies.  His  taxonomic 
ideas  on  the  subfamily  were  recorded  in 
the  regional  catalogs  put  out  by  him  and 
his  collaborators  (Townes  et  al  1961; 
Townes  et  al  1965;  Townes  &  Townes 
1966;  Townes  &  Townes  1973).  These 
groupings  and  generic  concepts  are  often 
at  odds  with  those  of  other  authors,  most 
notably  Heinrich,  who  published  mono- 
graphs on  the  ichneumonines  of  the  Ne- 
arctic,  sub-Saharan  Africa,  and  Southeast 
Asia.  The  situation  is  especially  confusing 
because  Townes  based  family-group 
names  on  the  oldest  included  genus  and 
refused  to  recognize  the  validity  of  cer- 
tain Opinions  issued  by  the  International 
Commission  for  Zoological  Nomencla- 
ture (hereafter  referred  to  as  "the  Com- 
mission") (Townes  1969).  While  Heinrich 
endeavored  to  use  nomenclature  in  ac- 
cordance with  the  International  Code  of 
Zoological  Nomenclature  (International 
Commission    for    Zoological    Nomencla- 


ture 1985;  hereafter  referred  to  as  "the 
Code"),  he  sometimes  used  junior  syn- 
onyms of  his  ow^n  authorship  in  prefer- 
ence to  older  names  of  other  authors  or 
failed  to  cite  proper  dates  and  authorship 
for  family-group  names  (cf.  Protichneu- 
monini  Heinrich,  1934  vs.  Heresiarchini 
Ashmead,  1900,  and  Ichneumonini  Ash- 
mead,  1895  instead  of  Ichneumonini  La- 
treille,  1802  (Heinrich  1977:  10,  57). 

TOWNES  AND  OPINIION  159 

Townes  (1957,  1969)  wrote  at  length 
about  his  reasons  for  considering  certain 
Opinions  of  the  Commission  invalid.  The 
following  summarizes  Townes'  general 
views:  "There  is,  however,  a  spurious 
"Opinion  of  the  International  Commis- 
sion" (no.  178)  that  was  published  under 
the  personal  responsibility  of  Francis 
Hemming,  without  the  knowledge  or 
consent  of  the  International  Commission, 
and  without  an  official  vote  of  the  Com- 
mission on  the  matter.  Citations  of  this 
"Opinion"  have  been  misleading. 
(Townes  &  Townes  1981).  Most  hymen- 
opterists,  outside  of  ichneumonid  spe- 
cialists, do  not  accept  Townes'  arguments 
(Day  1981  is  an  exception).  Townes'  ex- 
tensive ichneumonid  work  has  lead  many 


286 


Journal  of  Hymenoptera  Research 


students  of  that  group  to  follow  his  no- 
menclature. We  do  not,  however,  accept 
Townes'  position  or  his  supporting  ver- 
sion of  history;  a  detailed  account  of  our 
view  is  given  in  Appendix  I.  Opinion  159, 
which  deals  with  application  of  the  ich- 
neumonid  names  Ichneumon,  Pimpla,  and 
Ephialtes,  is  considered  by  us  to  be  a  valid 
publication  of  the  Commission. 

TRIBAL  CONCEPTS 

Correct  nomenclature  for  family-group 
taxa  in  Ichneumonidae,  excluding  Ichneu- 
moninae,  has  been  published  by  Fitton  & 
Gauld  (1976;  1978).  We  here  present  the 
results  of  our  investigations  of  ichneu- 
monine  family-group  names,  using  the 
Code  to  determine  the  validity  of  the  46 
names. 

As  mentioned  above,  Townes  and  Hein- 
rich  disagreed  on  the  matter  of  tribal 
boundaries  and  composition  (Gauld  1984: 
184-185).  In  the  absence  of  a  worldwide 
monographic  treatment  of  the  subfamily, 
we  have  for  the  most  part  adopted  the 
tribal  classification  used  by  Gauld  (1984). 
Townes  had  intended  to  treat  the  subfam- 
ily as  a  fifth  volume  of  his  series  of  generic 
monographs  but  funding  and  health  prob- 
lems interfered.  Townes'  thoughts  on  ich- 
neumonine  classification  are  reflected  in 
his  regional  catalogs  and  in  the  arrange- 
ment of  the  American  Entomological  In- 
stitute collection. 

Before  proceeding  with  a  listing  of  the 
family-group  names,  we  give  a  brief  con- 
sideration of  problems  in  certain  tribes 
and  subtribes. 

1.  Alomyini-Phaeogenini-Heteris- 
chnini.  Confusion  reigns  concerning  the 
composition  of  the  tribe.  European  au- 
thors (Perkins  1959;  Diller  1981;  Rasnit- 
syn  &  Siitan  1981)  have  traditionally 
placed  Alomya  in  a  tribe  or  subfamily  of 
its  own.  Townes  consistently  put  it  with 
the  group  of  genera,  centered  about 
Phaeogenes,  that  previously  cited  workers 
referred  to  as  Phaeogenini.  Townes  be- 
lieved Alomya  to  be  closely  related  to  Cen- 


ieterus  and  Colpognathus,  genera  placed 
by  others  in  Phaeogenini  (Townes,  pers. 
comm.  to  DBW).  Moreover,  Townes 
placed  the  genera  Heterischnus  and  Liisius 
in  a  separate  tribe,  Heterischnini.  Perkins, 
Diller,  and  Rasnitsyn  &  Siitan  have  all 
maintained  Heterischnus  in  the  Phaeogen- 
ini. Wahl  is  examining  the  relationships 
of  these  genera  as  part  of  a  forthcoming 
revision;  preliminary  results  indicate  that 
all  the  aforementioned  genera  constitute 
a  monophyletic  group,  and  they  are  here 
treated  as  one  tribe.  With  the  inclusion  of 
Alomya,  the  correct  tribal  name  is  Alo- 
myini;  Townes  &  Townes  (1951:  276)  hav- 
ing made  a  first-revisor  decision  between 
the  Forster  names  Alomyoidae  and 
Phaeogenoidae  (published  simultaneous- 
ly in  1869).  The  subtribal  divisions  of 
Diller  (1981;  1994)  are  not  used  here, 
pending  further  study. 

2.  Platylabini-Eurylabini-ZimmeriinL 
Townes  (Townes  et  al  1961;  Townes  et  al 
1965;  Townes  &  Townes  1973)  consistent- 
ly placed  the  genera  Eurylabus  and  Cotih- 
eresiarches  (=  Zimmeria)  in  the  Platylabini. 
Other  authors  (Heinrich  1967,  1974;  Per- 
kins 1959;  Rasnitsyn  &  Siitan  1981)  put 
them  in  the  Eurylabini  and  Zimmeriini, 
respectively.  When  Heinrich  (1934)  first 
described  the  Eurylabini,  he  recognized 
the  Eurylabus  group  and  the  Goedartia 
group.  Townes  (Townes  et  al  1961)  broke 
up  the  tribe,  placing  Eurylabus  in  the  Pla- 
tylabini and  the  Goedartia  group  in  its 
own  tribe,  Goedartiini.  Neither  Eurylabus 
nor  Cotiheresiarches  exhibits  the  character- 
istic small  convex  clypeus  of  Platylabus 
and  its  relatives,  and  only  Cotihersiarches 
has  a  flattened  petiole.  Until  detailed 
studies  are  made  on  platylabine  generic 
relationships,  we  believe  it  best  for  now 
to  recognize  Eurylabini  and  Zimmeriini 
as  separate  tribes. 

