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Qt(Q 

GZOO 


HARVARD  UNIVERSITY 


LIBRARY 

OF  THE 


Museum  of  Comparative  Zoology 


E.D.-  Q<yi4 


Quaestiones 


6? 


entomologicae 


COMP-  ZOOU. 

’ LIBRARY 


197?- 


harvard 

UNW&R^iT^ 


A periodical  record  of  entomological  investigations, 
published  at  the  Department  of  Entomology, 
University  of  Alberta,  Edmonton,  Canada. 


VOLUME  VIM 


1972 


11 


CONTENTS 


Editorial  — One  Eye  911,  the  Pot 1 

Rosenberg  — A chironomid  (Diptera)  larva  attached  to  a libellulid  (Odonata)  larva  ....  3 

Gooding  — Digesthp-processes  of  haematophagous  insects  I.  A literature  review 5 

Craig  - Rapid  orientation  of  wax  embedded  specimens  61 

Book  review 63 

Announcement Y 64 

Editorial  — Dissection  of  Science 65 

Griffiths  — Studies  on  boreal  Agromyzidae  (Diptera).  I. 

Phytomyza  miners  on  Saxifragaceae 67 

Schaaf  — The  parasitoid  complex  of  Euxoa  ochrogaster  (Guenee) 

(Lepidoptera:  Noctuidae) 81 

Kevan  — Collembola  on  flowers  on  Banks  Island,  N.  W.  T 121 

Book  review 123 

Book  review 124 

Book  review 125 

Announcement 127 

Book  review 129 

Whitehead  — Classification,  phylogeny,  and  zoogeography  of 

Schizogenius  Putzeys  (Coleoptera:  Carabidae:  Scaritini) 131 

Announcements  349 

Evans  - A temperature  controlled  capacitance-type  actograph 

for  cryptozoan  arthropods 351 

Doyen  — Familial  and  subfamilial  classification  of  the  Tenebrionoidea  (Coleoptera) 

and  a revised  generic  classification  of  the  Coniontini  (Tenebrionidae) 357 

Griffiths  — Studies  on  boreal  Agromyzidae  (Diptera).  II.  Phytomyza  miners 

on  Senecio,  Petasites  and  Tussilago  (Compositae,  Senecioneae) 377 

Announcement 406 

Supplement  — Proceedings  of  a Symposium  organized  by  the  Department  of  Entomology, 
University  of  Alberta  on  the  Occasion  of  the  50th  Anniversary  of  its  Foundation 


INDEX 


Abasa,  R.  O.,  (see  Langley,  P.  A.) 

6,  10,  42,55 
Acanthia  lectularia , 56 
Acanthocheilonema  perstans,  54 
Actebia  fennica,  91,  97,  98 
Aculichneumon,  118 
Adenostyles,  382 
Adephaga,  345 
Adesmia,  361,  370 
Adesmiini,  357,  360,  364,  371,  376 
Adler,  S.,  10,  11,  14,  17,27,48 
Aedes,  25,  30,  45 

aegypti,  6,  7,  9,  11,  12,  14,  17,  18,  19, 
20,21,23,24,  25,26,  29,30,31, 
33,  34,  35,  36,  37,  38,  40,  41,  42, 
43,44,  45,46,  48,49,50,51,52, 
53,54,  56,57,58,60,  124,  125 
africanus,  124 
albimanus,  1 1 
albopictus,  7,  44,  45 
argenteus,  1 1 
atropalpus,  7,  41,  54 
australis,  30,  33 
calopus,  14 
cantans,  6,  9,  20 
cell  lines,  125 
cinereus,  6,  9,  20,  44 
detritus,  6,  9,  14,  20 
dorsalis,  20 
geniculatus,  20 
hexodontus,  9,  29,  49 
infirmatus,  9 
notoscriptus,  30,  33 
punctor,  6,  9,  20 
quadrimaculatus , 7 
rusticus,  14 
scutellarus,  30,  33 
simpsoni,  124 
sollicitans,  7,  9,  45 
(Stegomyia)  aegypti,  49 
sticticus,  9 
stimulans,  18 
sub  alba  t us,  1 
taeniorhynchus,  7,  9 
togoi,  23 
trichurus , 30 
triseriatus,  7,  9 
vexans,  1,9,  14 
Aeolothripidae,  123 


Aeolothripinae,  123 
Aeolothripini,  123 
Aeolothripoidea,  123 
Agamermis,  83,  112,  113,  115,  116 
Agromyza  reptans,  76 
rufipes,  76 
Agromyzid,  394 
Agromyzidae,  76,  395,  396 
boreal,  67-75,377-394 
larval  host-plants,  67,  70 
Agroperina  dubitans , 95 
Agrotis  gladiaria,  104,  107 

orthogonia,  84,  85,  87,  90,  94,  102,  104, 
107,  110 
verier abilis,  1 10 
ypsilon,  90,  107 
Akidini,  376 

Akov,  S.,  19,  23,  29,  31,  36,  40,  41,  48 
Alaephus,  36 1 

Alleculidae,  358,  359,  360,  361, 362,  375, 
376 

Alleculinae,  362 
Allen,  H.  W.,  87,  93,  117 
Allen,  J.  R.,  (see  Gosbee,  J.)  17,  52 
allopatric  populations,  139 
Alphitobius  piceus,  353,  355 
Alysiinae,  394 
Amathes  smithi,  1 10 
Ambly teles  subfuscus,  98 
amphibius  subgroup,  320,  340 
Amphipods,  353 

Anderson,  J.  R.,  6,  10,  22,  32,  49 
Andropolia,  91 
contacta,  91 
vancouvera,  91 
Anectus,  373,  374 
vestitus,  373 
Anisopygys,  1 18 

Anopheles,  18,  20,  24,  45,  49,  55,  56,  58 
aconitus,  20 
albimanus,  7,  9,  54 
aquasalis,  30 
aztecus,  1 
bifurcatus,  44 
claviger,  15,  20 
crucians,  15 
culicifacies,  20 
freeborni,  7,  17,  43 
funestus , 18,  19,  53 


IV 


Anopheles  (continued) 

gambiae,  18,  19,  23,  24,  25,  53,  124 
jamesii , 15 
labranchiae,  46 

labranchiae  atroparvus,  15,  23,  24, 

25,  29 
litoralis,  9 
ludlowae,  9,  21 
maculatus , 9,  15 

maculipennis , 11,  13,  15,  19,  20,  24, 
29,  33,  38,  39,  41,  43,  44,  45,  49, 
50,  53,  56,  58,  60 
maculipennis  atroparvus,  23 
maculipennis  maculipennis , 15 
maculipennis  messeae,  15 
minimus , 9 
plumbeus,  13,  15 
punctipennis,  15 
punctulatus,  19 

quadrimaculatus,  7,  9,  13,  15,  17, 

20,  21,  31,  33,  45,  46,  54,  56 
rossi,  15 

sacharovi,  29,  43 

stephensi,  15,  23,  24,  25,  26,  43,  44, 
52 

subpictus,  15 
superpictus,  29,  43 
tarsimaculatus,  11,  57 
vagus,  18 

Anophelines,  50,  57 
ants,  129 

Antherea  eucalypti,  125 
Antro forceps,  345 
bolivari,  196,  197 
Apanteles,  102,  103 

acronyctae,  102,  103-104,  112,  113 
griffini,  102,  104,  108,  113,  116 
laeviceps,  102,  105,  113,  114,  115, 
116 

Apaulina  avium,  17 
Aphaniptera,  6 
Aphids,  126 
Aphroteniinae,  345 
Apis  mellifera,  49 
aquatic  invertibrates,  4 
Arachnids,  51 

Ar chips  argyrospilus,  102,  119 
arechavaletae  group,  171,  192-194,  314, 
317,  323 


arechavaletae  (continued) 
lineage,  314,  323 

-truquii-capitalis  lineage,  311,  314,  315 
arimao  subgroup,  321,  339,  340 
Armigeres  subalbatus,  9,31 
Army  worm,  117,  120 
wheat-head,  120 
Arnal,  A.,  21,  26,  49 

Arnaud,  P.  H.,  (see  Sabrosky,  C.  W.)  84,  87, 
90,  119 

Arnett,  R.  H.,  358,  361,  374 
Artemisia,  381 

Arthropod,  5,  44,  59,  130,  375,  376 
cell  cultures,  125-127 
cryptozoan,  351-353 
hosts,  55 
tissue,  126 
Asidinae,  358,  362 
Asidini,  358,  360,  364,  375 
Atkinson,  N.  J.,  (see  King,  K.  M.)  82,  83, 

84,  85,  87,  90,  91,  94,  95,  98,  101,  102, 
107,  110,  114,  118 

Azambuja,  C.  E.  A.,  (see  Rachou,  R.  G.)  9, 
57 

Baart,  E.  E.,  (see  Grobbelaar,  J.  H.)  351, 
352,  353 

Bacillus  pestis,  49 
Bacot,  A.  W .,  7,  44,  49 
Bailey,  L.,  12,  49 
Baker,  A.  W.,  91,  117 
Ball,  G.  E.,  287,  307,  311,  331,  335,  345 
Baptist,  B.  A.,  13,  14,  17,  49 
Barlow,  C.  A.,  9,  49 
Barr,  T.  C.,  197,  345 
Barrington,  E.  J.  W.,  5,  49 
basalis  group,  165,  166,  170,  184-186,  312, 
315,  317,  322,  323 
lineage,  314,  315,  323 
-truquii  lineage,  315 
Bastide,  P.,  (see  Combre,  A.)  41,  50 
Bates,  H.  W„  133,  158,  173,  182,  191,  192, 
204,  216,  231,  232,  254,  256,  345 
Bates,  M.,  11,  21,  29,  43,  46,  49 
Beatty,  H.  A.  (see  O’Connor,  F.  W.)  6,  9,  56 
Becla,  F.  (see  Kryriski,  S.)  6,  8,  23,  54 
bee,  129 
fly,  1 17 

Beesley,  W.  N.,  (see  Kershaw,  W.  E.)  54 
Beiger,  M„  381,  383,  387,  394 


V 


Belicek,  J.,  63-64 
Bell,  R.  T.,  143,345 
Bembidiine,  335 
Bennett,  G.  F.,  6,  9,  10,  49 
Berecyntus,  107 
bakeri,  107 

bakeri  var.  arizonensis,  107 
bakeri  var.  bakeri,  107 
bakeri  var.  euxoae,  1 07 
bakeri  var.  gemma,  107,  118,  119 
Berg,  V.  L.,  93,  117 
Bertram,  D.  S.,  23,  24,  49 
Beytout,  D.,  (see  Combre,  A.)  41,  50 
Bier,  M.,  (see  Buck,  F.  F.)  36,  50 
Bird,  R.  C.  (see  Bertram,  D.  S.)  23,  24,  49 
Bishop,  A.,  33,  45,  49 
Bishop,  F.  C.,  22,  49 
Blackflies,  17,  31,  38,  47,  51,  52,  60 
Blackwelder,  R.  E.,  141,  302,  306,  345 
Blaisdell,  F.  E.,  359,  361,  370,  373,  374 
Blaps,  359 
Blaptini,  359,  360 
Blatchley,  W.  S.,  141, 345 
blood-sucking  insects,  47,  48,  50,  52 
digestive  physiology  of,  5-48 
Blowfly,  5 1 
larvae,  36 

Blumberg,  D.  R.,  358,  374 
Bock,W.J.,  335,345 
Bohart,  G.  E.,  93,  117 
Boissezon,  P.,  26,  49 
Bombardier  beetles,  346 
Bombyliid,  93,  112,  116 
immature,  93 
parasite,  1 17 
pupae,  1 17 

Bombyliidae,  93-95,  1 13,  1 17,  1 18,  1 19 
Bombyx  mori,  126 

Bonnetia  comta,  87-90,  92,  94,  113,  116 
Boorman,  J.  P.  T.,  6,  9,  12,  18,  49 
Boros,  358 

Boving,  A.  G.,  361,  374 
Bowman,  L.,  (see  Hudson,  A.)  18,  54 
Brachinida,  302,  305,  308,  346 
Brachinus,  131,  132,  307,  308,  309, 

328,  329,  330,  331,  334 
conformis,  308 
cyanipennis,  308 
medius,  308 


Brachinus  (continued) 
oaxacensis,  308 
ovipennis,  308 
patruelis,  308 
tenuicollis,  308 
Brachycera,  6 
brachypterous,  143 

Braconidae,  102-107,  1 13,  1 17,  1 19,  394 
Branchini,  361,  362,  363,  364,  367 
classification  of,  370-374 
Bronchus,  363,  369,  373,  374 
floridanus,  363,  364,  365,  366,  369,  373 
woodii,  363,  364 
Breigel,  H.,  33,  49 

brevisetosus  group,  168,  206-208,  302,  31 1, 
317,318,320,  341 
Brimley,  C.  S.,  141,  345 
Brooks,  A.  R.,  84,  87,93,94,  95,  112,  117 
Brown,  K.  W.,  358,  375 
Brown,  W.  L.  Jr.,  287,  292,  336,  345 
Brundin,  L.,  302,  303,  304,  345 
Buck,  A.  de,  13,  15,  16,  19,  38,  39,  45,  49, 

50 

Buck,  F.  F.,  36,  50 
bugs,  14 

blood-sucking,  7 
Buhr,  H.,  383,387,392,394 
Bull,  C.  G.,  29,  50 
Bursell,  E.,  34,  50 

(see  Rajagopal,  P.  K.)  7,  57 
Biittiker,  W.,  20,  50 
Buxton,  P.  A.,  5,  6,  8,  10,  50 
(see  Weitz,  B.)  28,  29,31,32,59 
Cabasso,  V.  22,  50 

Caires,  P.  F.,  de,  (see  Micks,  D.  W.)  33,  56 
Calliphora  erythrocephala,  5 1 
Caly corny za,  378 
Camin,  J.H.,  141,315,345 
Campoletis,  100 

atkinsoni,  98-100,  105,  113,  114,  115,  116 
Campos,  M.,  (see  Freitas,  J.  R.)  34,  52 
Capek,  M.,  102,  117 

capitalis  group,  171,  198,  302,  31  1,  317,  323, 
327 

lineage,  311,  314,  315,  322,  323 
Caprifoliaceae,  69 
Carabici,  348 
carabid  beetles,  139,  353 
Carabidae,  1 18,  345,  346,  347,  348 


V 


Carabinae,  346 

carinatus  group,  170,  189-191,  317,  323 
lineage,  323 

Cartwright,  E.,  (see  Friend,  W.  G.)  7,  52 
Casey,  T.  L.,  358,  361,  362,  370,  371, 
373, 375 

Castelnau  (de  Laporte),  F.  L.  N.  C.,  133, 
345 

Cediopsylla  inaequalis  inaequalis,  22 
Centronopus,  359 
Ceratophyllus  fasciatus,  28,  56 
Ceratopogonidae,  10,  52,  54,  55,  56,  58 
cesium  tagging,  6 
Chalcidoidea,  1 1 9 
Chamberlain,  R.  W.,  44,  50 
Champion,  G.  C.,  362,  373,  375 
Champlain,  R.  A.,  11,  17,  42,  50 
Chao,  J.,  21,  50 

Chapman,  H.  C.,  (see  Woodard,  D.  B.)  9, 

59 

Chapman,  R.  F.,  109,  117 
Chasmias,  1 1 8 

Chellappah,  W.  T.,  (see  Zaman,  V.)  31, 

60 

Chilopods,  353 
Chironomid  larva,  3,  4 
midges,  345 
Chironomidae,  4 
Chirothripini,  123 
Chorizagrotis,  1 10 

auxiliaris,  95,  98,  104,  107,  1 10 
thanatologia,  95,  1 10 
Choy,  C.  T.  H.,  (see  Friend,  W.  G.)  7,  52 
Christophers,  Sir  S.  R.,  5,  1 1,  50 
Chromatomyia,  69 
Chrysanthemum  leucanthemum , 392 
Chrysops,  11,  1 7,  54 
dimidiata,  10,  54 
silacea,  10,  28,  50,  52,  54,  59 
Cibdelis  blaschkei , 359 
Cicindela,  348 

duodecimguttata , 346 
maritima,  346 
oregona,  346 
Cicindelidae,  345 
Cicindelinae,  347 
Cimex , 33 

hemipterus,  8,  14,  59 
lectularius,  6,  8,  14,  17,  23,  32,  35, 


Cimex  (continued) 

36,44,  49,54,57,58 
rotundatus , 14 
Cirphis , 91 
Cistelidae,  362 
Cistelides,  362 
cladistic  classifications,  306 
Clausen,  C.  P.,93,  94,  117 
Clements,  A.  N.,  5,  50 
Cleptoria,  302,  309,  342,  346 
abbotti,  309 
bipraesidens,  309 
divergens,  309 
macra,  309 
rileyi , 309 
Clivina , 347,  348 
amphibia , 236 
frontalis , 236 
lineolata,  246 
sulcata,  179 
Clivinina,  347 

Cnephia  dacotensis,  36,  61,  62 
ornithophilia,  27 
cockroach,  125,  353 
american,  55 

Coelini,  357,  360,  361, 362,  363,  364,  367, 
370 

classification  of,  370-374 
Coelometopini,  360,  361 
Coelomorpha,  369,  371, 373 
maritima,  363,  369,  372,  373 
Coelosattus,  370,  371,  373 
fortineri,  363,  368,  369,  372,  373 
Coelotaxis,  370,  373 
punctulata,  363,  369,  372,  373 
Coelus,  363,  369,  371,  373,  374 
ciliaris,  369 
ciliatus,  363,  373 
globosus,  363,  372 
re  mot  us,  363 

Coleoptera,  93,  118,  131-344,  345,346, 
347,  348,  357,  367,  369,  374,  375, 

376 

Collembola,  121 
Combre,  A.,  41 , 50 
Compositae,  377,  390 
Coniontellus,  370,  373 
inflatus,  363 
obesa,  369 


Vll 


Coniontides,  370,  373 
latus,  362,  369,  372 
Coniontinae,  362,  375 
Coniontini,  376 

generic  classification  of,  357 -37 4 
Coniontis , 363,  370,  371,  373,  374 
hoppingi,  372 
lata , 313 
obesa , 373 

viatica,  362,  369,  373 
Conipinus,  373 
Conisattus,  370,  373 
rectus,  373 
Conistra  devia,  91 
Cook,  W.  C.,  107,  110,  117 
Copablepharon  viridisparsa,  90 
Copidodoma  bakeri,  82,  107-1 12,  113, 
114,  115,  116 
Copidosoma  gelechiae,  1 1 8 
Corbet,  P.  S.,  4 

Comet,  M.,  (see  Mattern,  P.)  30,  56 
Cornwall,  J.  W.,  13,  14,  15,  16,  17,  39, 

50 

Corynothrix  borealis,  1 2 1 
Cossyphini,  359,  361 
Cragg,  F.  W.,  27,  28,  50 
Craig,  D.  A.,  61-62 

Craighead,  F.  C.,  (see  Boving,  A.  G.)  361, 
374 

Cratichneumon,  118 
crenulatus  group,  131,  132,  148,  149- 
153,  165,312,317,322,323,327, 
337 

lineage,  312,  323 

-quinquesulcatus-tenuis  lineage,  314 
Crewe,  W.,  28,  50,  (see  Gordon,  R.  M.) 

10,  52,  (see  Kershaw,  W.  E.)  54 
Crosskey,  R.  W.,  10,  50,  (see  Lewis,  D.  J.) 
50 

Crovello,  T.  J.,  (see  Sokal,  R.  R.)  302, 
306,  348 

Crowson,  R.  A.,  302,  303,  346,  357, 

361,  375 
Cmciferae,  121 
Cry  modes  devastator,  1 10 
Cryptoglossini,  360,  364 
Csiki,  E.,  141,265,346 
Ctenichneumon,  118 
Ctenophthalmus,  28 


Culex,  45 

fatigans,  18,  52,  54,  56 
pipiens,  7,  11,  13,  15,  17,21,23,  26,31, 
33,39,  43,45,49,54 
pipiens  fatigans , 57,  124 
pipiens  molestus,  30 
pipiens  pallens,  7,  12 
pipiens  quinquefasciatus,  6,  7,  9,  15,  18, 

20,  21,  29,  30,  31, 33,  34,  35,  36,  37, 

39,  40,  46,  48 
restuans,  16 
salinarius,  9,  16 
tarsalis,  21,30,  50,  124 
Culicidae,  49,51,52,54,58 
Culicoides  impunctatus,  21,  58 
nubeculosus,  11,  1 8,  26,  3 1 , 56 
obsoletus,  21, 26,  54,  58 
Culiseta  annulata,  9,  13,  16,  17,  20,  39,  58 
inornata,  9,  56,  124 
Cushman,  R.  A.,  101,  117 
Cutworm,  82,  83,  84,  87,  89,  91,  97,  98,  100, 
103,  104,  107,  117,  118,  119 
army,  1 19 
black  army,  9 1 
climbing,  114,  120 
ground,  81,  114 
larvae,  95,  97,  106 
pale  western,  1 1 9 
prairie,  106,  119 

red-backed,  81,82-83,  112,  114,  118 
western,  1 1 7 
Cychrini,  345 
Cyclorrhapha,  57,  76 
Dacoderidae,  359,  376 
Dacoderus,  358 
Darkling  beetle,  375 
Darling,  S.  T.,  20,  45,50 
Darlington,  P.  J.,  Jr.,  133,  264,  302,  303, 

304,  326,  346 

darlingtoni  group,  171,  196,  197,  302,  311, 
322,  327 

Dasgupta,  B.,  24,  50 
Dasybasis  froggatti,  29 
Davies,  D.  M.,  36,  39,  41,  43,  50,  (see  Yang, 
Y.  J.)  11,  17,  35,  36,  38,  39,  40,  41,  43,  60 
Davies,  J.  B.,  (see  Lewis,  D.  J.)  4 
Davis,  G.  E.,  23,29,51 
Davis,  W.  A.,  23,33,  51 
Day,  M.  F.,  5,  11,  12,  43,  5 1 , (see 


Vlll 


Day,  M.  F.  (continued) 

Waterhouse,  D.  F.)  5,  59 
Deegan,  T.,  (see  Kershaw,  W.  E.)  6,  10, 

54 

DeFoliart,  G.  R.,  (see  Anderson,  J.  R.) 

10,  49 

Dendrothripini,  123 
Denisova,  Z.  M.,  12,  51 
depressus  group,  135,  169,  171, 263-301, 
302,  31 1,  317,  318,  320,  321,  327, 
338,  339,  340,  344 
subgroup,  321,  340 

Detinova,  T.  S.,  5,  18,  19,  20,  23,41,51 

Deutzia , 67 

Devine,  T.  L.,  13,  51 

diala t us  subgroup,  320 

Diamanus  mo nt anus,  22 

Diamond-back  moth,  100 

Diaperini,  36 1 

Diphyus,  95,  97,  98,  99,  1 13,  1 14,  1 15, 
116 

Diplopods,  353 
Diprionidae,  1 1 8 

Diptera,  3,  4,  6,  46,  49,  50,  5 1 , 52,  54, 
55,  56,  57,  58,  59,  60,  61, 67,  69, 
76,83,93,  117,  118,  119,  120, 

377,  394,  395,  396 
cyclorrhaphous,  68 
Dirofilaria  immitus,  45,  46,  54,  60 
Discodemus,  373 
Dominick,  R.  B.,  63 
Doronicum  clusii,  362 
Doutt,  R.  L.,  81,  117 
Downe,  A.  E.  R.,  30,  31, 32,  51 
Doyen,  J.  T.  357-374,  375 
Drosophila , 125,  126 
cell  culture,  1 25 
Duncan,  J.  T.,  44,  5 1 
Dutky,  S.  R.,  (see  Schechter,  M.  S.)  351, 
353 

Ectopimorpha , 1 1 8 
Edman,  J . D. , 31,51 
Edrotes,  367,  375 
Edwards,  C.  R.,  63 
Ehrlich,  P.  R.,  302,  306,  346 
Eisner,  T„  358,  359,  375 
Elaphria  nucicolora , 91 
El e odes,  359 
longicollis , 375 


Eleodes  (continued) 
obsoleta,  31 A 
Eleodini,  359,  360,371 
Eligh,  G.  S.,  (see  West,  A.  S.)  5,  29,  59 
Ellipsoptera,  348 
elongatus-carinatus  lineage,  314 
elongatus  group,  170,  186-189,  317,  323 
lineage,  314,  323 
Emden,  F.  I.  van,  360,  375 
Encyrtidae,  107-112,  113 
Enright,  J.  T.,  353 
Entomobrya  comparata,  121 
Entomobryiidae,  121 
Entomochilus , 367,  370 
varius  laevis , 363 
entomophagous  groups,  81 
species,  81 
Epiphysa,  367,  370 
Epitragini,  359,  360 

Eppley,  R.  K.,  (see  Bohart,  G.  E.)  93,  117 
Erigorgus,  101,  102 
Erodiini,  360 
Erotidothripinae,  123 

Erwin,  T.  L.,  140,  307,  326,  328,  329,  330, 
331, 334,  335,  346,  (see  Ball,  G.  E.)  31 1, 
345 

Eschscholtz,  J.  F.,  373,  375 
Eucirrhoidea  pampina,  104 
Eupalamus , 1 18 
Eupsophulus , 361 
Eurychorini,  376 
Euryderus,  345 
Eusatti,  370 
Eusattodes,  373 

Eusattus , 363,  369,  370,  371,  373,  374,  376 
ciliatus,  370 
difficilis,  373 

dubius , 363,  366,  369,  370,  373 
erosus,  363,  369,  373 
laevis,  373 

muricatus , 363,  365,  369,  370,  371,  372, 
373 

puberulus,  370 

reticulatus,  362,  363,  366,  369,  372 
robustus,  363,  369,  373 
Eutanyacra , 1 1 8 

suturalis,  95,  97,  98,  99,  1 13,  1 15,  1 16 
Eutriatoma,  33 
maculatus , 53 


IX 


Euxoa,  94,  110,  118 
auxiliaris,  91,  119 
campestris,  1 1 2 
( chorizagrotis)  auxiliaris,  1 1 9 
dargo,  1 12 
detersa,  1 1 0 
divergens,  1 1 2 
excellans,  98 

flavicollis , 94,  95,  97,  98,  1 10 
intrita,  1 1 0 

messoria,  90,  98,  107,  1 10 
ochrogaster,  1 1 8 

the  parasitoid  complex  of,  81-117 
scandens,  95,  98,  1 10 
tesselata,  94,  95,  1 12 
tristicula,  90,  107,  110 
verticalis,  1 1 2 
Evans,  W.  A.  L.,  36,  51 
Evans,  W.  G.,  351-355 
Evarthrus,  131,  132,  302,  305,  307,  308, 
309,  328,  329,  330,  334,  346 
gravesi,  308 
hypherpiformis,  308 
Exephanes , 1 1 8 
Exopterygotes,  6 
eye  gnats,  1 1,  54 

Fall,  H.  C.,  133,  209,  254,  268,  346 
Fallis,A.  M.,  5,27,  51 
Fattig,  P.  W.,  141,239,  346 
Feltia  ducens,  94,  95,  98,  104,  107,  110, 
112 

subgothica,  107,  110 
Feng,  L-C.,  26,  51 

Ferguson,  M.  J.,  (see  Micks,  D.  M.)  21, 
46,  56 

Ferguson,  R.  B.,  63 

Ferreira  Neto,  J.  A.,  (see  Rachou,  R.  G.) 
9,  57 

ferrugineus  group,  131,  132,  167,  179- 
184,  312,  317,  322,  323,  327,  337, 
340 

lineage,  312,  314,  315,  323,  341 
filariasis,  59,  60 

Finlayson,  T.,  83,  95,  97,  99,  1 18 
Fisk,  F.  W.,  12,  26,  29,  33,  35,  36,  37, 
40,  4 1 , 42,  5 1 , (see  Patterson,  R.  A.) 
35,  57,  (see  Champlain,  R.  A.)  11,  17, 
42,  50 

fleas,  7,  22,  28,  33,  44,  53,  57 


Fletcher,  J.,  107,  1 18 
Flies,  7,  1 1,38,  42,378 
blood  meal  size,  10 
mining,  395 
muscoid,  1 18 
tachinid,  1 19 
Florence,  L.,  22,  .5 1 
Forbes,  63 
Foulk,  J.  D.,  10,  52 

Fraenkel,  G.,  (see  Galun,  R.)  43,  52,  (see 
Lipke,  H.)  5,  55 
Franclemont,  J.  G.,  63 
Franco,  L.  B.,  (see  Micks,  D.  W.)  33,  56 
Frank,  J.  H.,  83,  114,  118 
Franklinothripini,  123 
Freitag,  R.,  137,  308,  328,  329,  330,  331, 

335,  346,  (see  Lindroth,  C.  H.)  246,  347 
Freitas,  J.  R.,  6,  7,  9,  34,  52 
Frey,  R„  383,  394 

Freyvogel,  T.  A.,  17,  23,  24,  25,  43,  52,  (see 
Staubli,  W.)  23,  24,  58 
Frick,  K.  E.,  389,  394 
Friend,  W.  G.,  7,  8,  52 
Fuscigonia,  87 
(fuscicollis),  84 
Galum,  R.,  43,  52 

Gamal-Eddin,  F.  M.,  (see  Rostom,  Z.  M.  F.) 
17,57 

Gander,  E.,  23,  25,  52 
Garnham,  P.  C.  C.,  9,  52 
Gasterophilus  intestinalis,  16,  17,  34,  40,  58 
Gastrophilus  equi,  57 
Gebien,  H.,  358,  361, 362,  373,  375 
Gemminger,  M.,  141, 265,  346 
Genioschizus,  131,  132,  144,  148,  165,312, 
314,  316,  323,  337 
Gentianaceae,  69 
Gibson,  A.,  98,  107,  110,  118 
Gillett,  J.  D.,  124 
Gilmour,  D.,  5,  52 
Girault,  A.  A.,  110,  118 
Glossina,  11,  17,  1 8,  53,  60 
austeni,  6,  10,  17,  39,  42,  48,  53,  55 
brevipalpis , 10,  56 

morsitans , 7,  10,  11,  13,  16,  28,  3 1,  34,  38, 
39,42,  50,  54,  55 
morsitans  submorsitans,  1 7 
pallidipes,  59 
palpalis,  6,  10,  28,  47,  56 


X 


Glossina  (continued)  * 
submorsitans,  28,  34,  50 
swynnertoni , 22,  31,  59 
tachinoides,  6,  10,  11,  13,  16,  17,  18, 
28,  38,  50,  60 
Glugea  disstriae , 126 
Gnathocephalon,  1 1 9 
Goatly,  K.  D.,  (see  Jordan,  P.)  9,  54 
Gonia,  84-85,  94,  111,  112,  116,  117, 

119 

aldrichi,  85-86,  87,88,  113,  114,  115, 
116 

capitata,  84,  87,  112,  113 
( capitata,  sequax),  84 
fuscicollis,  87,  112,  113,  116 
sequax,  87,  88,  112,  113 
Goniderini,  360 
Goodchild,  A.  J.  P.,  8,  52 
Gooding,  R.  H.,  5-48,  52 
Gordon,  R.  M.,  10,  14,52 
Goring,  N.  L.,  (see  Downe,  A.  E.  R.)  30, 
51 

Gosbee,  J.,  17,  52 
Graham,  A.  R.,  82,  100,  106,  1 18 
Grapholitha,  91 
grasshopper,  egg  pods,  1 17 
grassworm,  1 17 
Gravenhorstia,  102 
propingua,  101-102,  105,  111,  113, 
116 

Greene,  C.  T.,  83,  87,  90,  118 
Griffiths,  G.  C.  D.,  67-75,  76,  377-394, 
395 

Griswold,  C.  L.,  (see  Schaffner,  J.  V.) 