3.  Goedartiini-Compsophorini.  When 
originally  proposed  by  Townes  et  al 
(1961),  Goedartiini  included  Goedartia, 
Maraces,  Charitojoppa,  and  Habrojoppa. 
Compsophorini  was  described  by  Hein- 


Volume  4,  1995 


287 


rich  (1967)  and  included  Compsophorus, 
Oxyjoppa,  Eccoptosagelliis,  Tosquinetia,  Ep- 
ijoppa,  Charitojoppa,  Hahrojoppa,  and  Pyr- 
amidamhlys;  Goedartia  was  explicitly  ex- 
cluded from  the  tribe.  Townes  (Townes  & 
Townes  1973)  synonymized  Heinrich's 
Compsophorini  under  Goedartiini.  Hein- 
rich  (1975)  placed  only  Goedartia,  Maraces, 
and  Pseudomaraces  in  Goedartiini,  retain- 
ing his  original  concept  of  two  tribes. 

Townes'  concept  of  Goedartiini  is  based 
on:  1)  a  wide  mandible  that  is  not  tapered 
apically,  2)  the  occipital  carina  that  meets 
the  hypostomal  carina  at  the  mandibular 
base,  and  3)  the  cell  l+2Rs  (areolet)  of  the 
fore  wing  that  is  pointed  above,  not  trun- 
cate (Townes  et  al  1961).  Heinrich  (1967) 
pointed  out  that  Goedartia  differs  from 
Compsophoriis  and  its  relatives  in  many  de- 
tails of  the  head  and  mesosoma,  including 
the  male  flagellum.  Until  fundamental 
studies  are  made  of  the  relationships  with- 
in the  stenopneusticine  Ichneumoninae,  it 
seems  best  to  maintain  Heinrich's  concept 
of  two  tribes. 

4.  Ichneumonini.  As  mentioned  previ- 
ously, we  accept  that  Opinion  159  of  the 
Commission  was  validly  issued.  Accord- 
ingly, Ichneumonini  is  the  correct  name 
for  the  tribe  that  Townes  called  Joppini. 
Gyrodontini  was  used  by  Carlson  (1979) 
due  to  Heinrich's  belief  that  Joppini  was 
strictly  a  Neotropical  group  (G.  Heinrich, 
pers.  comm.  to  R.W.  Carlson).  If  one  be- 
lieves the  Townes  position  to  be  correct,  it 
should  be  noted  that  Joppini  is  the  correct 
tribal  name.  The  Joppini  of  Townes  and 
Ichneumonini  of  Heinrich  are  essentially 
the  same  except  that  Heinrich  placed  Pseii- 
doplatylabus,  AcantJwjoppa,  and  related 
genera  in  the  Joppocryptini  (referred  to  as 
Acanthojoppini  in  Heinrich's  publications) 
and  placed  Tetragonochora  in  a  tribe  of  its 
own  (Heinrich,  1934). 

5.  Amblytelina.  Amblytelina  based  on 
Amblytelcs  Wesmael  is  a  junior  homonym 
of  Amblytelides  Blackburn,  1892  (Black- 
burn, 1892)  based  on  the  coleopterous  ge- 
nus Amblytelus  Erichson  (Carabidae).  Carl- 


son (1979)  emended  Amblytelina  to  Am- 
blytelesina  but  this  is  not  in  accordance 
with  the  Code,  as  such  cases  are  to  be  re- 
ferred to  the  Commission.  A  request  to  the 
Commission  is  being  prepared. 

6.  Protichneumonini-Heresiarchini. 
Townes  called  this  tribe  Ichneumonini, 
due  to  his  views  on  the  validity  of  Opin- 
ion 159.  Carlson  (1979)  and  Gupta  (1987) 
have  used  this  interpretation  as  well,  but 
all  other  authors  have  used  Protichneu- 
monini.  Depending  upon  the  treatment  of 
Heresiarches  and  related  genera,  however, 
Protichneumonini  may  not  be  the  correct 
name.  Perkins  (1959)  and  Rasnitsyn  &  Sii- 
tan  (1981)  maintain  Heresiarches  in  a  sep- 
arate tribe,  Heresiarchini.  Townes  always 
kept  the  genus  in  his  Ichneumonini  and 
Heinrich  did  likewise  in  the  equivalent 
Protichneumonini.  Heinrich  (Heinrich 
1960:  21)  discussed  the  situation  and  he 
later  recognized  the  group  as  a  subtribe  of 
Protichneumonini.  This  arrangement  ne- 
cessitates changing  the  tribal  name  to  Her- 
esiarchini, as  pointed  out  by  Perkins  (1959: 
28fn). 

7.  Trogini.  Trogini  based  on  Trogiis  Pan- 
zer is  a  junior  homonym  of  Trogidae 
MacLeay,  1819  (MacLeay,  1819),  based  on 
the  coleopterous  genus  Trox  Fabricius 
(Trogidae).  Carlson  (1979)  emended  Tro- 
gini to  Trogusini  but  this  is  not  in  accor- 
dance with  the  Code,  as  such  cases  are  to 
be  referred  to  the  Commission.  A  request 
to  the  Commission  is  being  prepared. 

Heinrich  placed  Callajoppm  and  related 
genera  in  the  Trogini  as  a  subtribe,  Calla- 
joppina.  Townes  instead  put  these  genera 
in  the  Heresiarchini,  restricting  Trogini  to 
the  parasitoids  of  Papilionidae  and  other 
Papilionoidea.  This  arrangement  is  fol- 
lowed here  but  it  should  be  noted  that  the 
Callajop^pa-group  is  part  of  a  continuum 
linking  certain  Heresiarchini  with  the  Tro- 
gini (Gauld  1984:  184-185;  Heinrich  1968: 
82,  1977:  284),  thus  making  the  placement 
of  this  group  in  one  tribe  or  another  a  sub- 
jective decision. 


288 


Journal  of  Hymenoptera  Research 


FAMILY-GROUP  NAMES  OF  THE 
ICHNEUMONINAE 

ICHNEUMONINAE  Latreille,  1802. 

1.  Alomyini  Forster,  1869  (incl.  Heterischnini 
of  Townes) 

Alomyoidae  Forster,  1869:  144,  194.  Type- 
genus:  Alomya  Panzer. 

Phaeogenoidae  Forster,  1869:  144,  191. 
Type-genus:  Phaeogenes  Wesmael. 

Dicaelotina  Holmgren,  1889:  343.  Type-ge- 
nus: Dicaelotus  Wesmael. 

Diadromina  Holmgren,  1889:  345.  Type-ge- 
nus: Diadromus  Wesmael. 

Epitomina  Holmgren,  1889:  347.  Type-ge- 
nus: Epitomus  Forster. 

Gnathoxina  Holmgren,  1889:  344.  Type-ge- 
nus: Gnathoxys  Wesmael. 

Herpestomina  Holmgren,  1889:  344.  Type- 
genus:  Herpestonius  Wesmael. 

Oronotina  Holmgren,  1889:  343.  Type-ge- 
nus: Oronotus  Wesmael. 

Stenodotina  Schmiedeknecht,  1903:  262. 
Type-genus:  Stenodontus  Berthoumieu. 

Heterischnini  Townes  et  al,  1961:  337,  458. 
Type-genus:  Heterischnus    Wesmael. 

Chauviniina  Diller,  1981:  95,  98.  Type-ge- 
nus: Chauinnia  Heinrich. 

Dicaelodontina  Diller,  1994:  126.  Type-ge- 
nus: Dicaelodontus  Diller. 

2.  Platylabini  Berthoumieu,  1904.  (=   Pristi- 
cerotini  of  Townes) 

Platylabini  Berthoumieu,  1904:  4.  Type-ge- 
nus: Plati/labus  Wesmael. 

[Pristaceratini  Townes  &  Townes,  1951:  280. 
Not  available  under  Article  13  of  the 
Code.  Incorrect  spelling.] 

Pristicerotini  Townes  et  al,  1961:  393,  458. 
Type-genus:  Pristiceros  Gravenhorst. 

3.  Eurylabini  Heinrich,  1934. 

Eurylabini  Heinrich,  1934:  64-67.  Type-ge- 
nus: Eurylabus  Wesmael. 

4.  Zimmeriini  Heinrich,  1934. 
[Zimmerini  Heinrich,  1934:  67.  Type-genus: 

{Zimmeria  Heinrich)  =  Cotiheresiarches  Te- 
lenga.  Incorrectly  formed  stem.] 
Zimmeriini  Heinrich;  Rasnitsyn  &  Siitan, 
1981:  510.  Justified  emendation. 

5.  Ceratojoppini  Heinrich,  1938. 
Ceratojoppini  Heinrich,  1938:  25, 121.  Type- 
genus:  Ceratojoppa  Cameron. 


6.  Ctenocalini  Heinrich,  1938. 

Ctenocalini  Heinrich,  1938:  25,  40-41.  Type- 
genus:  Ctenocalus  Szepligeti. 

7.  Goedartiini  Townes  et  al,  1961. 

Goedartiini  Townes  et  al,  1961:  399,  458. 
Type-genus:  Goedartia  Boie. 

8.  Compsophorini  Heinrich,  1967. 

[Compsophorina  Heinrich,  1962:  688.  Not 
available  under  Article  13  of  the  Code.] 

Compsophorini  Heinrich,  1967:  25.  Type- 
genus:  Compsophorus  Saussure. 

9.  Ischnojoppini  Heinrich,  1938. 

Ischnojoppini  Heinrich,  1938:  25, 117.  Type- 
genus:  Ischnojoppa  Kriechbaumer. 

10.  Listrodromini  Forster,  1869. 

Listrodromoidae  Forster,  1869:  144,  194. 
Type-genus:  Listrodromus  Wesmael. 

11.  Oedicephalini   Heinrich,   1934.    (=    Noto- 
semini  of  Townes) 

Oedicephalini  Heinrich,  1934:  67,  118-119. 

Type-genus:  Oedicephalus  Cresson. 
Notosemini  Townes  et  al,  1961:  338,  459. 

Type-genus:  Notosemus  Forster. 

12.  Ichneumonini  Latreille,  1802.  (=  Joppini  of 
Townes  in  part) 

(The  following  are  incertae  sedis  within  the 
tribe,  as  the  type  genera  were  never  for- 
mally assigned  to  a  subtribe  by  Heinrich: 

Joppinen  Kriechbaumer,  1898:  2.  Type-ge- 
nus: joppa  Fabricius. 

Merolidini  Brethes,  1909.  Type-genus:  Mer- 
olides  Brethes.  (The  type  and  sole  speci- 
men of  Merolides  arechavaletai  Brethes  is 
lost;  Townes  (1966;  pers.  comm.)  placed 
it  in  Joppini,  near  Limerodes.) 

Tetragonochorini  Heinrich,  1934:  64,  67. 
Type-genus:  Tetragonochora  Kriechbau- 
mer. (Townes  (Townes  &  Townes,  1966) 
placed  this  genus  in  his  Joppini.) 

a.  Ichneumonina  Latreille,  1802. 

Ichneumonides  Latreille,  1802:  318.  Type- 
genus:  Ichneumon  Linnaeus. 

[Pterocorminae  Heinrich,  1949:  256.  Not 
available  under  Article  13  of  the  Code.] 

b.  Gyrodontina  Schmiedeknecht,  1902. 

Gyrodontini  Schmiedeknecht,  1902:  9,  28. 
Type-genus:  Gyrodonta  Cameron. 


Volume  4,  1995 


289 


Cratichneumonina  Heinrich,  1967:  27. 
Type-genus:  Crat ichneumon  Thomson. 

[c.  Amblytelina  Viereck,  1918.] 

[Amblytelinae  Viereck,  1918:  74.  Type-ge- 
nus: Amblyteles  Wesmael.  Amblytelina 
based  on  Amhlyidcs  Wesmael  is  a  junior 
homonym  of  Amblytelides  Blackburn, 
1892  (Blackburn,  1892:  85),  based  on  the 
coleopterous  genus  Ambh/telus  Erichson 
(Carabidae)] 

[Amblytelesina  Carlson,  1979:  507.  Illegal 
emendation.  Article  55(b)  of  the  Code.] 

d.  Hoplismenina  Heinrich,  1967. 
Hoplismenina  Heinrich,  1967:  26.  Type-ge- 
nus: Hoplismenus  Gravenhorst. 

e.  Aethioplitina  Heinrich,  1967. 
Aethioplitina  Heinrich,  1967:  27.  Type-ge- 
nus: Acthioplites  Heinrich. 

13.  Joppocryptini  Viereck,  1918. 
Joppocryptinae  Viereck,  1918:  73.  Type-ge- 
nus: foppociyptus  Viereck. 

Acanthojoppini  Heinrich,  1934:  65,  67,  138. 
Type-genus:  {Acauthojoppa  Cameron)  = 
EccopHosage  Kriechbaumer.  Townes  (1966) 
placed  this  genus  in  his  Joppini. 

14.  Heresiarchini  Ashmead,  1900  (=  Ichneu- 
monini  of  Townes;  includes  part  of  Trogini 
of  Heinrich). 

a.  Protichneumonina  Heinrich,  1934. 
Protichneumonini  Heinrich,   1934:   66,  84. 

Type-genus:  Protichneumon  Thomson. 

b.  Heresiarchina  Ashmead,  1900. 
Heresiarchini  Ashmead,  1900:  567.  Type-ge- 
nus: Heresiarches  Wesmael. 

c.  Apatetorina  Heinrich,  1967. 
Apatetorina  Heinrich,  1967:  26,  50.  Type-ge- 
nus: Apatetor  Saussure. 

d.  Callajoppina  Heinrich,  1962. 
Callajoppina  Heinrich,  1962:  809-810.  Type- 
genus:  Callajoppa  Cameron. 