104,  119 

Grobbelaar,  J.  H.,  351,  352,  353 
Groschke,  F.,  384,  386,  394 
ground  beetles,  346,  347 
Grusz,  F.,  14,  39,  52 

Guardia,  V.  M.,  (see  Zeledon,  R.)  6,  8,  60 
Guedes,  A.  da  Silverra,  (see  Freitas,  J.  R.) 
6,  7,  9,52 

Guelmino,  D.  J.,  20,  53 
guinea  pig,  23,  50,  54,  55 
Guppy,  J.C.,  91,  104,  118 
Gupta,  V.  K.,  (see  Townes,  H.,  101,  102, 
119 

Guptavanij,  P.,  1 1,  53 
Gwadz,  R.  W.,  7,  9,  53 


Gymnopais,  61, 62 
Gyriosomus,  364,  367,  370 
modestus,  363 
Haematophagous  insects 

changes  in  gut  contents,  18-22 
digestive  enzymes  and  their  properties, 
35-40 

digestive  processes,  5-48 
distribution  of  meals  within  the  alimentary 
canal,  7-12 

enzyme  content  of  the  gut,  40-43 
histological  changes  in  the  gut  and  blood 
meal,  22-29 

relationship  of  digestive  processes  to 
vectoring  ability,  43-47 
salivary  glands  and  their  secretions,  12-18 
serological  and  chemical  analysis  of  gut 
contents  during  digestion  of  the  blood 
meal,  29-34 
size  of  blood  meal,  6-7 
Haematopinus  suis,  22 
Haldeman,  S.  S.,  133,  236,  346 
Halffter,G.,  327,  335,346 
Hall,  J.  C.,  (see  Painter,  R.  H.)  93,  94,  95,  1 19 
Halocoryza,  133,  140,  143,  144,  196,303, 
326,  335,348 
acapulcana,  145,  146 
arenaria,  144,  146,  348 
Hansens,  E.  J.,  (see  Davis,  W.  A.)  23,  33, 

51 

Hardwick,  D.  F.,  82,  118 
Hardy,  J.,  69,  76 

Harold,  E.  von,  (see  Gemminger,  M.)  141, 

265, 346 
Harpalini,  345 

Hatch,  M.  H.,  268,  269,  287,  294,  346 
Hawking,  F.,  (see  Yorke,  W.)  28,  60 
Hawkins,  R.  I.,  17,  39,  53,  (see  Hellmann,  K.) 

13,  14,  38,39,53 
Hayashi,  N.,  360,  375 
Hays,  K.  L.,  8,  53 
Heinrich,  G.  H.,  98,  118 
Heleidae,  56 
Helerothripidae,  123 
Heliophila  commoides,  91 
Heliothripinae,  123 
Hellmann,  K.,  13,  14,  38,  39,  53,  (see 
Hawkins,  R.  I.)  39,  53 
hemimetabolous  insects,  blood  meal  size,  8 


XI 


Heming,  B.  S.,  123,  127 
Hemipenthes,  93,  94 
Hemiptera,  6,  49,  52,  53,  57,  59 
Hendel,  F.,  72,  73,  76,  380,  381,  382, 
383,386,  391,392,  395 
Hennig,  W.,  140,  141,  302,  303,  304, 
305,  306,  346 
Heptagyinae,  345 

Hering,  E.  M.,  72,  73,  76,  387,  395,  (see 
Groschke,  F.)  384,  386,  390,  394 
Hering,  M.,  73,  76,  382,  383,  388,  395 
Herndon,  B.  L.,  31,  53 
Hershkovitz,  P.,  326,  327,  346 
Heterobionta,  360 
Heteromera,  376 
Heteroptera,  49 
Heterostylum  robustum,  1 17 
Heterotarsini,  360 
Heterothripini,  123 
Hewitt,  C.  G.,  22,  53 
Hippelates  pallipes,  1 1,  44,  54 
Hippobosca,  16 
Hippoboscidae,  29 
Hirudo  medicinalis,  53 
Hoare,  C.  A.,  28,  47,  53 
Hocking,  B.,  2,  65-66 
Hocking,  K.  S.,  18,  19,  53 
Hodges,  R.  W.,  63 
Hoffman,  R.  L.,  309,  342,  346 
Holdenried,  R.,  22,  53 
Holoubek,  K.,  (see  Schildknecht,  H.) 
359,  376 

Holstein,  M.,  32,  53 
Homoptera,  cell  culture,  125 
Homotherus,  1 18 
Honeybee,  49 
Hopkins,  D.  M.,  335,  347 
Horn,  G.  H.,  358,  361,  371,  373,  375, 
(see  LeConte,  J.  L.)  358,  36 1 , 362, 
371,  376 
horse  bot-fly,  58 
horse  flies,  12,  16,  51,  58 
Hosoi,  T.,  7,  12,  53 
house  fly,  36,  55 
House,  H.  L.,  5,  53 
Howard,  L.  M.,  9,  12,  18,  23,  24,  53 
Hoyer’s  medium,  135 
Huang,  C.  T.,  35,  36,  37,  38,  53 
Hubbs,  C.,  (see  Hubbs,  C.  L.)  136,  347 


Hubbs,  C.  L\,  136,347 

Hudson,  A.,  18,  41,  54,  (see  Orr,  C.  W.  M.) 

17,  56 

Hudson,  J.  E.,  124 
Huff,  C.  G.,  26,45,54 
Hull,  D.  L.,  302,  304,  305,  306,  347 
Hulten,  E.,67,  75,76,  378,  395 
Hunter,  T.  A.,  (see  Freyvogel,  T.  A.)  17,  43, 
52 

Hybomitra  affinis,  10 
frontalis,  10 
Hydrangea,  61 
Hydrangeaceae,  67 
Hydrocanthari,  348 

Hymenoptera,  93,  1 17,  1 18,  1 19,  120,394 
Hynes,  H.  B.  N.,  93,  118 
Hyperalonia  oenomaus,  117 
Hyponomer,  396 
Ichneumonid  parasitoids,  94 
Ichneumonidae,  95-102,  1 13,  1 17,  1 18,  1 19 
Ichneumonines,  1 1 6 
Ichneumonini,  95,  118 
Insect  Societies,  129 
Intertidal  beetles,  351 
Iso  par  ce,  63 
Isopods,  353 
Italodytes  stammeri,  196 
jacarensis  group,  170,  171-173,  312,  317,  323 
lineage,  323 

-optimus  lineage,  3 12,  3 14,  3 1 5 
Jackson,  C.  H.  N„  18,54 
Jacobson,  L.  A.,  82,  1 18 
Jacot-Guillarmod,  C.  F.,  123 
Jamnback,  H.,  26,  54 
Jaquet,  C.,  (see  Freyvogel,  T.  A.)  25,  52 
Jeffery,  G.  M.,  9,  54 
Johnson,  C.  G.,  6,  8,  54 
Jones,  R.  M.,  8,  54 
Jordan,  P.,  9,  54 

Kalra,  N.  L.,  (see  Wattal,  B.  L.)  8,  59 

Kaltenbach,  J.  H„  386,  395 

Kamal,  A.  S.,  (see  Rockstein,  M.)  17,  57 

Kartman,  L.,  44,  45,  54 

Keleynokova,  S.  I.,  360,  375 

Kelley,  M.  H.,  (see  Schubert,  J.  H.)  29,  57 

Kendall,  D.  A.,  358,  375 

Kershaw,  W.  E.,  6,  10,  54 

Kevan,  D.  K.  McE.,  (see  Kevan,  P.  G.)  121 

Kevan,  P.  G.,  121 


Xll 


King,  K.  M.,  82,  83,  84,  85,  87,  90,  91, 
94,  95,  98,  101,  102,  106,  107,  1 10, 
114, 118 

King,  W.  V.,  (see  Bull,  C.  G.)  29,  50 
Kirsch,  T.,  133,  198,347 
Koch,  C.,  358,  36 1 , 362,  364,  375 
Korschevsky,  R.,  360,  376 
Kramer,  H.,  (see  Schildknecht,  H.)  359, 
376 

Krynski,  S.,  6,  8,  23,  54 
Kuchta,  A.,  (see  Krynski,  S.)  6,  8,  23,  54 
Kult,  K.,  133,  142,  155,  165,  185,  188, 
197,  201,  202,  231, 232,  254,  270, 
347 

Kumm,  H.  W.,  11,  44,  54 

Kurten,  B.,  335,  347 

Kutuza,  S.  B.,  (see  Moloo,  S.  K.)  10,  56 

Lacinopolia  renigera,  110,  112 

Lacordaire,  J.  T.,  358,  362,  371,  376 

Lagriidae,  358,  359,  360,  361,  362,  375 

Lagriides,  362 

Lagriinae,  362 

Landry,  S.,  130 

Langley,  P.  A.,  6,  10,  34,  42,  54,  55 
Laphygma  frugiperda,  1 17 
Larvaevoridae,  120 
Lascoria  ambigualis,  91 
Laurel,  A.  G.,  9,  55 
Lavoipierre,  M.  M.  J.,  44,  55 
Lawrence,  J.  F.,  358,  376 
Lea,  A.  O.,  7,  9,  41,55 
leafhoppers,  126 

LeConte,  J.  L.,  133,  142,  150,  221,  229, 
236,  238,  246,  287,  347,  358,  361, 
362,  371,  373,  376 

Lee,  C.  U.  (see  MacGregor,  M.  E.)  11,  55 
Leiby,  R.  W.,  109,  118 
Leiochrini,  361,  362 
Leiochrodes,  36 1 
Leishmania  donovani,  27,  44,  56 
tropica,  44,  56 
Leng,  C.  W.,  141,265,347 
Leonard,  M.  D.,  141, 347 
Lepidichora  discoidalis,  376 
Lepidochorini,  360 
Lepidoptera,  59,  81-1 17,  118,  119 
cell  culture,  125 
larvae,  120 
Leptoconops,  18 


Leptoconops  (continued) 

(Holoconops)  becquaerti,  21,55 
kerteszi,  10,  52 
Lesquerella  arctica , 1 2 1 
Lester,  H.  M.  O.,  10,  1 1,  13,  16,  18,  38,  55 
Leucocytozoon  simondi,  49 
Leucophaea  maderae,  1 25 
Lewis,  D.  J.,  4,  1 1, 27,  47,  (see  Buxton,  P.  A.) 
6,  10,  55 

Libellulid  larva,  3,  4 
lice,  7,  14 
human,  5 1 

Lima,  m.  m.,  (see  Rachou,  R.  G.)  9,  57 
Limerodops,  118 
Lin,  S.,  36,  55 

Lindroth,  C.  H.,  133,  140,  142,  150,  191, 

209,  221,  229,  236,  238,  246,  254,  265, 
266,  267,  268,  269,  277,  287,  294,  347 
lindrothi  group,  131,  132,  167,  171,  198, 
199-202,  302,  31 1, 322,  327,  337,  338 
Lindsay,  D.  W.,  310,  347 
lineolatus  group,  1 69,  246-25 1 , 302,  3 1 1 , 

317,  318,  321,  341 
Linley,  J.  R.,  18,  21,  55 
Linnaemyia  comta,  1 17 
Linnaniemi,  W.  M.,  72,  76,  383,  395 
Linsley,  E.  G.,  (see  Mayr,  E.)  348 
Lipke,  H.,  5,  55 
Liriomyza,  378 
Lithophane  innominata,  91 
Lloyd,  L.,  16,  17,  39,  55,  (see  Lester,  H.  M. 

O.)  10,  11,  13,  16,  18,38,55 
Lloyd,  R.  B.,  32,  55 
Loa  loa,  50,  52,  54 
Locust,  desert,  1 18 

Lofy,  M.  G.,  (see  Templis,  C.  F.)  30,  58 
longipennis  group,  168,  252-263,  265,  302, 

31  1,317,  318,  320,  321,  327,  338,  340, 
344 

Loricerini,  345 
Lotmar,  R.,  28,  55 
louse,  23,  50 
clothes,  54 
hog,  22,  50 

Loxostege  sticticalis,  104 
Lumsden,  W.  H.  R.,  (see  Gordon,  R.  M.)  1 4,  52 
Lyneborg,  L.,  (see  Ryden,  N.)  392,  395 
Macfie,  J.  W.  S.,  (see  Yorke,  W.)  13,  14,  15, 

16,  17,  60 


xiii 


MacGregor,  M.  E.,  11,  12,21,34,55 
Maclnnes,  D.  G.,  (see  Hocking,  K.  S.)  18, 
19,  53 

MacKerras,  M.  J.,  19,  55 
Macroevolutionary  models,  345 
Macrolepidoptera,  119 
Maddrell,  S.  H.  P.,  7,  55 
Malacosoma  americanum,  126 
disstria,  126 
malaria,  50,  56 
parasite,  56,  57 

Maldonado-Koerdell,  M.,  326,  328,  334, 
347 

Mansonia  perturbans,  9,  30 
richiardii,  6,  9,  16,  20 
Marcuzzi,  G.,  360,  376 
Mariani,  M.,  46,  56 
Martin,  J.  L.,  102,  118 
Martin,  P.  S.,  293,  347 
Masseyeff,  R.,  (see  Mattern,  P.)  30,  56 
Mattern,  P.,  30,  56 
Matthews,  J.  V.,  (see  Hopkins,  D.  M.) 
335,347 

Maxwell,  C.  W .,  (see  Wood,  G.  W.)  102, 
120 

Mayr,  E.,  136,  137,  138,  139,  140,  141, 
302,  305,  306,  336,  347,  348 
McConnachie,  E.  W.,  (see  Bishop,  A.)  33, 
45,49 

McHenry,  F.,  (see  Eisner,  T.)  358,  375 
McKiel,  J.  A.,  (see  Wood,  G.  W.)  102,  120 
McKinley,  E.  B.,  14,  17,  56 
McMillan,  E.,  82,  107,  109,  1 10,  1 14, 

115,  118 

Megahed,  M.  M.,  7,  1 1,  12,  18,  26,  56 
Megasattus,  373 

Mehringer,  P.  J.,  (see  Martin,  P.  S.)  293, 
347 

Meijere,  J.  C.  H.  de,  72,  73,  76,  378,  387, 
388,391,392,395 

Meinwald,  J.,  (see  Eisner,  T.)  359,  375 
Melanagromyza,  378 
Melanesian  ant  fauna,  348 
Melanthripinae,  123 
Meliana  albilinea,  104,  120 
Mellanby,  K.,  6,  7,  10,  56 
Melophagus  ovinus,  29 
Memoria,  J.  M.  P.  (see  Rachou,  R.  G.)  9, 
57 


Menees,  J.  H.,  83,  119 
Merothripidae,  123 
Merothripinae,  123 
Merothripoidea,  123 
Mesothripidae,  123 
Metachaeta,  90 
helymus,  90 

Metcalf,  R.  L.,  13,  14,  15,  16,  17,  56 
Meteorus,  106 

dimidiatus,  106,  107,  112,  113 
vulgaris , 103,  106-107,  108,  113,  114,  115, 
116 

Micks,  D.  M. ,2 1,46,  56 
Micks,  D.  W.,  33,  56 
Microfilaria  bancrafti,  56 
Microfilariae,  52,  54 
Microgastrinae,  102 
Microplitus,  102 

kewleyi , 102,  104-106,  107,  113, 

116 

mitii  group,  69 
Miller,  L.  A.,  10,56 
Mimulus,  310 
guttatus,  347 
Minchin,  E.  A.,  28,  56 
Miners,  69 

leaf,  67-75,377-394 
on  Saxifragaceae,  67-75 
Phytomyza,  377-394,  395 
primrose-leaf,  76 
Mirov,  N.  T.,  201,338,348 
Mitella,  67,  70 
nuda,  75,  80 
Mites,  82,  353 
Mitzmain,  N.  B.,  18,  56 
Moloo,  S.  K.,  10,  56 
Molurini,  376 

Monommidae,  358,  359,  361, 362 
Monophyletic  group,  303 
Moore,  P.  J.,  (see  Kershaw,  W.  E.)  6,  10,  54 
Moran,  V.  C.,  (see  Grobbelaar,  J.  H.)  351, 
352,  353 

Morris,  G.  J.,  (see  Grobbelaar,  J.  H.)  351, 

352,  353 

Morrison,  F.  O.,  84,  119 
Mosher,  63 

mosquitos,  5,  7,  14,  15,  16,  18,  24,  29,  48, 
49,  50,  52,  55,  56,  57,  58,  59,  60,  124 
blood  meal  size,  9 


XIV 


mosquitos  (continued) 

British,  58 
mouthparts  of,  52 
West  African,  5 1 
yellow  fever,  50 
Most,  H.,  (see  Yoeli,  M.)  46,  60 
Mouchet,  J.,  (see  Combre,  A.)  41,  50 
Muir,  F.,  (see  Sharp,  D.)  359,  376 
Muirhead-Thomson,  R.  C.,  18,  56 
Munroe,  E.  G.,  63 

Murgatroyd,  F.,  (see  Yorke,  W.)  28,  60 
Musca  convexifrons,  16 
crassirostris,  16,  17,  39 
domestica,  44,  52 
nebulo,  16 
pattoni,  16 
vitripennis,  17 
Muscidae,  10 
Muscoid,  6,  16 

Mutchler,  A.  J.,  (see  Leng,  C.  W.)  141, 
347 

Mymarothripinae,  123 

Myser,  W.  C.,  (see  Devine,  T.  L.)  13,  51 

Napier,  L.  E.,  (see  Lloyd,  R.  B.)  32,  55 

Neamblymorpha,  118 

Nearctic,  133 

Nematocera,  6 

Nematoda  Merinthidae,  113 

Nematodes,  44,  55 

Neobrachinus,  308 

Neodiprion  sertifer,  1 1 8 

Neotropical,  133 

Nesostes,  373 

Netelia,  101,  105,  113,  116 
Nielson,  B.  O.,  (see  Ryden,  N.)  392,  395 
Nielson,  W.  T.  A.,  (see  Wood,  G.  W.)  102, 
104,  107,  110,  120 
Nilio,  361 

Nilionidae,  358,  359,  360,  361, 362 

Nilionides,  362 

Nilioninae,  362 

No  card  ia  rhodnii,  39 

Noctuid,  93 

hosts,  94,  95,  100 
larvae,  1 14 

Noctuidae,  81-117,  118,  120 
Nord,  F.  F.,  (see  Buck,  F.  F.)  36,  50 
Nosopsyllus  fasciatus,  28,  33 
Nowakowski,  J.  T.,  68,  76,  380,  386, 


Nowakowski,  J.  T.  (continued) 

387, 395 

Nunberg,  M.,  387,  395 
Nuttall,  G.  H.  F.,  14,  15,  17,  18,  22,  23,  43, 
56 

Nyctelia,  362,  363,  364,  367,  370 
varipes,  363 
Nycteliinae,  362 

Nycteliini,  357,  363,  364,  367,  371 
Nycterebosca  falcozi,  29 
Nycteribiidae,  29 
Nyctoporini,  360,  361 
obscurella  group,  380 

ocellatus  group,  171,  194-197,302,311,322 
ochrogaster  group,  1 18 
O’Connor,  F.  W.,  6,  9,  56 
Odonata,  3,  4, 
larvae, 4 

O’Gower,  A.  K„  5,  18,  20,  30,  33,  56 
Omophlinae,  358 
Onymacris,  376 
rugatipennis,  376 
Opadothripini,  123 
Opatrini,  360 
Ophiomyia , 378 

optimus  group,  131,  132,  167,  170,  173-179, 
189,  312,  315,  317,  322,  323,  327,  338 
lineage,  314,  323 
Orgichneumon,  118 
Ornithodorus,  56 
moubata,  23 
Ornithomyia,  29 
Orothripini,  123 

Orr,  C.  W.  M.,  17,  56,  (see  Hudson,  A.)  18, 

54,  56 

Ortholfersia  macleayi,  29 
O’Sullivan,  P.  J.,  (see  Roberts,  F.  H.  S.)  20,  57 
Owen,  W.  B.,  9,  56 
Oxinthas,  364,  369,  373,  374 
praocioides,  363,  369,  372,  373 
Packchanian,  A.  A.,  44,  56 
Painter,  R.  H.,  93,  94,  95,  119 
Palaeoclimatology,  347 
Palaeoecology,  347 
Palaeogeography,  347 
Palaeothripidae,  123 
Panchaetothripinae,  123 
Paniscus,  101 , 119 
Panstrongylus  megistus,  6,  7,  23,  34 


XV 


Paradis,  R.  O.,  102,  119 
Paraphytomyza,  394 
Parasitism,  53,  130 
Parasitoid,  81-117 
Paratany  tarsus,  3,  4 
Parker,  D.  D.,  22,  57 
Parr,  H.  C.  M.,  10,57 
Pascual,  R.,  (see  Patterson,  B.)  326,  348 
Pasteurella  tularensis,  22,  SI 
Pasturella  pestis,  44 
pathogenic  organisms,  5 
patristic  classifications,  306 
Patterson,  B.,  326,  348 
Patterson,  R.  A.,  35,  36,  57 
Patton,  W.  S.,  (see  Cornwall,  J.  W.)  13, 
14,  15,  16,  17,39,50 
Pawan,  J.  L.,  11,  57 
Peat,  A.  A.,  (see  Kumm,  H.  W.)  44,  54 
Peck,  O.,  107,  110,  112,  119 
Pediculus , 33 

humanus , 6,  8,  14,  18,  22,  23,  32,  33, 
35,36,  37,48,52,56 
humanus  corporus , 22,  50 
humanus  humanus,  22,  32 
Pedinini,  360 
Pedobionta,  360 
Pelecyphorus,  375 

Peridroma  margaritosa,  95,  104,  107 
saucia,  91,  107 
Perimylopidae,  376 
Perimylops,  358 
Periplaneta  americana,  37,  38 
Peris cepsia,  90 
carbonaria,  90 
helyma,  90 

helymus,  90-91,92,  113,  116 
laevigata,  90,91,92,  112,  113,  116 
sequax,  90 
Perlidae,  4 
Permothripidae,  123 
Petasites,  377,  378,  379,  380,  381, 383, 
385,  386,  388,  389,  390,  391,  393 
albus,  381, 383,  390,  392 
frigidus,  383,  384,  386,  389,  392,  393 
hybridus,  382,  383 
hyperboreus,  393,  403 
japonicus,  388,  389 
palmatus,  383,  384,  388 
(?  palmatus  x frigidus),  383 


Petasites  (continued) 
paradoxus,  383,  392 
sagittatus,  384,  385 
vitifolius,  384 
Petria,  358,  376 
Phalaenidae,  117,  118,  120 
Philadelphus , 67 

Philip,  C.  B.,  (see  Davis,  G.  E.)  23,  29,  5 1 
Philoematomyia  insignis,  16,  17 
Philolithus,  375 
Phlebotomus  argentipes,  32 
chinensis,  26,  27 
mongolensis,  26 
papatasi,  11,  14,  17,  27,  48 
squamirostris , 26 
Phorichaeta,  90 
sequax,  90 
Phthirus  pubis,  52 

Physogasterini,  357,  361,  363,  364,  367 
Phytagromyza,  394 

Phytomyza,  67,  69,  70,  71,  72,  75,  378,  390, 
394,  395 
agromyzina,  70 

aizoon,  67,  71,  73-74,  77,  78,  79 
alb  ice  ps,  381 

albiceps  group,  377,  378,  379,  380-388 
alpina,  377,  379,  380,  384-386,  387,  397, 
401, 404,  405 
aronici,  386 

atricornis,  76,  378,  388,  389,  395 
buhriella,  380,  390-391, 399 
burchardi,  378,  379 
ciliata,  392 

deirdreae,  67,  70-72,  73,  77,  78,  79,  80 
farfarae,  380,  388,  389,  391-392,  393,  400, 

402 

fuscula,  392 

hordeola,  379,  380,  388,  389 
hyperborea,  377,  378,  379,  380,  392-393, 
401 

hypophylla,  377,  379,  380,  393,  400,  401, 

403 

ilicis,  70 
involucratae , 70 
jacobaeae,  378 
lactuca,  379 
lappae,  387,  388 

377,  379,  393-394,  399 
m////,  70,  378 


XVI 


Phytomyza  (continued) 

mitellae,  67,  70,75,77,78,80 
nigra,  69 

notabilis,  390,  391 
notopleuralis,  70 
petasiti,  377 

ravasternopleuralis , 379,  380,  388,  398 
robustella  group,  377,  378,  388,  389, 
390-394 
rydeniana,  381 

saxifragae,  67,  70,  71,72-73,74,76, 
77,  78,  79,  80 

senecionella,  379,  380,  388,  389,  399 
senecionis , 379,  386-388,  398,  404 
senecionis  ravasternopleuralis,  388 
seneciovora,  378,  379 
syngenesiae,  76,  379,  388,  389,  390 
syngenesiae  group,  377,  378,  379,  380, 
388-389,  394 

tiarellae,  67,  70,  74-75,  77,  78,  80 
tussilaginis , 377,  380,  381-382,  385, 
386,  387,  388,  398,  401, 402,  405 
tussilaginis  kevani,  377,  380,  384,  405 
tussilaginis  petasiti,  380,  381, 382, 
383-384,  405 

tussilaginis  tussilaginis,  380,  382-383, 
405 

Phytomyzinae,  395 
Pick,  F.,  23,  57 
Pimeliini,  359,  361,  362,  376 
Pinus,  348 
strobus,  1 1 2 
Pippin,  W.  F.,  8,  57 
Pissodes  strobi,  110,  112,  119 
planthoppers,  126 
planulatus  group,  246 
subgroup,  320,  340 
Plasmodium,  19,  43,  44 
cathamerium,  21, 45,  54 
gallinaceum,  18,  37,  41 , 45,  46,  49, 

53,  58 

relictum,  21,  45,  46,  54,  56 
Platyscelini,  360 
Plecoptera,  4 

pluripunctatus  group,  136,  138,  167, 

206,  208-225,  302,  311, 317,  318, 
320,321,327,338,  339,  341,344 
Plutella  maculipennis,  100 
Pluto,  63 