[15.  Trogini  Forster,  1869.] 

[Trogoidae  Forster,  1869:  144,  188.  Type-ge- 
nus: Tragus  Panzer.  Trogini  based  on  Tra- 
gus Panzer  is  a  junior  homonym  of  Tro- 
gidae  MacLeay,  1819  (MacLeay,  1819: 
136),  based  on  the  coleopterous  genus 
Trox  Fabricius.  Although  Forster  cites 
Tragus  Gravenhorst,  Gravenhorst  includ- 


ed the  type  species  of  Tragus  Panzer  in 
his  treatment  of  the  genus  and  had  the 
same  generic  concept] 
[Trogusina  Carlson,  1979:  538.  Illegal  emen- 
dation. Article  55(b)  of  the  Code.] 

ACKNOWLEDGEMENTS 

Robert  Carlson's  generosity  in  sharing  his  list  of 
ichneumonid  family-group  names  is  gratefully  ac- 
knowledged. We  wish  to  thank  the  following  for  their 
advice  and  comments:  John  Barron,  Yves  Bosquet, 
Terry  Erwin,  Ian  Gauld,  Henri  Goulet,  CD.  Michener, 
Charles  Porter,  Alexandr  Rasnitsyn,  Michael  Sharkey, 
G.  Stuart  Walley,  and  Robert  Wharton. 

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APPENDIX  1 

Townes'  arguments  regarding  the  invalidity  of 
Opinion  159  were  put  forth  in  Townes  (1957,  1969); 
his  version  of  the  relevant  events  may  be  summarized 
as  follows  from  these  two  references  except  where 
otherwise  noted: 

1913 — At  the  Monaco  International  Congress  of  Zo- 
ology, the  Commission  was  delegated  authority  to 
suspend  the  Rules  and  declare  on  various  cases,  on 
the  authority  of  the  Congress. 
Suspensions  could  be  made  only  under  the  follow- 
ing procedures: 

a.  Not  less  than  one  year's  notice  that  such  a  sus- 
pension is  under  consideration  in  two  or  more 
of  five  specified  publications. 

b.  The  votes  of  the  Commission  is  unanimous  in 
favor  of  suspension. 

c.  If  the  vote  is  a  %  majority  of  the  full  Commis- 
sion, but  not  unanimous,  in  favor  of  suspension, 
the  Commission  must  report  the  facts  to  the  next 
Congress  (where  specified  arbitration  would  be 
made.) 

1935 — J.C.  Bradley  presented  a  list  of  generic  names 
at  the  Commission  meeting  in  Lisbon,  requesting 
suspension  of  the  Rules  in  these  cases.  Bradley  re- 
quested new  type-species  designations  for  Ichneu- 
mon, Pimpla,  and  Ephialtes.  Six  out  of  18  Commis- 
sioners attended  the  Lisbon  meeting  and  six  tem- 
porary Commissioners  were  appointed.  Ten  out  of 
12  were  present  when  the  Bradley  proposal  was 
presented  and  approved.  No  notice  of  intention  to 
consider  these  names  had  been  published  in  ad- 
vance of  the  Lisbon  meeting. 


1936 — When  it  was  pointed  out  that  the  required  ad- 
vance notice  had  not  been  published  regarding  the 
proposed  changes,  the  Commission  published  no- 
tices in  1936  that  it  would  officially  consider  Brad- 
ley's list  at  some  future  time. 

1939 — Karl  Jordan  and  Francis  Hemming  met  and 
made  a  decision  between  themselves.  The  au- 
thority for  their  action  was  that  upon  adjournment 
of  the  Lisbon  sessions,  they  were  to  "take  such 
other  action  as  might  appear  to  them  necessary  or 
expedient  ...  to  give  effect  to  the  decisions  reached 
by  the  Commission  at  the  Lisbon  Session  .  .  ." 
(Hemming  1945a).  "The  direction  upon  adjourn- 
ment of  the  Lisbon  meeting  of  the  Commissioners 
to  Jordan  and  Hemming  to  put  into  effect  the  de- 
cisions of  the  meeting  did  not  give  Jordan  and 
Hemming  power  to  make  the  decisions  ...  " 
(Townes  1957). 

1943 — Following  this,  Bradley's  proposals  never 
came  up  again  for  decision.  Townes  contacted 
James  L.  Peters,  the  acting  secretary  at  Lisbon  and 
asked  whether  any  further  action  had  been  taken 
on  the  Bradley  names.  Peters  replied  that  the  ques- 
tion of  the  ichneumonid  names  had  never  been  cir- 
culated. Furthermore,  he  stated  that  "no  vote  had 
been  taken  on  the  subject  of  these  three  names  .  .  ." 
(Townes  1969). 

Townes  thus  concluded  that  the  Commission  never 
took  action.  According  to  Townes,  Hemming  argued 
the  requirements  for  suspending  the  Rules  had  been 
met  by:  1)  a  vote  in  1935  before  publication  of  the 
cases,  2)  publication  in  1936  that  the  cases  would  be 
considered,  3)  the  fact  that  no  Commissioner  wrote 
him  after  1936  with  objections.  Townes  rejects  these 
arguments  because:  1)  the  vote  must  be  taken  after 
publication,  2)  only  12  Commissioners  were  in  Lisbon 
and  only  ten  in  the  actual  meeting,  while  the  require- 
ment is  for  a  unanimous  vote  by  all  18  Commission- 
ers, 3)  awaiting  unsolicited  letters  is  not  the  same  as 
an  actual  request  for  a  vote.  Finally,  Townes  noted 
that  the  Bradley  petition,  signed  by  59  hymenopter- 
ists,  was  circulated  in  1928  and  is  not  the  same  as 
that  presented  in  1935,  except  in  the  case  of  Cri/ptus. 
The  type-species  for  Icluwinnon,  Ephialtes,  and  Pimpla 
are  the  as  same  recognized  by  Townes. 

This  recounting,  however,  omits  crucial  informa- 
tion. While  the  Monaco  Congress  listed  the  condi- 
tions under  which  the  Rules  could  ordinarily  be  sus- 
pended. Secretary  C.W.  Stiles  {Compte  Rendu  1913: 
892-893)  pointed  out  the  following:  "Upon  a  unani- 
mous vote,  "By-Laws"  may  be  temporarily 
"suspended",  that  is  to  say,  they  may  be  set  aside 
and  the  bodv  takes  action  on  the  matter  under  con- 
sideration unrestricted  by  the  provisions  of  the  By- 
Laws;  and  such  action,  if  taken  under  a  "Special 
Rule"  framed  for  the  case  at  hand  or  without  refer- 
ence to  any  rules,  except  the  "Constitution"  and  rec- 
ognized "Parliamentary  Rules",  has  all  the  validity 


292 


Journal  of  Hymenoptera  Research 


of  an  action  taken  under  the  "By-Laws".  Thus,  if  the 
Congress  confers  upon  the  Commission  the  plenary 
power  to  suspend  the  Regies  in  any  given  case,  it 
practically  says  to  the  Commission:  "If  you  carry  out 
the  precautions  provided  for  in  the  Supplemental  Re- 
port, you  may  decide  any  given  case  arbitrarily  with- 
out reference  to  the  Regies  or  you  may  make  a  "Spe- 
cial Rule"  to  govern  that  particular  case,  and  this 
Congress  will  accept  your  decision  as  being  just  as 
authoritative  as  if  you  had  made  this  ruling  strictly 
in  accord  with  the  Code."  A  plan  of  this  kind  is  thor- 
oughly in  accord  with  recognized  parliamentary  cus- 
toms and  it  has  the  great  advantage  of  avoiding  the 
necessity  of  introducing  "ExcepHons"  to  the  Rules." 
This  is  the  background  against  which  the  events  of 
1935  occurred. 

Contrary  to  Townes'  statement,  Bradley's  petition 
of  1935  was  the  same  as  that  circulated  in  1928.  It 
was  presented  to  the  International  Committee  of  En- 
tomological Nomenclature  (ICEN)  in  Madrid  in  the 
week  preceding  the  opening  of  the  Twelfth  Interna- 
tional Congress  of  Zoology  in  Lisbon  (Hemming 
1943a).  The  petition  was  considered  and  a  number  of 
proposals  were  made  by  the  ICEN  concerning  reso- 
lution of  the  problems  (Hemming  1945a:  282). 

The  ICEN's  recommendations  for  this  and  other 
cases  were  laid  before  the  Commission  at  its  second 
meeting  on  16  Sept.  1935.  Because  it  had  not  been 
possible  for  the  Commission  to  meet  prior  to  the  Con- 
gress, there  were  only  7}/i  days  to  discuss  the  cases 
and  prepare  a  report  to  the  Congress.  President  Jor- 
dan felt  it  best  to  take  the  pending  cases  into  imme- 
diate consideration  and  "for  this  purpose  suspend  the 
By-laws  of  the  Commission  to  such  extent  as  might 
be  necessary  for  the  period  of  the  present  Congress." 
(Hemming  1943b:  10-11).  At  that  meeting,  10  out  of 
11  Commissioners  present  in  Lisbon  were  in  atten- 
dance; Walter  Arndt,  an  alternate,  did  not  attend  the 
five  Commission  meetings  and  Teiso  Esaki  was  elect- 
ed to  the  Commission  at  this  particular  session.  The 
attending  Commissioners  passed  the  usual  parlia- 
mentary unanimous  consent  to  suspend  the  By-laws 
(note  this  was  not  an  emendaiion  of  the  By-Laws, 
which  would  have  required  an  affirmative  vote  of  12 
members  (ICZN  By-laws,  Article  VIII,  Sect.  I)).  It  was 
agreed  to  give  immediate  consideration  to  all  sub- 
mitted cases  that  it  was  felt  had  reached  the  stage  at 
which  a  decision  could  be  made.  To  give  effect  to  this, 
use  would  be  made  of  the  By-law  suspension  that 
had  just  been  agreed  upon.  Since  this  involved  vio- 
lating the  normal  procedures  for  "suspension  of  the 
Regies"  in  that  the  prescribed  advertisement  had  not 
been  published  one  year  previously,  the  various  cases 
and  the  Commission's  decisions  would  be  advertised 
as  soon  as  possible  after  the  conclusion  of  the  Lisbon 
meetings  and  no  Opinions  would  be  published  until 
after  one  year  from  the  date  of  the  advertisement's 
dispatch.  Nowhere  does  Townes,  in  his  recounting. 


mention  that  the  Commission  suspended  its  By-laws, 
as  provided  by  parliamentary  procedures. 

At  the  Commission's  third  session,  it  agreed  "un- 
der suspension  of  the  Rules"  to  set  aside  current  type 
designations  for  the  names  on  the  Bradley  list  and 
declare  new  ones.  For  ichneumonids,the  Commission 
agreed  to  reject  EpMaltes  Schrank  1802  and  to  desig- 
nate the  following  type  species: 

Ichneumon  L.:  Iclmeumon  extensorius  L. 

Pimpla  Fabricius:  Ichneumon  instigator  Fabricius 

Epiiialtes  Gravenhorst:  Ichneumon  manifestor  L. 

The  Commission  "came  to  the  conclusion  that  the 
more  radical  of  the  proposals  submitted  by  the 
[ICEN]  provided  the  most  satisfactory  solution  of  the 
difficulties  presented  by  the  present  case."  (Hemming 
1945a:  282).  Townes  neglected  to  mention  the  ICEN 
involvement  and  chose  to  portray  the  Commission 
decisions  as  part  of  an  illegal  and  arbitrary  process. 

The  Commission's  fifth  meeting  on  18  Sept.  1935 
saw  11  of  the  12  Commissioners  present;  it  was 
agreed  that  President  Jordan  "and  the  new  Secretary, 
when  elected,  should  be  authorized  to  make  such  ar- 
rangements, and  to  take  such  actions,  as  might  ap- 
pear to  them  necessary  or  expedient  .  .  ."  and  "to  se- 
cure the  due  publication  of  the  Opinions  agreed  upon 
from  time  to  time  by  the  Commission  at  their  Lisbon 
Session."  (Hemming  1943b:  48).  The  Commission  also 
unanimously  agreed  to  adopt  as  their  report  to  the 
Congress  the  draft  prepared  by  Hemming  (ibid.,  p. 
47).  This  report  (Hemming  1943c),  which  duly  noted 
the  suspension  of  the  By-laws,  was  approved  by  the 
Congress  (Anonymous,  1943). 

Notice  of  the  possible  suspension  of  the  Rules  was 
published  in  May  1936  (Stiles  1936).  Except  for 
Townes'  allegation,  there  is  no  evidence  that  the  ad- 
vertisement was  published  at  the  urging  of  scientists 
concerned  that  no  published  notices  had  been  made 
prior  to  the  Lisbon  sessions.  It  was  done  on  the  au- 
thority of  the  Commission  operating  under  the  sus- 
pension of  the  By-laws. 

After  the  1936  advertisement,  one  communication 
was  received  regarding  the  ichneumonid  names.  It 
bore  the  signature  of  S.A.  Rohwer  in  the  name  of  the 
Committee  on  Nomenclature  of  the  Entomological 
Society  of  Washington  (Hemming  1945a).  The  letter 
was  against  the  proposed  new  type-species;  append- 
ed to  the  letter  was  a  note  of  dissent,  in  favor  of  the 
proposed  nomina  conservanda,  by  R.A.  Cushman,  one 
of  the  prominent  ichneumonologists  of  the  day. 