Podonominae,  345 
Poecilanthrax,  93,  94,  119 

alcyon,  82,  94-95,  96,  111,  113,  116 
halcyon,  94 
lucifer,  93 

mllistonii,  95,  113,  116 
Polia  acutermina,  90 
adjuncta,  91 
purpurissata,  102,  120 
Polydesmida,  346 
Poly  gen  is  gwyni,  22 
Porosagrotis  orthogonia,  1 17 
Praocini,  357,  361,  362,  363,  364,  367,  371 
Praocis,  362,  367,  370,  371 
chiliensis,  363 
penai,  363,  365,  366,  368 
pilula,  363,371 

Prosimulium  decemarticulatum,  10,  27,  36, 

41 

fuscum,  17,  36,  39 
hirtipes,  27,  3 1 
Pro  to  calliphora  avium,  17 
Pseudaletia  unipuncta,  91, 95,  98,  104,  1 18 
Pseudambly teles,  98,  118 

subfuscus,  97,98,  112,  113,  116 
Pseudoambly  teles,  118 
Pseudoscorpions,  353 
Psorophora  confinnis,  9 
cyanescens,  9 
discolor,  16 
ferox,  9 

Pthirus  pubis,  14,  29 
Pulex  irritans,  22 
Puri,  I.  M.,  14,  57 

Putzeys,  J.  A.  A.  H.,  133,  140,  142,  150,  153, 
161,  165,  175,  176,  179,  184,  191,  192, 
197,  198,  202,  204,  206,  221,  229,  231, 
232,  236,  238,  246,  254,  256,  263,  265, 
287, 348 

Pycnocerimorpha,  360 
Pycnocerini,  360 

quadripunctatus  group,  170,  202-204,  302, 
311,322 

quinquesulcatus  group,  148,  149,  153-156, 
312,  317,  322,  323,  327 
lineage,  323 
-tenuis  lineage,  3 1 4 
Rachou,  R.  G.,  9,  57 
Rajagopal,  P.  K.,  7,  57 


XVII 


Rampazzo,  L.,  (see  Marcuzzi,  G.)  360, 
376 

rat-flea,  56 

Raven,  P.  H.,  (see  Ehrlich,  P.  R.)  302, 
306, 346 

Ray,  H.  N.,  (see  Dasgupta,  B.)  24,  50 
Reaumuria  (aldrichi),  84 
Reduviid,  56 
Reduviidae,  52 

Reid,  E.  T.,  (see  Lewis,  D.  J.)  4 
Reinhard,  H.  J.,  90,  91,  119 
Reinholz,  S.,  (see  Owen,  W.  B.)  9,  56 
Reitter,  E.,  358,  376 
Reynolds,  F.  H.  R.,  (see  St.  John,  J.  H.) 
44,  57 

Rhodnius,  32,  33 

prolixus,  7,  8,  13,  14,  17,  23,  32,  35, 
36,38,  39,44,50,52,  53,55,57, 

59 

Rhynchagrotis  cupida,  91 

Rhynchophora,  345 

Richards,  A.  G.,  (see  Lin,  S.)  36,  55, 

(see  Richards,  P.  A.)  28,  57 
Richards,  P.  A.,  28,  57 
Rickettsiae,  126 

Ringle,  D.  A.,  (see  Herndon,  B.  L.)  31,  53 
Roberts,  F.  H.  S.,  20,  57,  (see  MacKerras, 
M.  J.)  19,55 
Rockstein,  M.,  17,  57 
Rohdendorf-Holmanova,  E.  B.,  383,  395 
Rohlf,  F.  J.,  (see  Sokal,  R.  R.)  136,  348 
Rosenberg,  D.,  3-4 
Ross,  H.  H.,  328,  348 
Rostom,  Z.  M.  F.,  17,  57 
Roy,  D.  N.,9,  17,34,57 
Rozeboom,  L.  E.,  46,  57 
Russell,  P.  F.,  21,  57 
Ryden,  N.,  388,  392,  395 
Sabethes  belisarioi,  1 24 
Sabrosky,  C.  W.,  84,  87,  90,  1 1 9 
St.  John,J.H.,  44,  57 
sallei  group,  1 68,  225-23 1 , 302,  3 1 1 , 

317,  318,  320,  341 
Salmaciinae,  120 

Salpeter,  M.,  (see  Eisner,  T.)  358,  375 
Salpingidae,  359,  376 
sandflies,  14,  32,  5 1 
Sanjean,  J.,  83,  84,  1 19 
Sarcophaga , 119 


Sarcophaga  bullata,  52 
Sasakawa,  M.,  70,  71, 72,  76,  388,  395,  396 
Saxifraga,  67,  70,  71, 72 
ferruginea,  71 
fusca,  7 1 
hieracifolia , 71 
lyallii,  7 1 
nivalis , 71, 72 
paniculata,  73 
punctata , 71,  80 
rotundifolia,  72,  73,  80 
sachalinensis , 71 
Saxifragaceae,  67,  70,  76,  371 
Say,  T.,  133,  246,  348,  362,  373,  376 
Scaphinotus,  345 
petersi,  345 
Scaptia  gattata,  29 
jacksoniensis,  29 
Scaritini,  131-344 
Scaurini,  360,  361 
Schaaf,  A.  C.,  81-117 
Schaefer,  C.  W.,  42,  57 
Schaffner,  J.  V.,  104,  119 
Schechter,  M.  S.,  351,353 
Schildknecht,  H.,  359,  376 
Schistocera  gregaria,  118 
Schizogenius,  347,  348 

amphibius,  166,  168,  234,  236-238,  240, 
243,  244,  245,  267,  320,  324,  332,  340, 
344 

angusticollis,  131,  132,  192,  193 
apicalis,  263,  264,  265,  321,  322,  324, 

328,  340,  344 

arechavaletae,  131,  132,  171,  192-193, 
195,325 
arenarius,  133 

arimao,  169,  264,  265,  295,  297,  305,  321, 
324,  332,  340,  344 

auripennis,  131,  132,  167,  179,  182-184, 
187,214,325,332,  341 
banningeri,  171, 201-202,  203,  322,  325 
basalis,  170,  184,  185,  186,  187,  325 
bicolor , 131,  132,  170,  177-179,322,325 
brevisetosus,  131,  132,  168,  206-208,  226, 
227,  324,  332,  341, 344 
brittoni,  225 
canaliculatus , 171 
capitalis,  198,  325 
carinatus , 170,  189-190,  191,325 


XV111 


Schizogenius  (continued) 

cearaensis,  131,  132,  170,  184,  185, 
186,  187,322,325 
championi,  131,  132,  138,  270,  271, 
277 

chiapatecus,  332 

chiricahuanus,  131,  132,  169,  252, 
254,  257-258,  260,  261,  262,  321, 
324,  332,  340,  344 
classification  of,  131-344 
clivinoides,  170,  171,  176,322,325 
costiceps,  170,  171,  188,  195,  325 
costipennis,  131,  132,  170,  189,  190- 
191,  195,  325 

crenulatus,  131,  132,  144-148,  150, 
151,  152,  160,328,  332,  337 
crenulatus  chiapatecus,  149,  150,  152- 
153,  163,  164,325,329 
crenulatus  crenulatus,  149,  150-152, 
163,  164,  165,325,329 
darlingtoni,  197,  322,  325 
depressus,  169,  268,  270,  285,  286, 
287-294,  295,  298,  301,  321,  324, 
329,  332,  336,  338,  340,  344 
dilatus,  131,  132,  168,  231,  232-234, 
243,  244,  245,  320,  324,  329,  332, 
339,  344 

dyschirioides,  170,  175-176,  178,  322, 
325 

elongatus,  170,  188-189,  191,  195, 

325 

emdeni,  131,  132,  169,  263,  264,  265, 
270,  295,  297,  321,  324,  332,  340, 
344 

exaratus,  154 

falli,  131,  132,  169,  214,  270,  277, 
279,  281-285,  286,  287,  294,  295, 
298,  300,  301,  321,  324,  331,  332, 
335,  336,  344 

ferrugineus,  167,  179-181,  182,  183, 
184,  187,325,332,  337,338 
frontalis,  236,  238 
gracilis,  171 

grossus,  170,  176-177,  179,  322,  325 
impressicollis,  149,  161,  163,  164, 

322,  325 

impuncticollis,  131,  132,  149,  160, 
161-162,  163,  164,325 
interstriatus,  171,  197,  322,  325 


Schizogenius  (continued) 
jacarensis,  131,  132,  170,  172-173,  178, 
191,325 

janae,  149,  155-156,  163,  164,  165,  322, 
325 

kulti,  131,  132,  168,218,219,  220,  221, 
222,  223-225,  226,  227,  228,  320,  324, 
328,  329,  331,  332,  336,  339,  344 
leprieuri,  171 

lindrothi,  131,  132,  167,  199-201,  203, 
322,  325,  327,  331,  332,  337,  338 
lineolatus,  141,  169,  238,  240,  246-251, 
260,  261,  262,  266,  267,  324,  332,  341, 
344 

litigiosus,  169,  266,  268-269,  294,  295, 
297,  321,  324,  332,  336,  340,  344 
longipennis,  131,  132,  169,  232,  252,  253, 
254-257,  258,  260,  261,  263,  320,  321, 
324,  329,  332,  340,  344 
maculatus,  148,  149,  156,  162,  165,  325 
materials,  133-134 
methods,  135-144 

multipunctatus , 170,  184,  185,  187,  191, 
216-221,325 

multisetosus,  138,  168,  216,  221,  226,  227, 
228,  320,  324,  329,  331,  332,  339,  344 
negrei,  131,  132,  170,  184,  185,  186,  187, 
322,  325 

neovalidus,  131,  132,  169,  252-254,  255, 
257,  258,  260,  261,  320,  321,  324,  332, 
340,  344 

ocellatus,  131,  132,  166,  171,  196-197, 
203,  325 

ochthocephalus,  131,  132,  169,  264,  270, 
281,  283,  284,  285-287,  294,  295,  298, 
300,  301,  321,  324,  331,  332,  336,  340, 
344 

optimus,  145,  146,  147,  165,  167,  173- 
175,  176,  177,  178,  322,  325,  332 
ozarkensis,  131,  132,  168,  240,  241,  242, 

244,  245,  246,  305,  310,  320,  324,  329, 
332,  336,  340,  344 

pacificus,  131,  132,  169,  254,  257,  258- 
263,  320,  321,  324,  332,  340,  344 
peninsularis , 131,  132,  182,  184 
phylogeny,  302-325 

planulatus,  168,  238-240,  242,  243,  244, 

245,  246,  267,  305,  310,  320,  324,  329, 
332,  336,  340,  344 


XIX 


Schizogenius  (continued) 

planuloides,  131,  132,  168,  240,  241- 
246,  305,  310,  320,  324,  329,  332, 
344 

pluripunctatus,  165,  166,  167,  217, 
218,  219,  220,  221-223,  226,  227, 
228,  269,  315,  320,  324,  329,  332, 
336,  339,  344 

plurisetosus,  131,  132,  138,  167,  209, 
214-216,217,218,219,220,  221, 
224,  226,  227,  228,  320,  324,  329, 
332,  339,  344 
putzeysi,  171,  198,  325 
pygmaeus,  131,  132,  138,  169,  214, 
263,  265,  270-277,  278,  279,  281, 
285,  286,  288,  294,  295,  296,  297, 
298,  299,  300,  307,  310,  311,  321, 
322,  324,  327,  329,  331,  332,  336, 
340,  344 

quadripunctatus,  170,  202-204,  325 
quinquesulcatus,  149,  153-154,  155, 
163,  164,325 

reichardti,  13 1,  132,  171,  192,  193- 
194,  195,325 
riparius,  197,  325 

sallei,  145,  146,  147,  168,  229-230, 
242,  243,  244,  245,  324,  332,  341, 
344 

scopaeus,  131,  132,  138,  169,  242, 
270,  276,  278-281,  285,  294,  295, 
296,  297,  298,  299,  300,  307,  310, 
311,  321,  324,  329,  332,  336,  340, 
344 

sculptilis,  131,  132,  149,  156-158, 

160,  163,  164,  322,  325,  327,  328, 
331,  332,  337 
sellatus,  171,  202 
seticollis , 138,  209,  320,  332,  339, 

344 

seticollis  seticollis,  1 67,  209-2 12,  214, 
216,  221,  224,  226,  227,  228,  320, 
324,  329 

seticollis  vandykei,  131,  132,  167,  211, 
212-214,  226,  227,  228,  320,  324, 
329,  339 

simplex,  221,  223 

strigicollis,  142-143,  165-167,  171, 

191, 195, 325 
sulcatulus,  171 


Schizogenius  (continued) 
sulcatus,  179 

sulcifrons,  141,  169,  238,  240,  246,  251, 
265-268,  295,  297,  321, 324,  332,  336, 
340,  344 

suturalis,  131,  132,  149,  160,  162-165,  325 
szekessyi,  149,  155,  156,  163,  164,  165, 
322,325 

taxonomy,  142-301 

tenuis,  146,  147,  149,  156,  157,  158-161, 
162,  163,  164,  322,  325,  327,  332,  337 
tibialis,  131,  132,  136,  168,  197,  231,  232, 
234-236,  243,  244,  245,  304,  306,  307, 
309,  310,  311,  320,  324,  329,  332,  339, 
344 

tristriatus,  168,  225,  231-232,  234,  243, 
244,  245,  256,  264,  320,  324,  329,  332, 
339,  344 

tristriatus  longipennis,  254 
truquii,  167,  204-206,  226,  227,  324,  325, 
332,  338,339,  341,344 
validus,  131,  132,  253,  254,  257 
vandykei,  332,  344 
zoogeography,  326-344 
Schizophora,  68,  69 

Schoute,  E.,  (see  Buck,  A.)  15,  16,  19,  45, 

49 

Schubert,  J.  H.,  29,  57 

Schulze,  L.,  360,  376 

Schwardt,  H.  H.,  (see  Tashiro,  H.)  10,  58 

Scotogramma  trifolii,  97 

Sehgal,  V.  K.,  384,  388,  389,  396 

Sella,  M.,  18,20,57 

Senecio,  377,  378,  379,  385,  387,  388,  389 
alpinus,  379,  385,  386 
atropurpureus  tomentosus,  389 
congestus  var.  palustris,  389 
cruentus,  389 
doria,  389 
fluviatilis,  379,  387 
fuchsii,  379,  387 

jacobaea,  378,  379,  385,  386,  387,  389, 

404 

lugens,  379,  385,  386,  394,  404 
mikanioides,  389 
nemorensis,  379,  387,  404 
pauperculus,  379,  386 
sheldonensis , 379,  394 
squalidus,  389 


XX 


Senecio  (continued) 
subalpinus,  379,  387 
vernalis , 389 
vulgaris,  389 
yukonensis,  389 
Senecioneae,  377,  378,  382,  386 
Sergentomyia  squamirostris,  26,  27 
Sericothripina,  123 
Sericothripini,  123 
Service,  M.  W.,  6,  9,  20,21,58 
Shambaugh,  G.  F.,  36,  40,  41,  58,  (see 
Fisk,  F.  W.)  26,  36,40,41,43,51 
Sharp,  D.,  359,  376 

Shipley,  A.  E.,  (see  Nuttall,  G.  H.  F.)  15, 
17,  56 

Short,  J.  R.  T.,  102,  119 
Shute,  P.  G.,  13,  14,  15,  16,  44,58 
Silberman,  M.  L.,  (see  Hopkins,  D.  M.) 
335, 347 

Simmons,  J.  S.,  (see  St.  John,  J.  H.)  44, 
57 

Simpson,  G.  G.,  139,  140,  141, 327,  348 
Simuliid,  60,  61 
larvae,  4 

Simuliidae,  10,  49,  50,  51,  60 
Simulium,  55 
anatinum,  27 
aureum,  10,  27 
croxtoni  10,  27 
damnosum,  10,  1 1,  27,  50,  55 
griseicolle , 27 
latipes,  10,  27 
neavei,  27,  55 
parnassum , 3 1 
pupae,  4 

quebecense,  10,  27 
rugglesi,  10,  27,  36,  41, 49 
venustum,  11,  1 7,  27,  3 1 , 36,  39,  40, 
41 

vittatum , 3 1,  36,  43 
Siphunculata,  6 
Skopin,  N.  G.,  360,  361,376 
Smith,  D.  S.,  367,  376 
Smith,  E.  M.,  (see  Freyvogel,  T.  A.)  17, 
43,  52 

Smith,  J.  B.,  141,  348 
Smith,  J.  J.  B.,  (see  Friend,  W.  G.)  7,  52 
Sneath,  P.  H.  A.,  (see  Sokal,  R.  R.)  137, 
348 


Snodgrass,  R.  E.,  5,  58 
Snow,  S.  J.,  93,  110,  119 
Social  Evolution,  130 
Sokal,  R.  R.,  136,  137,  302,  306,  348,  (see 
Camin,  J.  H.)  141,315,345 
Somaticus,  376 

Somers,  G.  F.,  (see  Sumner,  J.  B.)  35,  58 
S^nderup,  H.  P.  S.,  388,  389,  396 
Spaelotis  clandestina,  104,  120 
Spelaeodytes  mirabilis,  196 
Spencer,  K.  A.,  70,  76,  378,  379,  381,  382, 
383,  384,  386,  390,  392,  396,  399 
Sphaeriontis,  373 
Sphingoidea,  63 
spider,  129 
lycosid,  83 
Spilichneumon,  118 

superbus,  95,  97-98,  99, 1 13,  1 14,  1 15,  1 16 
Spilman,T.  J.,  359,  376 
Spirochaeta  duttoni,  23 
Spodoptera  frugiperda,  93 
stable  fly,  49,  50,  58 
Stage,  H.  H.,  9,  58 

Stahler,  N.,  (see  Terzian,  L.  A.)  21, 45,  58 

Stary,  B.,  383,  387,  388,396 

Staubli,  W.,  23,  24,  58,  (see  Freyvogel,  T.  A.) 

23,24,  25,52 
Steffan,  A.  W.,  4 
Stegomyia  (Aedes)  aegypti,  57 
Steinheil,  E.,  133,  188,  348 
Stenopneusticae,  118 
Stenosini,  361 

Stephen,  W.  P.,  (see  Bohart,  G.  E.)  93,  117 
Stohler,  H.  R.,  23,  24,  46,  58 
Stomoxys  calcitrans,  6,  10,  11,  16,  17,  21, 

22,  28,  32,  35,  36,  42,  44,  49,  50,  53, 
55,  57 
indica,  16 
sitiens,  17 

Strickland,  E.  H.,  1,  82,  83,  84,  87,  89,  90, 
98,  101,  103,  106, 107,  110,  114,  119 
strigicollis-elongatus-carinatus  lineage,  311, 
314,  315 

strigicollis  group,  171,  191-192,317,323, 

327 

lineage,  314,  323 
-truquii  lineage,  314 
substriatus  group,  308 

Sudia,  W.  D.,  (see  Chamberlain,  R.  W.)  44,  50 


XXI 


Suenaga,  O.,  10,  58 
sulci  fro  ns  subgroup,  321 
Sullivan,  W.  N.,  (see  Schechter,  M.  S.) 
351,353 

Sumner,  J.  B.,  35,  58 
Suter,  J.,  (see  Staubli,  W.)  23,  24,  58 
Swartzwelder,  J.  C.,  (see  Zeldon,  R.)  6, 
8,  60 

Swellengrebel,  N.  H.,  (see  Buck,  A.)  15, 
16,  19,  45,49 
Sympetrum  internum , 3 
syngenesiae  group,  69 
Syngrapha  epigaea,  104,  107,  120 
Systoechus  somali,  1 1 8 
vulgaris , 93,  117 
Tabanid,  29 
Tabanidae,  10,  51,  56 
Tabanus,  1 1,  27,  28,  50 
albimedius,  1 6,  27 
quinquevittatus,  10 
septentrionalis,  10 
sulcifrons,  10 
Tachinid,  84 
hosts,  94 

Tachinidae,  83-92,  113,  117,  119 
tachyine  beetles,  346 
Tacky s,  335 

Taraxacum  officinale , 82 
Tashiro,  H.,  10,  58 
Tatchell,  R.  J.,  16,  17,40,58 
Taufflieb,  R.,  (see  Mattern,  P.)  30,  56 
Tawfik,  M.  S.,  8,  58 
Taylor,  R.  L.,  1 10,  1 12,  1 19 
Tempelis,  C.  H.,  (see  Anderson,  J.  R.)  6, 
10,  22,32,  49 
Templis,  C.  F.,  30,  58 
Tenebrio,  360 

Tenebrionid,  362,  367,  374,  375 
larvae,  94 

Tenebrionidae,  353,  375,  376 
Tenebrionidi,  376 
Tenebrioninae,  361,  362 
Tenebrionini,  360,  361 
Tenebrionoidea, 

familial  and  subfamilial  classification 
of,  357-374 

morphological  and  ecological  charac- 
teristics, 358-361 

Tenorio,  P.  A.,  (see  Wagner,  C.)  35,  36, 


Tenorio,  P.  A.  (continued) 

46,  59 

Tentyriidae,  357-374 

Tentyriinae,  358,  359,  360,  361,  362,  375 
Tentyriine,  360 
Tentyriini,  360,  364 

tenuis  group,  131,  132,  148,  156-171,312, 
317,322,323,327,337 
lineage,  323 
Tephritid  larva,  392 
Terebrantia,  123 
termites,  129 

Terzian,  L.  A.,  20,  21,  45,  58,  (see  Wagner, 
C.)  35,36,  46,  59 

Theodor,  O.,  (see  Adler,  S.)  10,  11,  14,  17, 
27,48 

Thompson,  W.  R.,  82,  119 
Thomson,  J.  D.,  (see  Minchin,  E.  A.)  28,  56 
Thrassis  bacchi  gladiolis,  22 
Thripidae,  123 
Thripinae,  123 
Thripoidea,  123 
Thurman,  D.  C.,  8,  58 
Thysanoptera,  123 
Tiarella,  67,  70 
trifoliata,  74,  80 
tick,  50,  125 
cultures,  126 
tiger  beetles,  3 1 5 
Tiphia,  larvae,  1 17 
Tolmiea,  67,  70 
menziesii,  74 

Torren,  G.  v.  d.,  (see  Buck,  A.)  19,  45,  50 

Tortricid,  118 

Tortricidae,  1 19 

Tortrix  alleniana,  102 

torvus  group,  308 

Tothill,  J.  D.,84,  91,  119 

Townes,  H„  101,  102,  119 

Townes,  M.,  (see  Townes,  H.)  101,  102,  1 19 

Trachynotina,  376 

Trainer,  D.  O.,  (see  Anderson,  J.  R.)  10,  49 
Treherne,  R.  C.,  110,  119 
Trembley,  H.  L.,  7,  12,  58 
Treponema  pertenue,  44 
Treto thorax,  358,  361 
Triatoma , 33,  44,  56 
dimiata,  6,  8,  60 
gerstaeckeri,  8,  44,  57,  58 


XXII 


Triatoma  (continued) 
heidemani,  44 

infestans,  6,  8,  14,  17,  32,  34,  52 
lectularia,  44 
maculata,  13,  14,  38 
pro tr acta,  44 
rubrofasciata,  13,  14,44 
sanguisuga,  8,  44,  53 
sanguisuga  texana,  8,  57 
uhleri , 44 
Triatomidae,  59 
Triatominae,  53,  57 
Tricholabus,  1 1 8 
Trichoptera,  348 
Triorophini,  361,  364 
Triplehorn,  C.  A.,  370,  373,  376 
tristia tus  group,  168,  231-246,  264,  302, 
311,  317,  318,  320,  327,  338,  339, 
344 

subgroup,  320 

truquii-capitalis , lineage,  314 
truquii  group,  167,  204-206,  302,  311, 
317,323,339,  341 

lineage,  131,  132,  311,  312,  314,  315, 
316-320,  322,  323,  324,  325,  328, 
335,  337,  338,  339,  340,  341,  344 
Trypanosoma  bocagei , 27 
brucei,  44 
evansi,  44 
gambiense,  44 
grayi,  47,  53 
hippicum,  44 
lewisi , 28,  56 

Tschirnhaus,  M.  von,  68,  69,  76 
tsetse  flies,  5,  13,  18,  28,  47,  48,  50,  54, 
55,  56,  57,  59 

Tuomikoski,  R.,  302,  303,  348 
Turner,  T.  B.,  (see  Kumm,  H.  W.)  44,  54 
Tussilago,  377,  378,  379,  380,  381,  388, 
389,390,  391 

farfara,  382,  383,  390,  391,  392,  402 
Ulomimorpha,  360 
Ulomini,  360 

Upmanis,  R.  S.,  (see  Yoeli,  M.)  46,  60 
Usinger,  R.  L.,  (see  Mayr,  E.)  348 
Uvarov,  B.  P.,  5,  59 
Uzelothripidae,  123 
Vance,  A.  M.,  101,  119 
Vanderplank,  F.  L.,  22,  59 


Van  Dyke,  E.  C.,  133,270,  348 
Venard,  C.  E.,  (see  Devine,  T.  L.)  13,  5 1, 

(see  Guptavanij,  P.)  11,  53 
Vickery,  R.  K.,  Jr.,  (see  Lindsay,  D.  W.)  310, 
347 

Viereck,  H.  L.,  104,  120 
Villa , 93 

alternata,  112,  113,  114,  116 
fulviana,  113,  116 
(Hemipenthes)  moroides,  94 
lateralis,  112,  113,  116 
moroides , 113,  116 
(Villa)  alternata,  94 
(Villa)  fulviana,  94 
(Villa)  lateralis,  94 
Vinson,  J.,  143,  348 
virus,  125-127 
Voigt,  G.,  387,  396 
Vonk,H.  J.,  5,59 
Wagner,  C.,  35,  36,  46,  59 
Wagneria,  90,  119 
Waldbauer,  G.  P.,  5,  59 
Walkden,  H.  H.,  104,  110,  120 
wasps,  129 

Waterhouse,  D.  F.,  5,  29,  59,  (see  Day,  M.  F.) 
5,43,51 

Watt,  J.  C.,  357,  358,  359,  360,  361, 376 
Wattal,  B.  L.,  8,  59 
wax  embedded  specimens, 
rapid  orientation  of,  61-62 
Webb,  D.  A.,  67,  73,  74,  76 
Webster,  R.  L.,  104,  119 
weevil,  white  pine,  112,  119 
Weis,  K.  FI.,  (see  Schildknecht,  H.)  359,  376 
Weiss,  E.,  125 

Weitz,  B.,  28,  29,  31,32,  59 
West,  A.  S.,  5,  29,  59,  (see  Downe,  A.  E.  R.) 
30,  51,  (see  Gosbee,  J.)  17,  52,  (see  Orr, 

C.  W.  M.)  17,  56 
Wharton,  R.  H.,  9,  59 
Whitehead,  D.  R.,  131-344,  348 
Whitehouse,  F.  C.,  97,  110,  120 
Wigglesworth,  V.  B.,  5,  7,  1 1,  16,  17,  23,  28, 
32,33,43,59 

Williams,  C.  A.,  Jr.,  29,  3 1 , 33,  59 
Williams,  P.,  (see  Kershaw,  W.  E.)  6,  10,  54 
Willis,  H.  L.,  315,348 
Wilson,  E.  O.,  129,  320,  321, 348,  (see 
Brown,  W.  L„  Jr.)  292,  336,  345 


Wistreich,  G.  A.,  (see  Chao,  J.)  21, 50 
Wolfe,  J.  A.,  (see  Hopkins,  D.  M.)  335, 
347 

Wood,  G.  W.,  102,  104,  107,  1 10,  120 
Woodard,  D.  B.,  9,  59 
Wright,  W.  R.,  11,  59 
Wuchereria  bancofti,  54,  59 
Xenopsylla  cheopis , 2,  22 
Xylophones  pluto,  64 
Xystodesmidae,  346 
Yaguzhinskaya,  L.  W.,  24,  60 
Yang,  Y.  J.,  1 1,  17,  35,  36,  38,  39,  40, 
41,  43,  60,  (see  Davies,  D.  M.)  36,  39, 
41,43,  50 

Yates,  W.  W.,  (see  Stage,  H.  H.)  9,  58 
Yoeli,  M.,  46,  60 

Yorke,  W.,  13,  14,  15,  16,  17,  28,  60 

Yponomeutid,  100 

Zaman,  V.,  3 1,  60 

Zeledon,  R.,  6,  8,  60 

Zoemer,  H.,  387,  396 

Zopheridae,  358 

Zopherosis,  361 

Zopherus,  358 

Zophosini,  357,  362,  363,  371 
Zophosis,  362,  364,  367,  369,  370 
plana,  363,  365,  366,  368 
reticulata , 373 

Zuniga,  A.,  (see  Zeledon,  R.)  6,  8,  60 
Zuska,  J.,  83,  120 


* 


* 

■ 


Quaestiones 


MUS.  COMP.  ZC-CL 
L.'SRARY 


FEB  4 W2 


. HARVARD 

entomologicae 


A periodical  record  of  entomological  investigations, 
published  at  the  Department  of  Entomology, 
University  of  Alberta,  Edmonton,  Canada. 