Copies  of  the  letter  were  immediately  sent  to  the 
other  Commissioners  but  no  member  expressed  him- 
self as  agreeing  with  the  Rohwer  point  of  view  (ibid., 
p.  286).  Jordan  and  Hemming  met  on  19  June  1939 
under  the  authority  of  the  Resolution  adopted  by  the 
Commission  on  18  Sept.  1935  (Hemming  1945b). 
They  took  note  that  no  Commissioner  had  responded 
to  the  Rohwer  communication.  In  their  opinion,  no 
new  facts  had  been  brought  forth  that  were  not  before 


Volume  4,  1995 


293 


the  Commission  in  1935.  Given  this,  they  "decided 
the  proper  course  .  .  .  was  to  give  effect  to  the  deci- 
sions in  this  matter  reached  by  the  International 
Commission  at  the  Lisbon  Session  .  .  .  and  therefore 
that  Opinions  should  be  issued  as  soon  as  possible." 
(Hemming  1945a:  287).  The  resulting  Opinion  159 
was  published  in  1945  (ibid.). 

Townes  was  preparing  to  publish  a  catalog  of  Ne- 
arctic  Ichneumonidae  (Townes  1944-1945)  when  he 
wrote  to  Peters  in  1943.  He  took  Peters'  advice  and 
used  names  based  on  priority;  they  both  apparently 
expected  the  matter  to  be  voted  on  by  the  Commis- 
sion, with  the  outcome  uncertain.  Jordan  and  Hem- 
ming, however,  were  given  the  authority  to  make  de- 
cisions by  virtue  of  the  Commission  acting  under  the 
valid  suspension  of  the  By-laws.  Townes'  objections 
to  Hemming's  listing  of  the  usual  procedures  for  sus- 
pending the  Rules  (i.e.,  one  year's  notice  of  suspen- 
sion in  two  or  more  of  five  journals,  etc)  are  irrele- 
vant. Hemming  apparently  used  these  stock  phrases 
as  boilerplate. 

At  this  point,  the  reader  might  wonder  why  such 
effort  has  been  spent  on  an  arcane  issue.  From  our 
viewpoints  as  ichneumonologists,  it  is  important  that 
Townes'  arguments  be  put  to  rest  in  order  to  attain 
a  stable  ichneumonid  nomenclature.  Outside  Ichneu- 
monidae, there  always  remains  the  possibility  that 
someone  will  read  Townes'  version  of  events  and 
raise  havoc  in  groups  that  heretofore  have  attained 
relative  nomenclatural  stability. 

In  summary,  rejection  of  Townes'  arguments  gives 
the  following  generic  names: 

Pimplinae 
Ephialtini  (=  Pimplini  of  Townes) 
ICZN  name  Townes  name 

Piuipla  Fabricius,  1804 


Pimplini  (=  Ephialtini  of  Townes) 

ICZN  name  Townes  name 


Epliialtes  Graven- 
horst,  1829 

(type-species:  Ich- 
neumon manifes- 
tor  L.) 


Apcchthis  Forster, 
1869 


Ephmltes  Schrank,  1802 


(type-species:  Ich-  (type-species:  Ichneumon 

neumon  rufatus  compunctor  L.) 

L.) 

Piitipla  Fabricius,  Coccygominnis  Saussure, 

1804  1892 

(type-species:  Ich-  (type-species:  Coccygomi- 

neumon  instiga-  mus  madecassus  Saus- 

tor  L.)  sure) 

Ichneumoninae 

Ichneumonini  (=  Joppini  of  Townes) 

ICZN  name  Townes  name 

Icluu'uinoti  L.,  1758  Pterocormus  Forster, 

1850 

(type-species:  Ich-  (type-species:  Ichneumon 

neumon  extenso-  latrator  Fabricius) 

rius  L.) 

Heresiarchini  (=  Ichneumonini  of  Townes,  Proti- 
chneumonini  of  Heinrich) 

ICZN  name  Townes  name 

Coelichneumon  Ichneumon  L.,  1758 

Thomson,  1893 

(type-species:  Ich-  (type-species:  Ichneumon 

neumon  comita-  comitator  L.) 

tor  L.) 


(type-species:  Ichneumon 
manifestor  L.) 


CONSTITUTION  AND  BYLAWS  OF  THE 
INTERNATIONAL  SOCIETY  OF  HYMENOPTERISTS 


CONSTITUTION 

ARTICLE  I 

Section  1.  This  organization  shall  be  known  as  the  International  Society  of  Hymen- 

opterists. 
Section  2.  The  objectives  of  the  Society  shall  be  to  encourage  scientific  research  and  to 

promote  the  diffusion  of  scientific  knowledge  concerning  Hymenoptera. 

ARTICLE  II 

Section  1.  The  membership  of  the  Society  shall  consist  of  Active  Members,  Student 
Members,  Emeritus  Members,  and  Sustaining  Members. 

Section  2.  Active  Members  shall  be  individuals  who  are  interested  in  Hymenoptera 
and  the  objectives  of  the  Society.  Each  Active  Member  shall  pay  to  the  So- 
ciety annual  dues  as  prescribed  in  the  Bylaws. 

Section  3.  Student  Members  shall  be  full-time  undergraduate,  or  part-time  or  full-time 
graduate  students  at  a  recognized  institution  of  higher  learning.  Each  Stu- 
dent Member  shall  pay  to  the  Society  annual  dues  as  prescribed  in  the  By- 
laws. To  qualify  for  Student  Membership,  a  student  must  have  his  or  her 
status  verified  by  a  faculty  member  at  his  or  her  institution.  Student  Mem- 
bers shall  have  all  the  rights  and  privileges  of  active  membership  except 
holding  office. 

Section  4.  Emeritus  Members  shall  be  members  who  have  retired  from  active  service, 
and  who  petition  the  Executive  Committee  for  a  change  in  classification. 
Election  to  Emeritus  Membership  requires  a  three-fourths  vote  of  the  Ex- 
ecutive Committee  present  or  voting  by  mail.  Emeritus  Members  shall  have 
all  rights  and  privileges  of  active  membership  except  holding  office,  and 
shall  be  released  from  payment  of  dues.  Emeritus  Members  may  subscribe 
to  any  journals  published  by  the  Society  with  the  payment  of  subscription 
fees. 

Section  5.  Sustaining  Members  shall  be  educational  or  commercial  institutions,  or  in- 
dividuals interested  in  the  promotion  and  advancement  of  the  study  of  Hy- 
menoptera, and  who  are  in  sympathy  with  the  objectives  of  the  Society. 
Annual  dues  shall  be  paid  as  prescribed  in  the  Bylaws. 

ARTICLE  III 

Section  1.  The  officers  of  the  Society  shall  consist  of  a  President,  a  President-Elect,  an 
Editor  (when  and  if  the  Society  publishes  a  journal),  a  Secretary,  and  a  Trea- 
surer. The  President-Elect  shall  serve  for  two  years,  and  then  automatically 
assume  the  office  of  President  for  two  years.  The  Editor,  Secretary,  and  Trea- 
surer shall  each  serve  for  four  years. 

Section  2.  The  elected  officers  shall  take  office  at  the  close  of  the  business  session  of 
the  next  scheduled  meeting  of  the  Society  following  their  election  and  shall 
hold  office  until  their  successors  have  been  elected  and  take  office. 

ARTICLE  IV 

Section  1.  The  President  shall  discharge  the  usual  duties  of  a  presiding  officer  at  the 
meetings  of  the  Society  and  the  Executive  Committee. 


Volume  4,  1995  295 

Section  2.  The  President-Elect  shall  assume  the  duties  of  the  President  in  the  event  of 
his  or  her  absence.  In  the  event  that  both  the  President  and  President-Elect 
are  absent  at  a  meeting.  The  President  may  appoint  another  member  of  the 
Executive  Committee  to  serve  as  presiding  officer. 

Section  3.  The  Editor  shall  assume  editorial  responsibility  for  any  journals  published 
by  the  Society.  He  or  she  shall  be  responsible  for  selecting  and  delegating 
the  duties  of  any  subject  editors. 

Section  4.  The  Secretary  shall  keep  the  records  of  the  Society.  He  or  she  shall  be  re- 
sponsible for  notifying  the  membership  of  meetings,  calls  for  papers,  dues, 
nominations,  annual  reports,  and  all  other  items  as  directed  by  the  Executive 
Committee.  He  or  she  shall  record  minutes  of  meetings  of  the  Society  and 
Executive  Committee,  and  maintain  the  record  of  names  and  address  of 
members. 

Section  5.  The  Treasurer  shall  have  custody  of  all  funds  of  the  Society  and  shall  make 
routine  disbursements  as  required.  Unusual  disbursements  and/or  invest- 
ments shall  be  made  only  with  the  concurrence  of  the  Executive  Committee. 
He  or  she  shall  keep  an  account  of  receipts  and  disbursements  and  those 
accounts  shall  be  audited  as  provided  by  the  Executive  Committee.  He  or 
she  shall  present  a  statement  of  the  receipts  and  expenditures  and  funds  for 
the  past  year  at  the  business  session  of  regular  meetings  of  the  Society. 

ARTICLE  V 

Section  1.  The  Executive  Committee  shall  consist  of  the  President,  President-Elect,  Ed- 
itor (if  elected).  Secretary,  and  Treasurer. 

Section  2.  The  Executive  Committee  shall  execute  and  administer  the  affairs  of  the 
Society  during  the  interval  between  the  regular  meetings.  It  shall  be  respon- 
sible for  authorizing  all  unusual  expenditures  by  the  Treasurer  and  shall  fill 
temporary  vacancies  among  the  officers. 

ARTICLE  VI 

Section  1.  The  Executive  Committee  shall  appoint  a  Nominating  Committee  of  at  least 
three  members  to  serve  for  one  year.  The  Nominating  Committee  shall  so- 
licit nominations  for  those  vacancies  on  the  Executive  Committee  that  will 
occur  in  the  following  year.  The  Nominating  Committee  shall  nominate  at 
least  two  persons  for  each  office  and  ascertain  whether  the  proposed  nom- 
inees will  accept  office  if  elected.  The  slate  of  nominees  will  be  made  avail- 
able to  the  membership  by  the  Secretary,  and  a  mail  ballot  will  be  taken. 
The  candidate  who  receives  the  majority  of  votes  cast  shall  be  declared  elect- 
ed. 

Section  2.  The  Executive  Committee  shall  have  the  authority  to  appoint  a  member  of 
the  Society  to  fill  the  remainder  of  the  term  of  any  officer  of  the  Society  who 
is  unable  to  continue  for  whatever  reason. 

Section  3.  Standing  committees  and  ad  hoc  committees  shall  be  appointed  by  the  Ex- 
ecutive Committee. 

ARTICLE  VII 

Section  L  The  Society  shall  hold  regular  meetings  as  provided  in  the  Bylaws.  The 
Executive  Committee  may  arrange  for  special  meetings  and /or  joint  pro- 
grams with  related  scientific  groups,  and  for  presentation  of  papers. 


296  Journal  of  Hymenoptera  Research 

ARTICLE  VIII 

Section  1.  Amendments  to  the  Constitution  of  the  Society  may  be  proposed  in  writing 
by  any  member  to  the  Executive  Committee.  Such  proposed  amendments 
as  are  approved  by  a  majority  vote  of  the  Executive  Committee  shall  be 
submitted  to  the  members  by  mail  ballot,  and  will  be  adopted  by  a  favorable 
vote  of  two-thirds  of  the  members  voting. 

ARTICLE  IX 

Section  1.  In  the  event  of  dissolution  or  termination  of  the  International  Society  of 
Hymenopterists,  title  to  and  possession  of  all  property  of  the  Society  shall 
pass  forthwith  to  such  organization,  dedicated  to  similar  purposes  and  qual- 
ified for  exemption  under  Section  501(c)(3)  of  the  Internal  Revenue  Code  of 
1954  (or  corresponding  provision  of  an  future  United  States  Internal  Reve- 
nue Law),  as  the  Executive  Committee  shall  deem  best  qualified  to  carry  on 
the  functions  of  the  Society. 

BYLAWS 

1.  Annual  dues  for  Active  Members  shall  be  $25.00  US  per  year.  Student  Members 
shall  pay  $15.00  US  per  year.  Sustaining  Members  shall  pay  $50.00  US  per  year. 
Dues  may  be  waived  upon  request  at  the  discretion  of  the  Treasurer. 

2.  These  Bylaws  may  be  amended  or  suspended  by  a  two-thirds  vote  of  the  members 
present  at  any  regular  meeting.  New  Bylaws  may  be  adopted  by  a  two-thirds  vote 
of  the  members  present  at  any  regular  meeting. 

3.  Regular  annual  meetings  shall  be  held  in  conjunction  with  the  annual  meetings  of 
the  Entomological  Society  of  America.  Regular  quadrennial  international  meetings 
shall  be  held  as  scheduled  when  and  where  by  the  Executive  Committee. 

4.  Newly  elected  members  of  the  Executive  Committee  shall  take  office  at  the  regular 
meeting  held  in  conjunction  with  the  annual  meeting  of  the  Entomological  Society 
of  America  following  their  election. 


INSTRUCTIONS  FOR  AUTHORS 
General  Policy 

The  Journal  of  Hymenoptera  Research  invites  papers  of  high  scienrific  quality  reporting  comprehensive  re- 
search on  all  aspects  of  Hymenoptera,  including  biology,  behavior,  ecology,  systematics,  taxonomy,  genetics, 
and  morphology.  Taxonomic  papers  describing  single  species  are  unlikely  to  be  accepted  unless  a  strong  case 
is  evident,  such  as  importance  in  economic  entomology  or  with  concurrent  biology  or  ecology.  Manuscript 
length  generally  should  not  exceed  50  typed  pages;  however,  no  upper  limit  on  length  has  been  set  for  papers 
of  exceptional  quality  and  importance,  including  taxonomic  monographs  at  generic  or  higher  level. 

All  papers  will  be  reviewed  by  at  least  two  referees.  The  referees  will  be  chosen  by  the  appropriate  subject 
editor.  However,  it  would  be  helpful  if  authors  would  submit  the  names  of  two  persons  who  are  competent 
to  review  the  manuscript. 

The  language  of  publication  is  English.  Summaries  in  other  languages  are  acceptable. 

The  deadline  for  receipt  of  manuscripts  is  1  February  of  each  year. 