VOLUME  VIII 


NUMBER  1 


JANUARY  1972 


QUAESTIONES  ENTOMOLOGICAE 


A periodical  record  of  entomological  investigation  published  at  the  Department  of 
Entomology,  University  of  Alberta,  Edmonton,  Alberta. 

Volume  8 Number  1 1 January  1972 


CONTENTS 


Editorial  - One  Eye  on  the  Pot  1 

Rosenberg  - A chironomid  (Diptera)  larva  attached  to  a libellulid  (Odonata)  larva  ....  3 

Gooding  - Digestive  processes  of  haematophagous  insects  I.  A literature  review 5 

Craig  - Rapid  orientation  of  wax  embedded  specimens  61 

Book  review 63 

Announcement 64 


Editorial  — One  Eye  on  the  Pot 

The  year  1972  marks  the  fiftieth  anniversary  of  the  establishment  of  the  Department  of 
Entomology  at  the  University  of  Alberta.  Anniversary  —the  turning  of  the  year—  is  perhaps 
an  unfortunate  term,  and  especially  so  in  relation  to  a university,  the  function  of  which  is 
the  unification  —or  turning  into  one-  of  knowledge.  For,  though  history  may  repeat  itself, 
it  is  to  be  hoped  that  a university  does  not.  Perhaps  it  is  the  celebration  of  centennials  and 
other  major  anniversaries  that  encourages  history  to  repeat  itself,  for  such  celebrations  are 
often  preoccupied  with  the  events  of  a hundred  years  earlier.  If  we  should  not  be  preoc- 
cupied with  the  past,  then,  trapped  in  the  tunnel  of  time,  our  only  remaining  option  is  to 
look  forwards.  But  gazing  into  crystal  balls  is  fraught  with  perils  for  these  without  an  anchor 
in  history,  so  perhaps  we  need  to  take  a quick  glance  backward  to  see  whence  we  have  come 
before  viewing  the  horizon  ahead  to  chart  a course  for  the  future.  Oh  for  a pair  of  compound 
eyes,  each  able,  though  plastered  on  to  the  head,  to  look  both  ways  without  asking  it  to 
turn  for  them.  Too  little  attention  has  been  paid  to  this  feature  of  the  structure  of  insects, 
in  attempts  to  understand  their  ability  to  stay  on  course  in  migration. 

In  1921  E.  H.  Strickland,  then  working  for  the  Canada  Department  of  Agriculture  at 
Lethbridge,  visited  Edmonton  to  consider  an  offer  made  to  him  by  the  University  of  Alberta. 
It  must  have  been  later  in  the  year  than  this  issue  of  Quaestiones  entomologicae  will  appear, 
for  the  then  Dean  of  Agriculture  took  him  out  on  to  some  rough  ground  at  the  north  end  of 
the  campus  overlooking  the  valley  of  the  North  Saskatchewan  River  —like  stout  Cortez— 
and  showed  him  the  place  where  a building  which  would  house  the  proposed  department  of 
entomology  was  to  be  built.  We  do  not  know  whether  Strickland  was  swayed  by  what  he 
saw  on  that  day,  but  in  April  1922  the  Department  was  created  and  twenty-five  years  later 
the  staff  increased  by  100  per  cent  and  he  and  I each  occupied  a roomlet,  at  either  end  of 
a modest  room  in  the  medical  building.  Our  supporting  staff  consisted  of  half  a stenographer 
and  $ 12  a month  worth  of  part  time  student  help.  Professor  Strickland’s  budget  for  entomo- 
logical books  for  the  library  in  1923  was  $20,  and  the  entire  budget  for  the  department  in 
1947-8  was  $9,401.  Although  nobody  saw  fit  to  calculate  it,  the  cost  of  a “teaching  unit” 
was  $18,  a fifth  of  the  cost  today;  we  now  teach  four  times  the  “units”  with  three  times 
the  staff  and  twenty  times  the  money.  In  1958,  four  years  after  Professor  Strickland  retired, 
the  Department  of  Entomology  moved  into  its  present  quarters  in  the  Agriculture  and  Bio- 
Sciences  building,  which  is  just  about  where  the  Dean  had  said  it  would  be.  Two  years  later 
these  quarters  proved  inadequate  and  a migratory  branch  of  the  department  finally  came  to 


2 


an  uneasy  rest  in  the  basement  of  Athabasca  Hall. 

So  much  for  the  backward  glance.  What  of  the  future?  When  the  first  issue  of  Quaestiones 
entomologicae  appeared  a caustic  reader  remarked  that  he  had  no  use  for  fly-by-night 
periodicals  which  usually  folded  up  in  five  years.  We  have  outlived  that  one.  Some  other  pen 
than  mine,  I predict,  will  write  an  editorial  for  the  volume  which  marks  the  centennial  of 
the  Department  of  Entomology.  The  editor  who  writes  it  may  well  have  something  caustic 
to  say  about  this  editorial  for  some  of  my  predictions  will  be  wrong;  but  if  this  one  is,  at 
least  he  will  not  say  so. 

It  is  customary  to  predict,  by  direct  extrapolation  of  a smoothed  curve  of  human  popula- 
tion, that  there  will  be  7.5  billion  of  us  by  the  year  2000.  With  rather  less  reliability  it  might 
be  predicted  that  by  2022  there  will  be  double  that  number.  But  the  curve  of  human  pop- 
ulation is  based  on  close  approximations  for  less  than  300  years  and  we  do  know  that 
Xenopsylla  cheopis  was  indirectly  responsible  for  putting  several  dents  in  the  curve  before 
that  time  and  glacial  epochs  probably  did  likewise.  Furthermore  at  least  some  drafts  of  the 
early  part  of  the  curve  have  been  based  on  several  species  of  hominid.  Add  to  this  Deevey’s 
(1)  elegant  demonstration  that  this  is  man’s  third  population  explosion  and  the  first  to 
have  its  roots  in  non-renewable  resources,  and  any  applied  entomologist  might  be  expected 
to  predict  an  end  to  the  outbreak  before  2022.  A successful  prediction  of  such  an  event 
should  lead  to  a return  to  favour  of  entomologists,  among  both  their  biological  colleagues 
and  the  population  at  large.  Indeed,  since  a reduction  in  the  number  of  insect  species  will 
almost  certainly  have  resulted  from  the  activities  of  the  peak  population  of  man,  perhaps 
the  economy  of  a declining  population  will  support  enough  taxonomists  to  catch  up  a bit 
with  the  task  of  describing  these  species. 

Sparked  by  the  shortage  of  food  and  specifically  of  animal  protein,  man  will  have  re- 
turned to  eating  insects.  His  smaller  population  will  have  been  able  to  retreat  from  mono- 
culture so  that  insect  pest  problems  will  have  been  reduced  —perhaps  to  a point  where 
eating  the  troublesome  species  will  provide  adequate  control.  Of  course  the  arts  of  the 
kitchen  should  have  so  progressed  that  even  a swarm  of  hoary  old  horseflies  might  be 
transformed  into  a delectable  dish,  and  we  can  but  imagine  the  succulent  delight  a skilled 
cook  could  render  from  the  abdomens  of  queen  termites  forcibly  fed  on  proof  copies  of 
last  quarter’s  Quaestiones  entomologicae.  Or  perhaps  by  then  it  will  be  last  month’s. 

Clearly,  as  compared  with  today,  the  relative  importance  of  the  positive  and  the  negative 
aspects  of  applied  entomology  will  be  reversed.  Benefactory  entomology  will  transcend  pest 
control. 

The  Department  of  Entomology  at  the  University  of  Alberta  will  be  re-united  under  one 
roof  -perhaps  that  of  Athabasca  Hall—  but  it  will  be  understood  that  the  roof  will  be 
demolished  in  the  following  year. 

And  so,  like  the  cross-eyed  cook  in  the  English  folk-song,  we  look  briefly  back  and 
soberly  forward  from  the  first  issue  of  volume  eight  —‘with  one  eye  on  the  pot  and  the 
other  up  the  chimney...’ 

(1)  Scientific  American  203:195,  1960 


Brian  Hocking 


A CHIRONOMID  (DIPTERA)  LARVA  ATTACHED  TO  A 
LIBELLULID  (ODONATA)  LARVA 


D.  ROSENBERG 
Department  of  Entomology 

University  of  Alberta  Quaestiones  entomologicae 

Edmonton  7,  Alberta  8 : 3-4  1972 


While  identifying  libellulid  larvae  collected  in  1966-1968  from  two  sloughs  located  about 
9 miles  southeast  of  Edmonton,  Canada,  I found  one  specimen  with  the  tube  of  a chirono- 
mid  larva  attached  to  its  prothorox.  (See  photograph  below;  inset  magnification  is  approxi- 
mately 35x).  The  libellulid  was  a Sympetrum  sp.  probably  internum  Montogomery  and  the 
chironomid  within  the  tube  was  a Par atany  tarsus  sp. 


4 


Rosenberg 


s The  association  is  probably  not  truly  phoretic  like  those  reported  by  Steffan  (1965a, 
1965b,  1967)  between  chironomid  larvae  and  other  aquatic  invertebrates,  mainly  insects, 
but  more  probably  is  accidental  like  those  reported  by  Lewis  et  al  (1960)  and  Corbet  (1962) 
between  immature  simuliids  and  odonate  larvae.  I have  reached  this  conclusion  because  only 
one  of  the  few  hundred  libellulid  larvae  I examined  carried  a chironomid  larva  and  because 
Paratany tarsus  sp.  larvae  are  one  of  the  most  common,  free  living  chironomid  larvae  in  the 
sloughs  I studied.  Such  associations,  however,  may  represent  a stage  in  the  development  of 
phoresis  and  as  Corbet  (1962)  wrote:  “further  attention  paid  to  anomalous  cases  of  this  kind 
might  throw  light  on  the  way  in  which  the  well-known  [examples  of]  phoresis  originated”. 

ACKNOWLEDGEMENTS 

I would  like  to  thank  P.S.  Corbet;  and  A.L.  Hamilton  and  O.A.  Saether  for  helping  in 
identifying  the  libellulid  and  the  chironomid  respectively  and  J.S.  Scott  for  the  photograph- 
ic work. 


REFERENCES 

Corbet,  P.S.  1962.  Observations  on  the  attachment  of  Simulium  pupae  to  larvae  of 
Odonata.  Ann.  Trop.  Med.  Parasit.,  56  (2):  136-140 
Lewis,  D.J.,  E.T.  Reid,  R.W.  Crosskey,  & J.B.  Davies.  1960.  Attachment  of  immature 
Simuliidae  to  other  arthropods.  Nature,  Lond.,  187  (4737):6 1 8-6 1 9 
Steffan,  A.W.  1965a.  On  the  epizooic  associations  of  Chironomidae  (Diptera)  and  their 
phyletic  relationships.  Proc.  12th  int.  Congr.  Ent.,  London  1964,  1:77-78 
Steffan,  A.W.  1965b.  Plecopteracoluthus  downesi  gen.  et  sp.  nov.  (Diptera:  Chironomidae), 
a species  whose  larvae  live  phoretically  on  larvae  of  Plecoptera.  Canad.  Ent.,  97  (12): 
1323-1344. 

Steffan,  A.W.  1967.  Larval  phoresis  of  Chironomidae  on  Perlidae.  Nature,  Lond.,  213 
(5078):846-847. 


DIGESTIVE  PROCESSES  OF  HAEMATOPHAGOUS  INSECTS 
I.  A LITERATURE  REVIEW 


R.  H.  GOODING 
Department  of  Entomology 
University  of  Alberta 
Edmonton  7,  Alberta 


Quaestiones  entomologicae 
8:  5-60  1972 


About  240  papers  published  between  1903  and  early  1971,  providing  information  on 
more  than  150  species  of  haematophagous  insects,  are  reviewed.  Aspects  of  digestive  physi- 
ology covered  are  size  of  the  blood  meals  and  their  distribution  within  the  alimentary  canal, 
properties  of  the  salivary  glands,  gross  and  histological  changes  in  the  gut  and  its  contents, 
the  enzyme  content  of  the  gut  and  the  properties  of  the  digestive  enzymes.  The  relationship 
of  digestive  processes  to  vectoring  ability  is  discussed  briefly. 

Environ  240  articles  publies  entre  1903  et  1971  donnant  des  informations  sur  plus  de 
150  especes  d’insectes  hematophages,  sont  revises  dans  cet  ouvrage.  Les  aspects  de  la  physi- 
ologic de  la  digestion  qui  sont  couverts  sont:  la  quantite  de  sang  que  contiennent  les  repas 
et  sa  distribution  dans  le  tractus  alimentaire,  les  caracteristiques  des  glandes  salivaires,  les 
changements  histologiques  et  Vensemble  des  changements  de  Tintestin  et  de  son  contenu, 
le  contenu  enzymatique  de  Tintestin  et  les  caracteristiques  propres  des  enzymes  digestives. 
La  relation  qui  existe  entre  le  processus  digestif  et  Vaptitude  quont  certains  insectes  a etre 
vecteur  de  maladies  est  decrite  brievement. 

The  primary  objectives  of  this  paper  are  to  summarize  the  knowledge  of  the  digestive 
physiology  of  blood-sucking  insects  and  to  indicate  some  unsolved  problems.  Aspects  con- 
sidered include  the  size  of  the  blood  meals,  the  fate  of  these  within  the  digestive  tract,  and 
the  sources  and  nature  of  the  enzymes  involved  in  their  breakdown.  Sources  of  the  blood 
meals  and  the  mechanisms  of  ingestion  and  absorption  are  not  considered  except  where 
these  appear  to  have  a bearing  on  digestion. 

A comparative  study  of  digestion  in  blood-sucking  insects  is  of  interest  primarily  for  two 
reasons.  First,  blood  feeding  has  evolved  several  times  in  the  insects  and  a comparison  of 
digestive  processes  may  reveal  that  different  species  have  overcome  problems  presented  by 
the  blood  meal  in  different  ways.  Conversely,  the  nature  of  their  food  may  have  resulted  in 
convergence  in  the  digestive  processes  of  different  haematophagous  species.  A second  and 
more  practical  reason  is  that  many  blood-sucking  arthropods  are  vectors  of  pathogenic 
organisms.  Since  most  of  these  pathogens  spend  some  time  in  the  gut  of  the  vector,  digestive 
processes  of  the  insect  could  influence  vectoring  ability.  The  possible  importance  of  the 
digestive  processes  of  insects  in  studies  of  host  relationships,  nuisance  created  by  blood- 
sucking insects,  or  the  efficiency  of  disease  transmission,  has  been  suggested  by  several 
authors  (West  and  Eligh,  1952;  O’Gower,  1956;  Detinova,  1962). 

There  are  several  general  reviews  of  digestive  physiology  of  insects  (Day  and  Waterhouse, 
1953a,  b,  c;  Gilmour,  1961;  House,  1965;  Uvarov,  1929;  Waterhouse,  1 957;  Waterhouse 
and  Day,  1953;  Wigglesworth,  1965).  Related  material  has  also  appeared  in  articles  by 
Barrington  (1962),  Fallis  (1964),  House  (1958,  1961,  1962),  Lipke  and  Fraenkel  (1956), 
Snodgrass  (1935),  Vonk  (1964),  Waldbauer  (1968)  and  Wigglesworth  (1952).  The  digestive 
physiology  of  mosquitoes  was  reviewed  briefly  by  Clements  (1963)  and  this  and  the  chapter 
on  digestion  in  Wigglesworth  (1965)  constitute  the  most  extensive  reviews  published  on 
digestion  by  haematophagous  insects.  Chapters  20  and  22  in  Christophers  (1960)  contain 
some  relevant  material  but  the  discussion  of  digestion  by  adult  mosquitoes  (pp.  707-708) 
is  very  brief.  Much  of  the  work  on  digestion  by  tsetse  flies  was  reviewed  by  Buxton  (1955). 


6 


Gooding 


Within  each  of  the  following  sections  an  attempt  has  been  made  to  arrange  the  material 
more  or  less  taxonomically;  the  exopterygotes  (Hemiptera  and  Siphunculata)  are  discussed 
first  followed  by  the  Diptera  (Nematocera,  Brachycera,  then  Muscoid  flies)  and  finally  the 
Aphaniptera.  However,  to  facilitate  comparisons  it  occasionally  has  been  necessary  to  devi- 
ate from  this  arrangement. 


SIZE  OF  THE  BLOOD  MEAL 

Several  methods  have  been  used  to  estimate  blood  meal  size.  The  simplest  and  most 
widely  employed  involves  weighing  individuals  before  and  immediately  after  feeding.  Some 
workers  have  weighed  batches  of  insects  before  and  after  feeding  to  obtain  the  average 
weight  of  the  meal,  particularly  in  studies  involving  very  small  insects,  or  in  those  under- 
taken with  inadequate  equipment.  A further  modification  has  been  to  compare  the  weights 
of  batches  of  fed  and  unfed  insects;  this  has  been  particularly  useful  when  dealing  with  in- 
sects which  will  not  readily  feed  under  laboratory  conditions,  the  data  being  obtained  from 
field-caught  insects.  Blood  meal  volumes  have  usually  been  estimated  by  dividing  the  weight 
of  the  meal  by  the  specific  gravity  of  the  blood.  However  in  at  least  one  case  the  volume  was 
estimated  by  mixing  midguts  of  blood-fed  insects  with  a known  volume  of  fluid  and  meas- 
uring the  resulting  volume  (O’Connor  and  Beatty,  1937).  This  method  gave  an  unusually  low 
estimate  of  meal  size.  One  difficulty  encountered  with  all  these  procedures  is  that  some 
species  defecate  during  or  immediately  after  feeding. 

The  first  material  defecated  is  generally  from  a previous  meal  or  the  serum  of  the  meal 
just  consumed  (Boorman,  1960).  Since  erythrocytes  are  rarely  defecated,  tagging  these  with 
radioactive  cesium  or  iron  and  then  comparing  radioactivity  in  the  fed  insect  with  aliquots 
of  the  host’s  blood  has  yielded  estimates  of  the  meal  size  which  are  usually  not  influenced 
by  defecation  during  the  act  of  feeding.  Using  the  cesium  tagging  method  the  blood  meal 
size  of  Aedes  aegypti  (L.)  was  estimated  to  be  4.21  p\  compared  with  2.47  to  2.71  jul  by  the 
gravimetric  method  (Boorman,  1960).  A similar  discrepancy  in  meal  size  was  obtained  with 
Culex  pipiens  quinquefasciatus  Say  (10  p\  by  Fe59  method,  and  3.3  /il  by  gravimetric  meth- 
od) but  not  with  Triatoma  infestans  (Klug)  or  Panstrongylus  megistus  (Burmeister)  (de 
Freitas  and  Guedes,  1961).  Chemical  determination  of  the  amount  of  hemoglobin  (by  con- 
version to  alkaline  hematin)  in  the  engorged  insect  compared  with  the  hemoglobin  content 
of  the  host’s  blood  (Kershaw  et  al,  1956)  also  yields  estimates  which  are  not  influenced  by 
defecation. 

In  some  insects  Mansonia  richiardii  (Ficalbi)  and  Aedes  cinereus  Meigen  there  is  a corre- 
lation between  pre-feeding  weights  and  the  amount  of  blood  ingested,  but  in  others  ( Aedes 
cantans  (Meigen),  Aedes  detritus  (Haliday)  and  Aedes  punctor  (Kirby),  there  is  not  (Service 
1968a).  This  may  explain  some  of  the  variation  in  meal  sizes  reported  by  different  workers 
for  the  same  insect  species.  Environmental  temperature  apparently  influences  the  amount  of 
blood  consumed  by  fifth  instar  Triatoma  dimiata  (Latreille)  (174.5  mg  at  26.5  C,  281.6  mg 
at  23  C)  but  not  by  other  instars  (Zeledon  et  al,  1970).  The  source  of  the  blood  meal  may 
influence  the  quantity  of  blood  ingested  by  Pediculus  humanus  L.  (Krynski  et  al,  1952), 
Cimex  lectularius  L.  (Johnson,  1937)  and  A.  aegypti  (Bennett,  1965).  The  physiological 
state  of  Stomoxys  calcitrans  (L.)  may  influence  the  size  of  the  meal  ingested  (Anderson  and 
Tempelis,  1970).  Glossina  austeni  Newstead  irradiated  with  10  krad  as  pupae  or  15  krad  as 
teneral  adults  consumed  as  much  blood  as  non-irradiated  flies  (Langley  and  Abasa,  1970). 
Environmental  humidity  does  not  influence  the  meal  size  of  Glossina  tachinoides  Westwood 
(Buxton  and  Lewis,  1934)  or  Glossina  palpalis  (Robineau-Desvoidy)  (Mellanby,  1936).  In 
the  latter  species  the  second  meal  is  larger  than  the  first  and  older  females  tend  to  take 


Haematophagous  insects 


7 


larger  meals  than  young  females  (Mellanby,  1936). 

The  blood  meal  sizes  for  several  species  of  insects  are  presented  in  Table  I.  The  values  re- 
corded are  the  averages  reported  in  the  literature;  where  a range  is  given  this  represents  the 
range  of  average  meal  sizes  reported  in  the  literature.  Nymphal  instars  are  indicated  by  Ro: 
man  numerals. 

The  quantity  of  blood  ingested  by  the  insect  is  influenced  by  two  physiological  factors:  a 
chemical  in  the  blood  which  stimulates  the  insect  to  continue  feeding  and  stretch  receptors 
in  the  abdomen  which  inhibit  further  feeding.  Adenosine-5'-  phosphate  and  related  com- 
pounds stimulate  Culex  pipiens  (issp)  pallens  Coquillett  to  engorge  upon  blood  (Hosoi, 
1959).  Diphosphates  and  triphosphates  of  cytosine,  guanine,  inosine,  and  uridine;  creatine 
phosphate,  sodium  pyrophosphate,  riboflavin-5 '-phosphate,  5'-adenylic  acid  and  3',  5'- 
cyclic  adenylic  acid  stimulated,  prolixus  to  engorge  (Friend,  1965;  Friend  and  Smith,  1971). 
Termination  of  feeding  by  R.  prolixus  nymphs  is  apparently  determined  by  stretch  receptors 
in  the  abdomen  (Maddrell,  1963).  Nymphs  whose  nerve  cord  is  severed  between  the  pro-, 
and  mesothoracic  ganglia  will  consume  much  larger  meals  than  normal  nymphs;  nymphs 
with  a fistula  in  the  midgut  (which  permits  draining  the  midgut  during  the  act  of  feeding) 
will  consume  more  blood  over  a longer  period  of  time  than  the  controls  (Maddrell,  1963). 
By  cutting  the  ventral  cord  of  A.  aegypti  between  various  ganglia  and  then  giving  the  mos- 
quitoes a blood  meal,  Gwadz  (1969)  demonstrated  that  the  quantity  of  blood  ingested  is 
determined  by  segmental  stretch  receptors.  Similar,  but  less  extensive,  experiments  led  to 
the  .same  conclusions  for  A.  taeniorhynchus,  A.  triseriatus,  A.  subalbatus,  C.  pipiens 
quinquefasciatus  and  A.  quadrimaculatus  (Gwadz,  1969).  Lea  (1967)  reported  that  ablation 
of  the  median  neurosecretory  cells  did  not  affect  the  amount  of  blood  ingested  by  A. 
taeniorhynchus,  A.  sollicitans  ox  A.  triseriatus  but  reduced  the  blood  meal  size  of  A.  aegypti 
by  35%. 

The  nature  of  the  hunger  mechanism  has  not  been  elucidated  but  it  has  been  suggested 
that  some  blood-sucking  insects  have  an  optimal  frequency  of  feeding  (Gooding,  1960). 
Such  an  optimum  could  result  from  an  interaction  of  meal  size  with  the  rates  of  digestion, 
absorption,  and  utilization  of  some  component  of  the  meal.  The  respiratory  rate  of  G. 
morsitans  reaches  a maximum  about  24  hours  after  feeding  on  a guinea-pig.  At  1 2,  24,  and 
48  hours  after  feeding  there  is  a high  correlation  between  the  meal  size  and  the  respiratory 
rate.  Rajagopal  and  Bursell  (1966)  interpreted  this  increased  oxygen  consumption  as  being 
linked  to  the  metabolic  processes  associated  with  digestion,  absorption,  deamination,  de- 
toxication, uric  acid  synthesis  and  excretion. 

DISTRIBUTION  OF  MEALS  WITHIN  THE  ALIMENTARY  CANAL 

In  the  lice,  fleas,  and  blood-sucking  bugs  the  digestive  tract  has  no  oesophageal  diverticula 
and  the  blood  meal  is  conveyed  directly  to  the  midgut.  Blood  is  stored  in  the  expanded  ante- 
rior part  of  the  midgut  in  bugs  and  is  digested  only  in  its  posterior  reaches  (Bacot,  1915; 
Wigglesworth,  1936). 

Flies,  however,  have  from  one  to  three  oesophageal  diverticula.  Mosquitoes  usually  dis- 
patch sugar  solutions  to  the  diverticula  and  blood  to  the  midgut.  The  subject  has  been  re- 
viewed by  Trembley  (1952)  and  Megahed  (1958).  Trembley  also  presented  data  on  the  dis- 
tribution of  blood  and  sugar  solutions  in  the  various  parts  of  the  alimentary  canal  of  9 
species  of  mosquitoes  (Anopheles  freeborni  Aitken,  Anopheles  aztecus  Hoffman,  Anopheles 
quadrimaculatus,  Anopheles  albimanus,  Aedes  albopictus  Skuse,  Aedes  aegypti,  Aedes  atro- 
palpus  (Coquillet)  Aedes  vexans,  and  Culex  pipiens  Linnaeus).  The  effects  of  interrupted 
feedings,  time  after  feeding  and  the  method  of  obtaining  the  blood  meal  (i.e.  from  a droplet 


Table  1A.  Blood  meal  sizes  of  some  hemimetabolous  insects. 


8 


Gooding 


* Indicates  a jul  value,  all  others  are  in  mg. 


Haematophagous  insects 


9 


Table  IB.  Blood  meal  size  of  female  mosquitoes. 


Species 

Meal  size 

Aedes  aegypti  (L.) 

2,1-  4.5  mg 

1.6-  4.2  Ml 

Aedes  can  tans  (M eigen) 

5.9  mg 

Aedes  cinereus  Meigen 

2.6  mg 

Aedes  detritus  (Haliday) 

3.5  mg 

Aedes  hexodontus  Dyar 

<5.5  mg 

Aedes  infirmatus 

4.9  mg 

Dyar  and  Knab 

Aedes  punctor  (Kirby) 

4.0  mg 

Aedes  sollicitans  Walker 

3.9-  8.5  mg 

Aedes  sticticus  (Meigen) 

2.1  mg 

Aedes  taeniorhynchus 

3.2-  3.8  mg 

(Wiedemann) 

Aedes  triseriatus  (Say) 

3.93  mg 

Aedes  vexans  (Meigen) 

2.1-  4.7  mg 

Anopheles  albimanus 

2.6  mg 

Wiedemann 

Anopheles  litoralis  King 

2.1  mg 

Anopheles  ludlowae  (Theobald) 

2.1  mg 

Anopheles  maculatus  Theobald 

2 mg 

Anopheles  minimus  Theobald 

1 mg 

Anopheles  quadrimaculatus  Say 

2.3-  5.5  mg 

Armigeres  subalbatus 

3.7  mg 

(Coquillett) 

Culex  pipiens  quinquefasciatus 

2.0-  5.0  mg 

Say 

1.0-10.  Ml 

Culex  salinarius  Coquillett 

2.1  mg 

Culiseta  annulata  (Schrank) 

7.0  mg 

Culiseta  inornata  (Williston) 

4.5  Ml 

Mansonia  perturbans  (Walker) 

4.3  mg 

Mansonia  richiardii  (Ficalbi) 

3.3  mg 

Psorophora  ciliata  (Fabricius) 

25.0  mg 

Psorophora  confinnis 

6.0  mg 

(Lynch  Arribalzaga) 

Psorophora  cyanescens 

9.2  mg 

(Coquillett) 

Psorophora  ferox  (Humboldt) 

5.1  mg 

References 

Garnham,  1947;  Gwadz,  1969; 
Howard,  1962;  Jeffery,  1956; 
Lea,  1967;  Roy,  1936 
Bennett,  1965;  Boorman,  1960; 

Jeffery,  1956 
Service,  1968a 
Service,  1968a 
Service,  1968a 
Barlow,  1955 

Woodard  and  Chapman,  1965 
Service,  1968a 

Lea,  1967;  Woodard  and  Chap- 
man, 1965 

Stage  and  Yates,  1936 
Gwadz,  1969;  Lea,  1967; 

Woodard  and  Chapman,  1965 
Gwadz,  1969 
Stage  and  Yates,  1936; 

Woodard  and  Chapman,  1965 
Jeffery,  1956 

Laurel,  1934 
Laurel,  1934 
Laurel,  1934 
Laurel,  1934 

Gwadz,  1969;  Jeffery,  1956; 

Woodard  and  Chapman,  1965 
Gwadz,  1969 

Rachou  et  al.,  1957;  Wharton, 
1960;  Jordon  and  Goatly, 
1962;  Gwadz,  1969 
O’Connor  and  Beatty,  1937; 

de  Freitas  and  Guedes,  1961 
Woodard  and  Chapman,  1965 
Service,  1968a 
Owen  and  Reinholz,  1968 
Woodard  and  Chapman,  1965 
Service,  1968a 

Woodard  and  Chapman,  1965 
Woodard  and  Chapman,  1965 

Woodard  and  Chapman,  1965 

Woodard  and  Chapman,  1965 


10 


Gooding 


Table  1C.  Blood  meal  sizes  of  some  flies.  Data  are  from  females  except  where  otherwise 
indicated. 


Species 

Meal  size 

References 

Ceratopogonidae 

Phlebotomus  papatasi  Scopoli 

0.1  mg 

Adler  and  Theodor,  1926 

Leptoconops  kerteszi  Kieffer 

0.23  mg 

Foulk,  1967 

Simuliidae 

Prosimulium  decemarti- 

2.2  Ml 

Bennett,  1963 

culatum  (Twinn) 

Simulium  aureum  Fries 

2.9  Ml 

Bennett,  1963 

Simulium  croxtoni 

3.26  Ml 

Bennett,  1963 

Nicholson  and  Mickel 

Simulium  damnosum  Theobald 

1.08  Ml 

Crosskey,  1962 

Simulium  latipes  (Meigen) 

2.65  Ml 

Bennett,  1963 

Simulium  quebecense  Twinn 

2.14  Ml 

Bennett,  1965 

Simulium  rugglesi 

1.36  mg 

Anderson  et  al. , 1962; 

Nicholson  and  Mickel 

1.94  m1 

Bennett,  1963 

Tabanidae 

Chrysops  dimidiata 

19.7  Ml 

Kershaw  et  al. , 1956 

Van  der  Wulp 

Chrysops  silacea  Austen 

24.2  - 30  Ml 

Gordon  and  Crewe,  1953; 
Kershaw  et  al. , 1956 

40  mg 

Kershaw  et  al. , 1954 

Hybomitra  frontalis  (Walker) 

27  mg 

Miller,  1951 

(=  T.  septentrionalis ) 

Hybomitra  affinis  (Kirby) 

156  mg 

Miller,  1951 

Tabanus  quinquevittatus 

71  mg 

Tashiro  and  Schwardt,  1949 

Wiedemann 

Tabanus  sulcifrons  Macquart 

344  mg 

Tashiro  and  Schwardt,  1949 

Muscidae 

Glossina  austeni  Newstead  (d) 

11.4  mg 

Langley  and  Abasa,  1970 

(9) 

19.9  mg 

Langley  and  Abasa,  1970 

Glossina  brevipalpis  (d) 

107.  mg 

Moloo  and  Kutuza,  1970 

Newstead 

Glossina  morsitans  (d) 

34  mg 

Lester  and  Lloyd,  1928 

Westwood  (9) 

37  mg 

Lester  and  Lloyd,  1928 

Glossina  palpalis  (d) 

22.74-  33.70  mg 

Mellanby,  1936 

(Robineau-Desvoidy)  (9) 

25.13-  41.67  mg 

Mellanby,  1936 

Glossina  tachinoides  (d) 

30  mg 

Lester  and  Lloyd,  1928 

Westwood  (9) 

28  mg 

Lester  and  Lloyd,  1928 

(?) 

8.94-  14.00  mg 

Buxton  and  Lewis,  1934 

Stomoxys  calcitrans  (L.)  (d) 

6.9  - 9.45  mg 

Anderson  and  Tempelis,  1970; 
Suenaga,  1965 

(9) 

10.5  - 16.43  mg 

- Anderson  and  Tempelis,  1970; 
Suenaga,  1965 

(?) 

25.8  mg 

Parr,  1962 

Haematophagous  insects 


11 


or  through  a membrane)  were  investigated.  The  results  showed  that  the  major  factor  in  de- 
termining the  destination  of  the  meal  in  the  alimentary  canal  was  its  composition.  Similar 
results  were  reported  for  Phlebotomus  papatasi  (Adler  and  Theodor,  1926),  Simulium  dam- 
nosum  (Lewis,  1953),  Simulium  venustum  Say  (Yang  and  Davies,  1968b),  Culicoides  nube- 
culosus  Meigen  (Megahed,  1956,  1958),  Chrysops  and  Tabanus  (Wigglesworth,  1931).  Al- 
though Anopheles  maculipennis  Meigen  generally  dispatched  blood  to  the  midgut  only, 
about  1/3  of  the  mosquitoes  also  had  some  blood  in  the  ventral  diverticulum  (Wright, 
1924).  It  has  been  claimed  that  A.  aegypti  but  not  A.  albimanus  (reported  as  Anopheles 
tarsimaculatus)  passed  blood  first  into  the  diverticula  and  then  within  half  an  hour  of  feed- 
ing into  the  midgut  (Pawan,  1937).  A honey  - citrated  blood  mixture  fed  to  A maculipennis, 
C.  pipiens,  and  A.  aegypti  (recorded  zsAedes  argenteus ) went  to  the  diverticula.  The  ventral 
diverticulum  filled  first,  followed  by  the  dorsal  diverticula  after  unusually  large  meals  (Mac- 
Gregor and  Lee,  1929).  Radiographic  studies  in  situ  of  the  distribution  of  sugar  solutions 
and  blood  meals  in  the  digestive  tract  of  A.  aegypti  showed  that  sugar  meals  could  be  moved 
from  one  diverticulum  to  another  and  that  during  ingestion  of  a blood  meal  the  hind  most 
part  of  the  midgut  filled  first  (Guptavanij  and  Venard,  1965). 

Among  some  flies  however,  ( Glossina  sp.,  Wigglesworth,  1931;  Stomoxys  calcitrans, 
Champlain  and  Fisk,  195 6;  Hippelates  pallipes  (Loew),  Kumm,  1932)  the  blood  meal  goes 
first  to  the  oesophageal  diverticulum  and  then  to  the  midgut.  In  the  eye  gnat,  H.  pallipes, 
only  a small  quantity  of  blood  goes  directly  to  the  midgut.  Within  30  mins  of  feeding  the 
gnats  begin  transfering  blood  from  the  crop  to  the  midgut  and  by  6 hours  after  feeding 
about  half  of  the  meal  is  transferred.  The  blood  blackens  in  the  midgut  as  digestion  takes 
place.  By  2 days  after  feeding  the  crop  is  empty  (Kumm,  1932).  The  first  portion  of  a 
blood  meal  ingested  by  tsetse  flies  ( Glossina  morsitans  and  Glossina  tachinoides ) is  conveyed 
to  the  midgut  where  it  is  localized  in  3 regions,  a clot  forming  in  the  most  posterior.  Addi- 
tional blood  is  dispatched  to  the  crop  (esophageal  diverticulum)  but  it  does  not  clot  there. 
By  2.5  hours  after  feeding  most  of  the  blood  passes  from  the  crop  into  the  midgut  which  is 
now  more  or  less  continuously  full  of  blood.  The  blood  at  the  posterior  end  of  the  midgut 
clots  and  progressively  darkens  while  the  blood  in  the  anterior  of  the  midgut  does  not  clot 
and  is  bright  red.  By  24  hours  after  the  meal  the  crop  is  empty  and  the  blood  in  the  midgut 
is  again  divided  into  regions  by  folds  in  the  gut.  The  blood  in  the  anterior  part  of  the  midgut 
forms  a pasty  mass  as  a result  of  absorption  of  most  of  the  serum  but  a true  clot  does  not 
form.  The  blood  cells  in  this  region  of  the  midgut  are  contained  in  a mucilaginous  secretion 
from  the  gut  cells.  By  48  hours  the  blood  mass  in  the  gut  greatly  decreases  and  the  dark 
mass  progressively  contracts  posteriorly  (Lester  and  Lloyd,  1928). 

When  blood  only  is  fed  to  A.  aegypti  it  is  dispatched  to  the  midgut  in  all  individuals, 
traces  being  found  in  the  diverticula  of  only  6%  of  the  mosquitoes  (Day,  1954).  Increasing 
the  concentration  of  glucose  in  a blood-glucose  mixture  increases  the  frequency  with  which 
meals  go  to  the  diverticula.  At  concentrations  above  0.46  M glucose  all  mosquitoes  send  the 
meal  to  the  diverticula,  only  7 to  10%  also  convey  food  to  the  midgut.  These  results  indicate 
that  there  are  receptors  in  this  mosquito  for  both  glucose  and  some  component  of  blood. 
A.  aegypti  can  detect  sucrose  and  probably  arabinose,  mannose,  and  raffinose  but  not  lac- 
tose when  mixed  with  blood.  Although  both  plasma  and  erythrocytes  are  detected  solutions 
of  haemoglobin  and  albumin  with  glucose  go  mainly  to  the  diverticulum.  Mosquitoes  feeding 
on  very  dilute  erythrocyte  suspensions  in  sugar  dispatch  the  sugar  solution  to  the  diverticu- 
lum and  the  erythrocytes  to  the  midgut.  This  is  apparently  accomplished  by  a group  of 
spines  in  the  neck  of  the  diverticulum  which  are  capable  of  acting  as  a seiving  mechanism 
when  particles  are  sparse.  Experiments  with  red  cell  ghosts  and  fly  sarcosomes  in  water 
indicate  that  the  particulate  nature  of  the  erythrocytes  is  one  factor  in  blood  detected  by 


12 


Gooding 


the  mosquito.  Day  (1954)  explained  the  distribution  of  fluids  by  proposing  that  the  pit 
organs  in  the  buccal  cavity  detect  sugars  and  the  resulting  impulses  mediate  the  relaxation  of 
the  diverticula  sphincter  muscles.  Similarly,  the  papillar  sense  organs  detect  components  of 
the  blood  and  relaxation  of  the  cardiac  sphincter  permits  blood  to  enter  the  midgut. 

Hosoi  (1954)  reported  upon  the  mechanism  by  which  Culex  pipiens  (issp)  pallens  distrib- 
utes fluids  to  the  midgut  and  diverticula.  As  in  other  mosquitoes,  sugary  solutions  go  to  the 
diverticula  and  blood  to  the  midgut.  Dilute  suspensions  of  erythrocytes  in  saline  go  to  the 
midgut  but  if  glucose  is  added  to  the  meal  there  is  an  increased  tendency  to  dispatch  the 
meal  to  the  diverticula.  Replacing  the  erythrocytes  with  other  particulate  matter  in  a 5% 
glucose  solution  generally  results  in  the  meal  being  dispatched  to  the  diverticula  - thus 
indicating  that  the  particulate  nature  of  a blood  meal  is  not  the  only  stimulus  for  dis- 
patching food  to  the  midgut.  Hosoi  suggested  that  erythrocytes  have,  adsorbed  to  their 
surfaces,  substances  which  are  responsible  for  stimulating  certain  sense  organs.  Sensory 
receptors  on  the  labium  respond  to  glucose  but  not  to  erythrocytes  while  the  reverse  is  true 
for  receptors  on  the  fascicle.  However  the  existence  of  other  sense  organs  capable  of  differ- 
entiating the  meal  composition  was  not  excluded. 

Theories  concerning  the  biological  importance  of  the  retention  of  sugars  in  the  diverticula 
were  reviewed  by  Trembley  (1952),  Megahed  (1958),  and  Christophers  (1960,  p.  489).  Two 
of  these  theories  pertain  to  digestive  physiology.  One  is  that  the  carbohydrate  meal  is  stored 
in  the  diverticula  so  that  the  hunger  mechanism,  which  is  presumed  to  originate  in  the  mid- 
gut, is  not  interfered  with,  and  the  mosquito  is  thus  always  ready  to  take  blood  (Day,  1954). 
The  other  is  that  by  storing  carbohydrate  solutions  in  impervious  structures  the  mosquito 
carries  with  it  a supply  of  water  which  may  be  passed  to  the  midgut  for  absorption  as 
needed.  If  the  second  theory  is  correct,  then  the  mosquito’s  physiology  is  adapted  to  con- 
serving water  from  a carbohydrate  meal  while  disposing  of  much  of  the  water  in  a blood 
meal  (Boorman,  1960;  Howard,  1962).  Denisova  (1949)  investigated  the  function  of  horse- 
fly diverticula  by  injecting  water  and  salt  solutions  into  the  flies.  She  concluded  that  the 
diverticula  store  water  which  is  supplied  in  small  amounts  to  the  midgut  minimizing  rapid 
drops  in  haemolymph  osmotic  pressure.  However,  it  seems  doubtful  that  nectar  solutions, 
with  their  high  sugar  content,  would  significantly  lower  the  osmotic  pressure  of  insect 
haemolymph  if  dispatched  directly  to  the  midgut. 

The  inhibition  of  honeybee  proteolytic  enzymes  by  honey  (Bailey,  1952)  suggests  that 
nectar  may  contain  substances  which  inhibit  insect  proteinases.  The  diverticula  of  insects 
which  consume  both  nectar  and  blood  may  thus  function  as  a mechanism  for  separating 
inhibitors  present  in  one  type  of  meal  from  the  digestive  enzymes  required  to  digest  another 
type  of  meal.  MacGregor  (1930)  stated  that  “Poisonous  fluids  invariably  enter  the  diverticu- 
la” of  mosquitoes  and  gave  as  an  example  the  ingestion  of  20%  formalin.  The  mosquitoes 
died  immediately  after  the  diverticula  filled  and  traces  of  formalin  passed  into  the  midgut. 
This  was  interpreted  as  indicating  that  absorption  did  not  take  place  from  the  diverticula. 

THE  SALIVARY  GLANDS  AND  THEIR  SECRETIONS 

Since  saliva  is  usually  the  first  insect  secretion  to  which  the  blood  meal  is  exposed  there 
has  been  considerable  interest  in  the  effects  of  saliva  upon  blood.  Bates  (1949)  suggested 
that  anticoagulins  and  haemagglutinins  from  the  salivary  glands  of  mosquitoes  assist  in  the 
preliminary  breakdown  of  the  blood  meal.  Fisk  (1950)  suggested  that  coagulation  or  agglu- 
tination of  the  blood  meal  denatures  its  proteins  sufficiently  to  permit  attack  by  mosquito 
proteinases.  Regrettably  these  suggestions  have  not  been  investigated. 

Aedes  aegypti  feeding  upon  suckling  mice  left  an  average  of  4.7  pg  of  saliva  in  the  mice 


Haematophagous  insects 


13 


during  consumption  of  a meal  (Devine  et  al,  1965).  Probably  because  of  the  minute  amount 
of  saliva  secreted  by  blood-sucking  insects,  most  investigators  have  used  homogenates  of  the 
salivary  glands  to  test  for  a variety  of  enzymes,  haemolysins,  agglutinins,  and  anticoagulins. 
The  most  commonly  encountered  are  agglutinins  and  anticoagulins;  a summary  of  their 
occurrence  among  the  blood-sucking  insects  is  given  in  table  2. 

The  ability  of  salivary  gland  emulsions  to  cause  agglutination  sometimes  depends  upon 
the  source  of  the  erythrocytes.  A.  maculipennis  salivary  glands  agglutinate  erythrocytes 
from  man,  donkey,  rabbit,  and  dogs,  but  not  those  from  mice,  guinea-pigs,  or  monkey 
(York  and  Macfie,  1924;  confirmed  for  white  mice  by  Shute,  1935).  The  agglutinin  from  A. 
quadrimaculatus  is  effective  against  red  blood  cells  of  man,  mule,  cow,  pig,  dog,  rabbit, 
guinea-pig,  rat,  and  mouse  but  not  chicken  or  turtle  (Metcalf,  1945). 

Agglutinins  are  restricted  to  the  median  acinus  of  the  salivary  glands  of  A.  maculipennis 
(de  Buck  1937)  and  A.  quadrimaculatus  (Metcalf,  1945)  but  occur  in  all  three  acini  of  C. 
annulata  salivary  glands  (de  Buck  1937).  Solutions  of  these  agglutinins  are  heat  labile  (York 
and  Macfie,  1924;  de  Buck  1937;  Metcalf,  1945).  The  agglutinin  from  A maculipennis  was 
reported  by  York  and  Macfie  (1924)  to  be  inactivated  by  desiccation,  but  by  de  Buck 
(1937)  to  be  stable  for  several  months  at  room  temperature  or  for  1 hour  at  99  C when 
dried.  Similar  properties  for  the  agglutinin  from  C.  annulata  were  reported  by  de  Buck 
(1937). 

Baptist  (1941)  reported  that  the  anticoagulin  in  R.  prolixus  salivary  glands  was  inacti- 
vated by  heating  to  70  C but  Hellmann  and  Hawkins  (1964)  reported  this  anticoagulin  was 
stable  at  60  or  80  C for  30  mins,  was  not  affected  by  0.1  N HC1  or  0.1  N NaOH  at  R.T.  or 
60  C for  30  mins  but  was  completely  destroyed  by  heating  to  100  C for  5 mins.  It  was  not 
precipitated  by  centrifuging  at  100,000  g for  30  mins  but  was  removed  from  solution  by 
dialysis  in  the  cold.  This  anticoagulin  (designated  Prolixin-S  by  Hellmann  and  Hawkins, 
1965)  did  not  inhibit  thrombin,  but  acted  mainly  upon  factor  VIII  (the  antihaemophilic 
factor).  Prolixin-S  retained  its  activity  upon  freezing,  freeze  drying  or  dialysis  but  was 
inactivated  by  trypsin. 

Anticoagulin  cannot  be  detected  in  R.  prolixus  midgut  immediately  after  feeding  but  is 
found  4 hours  later.  Fractionation  of  the  gut  contents  indicate  that  the  anticoagulin  is 
probably  present  but  inhibited  by  some  component  of  the  blood  meal  (Hellmann  and 
Hawkins,  1965).  A salivary  gland  anticoagulin  from  T.  rubofasciata  is  inactivated  by  normal 
rabbit  serum  but  not  by  serum  from  a rabbit  used  as  a host  for  one  year  (Cornwall  and 
Patton,  1914).  A salivary  gland  anticoagulin  from  T.  maculata  is  not  an  anti-thrombin  and  is 
slightly  less  heat  stable  than  that  of  R.  prolixus  (Hellmann  and  Hawkins,  1966). 

Anticoagulin  occurs  in  all  three  acini  of  the  salivary  glands  of  C.  pipiens  (de  Buck  1937), 
A.  quadrimaculatus  (Metcalf,  1945),  A.  plumbeus  and  A.  maculipennis  (de  Buck,  1937). 
However,  in  the  latter  species  its  concentration  in  the  lacteral  acini  is  very  low.  The  antico- 
agulin from  A.  maculipennis  is  stable  at  R.T.  for  several  months  when  dried  and  is  heat 
stable  as  either  a desiccated  preparation  (99  C for  1 hr)  or  saline  solution  (100  C for  35 
mins)  (de  Buck,  1937).  Anticoagulins  from  C.  pipiens  and  C.  annulata  are  stable  when 
desiccated  but  are  more  heat  labile  than  the  anticoagulin  from  A.  maculipennis  (de  Buck 
1937).  The  A.  quadrimaculatus  anticoagulin  is  thermostable  (Metcalf,  1945). 

The  anticoagulins  from  the  salivary  glands  of  G.  tachinoides  and  G.  morsitans  are  non- 
dialyzable  and  that  of  the  former  species  is  not  markedly  affected  by  0. 1 N KOH  or  0. 1 N 
HC1.  They  are  stable  at  temperatures  up  to  90  C but  at  this  temperature  lose  half  their 
activity  in  15  mins  and  all  activity  in  30  mins;  at  100  C all  activity  is  lost  in  15  mins.  From 
experiments  with  fibrinogen  solutions  and  citrated  blood,  Lester  and  Lloyd  (1928)  sug- 
gested that  the  tsetse  fly  anticoagulin  is  an  antikinase.  They  found  that  although  the  delay 


Table  2.  Agglutinins,  coagulins  and  anticoagulins  in  some  blood-sucking  insects. 


14 


Gooding 


Present  +,  not  found  — , those  not  checked  for  are  left  blank;  1 present  in  head  and  thorax,  2 present  in  abdomen. 


Table  2 (continued) 


Haematophagous  insects  15 


Table  2 (continued) 


16 


Gooding 


Haematophagous  insects 


17 


in  clotting  of  sheep’s  blood  is  a function  of  the  number  of  glands  present,  a simple  straight 
line  relationship  is  not  obtained. 

The  anticoagulin  in  the  salivary  glands  of  M.  crassirostris  (reported  as  Philoematomyia 
insignis)  is  only  slightly  inactivated  by  heating  to  100  C for  10  mins.  This  anticoagulin  is 
probably  non-antigenic  in  rabbits  and  rats.  However  normal  sera  of  calf,  rat,  and  rabbit 
contain  one  or  more  substances  which  inactivate  the  anticoagulin.  Unlike  sodium  citrate 
inhibition  of  coagulation,  the  anticoagulin  from  M.  crassirostris  is  not  overcome  by  addition 
of  CaCl2 . Anticoagulin  activity  of  female  salivary  glands  is  greater  than  that  of  males.  Newly 
emerged  adults  appear  to  have  less  anticoagulin  than  older  adults  which  have  had  an  oppor- 
tunity to  feed  (Cornwall  and  Patton,  1914). 

Hemolytic  activity  of  salivary  gland  emulsions  has  never  been  demonstrated,  although  it 
has  been  tested  for  in  C.  pipiens  (Nuttall  and  Shipley,  1903;  Yorke  and  Macfie,  1924),  A. 
aegypti  (Yorke  and  Macfie,  1924;  McKinley,  1929),  C.  annulata,  and  G.  tachinoides  (Yorke 
and  Macfie,  1924),  and  P.  papatasi  (Adler  and  Theodor,  1926). 

Although  salivary  glands  have  frequently  been  examined  for  digestive  enzymes  these  have 
rarely  been  demonstrated.  Salivary  glands  of  C.  lectularius,  R.  prolixus,  and  T.  infestans  do 
not  have  demonstrable  amounts  of  protease,  lipase,  invertase,  or  amylase  (Baptist,  1941). 
Although  esterases  occur  in  the  salivary  gland  tissues  of  Anopheles  freeborni  and  A.  aegypti 
they  are  not  demonstrable  in  the  salivary  secretions  (Frevogel,  Hunter  and  Smith,  1968). 
Roy  (1937)  examined  the  salivary  glands  of  G.  intestinalis  and  found  no  amylase,  lipase, 
maltase,  lactase,  pepsin  or  trypsin,  but  he  did  find  a milk-clotting  (proteolytic)  enzyme. 
The  existance  of  this  milk  clotting  enzyme  was  confirmed  by  Tatchell  (1958)  who  also 
demonstrated  invertase,  maltase,  and  amylase;  the  later  with  a pH  optimum  at  6.  Wiggles- 
worth  (1929)  reported  that  Glossina  morsitans  submorsitans  Newstead  and  G.  tachinoides 
salivary  glands  did  not  contain  amylase,  invertase,  maltase,  lactase,  trypsin,  pepsin,  or  pepti- 
dase. However,  the  salivary  glands  of  Glossina  austeni  contain  a factor  which  activates 
plasminogen  (Hawkins,  1966).  This  or  a similar  material  is  also  found  in  the  crop,  midgut, 
and  hindgut  where  presumably  it  contributes  to  the  lysis  of  the  blood  clot.  A weak  amylase 
occurs  in  S.  calcitrans  salivary  glands  (Champlain  and  Fisk,  1956).  However,  Rostonand 
Gamal-Eddin  (1961)  found  no  evidence  for  amylase  or  proteinase  (active  at  pH  6,  7,  or  8.3) 
in  the  salivary  glands  of  S.  calcitrans,  Stomoxys  sitiens  Rondani,  or  Musca  vitripennis 
Meigen.  The  salivary  glands  of  larvae  of  Protocalliphora  avium  Shannon  and  Dobroscky 
(reported  as  Apaulina  avium ) contain  butyrase  and  a weak  maltase  (Rockstein  and  Kamal, 
1954). 

Metcalf  (1945)  found  no  evidence  of  protease,  lipase,  amylase,  or  lecithinase-A  in  the 
salivary  glands  of  A.  quadrimaculatus.  Wigglesworth  (1931)  reported  that  Chrysops  and 
Glossina  salivary  glands  contained  a lipase.  No  invertase  occurs  in  the  salivary  glands  of 
Prosimulium  fuscum  Syme  & Davies,  or  Simulium  venustum  (Yang  and  Davies,  1968c). 

No  cytological  changes  were  detected  in  the  salivary  glands  of  A.  aegypti  immediately 
after  feeding  on  sugar  or  human  blood  but  histochemical  changes  observed  in  the  glands 
during  the  24  hours  following  a blood  meal  led  to  the  conclusion  that  “feeding  depletes  the 
glands  and  that  this  depletion  leads  to  the  resynthesis  of  secretory  products”  (Orr,  Hudson 
and  West,  1961). 

The  salivary  glands  of  female  blackflies  (probably  S.  venustum ) contain  a Periodic  acid- 
Schiff  positive  material  prior  to  the  time  a blood  meal  is  taken.  This  material  is  present  in 
only  small  amounts  in  the  glands  immediately  after  feeding,  but  reappears  during  the  next 
96  hours  (Gosbee,  Allen,  and  West,  1969). 

A major  function  of  anticoagulins  in  saliva  appears  to  be  the  prevention  of  premature 
clotting  of  the  blood  meal.  Lloyd  (1928)  reported  that  removal  of  the  salivary  glands  of  G. 


18 


Gooding 


tachinoides  did  not  prevent  feeding  on  humans,  but  sooner  or  later  the  flies  died  with  a 
blood  clot  in  the  proboscis  or  oesophagus.  Aedes  stimulans  (Walker)  whose  salivary  ducts 
had  been  cut  ingested  blood  in  a normal  manner  and  developed  eggs.  No  observable  differ- 
ence between  the  blood  clot  in  the  midgut  of  operated  and  control  mosquitoes  could  be 
detected  (Hudson,  Bowman,  and  Orr,  1960). 

GROSS  CHANGES  IN  QUANTITY  & QUALITY  OF  THE  GUT  CONTENTS. 

Mosquitoes  feeding  on  chicks  with  heavy  Plasmodium  gallinaceum  Brumpt  infections  may 
defecate  20  to  50%  of  the  meal  within  a few  hours  (Howard,  1962).  The  discharge  of  blood 
during  and  following  feeding  seems  to  favour  the  elimination  of  gametocytes  thus  reducing 
the  intensity  of  the  infection  in  the  mosquitoes  (Mitzmain,  1917).  When  undisturbed  Ano- 
pheles gambiae  Giles  and  Anopheles  funestus  Giles  will  engorge  until  blood  is  passed  from 
the  anus;  sometimes  the  amount  of  blood  passed  is  at  least  half  the  volume  ingested 
(Hocking  and  Maclnnes,  1949).  Neither  the  coagulation  time  nor  the  blood  corpuscles  are 
affected  by  this  rapid  passage  through  these  mosquitoes.  The  fluid  defecated  by  Aedes 
aegypti  during  and  just  after  feeding  contains  uric  acid,  simple  proteins  or  amino  acids,  and 
occasionally  red  blood  corpuscles  but  no  reducing  sugars  (Boorman,  1960).  The  volume  of 
the  fluid  passed  is  approximately  1.5  p\;  some  of  this  comes  from  the  serum  while  the 
remainder  (actually  the  first  few  drops  passed)  containing  uric  acid,  is  probably  present  in 
the  hind  gut  at  the  time  of  feeding. 

The  discharge  of  material  from  the  anus  during  and  just  after  feeding  has  been  observed  in 
Pediculus  humanus  (Nuttall,  1917b),  tsetse  flies  (Lester  and  Lloyd,  1928),  and  Culicoides 
nubeculosus  (Megahed,  1958). 

The  relatively  great  changes  in  the  size  and  shape  of  the  abdomens  of  some  blood-sucking 
insects  occurring  during  feeding  and  the  translucence  of  the  abdominal  pleura  have  permit- 
ted observations  on  digestion  with  no  more  than  a dissecting  scope  or  hand  lens.  Frequently, 
these  observations  could  be  made  in  more  detail  by  simply  dissecting  the  insect  and  without 
the  use  of  elaborate  histological  techniques. 

Sella  (1920a)  divided  the  digestion  of  blood  and  the  development  of  the  ovaries  into  7 
stages.  These  stages  were  summarized  by  Detinova  (1962,  p.  57)  who  also  reviewed  aspects 
of  blood  digestion  relating  to  age-grouping  methods  and  studies  of  ovarian  development. 
Modification  of  Sella’s  (1920a)  method  have  been  developed  by  several  workers  (Hocking 
and  Maclnnes,  1949  for  Anopheles',  Jackson,  1933  for  Glossina;  Linley,  1965  for  Leptoco- 
nops ). 