Format  and  Preparation 

Three  copies  of  each  manuscript,  including  copies  of  illustrahons,  should  be  submitted  on  letter  size  or 
A4  paper,  double  spaced,  with  at  least  25  mm  margins  on  all  sides.  On  the  upper  left  of  the  title  page  give 
name,  address  and  telephone  and  fax  numbers  of  the  author  to  whom  all  correspondence  is  to  be  sent. 

The  paper  should  have  a  concise  and  informative  title,  followed  by  the  names  and  addresses  of  all  authors. 
The  sequence  of  material  should  be:  title,  author(s),  abstract,  text,  acknowledgments,  literature  cited,  appendix, 
figure  legends,  figure  copies  (each  numbered  and  identified),  tables  (each  numbered  and  with  heading).  Each 
of  the  following  should  start  a  new  page:  (1)  title  page,  (2)  abstract,  (3)  text,  (4)  literature  cited,  (5)  figure 
legends,  (6)  footnotes. 

Following  acceptance  of  the  manuscript,  the  author  should  provide  the  editor  with  one  copy  of  the  manu- 
script accompanied  by  a  copy  on  diskette  using  DD,  double  sided  computer  diskettes  -  IBM  compatible  MS 
DOS  5.25  inch  or  IBM  and  Macintosh  3.5  inch  diskettes.  (Authors  who  do  not  have  access  to  a  computer 
should  submit  three  copies  of  the  manuscript.)  The  paper  may  be  submitted  in  most  PC  and  Mac  word 
processor  programs  such  as  Microsoft  Word,  FullWrite  Professional,  WordPerfect,  WriteNow,  Nisus,  Mac- 
Write,  or  MacWrite  II.  If  possible,  all  words  that  must  be  italicized  should  be  done  so,  not  underscored.  Use 
of  the  words  male  and  female  is  preferred  to  sex  symbols.  Tables  may  be  formatted  in  a  spread  sheet  program 
such  as  MS  Works  or  MS  Excel.  Text  should  be  double  spaced  typing,  with  25  mm  left  and  right  margins. 
Tables  should  be  put  in  a  separate  file.  Diskettes  should  be  accompanied  by  the  name  of  the  software  program 
used  (e.g.,  WordPerfect,  Microsoft  Word).  Authors  should  keep  backup  copies  of  all  material  sent  to  the  Editor. 
The  Society  cannot  be  responsible  for  diskettes  or  text  mislaid  or  destroyed  in  transit  or  during  editing. 

Illustrations  should  be  planned  for  reduction  to  the  dimension  of  the  printed  page  (14.8  x  21.8  cm,  column 
width  7.0  cm)  and  allow  room  for  legends  at  the  top  and  bottom.  Do  not  make  plates  larger  than  14  X  18  in. 
(35.5  X  46  cm).  Individual  figures  should  be  mounted  on  a  suitable  drawing  board  or  similar  heavy  stock. 
Photographs  should  be  trimmed,  grouped  together  and  abutted  when  mounted.  Figure  numbers  should  be 
on  the  plate,  but  it  is  strongly  recommended  that  names  be  included  after  the  numbers  (e.g..  Fig.  2,  texanus). 
Include  title,  author(s)  and  address(es),  and  illustration  numbers  on  back  of  each  plate.  Original  figures  need 
not  be  sent  until  requested  by  the  editor,  usually  after  the  manuscript  has  been  accepted.  Reference  to  figures/ 
tables  in  the  text  should  be  in  the  style  "(Fig.  I)"  "(Table  1)".  Measurements  should  be  in  the  metric  system. 

All  papers  must  conform  to  the  International  Code  of  Zoological  Nomenclature.  The  first  mention  of  a  plant 
or  animal  should  include  the  full  scientific  name  including  the  authority.  Genus  names  should  not  be  abbre- 
viated at  the  beginning  of  a  sentence.  In  taxonomic  papers  type  specimens  must  be  clearly  designated,  type 
depositories  must  be  clearly  indicated,  and  new  taxa  must  be  clearly  differentiated  from  existing  taxa  by 
means  of  keys  or  differential  diagnoses.  Authors  are  required  to  deposit  all  type  material  in  internationally 
recognized  institutions  (not  private  collections).  Voucher  specimens  should  be  designated  for  specimens  used 
in  behavioral  or  autecological  studies,  and  they  should  be  deposited  similarly. 

Acceptance  of  taxonomic  papers  will  not  require  use  of  cladistic  methods;  however,  authors  using  them 
will  be  expected  to  specify  the  phylogenetic  program  used  (if  any),  including  discussion  of  program  options 
used.  A  data  matrix  should  be  provided  if  the  subject  is  complex.  Cladograms  must  be  hung  with  characters 
and  these  should  include  descriptors  (not  numbers  alone)  when  feasible.  The  number  of  parsimonious  clado- 
grams generated  should  be  stated  and  the  reasons  for  the  one  adopted.  Lengths  and  consistency  indices  should 
be  provided.  Adequate  discussions  should  be  given  for  characters,  plesiomorphic  conditions,  and  distributions 
of  characters  among  outgroups  when  problematical. 

References  in  the  text  should  be  (Smith  1999),  without  a  comma,  or  Smith  (1999).  Two  articles  by  a  single 
author  should  be  (Smith  1999a,  1999b)  or  Smith  (1999a,  1999b).  For  papers  in  press,  use  "in  press",  not  the 
expected  publication  date.  The  Literature  Cited  section  should  include  all  papers  referred  to  in  the  paper. 
Journal  names  should  be  spelled  out  completely  and  in  italics. 

Charges 

Publication  is  free  to  members  of  the  International  Society  of  Hymenopterists.  At  least  one  author  of  the 
paper  must  be  a  member.  Reprints  are  charged  to  the  author  and  must  be  ordered  when  returning  the  proofs; 
there  are  no  free  reprints.  Author's  corrections  and  changes  in  proof  are  also  charged  to  the  author.  Color 
plates  will  be  billed  at  full  cost  to  the  author. 

All  manuscripts  and  correspondence  should  be  sent  to: 

Paul  M.  Marsh,  Editor 

P.  O.  Box  384 

North  Newton,  Kansas  67117 

(316)  284-0990 


CONTENTS 
(Continued  from  front  cover) 


QUICKE,  D.  L.  J.  and  R.  A.  KRUFT.   Latitudinal  gradients  in  North  American  braconid 

wasp  species  richness  and  biology    194 

QUICKE,  D.  L.  J.,  M.  FITTON  and  J.  HARRIS.  Ovipositor  steering  mechanisms  in  braconid 

wasps    110 

SHARKEY,  M.  J.  and  D.  H.  JANZEN.  Review  of  the  world  species  of  Sigalphus  (Hymenop- 

tera:  Braconidae:  Sigalphrnae)  and  biology  of  Sigalphus  romeroi,  new  species     ....       99 

WAHL,  D.  B.  and  W.  R.  M.  MASON.  The  family-group  names  of  the  Ichneumoninae  (Hy- 

menoptera:  Ichneumonidae)     285 

Constitution  and  Bylaws  of  The  International  Society  of  Hymenopterists    294 


Heckman 


BINDERY,         INC 
.,-  Boimd-ToPleise' 


OCT  01 

N.  MANCHESTER.  INDIANA  46962