Blood  ingested  by  Culex  pipiens  quinquefasciatus  (C.  fatigans)  turns  black  within  6 hours 
and  the  abdomen  is  3/4,  2/3,  1/2,  slightly  less  than  1/2,  and  less  than  1/3  distended  at  12, 
24,  36,  48  and  60  hours  after  a blood  meal  (O’Gower,  1956).  In  the  tropics  Anopheles 
species  generally  take  2 days  to  digest  a blood  meal  and  develop  eggs  but  Anopheles  vagus 
Donitz  requires  only  24  hours  to  accomplish  these  same  processes  in  Assam,  India  (Muirhead- 
Thomson,  1951). 

Chicken  blood  ingested  by  A.  aegypti  clots  in  about  30  minutes  compared  to  25  min- 
utes for  the  same  volume  on  a glass  slide  (Howard,  1962).  By  45  minutes  a clear  yellow- 
ish border  forms  around  ingested  blood  and  this  adheres  to  the  surfaces  of  both  the  midgut 
cells  and  the  blood  meal.  This  adherence  persists  for  about  12  hours  by  which  time  the 
peritrophic  membrane  is  being  formed.  By  24  hours  the  meal  becomes  a comparatively  firm 
clot,  and  although  its  volume  decreases  its  consistency  does  not  change  after  that  time.  By 
72  hours  after  feeding  the  midgut  is  usually  empty. 

Gross  changes  in  the  appearance  of  the  blood  meal  of  A.  aegypti  fed  on  a rat  have 


Haematophagous  insects 


19 


been  reported  by  Akov  (1965).  The  blood  is  bright  red  immediately  after  ingestion.  As 
digestion  proceeds  inwards  from  the  periphery  the  blood  meal  turns  brown.  By  24  hours 
the  meal  is  brown  except  for  the  center.  Akov  considered  digestion  to  be  complete  and 
elimination  of  the  residue  to  begin  when  the  color  of  the  blood  was  brown  throughout. 
On  this  basis  digestion  is  complete  and  defecation  begins  by  36  hours.  The  size  of  the 
blood  clot  decreases  and  the  midgut  wall  becomes  folded.  By  42  hours  after  feeding  2/3 
of  the  females  have  empty  midguts  and  1/3  contain  small  residues  of  blood.  By  48  hours 
about  1/4  of  the  mosquitoes  contain  residues  in  the  midgut  or  hindgut.  Elimination  of 
blood  residues  is  complete  in  all  mosquitoes  by  54  hours  after  feeding. 

Feeding  increasing  concentrations  of  5-fluorouracil  (5-FU)  to  A.  aegypti  for  2 days 
prior  to  a blood  meal  results  in  an  increasing  tendency  for  the  mosquitoes  to  retain  un- 
digested blood  in  the  midgut  (Akov  1965).  However,  the  rate  of  digestion  of  a subsequent 
blood  meal  is  normal.  Similar  results  are  produced  with  A,  aegypti  (1)  fed  metepa,  apholate, 
or  tepa  for  2 days  prior  to  a blood  meal,  (2)  held  in  contact  with  a metepa  treated  glass 
surface  or  (3)  irradiated  with  from  2,000  to  32,000  r (from  a Co60  source)  (Akov,  1966). 
Irradiation  of  the  mosquitoes  does  not  influence  the  rate  at  which  Evan’s  blue  stained 
Dextran  passes  from  the  midgut.  This  indicates  that  the  delay  in  emptying  the  midgut  of  a 
blood  meal,  by  A.  aegypti  treated  with  irradiation  or  chemosterilants,  is  due  not  to  an  effect 
upon  the  gut  mobility  but  upon  the  rate  at  which  the  digestion  products  are  absorbed  and 
utilized  (Akov,  1966).  In  mosquitoes  of  the  same  age,  the  evacuation  rate  is  lower  during 
the  second  meal  than  during  the  first. 

There  are  individual  differences  in  the  rate  of  digestion  by  Anopheles  punctulatus  Donitz 
(MacKerras  and  Roberts,  1947).  In  this  species  about  25%  of  the  females  still  have  blood  in 
the  midgut  72  hours  after  feeding.  MacKerras  and  Roberts  suggested  that  if  a large  amount 
of  blood  is  present  at  48  hours  the  Plasmodium  Marchiafava  and  Celli  ookinetes  may  be 
imprisoned  in  this  mass.  The  mean  time  for  gut  emptying  is  48  hours  for  A.  gambiae  and  60 
hours  for  A.  funestus  with  considerable  variation  in  the  rate  of  digestion  in  both  species 
(Hocking  and  Maclnnes,  1949).  In  A.  gambiae  evacuation  time  varies  from  24  to  72  hours; 
in  A.  funestus  from  24  to  96  hours.  Anopheles  maculipennis  whose  ovaries  are  developing 
digest  their  blood  meal  in  73.4  to  87.1  hours,  while  parous  females  whose  ovaries  are  not 
developing  require  57.7  to  60  hours  (Detinova,  1962).  These  findings  led  Detinova  to  sug- 
gest that  neuro-hormonal  regulation  of  digestion  may  occur  in  mosquitoes.  There  is  a slight 
but  significant  decrease  in  the  digestion  rate  with  aging  and  Detinova  (1962)  reported  that 
it  takes  longer  to  digest  the  first  meal  than  to  digest  subsequent  meals.  This  latter  finding  is 
the  opposite  of  that  reported  for  A.  aegypti  (Akov,  1966). 

Digestion  of  blood  is  quite  different  in  “long  winged”  and  “short  winged”  A.  maculi- 
pennis when  they  are  overwintering  (de  Buck,  Shoute,  and  Swellengrebel,  1932).  For  at 
least  the  first  24  hours,  the  blood  ingested  by  the  “long  winged”  females  is  divided  into 
an  anterior  translucent  half  and  a posterior,  opaque,  cellular  mass.  The  erythrocytes  show 
very  little  agglutination  when  the  midgut  is  dissected  in  saline.  At  24  to  26  C,  digestion 
of  the  meal  proceeds  very  slowly,  requiring  5 or  6 (occasionally  10)  days  before  the  red 
colour  (indicating  undigested  blood)  disappears.  In  the  “short-winged”  females  the  sequence 
of  events  is  quite  different.  Within  half  an  hour  of  feeding  the  serum  is  absorbed  from  the 
lumen  of  the  midgut  leaving  the  cells  in  close  contact  with  the  gut  epithelium.  There  is  a 
marked  agglutination  of  red  cells  and  the  digestion  of  the  meal  proceeds  rapidly  if  the 
temperature  is  sufficiently  high.  If  given  more  than  one  meal,  a certain  portion  of  the  “long 
winged”  population  digests  blood  in  the  same  manner  as  the  “short-winged”  females  (de 
Buck,  Torren,  and  Swellengrebel,  1933).  The  differences  in  digestion  by  “long-winged”  and 
“short-winged”  A.  maculipennis  are  not  observed  during  the  summer. 


20 


Gooding 


Variations  in  digestion  rate  occur  in  Anopheles  claviger.  These  are  correlated  with  the 
season  (Sella,  1920a)  and  with  environmental  temperature  (Sella,  1920b).  In  A.  maculi- 
pennis  the  rate  of  digestion  of  the  blood  meal  slows  down  as  the  temperature  falls  in  the 
autumn,  (Guelmino,  1951).  The  unchanged  condition  of  the  red  cells  suggests  that  diges- 
tion and  absorption  in  the  fall  are  limited  to  water  soluble  materials  that  can  be  used  to 
develop  adipose  tissue  (Guelmino,  1951). 

Detinova  (1962  p.  57),  reviewing  some  aspects  of  the  digestive  process  in  A.  maculi- 
pennis,  stated  that  “if  the  temperature  rises  to  the  optimum  the  speed  of  the  processes 
increases,  but  at  temperatures  above  the  optimum  they  slow  down”. 

Culiseta  annulata  taken  from  the  Poole  area  of  Dorset,  southern  England  require  4 weeks 
to  digest  a blood  meal  “at  temperatures  experienced  in  November  - February”  (Service, 
1968b).  Other  species  (A.  claviger,  C.  annulata,  Mansonia  richiardii,  Aedes  dorsalis  (Meigen), 
Aedes  geniculatus  (Olivier),  Aedes  detritus,  Aedes  punctor,  Aedes  cantans,  and  Aedes  cine- 
reus)  in  this  same  area  (Brownsea  Island,  Dorset)  take  5 to  8 days  to  digest  a meal  of  human 
blood  during  May  to  August  or  September  (Service,  1968a). 

Australasian  anophelines  complete  digestion  of  a blood  meal  in  3 to  4 days  under  summer 
conditions  (Roberts  and  O’Sullivan,  1949).  Buttiker  (1958)  observed  Anopheles  culicifacies 
Giles  and  Anopheles  aconitus  Donitz  in  several  locations  in  southeast  Asia  which  were  in  a 
quiescent  state  and  whose  midguts  contained  a blood  meal  which  was  “dark  red,  coagulated, 
very  hard  and  almost  completely  dessiccated.” 

The  digestion  of  blood  by  a mosquito  may  be  influenced  by  other  materials  fed  to  the 
mosquito.  When  various  species  of  Anopheles  are  fed  alternately  upon  gametocyte  carri- 
ers and  bananas,  many  mosquitoes  die  with  undigested  masses  of  blood  in  their  midguts 
(Darling,  1910).  Some  A.  aegypti  fed  upon  CaCl2,  MgCl2,  or  either  of  these  plus  oxyte- 
tracycline  before  and  after  ingesting  blood  from  a chicken,  still  have  a residue  of  blood  in 
the  midgut  four  days  later  when  held  at  26.5  C and  75%  R.H.,  conditions  under  which 
digestion  is  normally  complete  in  3 days  (Terzian,  1958).  In  mosquitoes  with  inhibited 
digestive  processes  these  undigested  residues  are  about  1/3  the  size  of  the  original  clot, 
are  orange-red,  contain  no  intact  red  cells  and  give  a positive  heme  test  (benzidine  reac- 
tion). (The  positive  benzidine  test  indicating  the  presence  of  intact  heme  groups  is  not 
surprising  in  view  of  O’Gower’s  (1956)  finding  that  mosquito  feces  give  a positive  benzi- 
dine test).  Inhibition  of  digestion  can  be  prevented  by  adding  the  chelating  agent  ethylene 
diamine  tetraacetic  acid  to  the  salt-antibiotic  mixture.  Since  mosquitoes  with  a large  residue 
of  undigested  blood  are  able  to  produce  large  numbers  of  eggs,  Terzian  (1958)  concluded 
that  it  was  “reasonable  to  assume  that  only  digestion  of  the  hemoglobin  fraction,  of  all  the 
fractions  contained  in  the  original  blood  meal,  is  affected  by  the  cations,  or  cation  antibiot- 
ic mixture.”  The  validity  of  this  assumption  is  questionable  in  view  of  the  finding  that  the 
in  vitro  activity  of  A.  aegypti  proteinase  upon  both  hemoglobin  and  serum  albumin  is 
inhibited  by  CaCl2,  MgCl2,  and  MnCl2  (Gooding,  1966a).  Terzian  (1963)  expanded  the  in 
vivo  inhibition  studies  to  include  both  A.  aegypti  and  Anopheles  quadrimaculatus,  4 cations, 
and  4 antibiotics.  Digestion  of  blood  is  inhibited  to  varying  degrees  in  both  species  by  the 
presence  of  calcium,  magnesium,  manganese  or  iron  in  sugar  solutions  consumed  prior  to  the 
blood  meal.  The  action  of  the  antibiotics  is  very  complex.  Oxytetracycline  by  itself  has  very 
little  effect  on  digestion  by  A.  aegypti  but  potentiates  the  action  of  calcium,  magnesium, 
and  manganese.  It  supresses  inhibition  by  iron.  However,  oxytetracycline  has  a marked 
inhibitory  effect  on  A.  quadrimaculatus  digestion,  potentiates  manganese  only  and  inhibits 
the  effect  of  iron.  Penicillin  reduces  the  effects  of  calcium  and  magnesium  upon  A.  aegypti, 
but  has  itself  an  inhibitory  effect  on  A.  quadrimaculatus  digestion.  A mixture  of  chloram- 
phenicol and  dihydrostreptomycin  in  the  diet  of  C.  pipiens  quinquefasciatus  slows  down 


Haematophagous  insects 


21 


digestion  of  canary  blood  infected  with  Plasmodium  relictum  Grassi  and  Feletti  to  such  an 
extent  that  about  1/3  of  the  mosquitoes  still  have  blood  in  the  midgut  after  7 days  (Micks 
and  Ferguson,  1961).  The  results  of  these  experiments  with  antibiotics  are  interesting  in 
view  of  Arnal’s  (1950)  claim  that  digestion  of  blood  is  initiated  in  Culex  pipiens  by  bacteria 
which  cause  the  haemolysis  of  the  red  blood  cells. 

Terzian  and  Stahler  (1964)  confirmed  Terzian’s  (1963)  findings  and  extended  the  ob- 
servations as  indicated  below.  A.  aegypti  completes  digestion  of  a meal  of  chicken  blood 
in  about  72  hours  at  26.7  C and  75%  R.H.  while  A.  quadrimaculatus  required  “at  least 
96  hours”.  Neither  NaCl  nor  KC1  at  concentrations  up  to  0.3  M have  any  inhibitory  ef- 
fect upon  digestion.  Penicillin  suppresses  the  inhibitory  effects  of  oxytetracycline  and 
neomycin  in  both  species.  Streptomycin  inhibits  digestion  in  A.  aegypti;  does  not  interact 
with  oxytetracycline,  but  is  suppressed  by  penicillin.  Neomycin  inhibits  digestion  by  A. 
aegypti  and  A.  quadrimaculatus  and  enhances  the  effects  of  calcium  and  magnesium  mark- 
edly in  the  former  but  only  slightly,  of  at  all,  in  the  latter.  Neomycin  enhances  the  inhibitory 
effect  of  manganese  in  both  species.  Iron  suppresses  the  inhibitory  effect  of  neomycin  in  A. 
aegypti  but  the  inhibition  by  iron  and  neomycin  are  additive  in  A.  quadrimaculatus.  The 
inhibitory  effects  of  the  cations  upon  digestion  in  these  mosquitoes  led  Terzian  and  Stahler 
(1964)  to  conclude  that  “the  process  of  blood  digestion  is  fundamentally  the  same  in  both 
species”.  However  they  suggested  that  the  effects  of  the  antibiotics  upon  a given  physiolog- 
ical process  vary  with  the  species.  Apparently  these  mosquitoes  are  able  to  produce  viable 
eggs  despite  the  inhibition  of  the  digestion  (Terzian  and  Stahler,  1964).  This  led  Terzian  and 
Stahler  to  conclude  that  the  “effective  inhibitory  compounds  do  not  interfere  with  the  di- 
gestion or  absorption  of  the  plasma  fraction  of  the  blood  but  rather  interfere  with  some  one 
phase  of  the  digestion  of  hemoglobin”.  This  is  essentially  the  same  conclusion  reached 
earlier  by  Terzian  (1958)  although  no  critical  experiments  were  done  to  see  if  digestion  of 
the  serum  proteins  had,  in  fact,  taken  place. 

Usually  only  female  mosquitoes  bite,  although  biting  males  have  also  been  recorded  (re- 
viewed by  Bates,  1949,  p.  79).  Whether  or  not  the  blood  ingested  by  males  is  digested  may 
vary  with  the  species.  Chao  and  Wistreich  (1959)  referred  to  the  results  of  unpublished 
experiments  indicating  that  male  Culex  tarsalis  Coquillett  could  not  digest  force-fed  blood 
meals  and  died  shortly  after  the  experimental  meal.  However,  Russell  (1931)  was  able  to 
induce  C.  pipiens  quinquefasciatus,  A.  aegypti  and  Anopheles  ludlowae  males  to  take  blood 
meals  which  were  dispatched  to  the  midgut.  In  one  male  C.  pipiens  quinquefasciatus  a 
single  oocyst  of  Plasmodium  cathemerium  Hartman  developed.  No  mention  was  made  of 
unduly  high  mortality  among  the  males.  Males  of  A.  aegypti  and  C.  pipiens  quinquefasciatus 
force-fed  repeatedly,  digested  these  blood  meals  in  the  same  time  as  did  the  females  (Mac- 
Gregor, 1931). 

Twenty-one  of  23  blood-fed  Leptoconops  (Holoconops)  becquaerti  (Kieffer)  held  at  29.4 
C emptied  their  guts  by  40  hours  after  the  meal.  At  36.7  C 1 midge  of  9 had  an  empty  gut 
at  24  hours  and  by  28  hours  4 of  the  9 midges  had  empty  guts  (Linley,  1965).  Service 
(1968c)  estimated  the  time  for  digestion  of  human  blood  by  two  species  of  midges  ( Culi - 
coides  impunctatus  Goetghebuer  and  Culicoides  obsoletus  (Meigen)  under  field  conditions 
in  Dorset  (southern  England).  During  April  (mean  temperature  8.9  C)  C.  obsoletus  took  an 
average  of  8.09  ± 0.63  days  to  digest  its  meal  while  in  May  (mean  temperature  1 1.6  C)  and 
June  (mean  temperature  15.9  C)  the  times  were  5.79  ± 0.31  and  5.14  ± 0.39  days  respectively 
For  C.  impunctatus  the  time  in  June  was  5.15  ± 0.33  days,  in  July  (15.6  C)  5.27  ± 0.66 
days;  August  (15.4  C)  5.09  ± 0.22  days,  in  September  (15.8  C)  7.38  ± 1.05  days. 

Stomoxys  calcitrans  which  are  hungry  at  the  time  they  are  offered  blood  will  gorge  until 
the  abdomen  is  “not  only  more  than  twice  its  usual  depth,  but  is  also  about  half  as  broad 


Gooding 


again  as  the  normal  breadth”  (Hewitt,  1914).  Within  half  an  hour  the  abdomen  may  return 
to  its  normal  size  and  by  2 hours  the  red  color  of  the  gut  contents  will  no  longer  be  visible 
externally.  Brown  feces  are  first  passed  about  6 hours  after  a blood  meal  and  these  were 
interpreted  by  Hewitt  as  being  excretion  of  digested  blood.  Defecation  of  brown  material 
usually  ends  about  72  hours  after  the  blood  meal  and  this  time  was  interpreted  as  the  period 
necessary  for  digestion  of  the  blood  meal.  Digestion  time  varies  from  50  to  95  hours  and 
depends  in  part  upon  the  size  of  the  meal  ingested.  From  the  number  and  size  of  fecal  de- 
posits Hewitt  concluded  that  digestion  is  most  rapid  26  to  52  hours  after  feeding.  Bishop 
(1913)  reported  that  the  rate  of  digestion  by  S.  calcitrans  is  affected  by  weather.  On  the 
basis  of  the  gross  appearance  of  the  abdomen  at  24  hours  after  a blood  meal  5 of  8,  at  46 
hours  9 of  10,  and  at  70  hours  8 of  8 wild  caught  male  S.  calcitrans  had  digested  their  meals 
(Anderson  and  Tempelis,  1970).  The  corresponding  figures  for  females  were  0 of  20,  5 of 
20,  and  7 of  24.  Anderson  and  Tempelis  (1970)  summarized  previous  reports  of  the  digestive 
rate  of  S.  calcitrans  and  concluded  that  “the  host  source  of  the  blood  meal  and  the  temper- 
ature at  which  flies  are  held  both  also  affect  digestion  rates”.  They  also  reported  (but 
without  presenting  the  data)  that  with  nulliparous  S.  calcitrans  “the  time  elapsing  between 
ingestion  and  digestion  varies  according  to  which  state  of  a gonotrophic  cycle  she  is  in”. 

Vanderplank  (1947),  stated  that  with  Glossina  swynnertoni  Austen  age  can  probably 
affect  the  duration  of  the  hunger-cycle  and  “young  flies  in  the  laboratory  take  smaller  meals 
and  digest  them  quickly”. 

Small  clear  globules  of  unknown  composition  appear  in  the  ventriculus  of  Diamanus 
montanus  (Baker)  and  Xenopsylla  cheopis  (Rothschild)  (but  not  Polygenis  gwyni  (Fox), 
within  an  hour  of  a blood  meal  but  usually  disappear  within  a day  (Holenreid,  1952).  Unfed 
P.  gwyni  (but  notD.  montanus  or  X.  cheopis)  have  bubbles  in  the  ventriculus  which  disappear 
1 to  24  hours  after  a blood  meal.  The  flea  ventriculus  is  swollen  and  bright  red  just  after  a 
blood  meal  but  shrinks  and  darkens  as  digestion  proceeds  and  in  D.  montanus  is  devoid  of 
blood  residue  1 to  10  days  after  the  meal.  None  of  these  species  defecate  undigested  blood. 
Parker  (1958)  observed  that  fleas  ( Cediopsylla  inaequalis  inaequalis  (Baker),  Thrassis  bacchi 
gladiolis  Jordan  and  Pule x irritans  L.)  fed  on  a variety  of  small  mammals  infected  with 
Pasteurella  tularensis  (McCoy  & Chapin)  generally  clear  their  digestive  tracts  of  the  bulk  of 
the  blood  meal  by  36  hours. 

Nuttall  (1917b)  reported  that  the  rate  of  digestion  by  P.  humanus  is  influenced  by  tem- 
perature. The  ingested  blood  remains  red  and  the  abdomen  swollen  for  4 days  when  the 
insects  are  kept  at  12  C.  At  31-37  C the  size  of  the  abdominal  contents  decreases  rapidly  in 
a few  hours  and  as  the  blood  is  digested  it  turns  from  red  to  reddish-brown  and  finally  to 
black. 


HISTOLOGICAL  CHANGES  IN  THE  GUT  AND  BLOOD  MEAL 

Secretory  and  absorptive  cells  are  not  differentiated  in  the  midgut  of  the  hog  louse, 
Haematopinus  suis  (L.)  and  these  activities  apparently  are  carried  out  by  all  gut  cells  (Flo- 
rence, 1921).  Within  an  hour  of  feeding,  erythrocytes  are  vacuolated  but  leucocytes  and 
platelets  are  not.  Platelets  are  destroyed  after  2 hours.  The  staining  properties  of  leucocyte 
cytoplasm  is  slightly  affected  within  2 hours  of  feeding  and  by  6 hours  after  the  meal  the 
nuclei  begin  to  disintegrate.  By  8 hours  the  blood  is  an  amorphous  mass. 

Pediculus  humanus  humanus  L.  (=  Pediculus  humanus  corporis  de  Geer)  at  30-32  C com- 
pletely destroy  the  erythrocytes  within  4 hours  of  feeding  on  humans  (Cabasso,  1947). 
However,  when  lice  are  fed  on  guinea  pigs  the  erythrocytes  remain  intact  for  48  hours,  about 
which  time  the  lice  die,  many  with  ruptured  guts.  The  mortality  rates  of  unfed  lice  and 


Haematophagous  insects 


23 


those  fed  human  or  guinea  pig  blood  indicate  that  those  fed  guinea  pig  blood  starve  to  death. 
Lice  consuming  a single  meal  on  guinea  pigs  and  subsequently  feeding  on  man  live  a normal 
life  span.  Cabasso  concluded  that  P.  humanus  can  not  digest  guinea  pig  blood.  Krynski, 
Kuchta  and  Becla  (1952)  claimed  that  guinea  pig  erythrocytes  rapidly  haemolyzed  within 
the  gut  of  P.  humanus.  Using  a hanging-drop  technique  they  showed  that  the  haemoglobin 
crystallized  in  “trigonal  pyramids  of  various  size”.  Crystal  formation  began  during  the 
feeding  period  and  within  6 hours  the  gut  was  filled  with  crystals  which  mechanically  dam- 
aged the  midgut  epithelium.  Guinea  pig  haemoglobin  also  crystallized  in  the  gut  of  Cimex 
lectularius  and  Ornithodorus  moubata  (Murray)  but  in  these  species  the  crystals  decomposed 
without  injuring  the  gut  wall  (Krynski  et  al,  1952). 

Davies  and  Hansens  (1945)  proposed  the  hypothesis  that  “the  digestive  enzymes  of  the 
louse  were  immunologically  specific  and  developed  to  act  upon  the  blood  taken  by  the 
young  insect  in  its  first  meal”.  This  hypothesis  was  tested  by  rearing  P.  humanus  for  10  days 
on  either  a man  or  a rabbit  and  then  transferring  half  of  each  group  to  another  man  or  an- 
other rabbit.  Mortality  during  the  next  9 days  was  independent  of  whether  the  lice  had 
switched  host  species  and  thus  the  data  did  not  support  the  hypothesis. 

The  rate  at  which  mouse  erythrocytes  and  Spirochaeta  duttoni  (Novy  and  Knapp)  are 
destroyed  in  the  gut  of  C.  lectularius  depends  upon  temperature.  At  12,  14,  16,  20  and  24  C 
erythrocytes  are  still  intact  at  278,  7122  (sic),  42,  31,  and  8-24  hours  after  ingestion  of  the 
blood  meal  (Nuttall,  1908). 

Erythrocytes  from  a sickle-cell  anemia  patient  all  exhibited  the  sickling  form  after  24 
hours  in  the  midgut  of  Panstrongylus  megistus  compared  with  only  1/3  “sickle-form”  in  a 
sealed  control  (Pick,  1955).  Haemolysis  began  in  3 days  and  was  complete  by  6 days  after 
feeding.  By  15  days  crystals  of  sickle-cell  haemoglobin  were  observed.  Crystals  of  normal 
hemoglobin  were  never  observed  in  the  gut  of  P.  megistus. 

Blood  meals  are  stored  in  the  expanded,  anterior  portion  of  the  midgut  (‘stomach’)  of 
Rhodnius  prolixus  (Wigglesworth,  1936).  Here  the  blood  cells  remain  intact  for  several  days 
and  the  haemoglobin  red  for  several  weeks  indicating  that  no  digestion  is  taking  place 
(Wigglesworth,  1936).  In  the  narrow,  posterior  portion  of  the  midgut  (‘intestine’)  the  blood 
turns  dark  brown  or  black  indicating  that  this  is  the  site  of  digestion. 

Digestion  of  the  blood  meal  by  mosquitoes  begins  at  the  outer  edge  of  the  meal  and  pro- 
ceeds inward  (Davies  and  Philip,  1931).  This  occurs  in  Culex  pipiens  (Huff,  1934),  Aedes 
aegypti  (Stohler,  1957;  Howard,  1962;  Akov,  1965;  Freyvogel  and  Staubli,  1965;  Gander, 
1968),  Anopheles  stephensi,  Anopheles  gambiae  and  Anopheles  labranchiae  atroparvus  (re- 
ported as  A.  maculipennis  atroparvus ) (Freyvogel  and  Staubli,  1965). 

Before  engorgement  the  midgut  epithelial  cells  of  A.  aegypti  are  columnar;  during  engorge- 
ment they  become  squamous  with  convex  internal  borders  (Howard,  1962).  As  digestion 
proceeds,  the  cells  return  to  their  original  shape.  These  observations  were  confirmed  by 
Freyvogel  and  Staubli  (1965)  and  were  extended  to  A.  stephensi,  A.  gambiae  and  A.  la- 
branchiae  atroparvus  (Staubli,  Freyvogel  and  Suter,  1966).  Although  the  shape  of  the 
epithelial  cells  changes  in  a reversible  manner  the  shape  of  the  tracheoles  serving  the  midgut 
cells  is  irreversibly  changed  from  a tight  spiral  before  the  first  meal  to  slightly  curved  after 
the  meal  (Detinova,  1962). 

A whorled  granular  endoplasmic  reticulum,  is  present  near  each  nucleus  in  the  midgut 
cells  of  fasting  or  sugar-fed  A.  aegypti.  During  the  ingestion  of  blood  (by  A.  aegypti  and 
Aedes  togoi  (Theobald),  these  whorls  unfold.  The  whorls  reform  upon  completion  of  diges- 
tion, and  the  endoplasmic  reticulum  may  be  involved  in  the  secretion  and  transport  of 
proteolytic  enzymes  (Bertram  and  Bird,  1961).  These  changes  in  A.  aegypti  were  confirmed 
by  Staubli,  et  al  (1966).  In  unfed  A.  labranchiae  atroparvus,  A.  gambiae  and  A.  stephensi  the 


24 


Gooding 


whorls  found  in  A.  aegypti  are  replaced  by  apical  granules  which  disappear  at  the  time  secre- 
tions are  detectable  in  the  midgut  lumen.  The  large  globules  of  RNA-  containing  material 
found  between  the  nuclei  and  the  lumen  borders  of  midgut  epithelial  cells  of  unfed  A. 
aegypti  by  Dasgupta  and  Ray  (1955)  may  be  the  whorls  of  endoplasmic  reticulum  reported 
by  the  workers  cited  above. 

Staubli  et  al  (1966)  suggested  four  possible  functions  for  the  whorled  endoplasmic  reticu- 
lum. First,  the  midgut  secretions  (e.g.  digestive  enzymes)  could  be  synthesized  and  stored  in 
the  whorls  prior  to  ingestion  of  the  blood  meal,  although  the  appearance  of  the  whorls  was 
not  consistent  with  this.  Second,  immediately  after  feeding  the  whorls  could  rapidly  synthe- 
size the  secreted  material.  Since  the  whorls  break  down  into  vesicles  (Staubli  et  al,  1 966) 
within  9 minutes  of  feeding  it  was  suggested  that  synthesis  was  completed  by  this  time. 
Third,  synthesis  of  the  secretory  material  could  take  place  on  the  vesicles  which  arise  from 
the  whorls  and  fourth,  the  endoplasmic  whorls  could  be  concerned  rather,  with  the  absorp- 
tion process.  The  first  and  second  and,  possibly,  the  second  and  third  suggestions  could  be 
distinguished  from  each  other  by  experiment.  However,  as  far  as  I know  the  critical  experi- 
ments on  mosquitoes  have  not  been  done. 

In  Anopheles  maculipennis  the  anterior  midgut  cells  secrete  a fairly  large  amount  of  a 
mucous-like  material  within  7 minutes  of  a blood  meal.  Although  this  mucous  forms  a plug 
at  both  ends  of  the  stomach  and  often  completely  surrounds  the  meal  the  mucous  “does  not 
appear  to  exert  any  important  effect  on  digestion,  as  the  erythrocytes  in  its  vicinity  are 
hardly  broken  down  at  all”  (Freyvogel  and  Staubli,  1965).  Other  species  of  Anopheles  appear 
to  produce  a smaller  quantity  of  mucus  a little  later  than  A.  maculipennis. 

In  mouse-fed  A.  aegypti  the  midgut  epithelium  produces  a granular  secretion  for  up  to  15 
hours  after  feeding  and  the  peritrophic  membrane  (PM)  is  apparently  formed  from  this 
(Bertram  and  Bird,  1961).  However,  with  chicken-fed  A.  aegypti , secretions  in  the  form  of 
discrete  hemispherical  droplets  on  the  internal  border  of  the  cells  first  appear  about  1 2 hours 
after  engorgement  (Howard,  1962).  These  droplets  increase  to  a maximum  size  at  about  40 
hours,  after  which  each  appears  to  be  attached  to  a cell  by  a stock.  In  contrast  Dasgupta  and 
Ray  (1955)  observed  in  blood  fed  A.  aegypti  a holocrine  secretion  which  disintegrated 
when  discharged  into  the  gut  lumen.  Howard  (1962)  interpreted  the  droplets  he  observed  in 
A.  aegypti  as  the  substance  from  which  the  PM  forms.  The  PM  first  appears  about  1 2 hours 
after  the  mosquitoes  feed.  This  membrane  increases  in  size  until  about  36  hours,  and  by  48 
hours  is  rather  brittle.  As  the  amount  of  blood  in  the  midgut  decreases  the  PM  is  fragmented 
by  the  contraction  of  the  midgut  muscles.  In  A.  aegypti  the  PM  forms  several  hours  after 
ingestion  of  the  blood  meal  but  before  digestion  begins  (Stohler,  1957).  As  digestion  pro- 
ceeds the  PM  becomes  harder  and  more  brittle  but  subsequently  softens  and  as  the  meal  is 
digested  the  PM  adheres  to  the  blood  meal,  not  the  midgut  cells. 

Yaguzhinskaya  (1940)  demonstrated  the  presence  of  a chitin-containing  PM  in  blood-fed 
A.  maculipennis . The  membrane  forms  after  a blood  meal  and  is  occasionally  open  at  the 
posterior  end.  The  remnants  of  the  membrane  are  defecated  after  the  meal  is  digested. 

A chitin  containing  PM  forms  around  the  blood  meal  in  the  mosquito  midgut  (Waterhouse, 
1953a).  If,  after  partial  digestion  of  the  first  blood  meal,  a second  meal  is  taken,  the  second 
meal  surrounds  the  first  and  a second  peritrophic  membrane  is  formed  around  the  entire 
mass  of  blood. 

The  development  of  the  PM  has  been  studied  in  A.  aegypti,  A.  stephensi,  A.  labranchiae 
atroparvus,  and  A.  gambiae  (Freyvogel  and  Staubli,  1965).  In  the  first  two  species  age  and 
number  of  blood  meals  have  no  effect  on  the  development  of  the  PM.  However,  some 
specimens  do  not  develop  a complete  PM  and  in  these  digestion  is  usually  abnormal.  In  A. 
aegypti  and  A.  stephensi  feeding  upon  man,  guinea  pigs,  rabbits  or  chicken  the  source  of  the 


Haematophagous  insects 


25 


blood  meal  does  not  affect  the  formation  of  the  PM.  A.  gambiae  formed  a PM  but  A. 
labranchiae  atroparvus  produce  nothing  more  than  a viscous  material  surrounding  the  blood 
meal.  However,  unlike  the  A.  stephensi  which  lack  a PM,  A.  labranchiae  atroparvus  digest 
the  blood  meal  in  a normal  manner.  In  A.  aegypti  the  PM  forms  in  5 to  8 hours  after  the 
meal  and  remains  until  digestion  is  practically  complete  (48  hrs).  In  A.  gambiae  PM  forma- 
tion requires  at  least  1 3 hours  but  may  persist  up  to  60  hrs.  The  corresponding  times  for  A. 
stephensi  are  32  and  72  hours.  The  membrane  in  Aedes  spp.  passes  through  stages  described 
as  viscous,  elastic,  solid  and  finally  fragile,  but  in  anophelines  it  never  develops  beyond  a 
delicate  membrane. 

Three  species  (A.  aegypti,  A.  gambiae,  A.  stephensi ) which  normally  form  a PM  do  not  do 
so  completely  if  they  ingest  only  a small  quantity  of  blood  (Freyvogel  and  Staubli,  1965). 
Mosquitoes  feeding  upon  a chicken  injected  with  heparin,  or  upon  defibrinated  blood,  form 
a PM.  A.  aegypti  and  A.  gambiae,  feeding  upon  guinea-pig  serum,  form  the  membrane  but  A. 
stephensi  usually  do  not.  When  A.  aegypti  are  given  an  incomplete  meal  on  guinea  pigs, 
followed  after  1 0 hours  by  a meal  on  chickens,  they  form  a PM  around  each  meal.  The  PM 
around  the  anterior  (chicken  blood)  meal  is  thinner  than  around  the  posterior  meal. 

Ringer’s  solution  will  not  dissolve  the  A.  aegypti  PM  but  does  dissolve  those  of  A.  gambiae 
and  A.  stephensi  (Freyvogel  and  Staubli,  1965).  However  Van  Wisselingh’s  chitosan-iodine 
test  is  positive  for  PM  of  all  three  species. 

A.  aegypti  consuming  less  than  0.1  mg  of  guinea-pig  blood  form  no  PM  and  they  must 
ingest  at  least  0.5  mg  before  forming  a complete  PM  (Freyvogel  and  Jaquet,  1965).  Howev- 
er, there  is  no  correlation  between  blood  meal  size  and  the  condition  of  the  PM  in  A. 
stephensi  and  probably  not  in  A.  gambiae.  Both  A.  aegypti  and  A.  gambiae  produce  a PM 
when  they  are  given  an  enema  of  physiological  saline  or  air.  The  PM  formed  in  both  species 
after  a meal  of  blood  or  serum  reacts  positively  to  Van  Wisselingh’s  chitosan-iodine  test. 
However  Freyvogel  and  Jaquet  reported  that  the  results  of  this  test  on  the  PM  formed  after 
a saline  or  air  enema  were  inconclusive. 

Formation  of  the  PM  and  digestion  of  the  blood  meal  in  A.  aegypti  and  A.  stephensi  have 
been  studied  in  frozen  sections  by  Gander  (1968).  The  PM  of  these  species  have  different 
structural  features:  in  A.  aegypti  it  is  laminar  while  in  A.  stephensi  it  consists  of  a granular 
material  imbedded  in  a Periodic  acid-Schiff  (PAS-)  positive  substance.  In  their  initial  stages 
of  formation  the  PM  of  both  species  contain  Periodic  acid-Schiff  positive  material  but  during 
blood  digestion  this  material  disappears  completely  from  the  A.  aegypti  PM  and  partially 
from  that  of  A.  stephensi.  Histochemical  tests  demonstrate  the  presence  of  carbohydrates 
and  lipids  in  both  PM’s.  In  A.  aegypti  cells  throughout  the  midgut  epithelium  undergo  an 
apocrine  secretion  while  in  A.  stephensi  there  is  a modified  merocrine  secretion  proceeding 
from  the  posterior  to  the  anterior  end  of  the  midgut.  Gander  (1968)  divided  blood  meal 
digestion  by  A.  aegypti  and  A.  stephensi  into  2 phases.  Early  in  phase  I the  midgut  secretes 
carbohydrates  and  lipids.  In  the  first  10  hours  after  a blood  meal,  carbohydrates  are  not 
detectable  in  the  epithelial  cells  of  A.  aegypti  but  can  be  found  in  those  of  A.  stephensi. 
Enzymes  are  probably  secreted  also  during  this  phase  but  only  erythrocytes  at  the  very  edge 
of  the  meal  show  signs  of  breakdown.  Phase  I ends  when  the  PMs  form,  16  hours  in  A. 
aegypti  and  30  hours  in  A.  stephensi  after  feeding,  and  no  further  secretion  by  the  midgut 
epithelial  cells  occurs.  Formation  of  lipid  droplets  within  the  blood  meal  and  accumulation 
of  these  on  the  lumen  side  of  the  PM  marks  the  beginning  of  Phase  II.  Digestion  of  the 
blood  meal  proceeds  inward  from  the  periphery  and  the  epithelial  cells  accumulate  carbohy- 
drates and  lipids.  In  A.  aegypti  the  peak  of  lipid  absorption  occurs  before  the  peak  of 
carbohydrate  absorption  while  the  reverse  is  true  for  A.  stephensi.  Peroxidases  occur  in  the 
midgut  epithelial  cells  of  both  species  of  mosquitoes  during  digestion  of  blood;  their  concen- 


26 


Gooding 


tration  remains  constant  in  A.  stephensi  but  in  A.  aegypti  reaches  a maximum  40  hours 
after  feeding.  Gander  felt  that  there  was  a descrepency  between  his  observation  that  diges- 
tion proper  did  not  begin  in  A.  aegypti  until  about  16  hours  after  feeding,  and  the  results 
of  Fisk  and  Shambaugh  (1952)  and  Gooding  (1966b)  which  showed  considerable  proteinase 
in  the  gut  by  this  time.  He  suggested  that  this  discrepency  was  connected  with  the  presence 
of  trypsin  inhibitors  within  the  blood  meal. 

Digestion  of  blood  by  C.  pipiens  was  studied  by  deBoissezon  (1930a,  1930b),  Huff  (1927, 
1934)  and  Arnal  (1950)  using  histological  techniques.  Huffs  observations  on  C.  pipiens  are 
similar  to  those  made  on  A.  aegypti.  The  rate  of  digestion  depends  upon  the  amount  of 
blood  ingested  and  upon  ambient  temperature  (deBoissezon,  1930a,  1930b).  Hemolysis  of 
the  erythrocytes  is  followed  by  crystallization  of  their  hemoglobin.  The  hemoglobin  crystals 
are  dissolved  by  the  digestive  juices  and  absorbed  and  digested  by  cells  in  the  floor  of  the 
wide  part  of  the  midgut.  Cells  in  the  anterior,  narrow  part  of  the  midgut  produce  a vitreous 
secretion  from  the  nucleolar  region  and  a granular  secretion  from  their  cytoplasm.  Cells  in 
the  wide  part  of  the  midgut  secrete  vesicles  which  occasionally  included  their  nucleolei.  In 
C.  pipiens  Arnal  (1950)  observed  merocrine  secretion  in  two  regions  of  the  midgut  during 
fasting  and  three  types  of  secretion  after  a blood  meal.  He  concluded  that  digestion  of  blood 
was  initiated  by  symbiotic  bacteria  which  penetrated  the  blood  meal,  caused  the  red  cells, 
but  not  the  leucocytes,  to  swell  and  eventually  to  lyse  and  which  also  prevented  clotting. 
Simultaneously,  the  midgut  cells  began  secreting.  The  cells  in  the  narrow,  anterior  region 
released  granules,  those  at  the  beginning  of  the  wide  portion  of  the  midgut  released  vacuoles 
and  holocrine  secretion  occurred  in  the  cells  in  the  floor  of  the  widest  part  of  the  midgut. 
Arnal  stated  that  the  pH  during  digestion  was  6.5  to  7,  and  speculated  that  the  secretions 
observed  were  trypsins  capable  of  acting  in  a slightly  acid  medium.  Stroma  and  the  leuco- 
cytes resisted  digestion,  but  the  bacteria  apparently  did  not  as  they  disappeared.  Iron  was 
detected  in  the  young  cells  in  the  floor  of  the  midgut  with  the  Liesegang  technique.  As 
absorption  took  place,  the  midgut  contents  thickened,  and  the  haemoglobin  crystallized  in 
the  midgut  lumen.  Haemoglobin  could  not  be  detected  in  the  hindgut. 

In  Culicoides  nubeculosus  the  midgut  epithelium  of  the  unfed  midge  has  columnar  cells 
which  become  more  or  less  cuboidal  on  ingestion  of  a blood  meal  (Megahed,  1956).  A PM, 
not  present  in  the  unfed  insect,  forms  within  5 hours  of  feeding  on  blood.  The  membrane 
varies  in  thickness  and  appearance  in  different  parts  of  the  stomach,  having  villi-like  struc- 
tures in  some  regions  and  a laminar  appearance  in  others.  The  PM  is  apparently  secreted  by 
the  midgut  epithelial  cells.  By  24  hours,  the  PM  develops  a perforation  at  its  posterior  end 
through  which  material  may  pass  from  the  midgut  to  the  hindgut.  The  PM  completely  sur- 
rounds the  blood  meal  except  at  its  posterior  end  but  a partial  disintegration  of  one  or  more 
layers  is  evident  in  some  regions.  By  48  hours  the  blood  meal  is  almost  completely  digested 
but  the  gut  still  contains  haematin  and  some  evidence  of  the  PM.  After  72  hours  the  midgut 
is  empty  of  both  blood  residues  and  the  remains  of  the  PM. 

During  the  first  2 days  after  Culicoides  obsoletus  feed  on  human  blood  the  gut  contents 
solidify  and  become  opaque  but  change  little  in  volume  (Jamnback,  1961).  Undigested 
blood  and  small  black  pigment  granules  and  rods,  presumably  digestion  products,  occur  in 
the  gut  by  the  third  day.  By  the  fourth  day  the  blood  meal  is  completely  digested  and  the 
gut  is  empty  when  the  midges  are  held  at  21  C. 

Feng  (1951)  examined  the  formation  of  the  PM  in  Phlebotomus  mongolensis  Sinton  and 
Phlebotomus  chinensis  Newstead  fed  on  Chinese  hamsters  and  in  Sergentomyia  squamirostris 
(Newstead)  (reported  as  Phlebotomus  squamirostris ) fed  on  a toad.  He  studied  also  the  influ- 
ence of  the  PM  upon  establishment  of  trypanosomes  in  these  sandflies.  In  P.  mongolensis  the 
PM  completely  envelopes  the  blood  meal  and  is  very  tough.  As  in  mosquitoes,  digestion  of 


Haematophagous  insects 


27 


the  blood  meal  begins  at  the  periphery  and  progresses  inward.  Digestion  of  the  blood  meal 
requires  5-6  days  and  as  material  is  digested  and  absorbed,  the  PM  shrinks  to  a small  spindle 
which  is  passed  complete  into  the  hindgut.  Leishmania  donovani  Laveran  and  Mesnil  flagel- 
lates live  only  within  this  peritrophic  sac  and  pass  into  the  hindgut  within  it.  In  P.  chinensis 
a PM  is  formed  but  begins  to  break  down  3 days  after  feeding,  fragments  of  it  passing  into 
the  hindgut  with  the  blood  meal  residue.  Digestion  of  a blood  meal  in  P.  chinensis  takes 
about  7 days.  The  disintegration  of  the  PM  releases  the  flagellates  which  move  forward  and 
establish  themselves  in  the  proventriculus.  Ultimately  they  migrate  forward  to  the  mouth 
parts.  The  PM  of  S.  squamirostris  appears  to  be  open  at  the  posterior  end.  Digestion  of  a 
blood  meal  by  this  species  is  complete  within  3 days.  Crithidia  of  Trypanosoma  bocagei 
Franpa  leave  the  midgut  through  the  open  posterior  end  of  the  PM  and  establish  themselves 
in  the  hindgut. 

In  Phlebotomus  papatasi  digestion  is  very  slow  and  haemolysis  of  the  erythrocytes  takes 
place  3 or  4 days  after  feeding  (Adler  and  Theodor,  1926).  “Unaltered  haemoglobin  is  never 
found  in  the  epithelial  cells  of  the  stomach  but  it  is  passed  in  the  feces”  and  it  was  con- 
cluded that  it  is  the  plasma  which  is  the  essential  component  of  blood  and  not  the  erythro- 
cytes. A PM  is  present  a day  or  two  after  a blood  meal. 

Peritrophic  membranes  occur  in  Simulium  anatinum  Wood,  Simulium  rugglesi,  Simulium 
aureum,  Simulium  latipes,  Simulium  quebecense,  Simulium  croxtoni,  Simulium  venustum, 
Prosimulium  decemarticulatum,  Prosimulium  hirtipes  (Fries)  and  Cnephia  ornithophilia 
Davies,  Peterson  and  Wood,  (Bennett  and  Fallis,  unpublished  work  cited  by  Fallis,  1964); 
Simulium  griseicolle  Becker,  and  Simulium  damnosum  (Lewis,  1950),  and  Simulium  neavei 
Roubaud  (Lewis,  1960).  In  S.  damnosum  the  PM  gives  a positive  chitosan  test  (Lewis,  1950, 
1953)  and  is  formed  after  ingestion  of  blood  but  not  sugar  (Lewis,  1953).  Flies  interrupted 
during  feeding  have  blood  in  both  the  tubular  (anterior)  portion  and  the  expanded  (posteri- 
or) portion  of  the  midgut.  Engorged  flies  have  all  the  blood  in  the  posterior  part  of  the 
midgut.  During  consumption  of  a blood  meal  some  of  the  contents  of  the  crop  apparently 
pass  into  the  anterior  part  of  the  midgut.  A delicate  membrane  forms  within  half  a minute 
of  completion  of  engorgement  and  this  membrane  is  quite  distinct  by  30  minutes  after 
feeding.  By  an  hour  after  engorgement  the  laminar  nature  of  the  PM  is  evident,  particularly 
in  the  knob  of  the  membrane  at  the  posterior  end  of  the  midgut.  The  membrane  gradually 
turns  yellow  and  then  brown.  Between  24  and  72  hours  after  the  meal,  the  blood  mass  de- 
creases in  size  and  the  PM  breaks  up  (Lewis,  1953).  In  blackflies,  digestion  proceeds  from 
the  periphery  toward  the  centre  of  the  blood  meal  (Fallis,  1964,  citing  unpublished  work  of 
Bennett  and  Fallis).  Cells  at  the  centre  of  the  blood  mass  may  remain  intact  for  more  than 
48  hours. 

“Resting”  cells  are  columnar  in  the  ‘stomach’  portion  of  the  midgut  of  Tabanus  albime- 
dius  (and  other  Tabanus  spp.)  but  are  converted  to  flattened  pavement  epithelium  when  the 
midgut  fills  with  blood  (Cragg,  1 920).  Secretory  cells  casting  off  large  droplets  are  seen  in 
the  midgut  most  frequently  during  the  5 minutes  after  feeding  and  are  rarely  found  more 
than  1 hour  after  a meal.  The  digestive  substances  acting  on  the  erythrocytes  cause  the 
formation  of  dark  pigments,  beginning  at  the  surface  of  the  blood  meal.  One  day  after  a 
blood  meal  and  later,  the  epithelial  cells  become  columnar  again  and  secrete  minute  droplets 
of  undetermined  fate.  Cells  in  the  anterior  portion  of  the  midgut  (cardia)  secrete  continu- 
ously. Columnar  cells  reform  as  digestion  of  the  meal  proceeds.  There  is  no  PM.  Red  blood 
cells  are  normal  for  a short  time  after  ingestion  but  soon  become  distorted,  shrunken,  and 
poorly  stained.  Stroma  are  detectable  for  24  hours  in  the  gut.  Pigments  form  early  and  by  2 
hours  the  stomach  contents  are  a purple,  tarry  mass.  About  8 hours  after  feeding  residue 
from  the  meal  begins  to  pass  into  the  hindgut. 


28 


Gooding 


No  PM  forms  in  the  gut  of  Chrysops  silacea  (Wigglesworth,  1931;  Crewe,  1961)  and  ob- 
servations of  the  gut  histology  of  this  species  made  at  various  times  after  a blood  meal 
“agree  exactly  with  those  of  Cragg  (1920)  on  Tabanus"  (Wigglesworth,  1931). 

The  midguts  of  Glossina  palpalis,  Glossina  submorsitans  and  Glossina  tachinoides  can 
each  be  divided  into  3 sections  on  a histological  basis  (Wigglesworth,  1929).  In  all  species 
the  anterior  half  has  an  irregular  columnar  epithelium  and  includes  a narrow  band  of  giant 
cells  containing  bacteroids.  In  this  region  the  blood  is  concentrated  by  removal  of  water  but 
there  is  no  digestion  of  the  blood  components.  In  the  next  region  of  the  midgut  there  are 
large,  deeply  staining  cells  which,  during  digestion  of  blood,  produce  and  release  large 
vacuolated  buds  of  cytoplasm  from  their  apical  surfaces;  these  buds  later  disintegrate  in  the 
lumen  of  the  midgut.  Blood  in  this  region  turns  dark  and  becomes  amorphous.  The  posterior 
region  of  the  midgut  has  regular,  columnar  epithelial  cells  which  become  vacuolated  late  in 
the  digestive  process  - probably  indicating  a role  for  them  in  absorption.  A PM  surrounds  the 
blood  meal  and  it  is  secreted  by  cells  of  the  proventriculus.  Hoare  (1931)  confirmed  in  G. 
palpalis , the  existence  of  a PM  consisting  of  a continuous,  open-ended  cylinder  reaching 
from  the  proventriculus  to  the  hind  gut  with  new  material  being  secreted  at  its  anterior  end 
each  time  a blood  meal  is  consumed.  Yorke,  Murgatroyd  and  Hawking  (1933)  also  reported 
PM’s  surrounding  the  blood  meals  of  Glossina  morsitans  and  G.  palpalis.  Weitz  and  Buxton 
(1953)  cited  unpublished  observations  of  Jackson  indicating  that  blood  remains  microscop- 
ically recognizable  longer  in  laboratory  held  tsetse  flies  than  in  marked  flies  in  the  field. 

The  anterior  part  of  the  midgut  of  Stomoxys  calcitrans  consists  of  a blood  reservoir  with 
columnar  epithelial  cells  (Lotmar,  1949).  Although  these  cells  secrete  material  (probably 
anticoagulins),  the  blood  cells  in  a meal  remain  unchanged  and  no  digestion  takes  place. 
Digestion  proceeds  as  the  blood  moves  posteriorly  through  the  digestive  region  of  the  midgut 
and  cyclic  changes  in  merocrine  secretion,  absorption  and  cell  regeneration  occur.  The  for- 
mation of  fat  globules  in  isolated  epithelial  cells  is  observed  1 to  2 hours  after  ingestion  of 
blood  and  these  cells  become  more  numerous  as  digestion  proceeds.  Digestion  is  more  or  less 
complete  in  24  hours. 

Minchin  and  Thomson  (1915)  described  the  histological  changes  occurring  in  the  midgut 
of  the  flea  Nosopsyllus  fasciatus  (Bose)  (reported  as  Ceratophyllus  fasciatus ) during  diges- 
tion of  Trypanosoma  lewisi  (Kent)  infected  blood  meals.  After  the  fleas  feed,  the  midgut 
cells  are  flattened  but  become  columnar  as  the  meal  is  digested.  Within  a few  hours  of 
feeding  the  red  blood  cells  break  down  and  by  24  hours  the  blood  meal  is  viscous  and  brick- 
red  and  contains  large  “grains”.  By  48  hours  the  stomach  contents  are  watery  and  brownish- 
black  and  contain  fewer  smaller  “grains”.  By  use  of  an  iron-haematoxylin-Lichtgrun-picric 
acid  combination  the  stomachs  of  the  fleas  may  be  divided  into  2 classes  - a grey-black  series 
with  a greenish  tinge  and  a bright  lemon-yellow  series.  In  the  grey-black  series  there  are 
many  grains  and  spherules  suspended  in  the  greenish  “coagulated  albuminous  matrix”  by 
18-24  hours  after  the  meal.  The  centre  of  the  gut  contents  lacks  the  coarse  grains  and  is 
clear.  The  grains  become  smaller  as  digestion  proceeds  and  by  36  hours  only  greenish-grey 
debris  next  to  the  epithelial  cells  remains.  Leucocytes  are  recognizable  24  hrs,  but  not  36 
hours  after  the  blood  meal.  Minchin  and  Thomson  concluded  that  “digestion,  or  more  prob- 
ably the  passage  backwards  toward  the  rectum  of  the  undigestible  remnants,  of  the  blood- 
debris  appears  to  proceed  from  the  center...  towards  the  periphery”.  The  stomachs  of  the 
yellow  series  contain  a closely  packed  granular  material  and  this  and  the  matrix  are  stained 
by  the  picric  acid.  Digestion  in  this  series  is  slower  than  in  the  grey-black  series  and  Minchin 
and  Thomson  considered  the  yellowish  stomachs  to  be  abnormal. 

The  midgut  epithelium  of  adult  fleas  ( Ctenophthalmus  Kolenati  sp)  has  intranuclear  crys- 
tals in  about  10%  of  the  cells  (Richards  and  Richards,  1969).  The  existence  of  these  crystals 


Haematophagous  insects 


29 


in  the  blood-feeding  adult  but  not  in  the  scavenging  larva  led  to  the  suggestion  that  they  are 
derived  from  the  hemoglobin  of  the  blood  meal. 

Waterhouse  (1953b)  presented  data  on  PM’s  based  on  a rather  extensive  survey  of  insect 
midguts.  He  classified  the  PM’s  as  type  I,  if  they  consisted  of  1 or  more  layers  “produced 
mainly  or  entirely  by  a ring  of  cells  at  the  anterior  end  of  the  midgut”  and  as  type  II,  if  they 
consisted  “typically  of  a series  of  thinner,  coaxial  layers  and  arises  by  periodic  delamination 
from  the  surface  of  the  striated  border  of  a layer  of  material  secreted  from  the  whole  midgut 
epithelium”.  Type  II  PM’s  occur  in  adult  mosquitoes  (A.  aegypti,  Culex  pipiens  quinque- 
fasciatus ) and  tabanids  ( Dasybasis  froggatti  (Ric.),  Scaptia  jacksoniensis  (Guer.)  and  Scaptia 
gattata  (Don)  while  type  I PM’s  are  found  in  the  Nycteribiidae  ( Nycterebosca  falcozi  Jobl.) 
and  some  Hippoboscidae  ( Ortholfersia  macleayi  Leach  and  Ornithomyia  Latreille  sp  but  not 
Melophagus  ovinus  (L)  ). 

SEROLOGICAL  AND  CHEMICAL  ANALYSIS  OF  GUT  CONTENTS 
DURING  DIGESTION  OF  THE  BLOOD  MEAL 

The  precipitin  technique  has  been  used  most  frequently  in  host  preference  studies  of  mos- 
quitoes but  has  also  yielded  data  on  digestion  rates.  Most  of  the  latter  demonstrate  that 
after  a certain  length  of  time  depending  on  environmental  conditions,  the  midgut  contents 
do  not  give  a positive  reaction  (Bull  and  King,  1923;  Davis  and  Philip,  1931;  Weitz  and 
Buxton,  1953;  West,  1950).  Bates  (1949,  p.  90)  cites  unpublished  observations  of  Balfour  on 
digestion  rates  in  Anopheles  superpictus  Grassi,  Anopheles  maculipennis  and  Anopheles 
sacharovi  Favre  in  Greece.  The  latter  two  species  were  similar  and  100%,  91%  and  79%  of 
the  mosquitoes  gave  positive  tests  after  2,  12,  and  14  hours  respectively.  A.  superpictus 
digested  blood  more  rapidly  and  96%,  72%  and  39%  gave  positive  reactions  after  2,  12,  and 
14  hours. 

Schubert  and  Kelley  (1950)  correlated  the  appearance  of  the  blood  meal  with  the  precip- 
itin reaction.  Aedes  aegypti  were  divided  into  three  groups  17  hours  after  feeding  on  a bird 
(species  not  stated).  Of  the  mosquitoes  containing  digested  and  haemolized  blood  67%  gave 
positive  precipitin  tests,  83%  of  the  partially  fed  mosquitoes  were  positive  and  96-100%  of 
the  fully  engorged  mosquitoes  were  positive. 

West  and  Eligh  (1952)  studied  the  digestion  rates  in  A.  aegypti  under  laboratory  condi- 
tions and  in  Aedes  hexodontus  under  field  conditions  with  the  precipitin  test.  They  showed 
that  digestion  in  A.  aegypti  held  in  total  darkness  occurred  more  rapidly  at  higher  tempera- 
tures (6  to  27  C).  The  rate  of  digestion  of  guinea-pig  serum  by  A.  aegypti  has  a Q10  = 2.0  in 
the  temperature  range  20  to  30  C (Williams,  1953).  West  and  Eligh  suggested  that  the  rate  of 
digestion  could  be  influenced  by  light,  and  by  the  species  of  mosquito  and  host.  It  was 
pointed  out  that  serological  techniques  indicate  alteration  of  the  blood  meal  proteins  and 
not  completion  of  digestion,  and  West  and  Eligh  (1952)  suggested  that  measurement  of 
protease  activity  as  done  by  Fisk  (1950)  “might  give  a more  accurate  indication  of  comple- 
tion of  blood  digestion  than  would  any  other  known  method”.  In  this  respect  the  results  of 
Akov  (1965)  are  interesting.  She  found  that  in  untreated  A.  aegypti  as  well  as  in  those 
treated  with  5-FU  that  there  was  a good  correlation  (coef.  corr.=  0.872)  between  the 
amount  of  proteinase  in  the  midgut  and  average  stage  of  development  of  the  ovaries  in  the 
mosquito.  Confirmation  of  this  was  obtained  by  feeding  mosquitoes  citrated  sheep  blood 
containing  1.25  pg  crystalline  soybean  trypsin  inhibitor/2  mg  blood;  this  inhibited  both 
midgut  proteinase  and  the  development  of  the  ovaries. 

Weitz  and  Buxton  (1953)  ran  precipitin  tests  at  irregular  intervals  on  mosquitoes  kept  at 
25  C and  80%  R.H.  Two  species,  Anopheles  labranchiae  atroparvus  and  A.  aegypti,  fed  on 


30 


Gooding 


man,  were  all  positive  after  16  hours  and  9%  and  27%  were  positive  after  3 days.  All  were 
negative  on  the  fourth  and  fifth  day.  All  Culex  pipiens  (issp)  molestus  Forskal  fed  on  man, 
gave  positive  precipitin  reactions  24  hours  after  the  blood  meal  but  all  were  negative  on  the 
third  and  fifth  days.  The  percentage  of  ox-fed  Anopheles  aquasalis  Curry  giving  a positive 
precipitin  test  also  declined  as  digestion  proceeded  (95%  after  16  hours,  26%  at  20  hours,  4% 
at  30  hours,  and  0%  at  40  hours). 

Differences  in  the  rate  of  digestion  of  human  blood  by  five  species  of  mosquito  were 
observed  when  they  were  held  under  identical  conditions  of  temperature  (27  C),  photoperi- 
od (ratio  of  light  to  dark  was  1:1),  and  humidity  (saturation  deficit  was  2±1  mm  Hg) 
(O’Gower,  1956).  The  time  required  for  half  of  the  mosquitoes  to  complete  digestion  to  a 
point  where  the  precipitin  test  was  negative  was  3 1 hours  for  Aedes  scutellarus  (Walker),  36 
hours  for  Aedes  notoscriptus  (Skuse),  38  hours  for  A.  aegypti,  46  hours  for  Culex  pipiens 
quinquefasciatus  and  48  hours  for  Aedes  australis  (Erichson).  Since  the  intraspecific  varia- 
tion in  the  size  of  females  was  almost  as  great  as  the  interspecific  variation,  O’Gower  felt 
that  “the  different  rates  of  digestion  of  human  blood  by  the  species  of  mosquitoes  tested 
would  seem  to  be  due  to  specific  differences  in  the  digestive  processes  and  not  to  specific 
difference  in  the  size  of  the  adults”.  O’Gower  investigated  the  effect  of  photoperiod  on  A. 
notoscriptus.  Mosquitoes  were  held  under  five  conditions,  ranging  from  continuous  light  to 
continuous  dark  and  precipitin  tests  were  run  at  36  and  40  hours  after  the  blood  meal.  As 
the  ratio  of  dark  to  light  increased  so  did  the  rate  of  digestion.  O’Gower  also  stated  (without 
supporting  data)  that  one  week  old  and  three  week  old  mosquitoes  (species  not  stated)  di- 
gested blood  at  the  same  rate. 

Downe,  Goring  and  West  (1963)  used  the  precipitin  test  to  study  the  effect  of  both  meal 
size  and  meal  source  on  the  rate  of  digestion  by  several  species  of  mosquito.  The  time  re- 
quired for  50%  or  100%  of  the  mosquitoes  to  completely  digest  or  denature  human  serum 
proteins  were  reported  for  A.  aegypti  and  Aedes  trichurus  (Dyar).  Using  these  criteria  it 
appeared  that  females  of  both  species  given  a small  blood  meal  (i.e.  where  ratio  of  weight  of 
blood  ingested  to  weight  of  mosquito  was  less  than  one)  digested  the  meal  much  more  rap- 
idly than  those  given  a large  blood  meal.  For  A.  aegypti  the  “50%  digestion  time”  and  the 
“100%  digestion  time”  for  small  blood  meals  were  16  and  36  hours  and  for  large  blood 
meals  were  40-44  hrs  and  52-56  hours.  The  corresponding  values  for  A.  trichurus  were  28 
and  48  hours,  and  64  and  76  hours. 

The  source  of  the  blood  meal  (man,  guinea  pig,  dog,  or  chicken)  had  little  effect  upon 
digestion  rates  in  several  species  of  Aedes,  but  did  affect  those  of  Mansonia  per  turbans 
(Downe,  Goring,  and  West,  1963).  In  this  species  the  “50%  digestion  time”  and  “100% 
digestion  time”  were  36  and  44  hours  for  chicken  blood,  48  and  56  hours  for  guinea  pig 
blood,  48  and  60  hrs  for  dog  blood  and  52  and  60  hours  for  human  blood. 

Templis  and  Lofy  (1963)  showed  that  Culex  tarsalis  digests  blood  meals  from  three  differ- 
ent species  of  bird  at  three  different  rates.  Positive  reactions  were  obtained  with  mosquitoes 
fed  on  all  three  species  of  birds  up  to  18  hours  after  feeding,  but  at  24  and  36  hours  the 
percentages  were  70  and  29  for  those  fed  on  the  white  crown  sparrow;  56  and  50  for  those 
fed  on  the  cow  bird;  and  100  and  73,  for  those  fed  on  the  English  sparrow. 

In  A.  aegypti  given  a small  meal  of  human  blood  followed  in  2 to  12  hours  by  a larger 
meal  of  guinea  pig  blood  digestion  of  the  human  blood  was  prolonged  (Downe,  1965).  This 
probably  occurred  because  the  human  blood  meal  was  surrounded  by  the  guinea  pig  blood 
meal  and  thus  protected  from  digestion,  which  proceeded  from  the  periphery  toward  the 
centre  of  the  midgut  contents. 

Using  agar  double  diffusion  and  immunoelectrophoretic  analysis  Mattern  et  al  (1967) 
found  that  laboratory  reared  C.  pipiens  quinquefasciatus  digested  human  albumin  within  24 


Haematophagous  insects 


31 


to  48  hours  but  retained  the  human  immunoglobulin,  IgG,  in  the  midgut  for  4 to  5 days. 
Wild  caught  mosquitoes  fed  on  man  and  then  kept  in  a tube  for  4 days  gave  the  same  results. 
On  the  other  hand,  wild  caught  mosquitoes  left  free  in  a room  for  48  hours  after  feeding  on 
man  were  negative  for  IgG  but  gave  strong  positives  for  albumin.  Even  though  a positive 
reaction  for  IgG  was  found  in  the  midguts  of  caged  C.  pipiens  quinquefasciatus  for  4 or  5 
days  the  appearance  of  a second  precipitin  band  indicated  that  digestion  of  IgG  began  about 
5 hours  after  the  mosquitoes  fed.  Using  the  agar  double  diffusion  technique  these  authors 
compared  the  IgG  digestion  products  in  the  mosquito  stomach  with  those  produced  by 
digestion  of  IgG  with  papain  and  trypsin.  They  concluded  that  “protein  cleavage  occurring 
in  the  stomach  of  mosquitoes  is  quite  different  from  that  produced  by  papain  or  trypsin”. 
All  these  results  were,  however,  based  on  a small  number  of  mosquitoes. 

Zaman  and  Chellappah  (1967)  studied  digestion  of  human  blood  by  Armigeres  subalbatus 
using  gel-diffusion  and  immunoelectrophoresis  and  concluded  that  the  serum  albumins  per- 
sisted in  the  midgut  longer  than  the  serum  globulins.  The  precipitin  band  for  the  former 
persisted  for  48  but  not  56  hours  while  for  the  latter  it  lasted  12  but  not  18  hours.  A similar 
pattern  of  digestion  was  obtained  with  A.  aegypti  digesting  guinea-pig  blood  (Williams,  1953). 

Herndon  and  Ringle  (1967)  used  the  double  diffusion  technique  in  microtubes  to  deter- 
mine the  length  of  time  host  antigens  were  detectable  in  the  midgut  of  blood-fed  A nopheles 
quadrimaculatus  and  Culex  pipiens.  Refrigerated  (6-9  C)  C.  pipiens  had  identifiable  antigens 
in  the  midgut  for  12  days  while  in  A.  quadrimaculatus  the  antigens  remained  identifiable  for 
only  1 week.  At  25-28  C the  antigens  were  identifiable  for  only  about  1 day. 

With  precipitin  tests  Edman  (1970a)  showed  that  2 and  4 day  old  A.  aegypti  digested 
human  blood  at  about  the  same  rate  (at  27  C and  70%  R.H.)  while  mosquitoes  6,  8,  and  10 
days  old  digested  it  at  a slower  rate.  There  was  no  difference  in  the  rate  of  digestion  by  8 
and  22  day  old  mosquitoes.  With  6 day  old  mosquitoes  digestion  was  slower  in  virgin  than  in 
mated  females.  Digestion  by  10  day  old  parous  females  was  slower  than  by  10  day  old  nul- 
liparous  females.  However  digestion  rates  of  the  second  blood  meal  in  10  day  old  mosquitoes 
were  the  same  as  of  the  third  blood  meal  in  1 8 day  old  mosquitoes.  This  last  finding  differs 
from  Akov’s  (1966)  for  the  rate  of  emptying  of  the  midgut  of  A.  aegypti.  Using  immunolo- 
gical techniques  and  antisera  of  high  titre,  Edman  (1970b)  found  no  consistent  differences 
in  the  rate  of  digestion  of  human  albumin,  7-globulin,  and  a-globulin  by  A.  aegypti.  Com- 
plete denaturation  of  the  proteins  occurred  between  60  and  66  hours  after  ingestion  of  the 
meal. 

Of  the  Culicoides  nubeculosus  fed  on  man,  80%  gave  positive  precipitin  reactions  after  24 
hours,  but  all  were  negative  after  3 days  (Weitz  and  Buxton,  1953).  In  the  same  study  it  was 
reported  that  laboratory  reared  Glossina  morsitans  fed  on  man,  ox,  sheep,  or  goat  were  all 
positive  for  2 days  and  on  the  third  day  gave  100%,  100%,  75%,  and  90%  positive  reactions 
respectively.  In  contrast  wild  caught  Glossina  swynnertoni  estimated  as  having  fed  on  mam- 
mals 3 days  earlier,  gave  only  28%  positives  and  those  estimated  as  having  fed  four  days 
earlier  gave  only  7%  positives. 

Downe  (1957)  used  serological  techniques  to  follow  the  digestion  of  horse  and  guinea-pig 
blood  (actually  the  serum)  by  several  species  of  black-flies  ( Simulium  venus turn,  Simulium 
vittatum  Zetterstedt,  Prosimulium  hirtipes  and  Simulium  parnassum  Malloch).  The  rates  of 
digestion  were  similar  in  all  these  species  and  were  not  markedly  affected  by  the  source  of 
the  blood.  At  19.4  - 21.1  C and  75  - 80%  R.H.  precipitin  reactions  were  obtained  in  nearly 
100%  of  the  insects  24  hrs  after  feeding.  This  declined  to  about  74%  by  32  hours,  40%  by 
40  hours,  and  to  3%  by  48  hours,  after  which  no  positives  were  obtained.  The  digestion  rate 
in  insects  maintained  under  field  conditions  was  retarded  at  lower  temperatures.  By  using 
the  Meyer  reduced  phenolphthalein  test  Downe  could  detect  blood  in  5 out  of  8 S.  venustum 


32 


Gooding 


50  hours  after  they  were  fed  on  a horse  but  similar  tests  on  1 1 S.  venustum  made  60  hours 
after  feeding  were  all  negative.  Downe  (1957)  stressed  that  the  serological  test  indicated 
only  when  serum  proteins  were  modified  and  not  when  the  blood  meal  was  actually  digested. 

Holstein  (1948)  studied  methods  of  producing  specific  antisera  and  presented  some  data 
on  the  length  of  time  after  a blood  meal  at  which  positive  precipitin  reactions  could  be 
detected.  For  Pediculus  humanus  positive  reactions  were  obtained  with  1/10  dilutions  of 
antisera  up  to  13  days,  with  1/100  up  to  10  days  and  with  1/1000  up  to  2 days  after  the  lice 
had  fed  on  humans.  However  it  is  not  clear  whether  the  lice  lived  for  the  full  1 5 days  of  the 
experiment  after  the  blood  meal.  For  Cimex  lectularius  positive  reactions  were  obtained  (di- 
lutions of  sera  given  in  brackets)  for  36  days  (1/10),  30  days  (1/100),  20  days  (1/1 ,000),  14 
days  (1/2,000),  1 1 days  (1/5,000),  6 days  (1/10,000),  and  1 day  (1/15,000).  Evidence  that 
C.  lectularius  digested  human  blood  slowly  was  obtained  by  Weitz  and  Buxton  (1953)  who 
found  100%  were  positive  after  5 days,  97%  positive  after  10  days,  40%  positive  after  20 
days,  and  22%  positive  after  30  days. 

Positive  precipitin  tests  for  human  blood  could  be  obtained  in  Phlebotomus  argentipes 
Annadale  and  Brunetti  for  up  to  8 days  after  they  had  fed  once  on  human  and  then  on 
mouse  blood  (Lloyd  and  Napier,  1930).  Although  the  number  of  sandflies  tested  at  each 
time  after  ingestion  of  the  meal  was  small,  the  results  indicated  that  the  rate  of  positive 
reactions  was  unaffected  by  the  duration  of  the  digestion  period!  This  suggests  to  me  that 
some  component  of  human  blood,  with  which  the  anti-serum  was  reacting,  was  not  digested 
by  these  sandflies. 

By  using  the  precipitin  test  on  male  Stomoxys  calcitrans  fed  an  unknown  volume  of 
citrated  human  blood  Anderson  and  Tempelis  (1970)  obtained  the  following  frequencies  of 
positives:  1 of  2 at  12  hours,  3 of  4 at  24  hours,  and  3 of  4 at  30  hours.  The  corresponding 
values  for  females  were:  2 of  2 at  12  hours,  4 of  5 at  24  hours,  and  2 of  3 at  30  hours.  In 
one  experiment,  the  weight  of  material  ingested  was  known:  males  consumed  6.9  mg  and 
females  10.5  mg  of  citrated  human  blood.  Precipitin  tests  on  these  gave  the  following  fre- 
quencies of  positive  for  males:  6 of  1 1 at  25  hrs,  5 of  9 at  30  hrs  and  0 of  10  at  33  hours 
after  feeding.  The  data  for  females  were  10  of  10  at  25  hours,  5 of  5 at  30  hours  and  9 of  10 
at  33  hours. 

By  spectroscopic  examination  of  the  gut  contents  of  Rhodnius  prolixus  a month  or  more 
after  feeding  on  a rabbit,  Wigglesworth  (1943)  found  evidence  of  oxyhaemoglobin,  methae- 
moglobin  and  traces  of  acid  haematin  and  concluded  that  “even  after  storage  for  this  length 
of  time  in  the  stomach  [=anterior  mid-gut]  digestion  of  haemoglobin  has  scarcely  begun”.  A 
similar  examination  of  the  “coiled  intestine”  [=posterior  mid-gut]  showed  that  oxyhaemo- 
globin occurs  only  in  the  region  near  the  stomach  whereas  acid  haematin  exists  throughout. 
The  haemoglobin  is  rapidly  digested.  The  black  residue  remaining  in  the  rectum  consists  of 
free  haematin.  Globulin  apparently  is  digested  leaving  the  unchanged  iron  porphyrin  which 
is  excreted.  Some  intact  haemoglobin  is  absorbed  from  the  digestive  tract,  but  this  is  appar- 
ently not  due  to  excessive  stretching  of  the  stomach,  since  bugs  given  a partial  meal  also 
absorb  haemoglobin.  Some  of  the  absorbed  haemoglobin  passes  into  the  haemolymph  and 
some  is  digested  in  the  midgut  cells  to  form  a modified  haem  pigment  and  free  iron. 

Digestion  in  the  ‘intestine’  of  Triatoma  infestans  is  reported  to  “follow  the  same  lines  as 
in  Rhodnius”  (Wigglesworth,  1943).  In  C.  lectularius  “Digestion  in  the  lumen  of  the  gut 
proceeds  as  in  Rhodnius  but  no  brown  or  green  pigments  can  be  seen  in  the  epithelium  of 
the  stomach  or  the  intestine”  (Wigglesworth,  1943). 

The  human  body  louse,  P.  humanus  humanus  digests  its  blood  meal  in  the  midgut  but 
does  pass  undigested  haemoglobin  in  the  feces  as  evidenced  by  the  presence  of  methaemo- 
globin  and  oxyhaemoglobin  in  the  excreta  (Wigglesworth,  1943).  A positive  reaction  with 


Haematophagous  insects 


33 


benzidine  occurs,  both  with  mosquitoes  (A.  aegypti,  A.  scutellaris,  A.  notoscriptus,  A.  aus- 
tralis and  C.  pipiens  quinquefasciatus)  90  hours  after  feeding  upon  human  blood  and  with 
the  material  defecated  by  blood  fed  mosquitoes.  Thus  O’Gower  (1956)  concluded  that  the 
haem  of  the  haemoglobin  was  not  broken  down  in  the  mosquito.  The  flea  Nosopsyllus 
fasciatus  apparently  passes  undigested  haemoglobin  in  the  feces  (Wigglesworth,  1943). 

Wigglesworth  (1943),  using  spectroscopic  techniques,  found  no  evidence  for  absorption  of 
undigested  haemoglobin  in  mosquitoes  (A.  maculipennis  and  A.  aegypti)  or  the  flea  (N. 
fasciatus ).  However  evidence  was  obtained  for  its  absorption  in  Rhodnius,  Triatoma,  Eutria- 
toma,  Cimex  and  Pediculus. 

The  hematin  crystals  in  the  feces  of  P.  humanus  fed  on  man  are  rhombic  plates  whereas 
those  in  feces  of  P.  humanus  fed  on  rabbits  are  smaller  and  cubic  (Davis  and  Hansens,  1945). 
Whether  this  difference  is  attributable  entirely  to  the  source  of  the  blood  or  to  possible  dif- 
ferences in  the  way  P.  humanus  digests  human  and  rabbit  blood  was  not  mentioned. 

Gooding  (1966b)  determined  the  amount  of  water  soluble  protein  in  the  midguts  of  A. 
aegypti  and  C.  pipiens  quinquefasciatus  fed  on  chickens.  Assuming  that  the  decline  in  midgut 
protein  content  represents  digestion  of  protein  by  the  mosquitoes  two  criteria  can  be  applied 
for  comparing  the  rates  of  digestion:  (1)  the  decrease  in  protein  content  in  mg/midgut/time 
interval  or  (2)  the  time  required  for  a certain  percentage  of  the  protein  in  the  meal  to  be 
removed.  Using  the  first  criterion  C.  pipiens  quinquefasciatus  digests  its  meal  faster  than  A. 
aegypti.  With  the  second  criterion  the  reverse  is  true.  The  time  required  for  a 50%  and  a 90% 
decrease  in  the  protein  content  of  the  midgut  was  approximately  18  and  40  hours  respec- 
tively for  A.  aegypti  and  23  and  48  hours  for  C.  pipiens  quinquefasciatus.  (Briegel  (1969) 
reported  that  half  the  protein  in  a blood  meal  disappears  from  the  midgut  of  Culex  pipiens 
within  24  hours  of  feeding  and  that  digestion  was  completed  by  72  to  96  hours).  These  ex- 
periments were  done  under  essentially  the  same  environmental  conditions  as  those  of 
O’Gower  (1956)  except  that  the  mosquitoes  were  fed  on  chickens  by  Gooding,  (1966b)  and 
on  humans  by  O’Gower,  (1956).  O’Gower  found  that  50%  of  the  A.  aegypti  and  of  the  C. 
pipiens  quinquefasciatus  gave  negative  precipitin  tests  by  38  and  46  hours  respectively. 
Gooding  estimated  that  90%  of  the  water  soluble  protein  had  been  removed  from  the  mid- 
guts of  A.  aegypti  and  C.  pipiens  quinquefasciatus  by  40  and  48  hours  respectively.  O’Gower 
in  fact  studied  the  denaturation  of  serum  proteins  and  Gooding  the  decline  of  total  water 
soluble  proteins  in  both  serum  and  hemoglobin.  Although  the  two  studies  were  done  with 
blood-meals  from  different  sources  the  results  were  consistant  since  in  vitro  studies  showed 
that  these  mosquitoes  hydrolyzed  serum  proteins  slower  than  hemoglobin  (Gooding,  1966a). 

During  the  digestion  of  guinea-pig  blood  by  A.  aegypti  the  ratio  of  protein  nitrogen  to 
total  nitrogen  in  the  midgut  declines  during  the  first  48  hours  after  feeding  indicating  diges- 
tion of  the  meal  (Williams,  1953).  However  during  this  period  there  is  no  decrease  in  the 
total  nitrogen  content  of  the  midgut  indicating  either  that  there  is  no  absorption  or  that 
nitrogenous  materials  are  being  secreted  into  the  midgut. 

Fisk  (1950)  estimated  the  pH  of  the  midguts  of  A.  aegypti  and  A.  quadrimaculatus  with 
indicators.  Stomachs  of  unfed  mosquitoes  of  both  species  had  a pH  of  6.5  while  those  of  A. 
aegypti  fed  on  human  blood  had  a pH  of  7.3.  With  a microelectrode,  Micks,  deCaires,  and 
Franco  (1948)  found  the  pH’s  of  the  stomachs  of  unfed  C.  pipiens,  C.  pipiens  quinque- 
fasciatus, A.  aegypti,  and  A.  quadrimaculatus  to  be  7.27,  7.43,  7.31,  and  7.59.  When  these 
species  were  fed  upon  chicks  the  pH  values  of  the  stomachs  were  7.52,  7.60,  7.67  and  7.75. 
Similar  measurements  of  blood  from  the  stomachs  of  A.  aegypti  fed  on  chickens  gave  values 
ranging  from  pH  7.47  to  pH  7.90  (Bishop  and  McConnachie,  1956).  Twenty-four  of  the  30 
samples  had  pH  values  between  7.60  and  7.76  and  there  was  no  correlation  between  the 
duration  of  the  digestion  (up  to  70  minutes)  and  the  pH  value.  From  these  pH  estimates,  it 


34 


Gooding 


may  be  inferred  that  the  proteinases  responsible  for  digestion  of  the  blood  meal  must  func- 
tion in  a slightly  alkaline  medium.  MacGregor  (1931)  reported  that  the  midgut  of  A.  aegypti 
and  C.  pipiens  quinquefasciatus  fed  upon  a solution  (pH  7)  of  bacto-peptone  and  B.D.H. 
universal  pH  indicator  had  a pH  of  approximately  3 to  4.  Roy  (1937),  examining  dissected 
Gasterophilus  intestinalis  larvae  with  indicators,  determined  the  pH  of  various  parts  of  the 
alimentary  canal  to  be:  salivary  glands  pH  7.1,  proventriculus  pH  7,  middle  part  of  midgut 
pH  7.4  and  hindgut  pH  6.8.  The  pH  of  the  midgut  of  blood  fed  Glossina  submorsitans  was 
about  6.6.  The  exact  mechanism  for  controlling  the  gut  pH  and  the  contribution  of  the 
buffering  capacity  of  the  ingested  blood  to  its  control  have  not  been  elucidated,  nor  has  the 
relationship  between  gut  pH  and  the  pH  optimum  of  all  the  enzymes  functioning  in  the  gut. 

DeFreitas  and  Campos  (1961)  studied  the  rate  of  elimination  of  Fe59  by  fifth  instar  and 
adult  T.  infestans  and  by  first  instar  Panstrongylus  megistus  fed  upon  a chicken  which  had 
had  Fe59  incorporated  into  its  haemoglobin.  The  results  indicated  the  rate  of  digestion  of 
haemoglobin  by  these  bugs.  There  was  little  or  no  excretion  of  Fe59  during  the  first  six  days 
after  feeding  on  the  radioactive  blood.  The  time  required  for  elimination  of  50%  of  the  Fe59 
was  40  days  for  1st  instar  P.  megistus,  31  days  for  5th  instar,  and  16  days  for  adult  T. 
infestans. 

The  major  nitrogenous  wastes  in  the  feces  of  G.  morsitans  are  uric  acid,  arginine,  histidine, 
and  hematin  (Bursell,  1965).  A reasonably  close  agreement  existed  between  the  quantities  of 
these  compounds  excreted  after  the  first  hunger  cycle  and  the  amount  which  would  theoret- 
ically be  produced  from  a blood  meal.  The  differences  between  these  amounts  during  the 
first  hunger  cycle  were  accounted  for  by  the  development  of  the  flight  muscles  during  this 
period.  Bursell  (1965)  also  found  a correlation  between  the  amount  of  blood  consumed  and 
the  amount  of  uric  acid  produced.  Thus  one  can  use  the  rate  of  excretion  to  estimate  the 
rate  of  digestion. 

Langley  (1966b)  used  this  technique  to  show  that  male  G.  morsitans  digests  chicken  and 
lizard  blood  faster  than  mammalian  blood.  However,  the  rates  for  digestion  of  blood  from 
rat,  guinea-pig,  sheep,  cow,  bushpig,  or  man  do  not  differ  significantly.  Digestion  of  impala 
blood  is  also  at  about  the  same  rate  as  that  of  other  mammals  (Langley,  1968a).  Laboratory 
reared,  non-teneral  males  feeding  on  guinea-pigs  digest  their  meal  more  slowly  than  field- 
caught,  non-teneral,  males.  Digestion  is  fastest  in  males  which  are  caught  after  they  have  fed 
upon  oxen  (Langley,  1966b).  These  differences  led  Langley  (1966b)  to  propose  that  the 
prefeeding  behaviour  of  the  flies  affected  the  subsequent  rate  of  digestion.  Non-teneral, 
field-caught  males  feeding  upon  guinea-pigs  digest  this  meal  at  the  same  rate  when  held  in 
continuous  light  as  when  held  in  total  darkness,  even  though  the  males  are  less  active  in  total 
darkness. 

Field-caught  male  and  female  G.  morsitans,  feeding  on  oxen,  digest  their  blood  meals 
more  rapidly  than  laboratory  reared  males  and  females  feeding  on  bovine  or  guinea-pig  blood 
(Langley,  1967a).  Laboratory-reared,  fertilized  females  digest  their  blood  meal  more  rapidly 
than  unfertilized  females.  Male  G.  morsitans  feeding  on  ox  blood  in  the  field,  excrete  their 
blood  meal  more  rapidly  than  field-caught  males  feeding  upon  guinea-pigs  in  the  laboratory 
(Langley,  1967c).  There  is  no  difference  between  rate  of  digestion  of  guinea-pig  blood  and 
cow  blood  (Langley,  1967b).  Field-caught  males  fed  several  times  on  guinea  pigs  in  the  labo- 
ratory digest  each  meal  more  slowly  than  the  preceeding  one  until  by  the  third  meal  the  rate 
of  digestion  is  only  slightly  greater  than  for  laboratory-reared  flies  (Langley,  1966c).  The 
digestion  rate  in  male  G.  morsitans  is  not  affected  by  the  sex  or  reproductive  condition  of 
the  guinea-pigs  upon  which  they  feed  (Langley,  1968b). 


Haematophagous  insects 


35 


DIGESTIVE  ENZYMES  AND  THEIR  PROPERTIES 

A variety  of  digestive  enzymes  have  been  found  in  blood-sucking  insects.  Interpretation 
of  the  literature  is  straight  forward  except  for  the  frequent  occurrence  of  the  word  “trypsin” 
to  describe  a proteinase  with  maximum  activity  in  an  alkaline  medium.  There  are  several 
proteinases  (carboxypeptidase,  amino-peptidase,  chymotrypsin,  etc)  which  are  not  readily 
distinguished  from  trypsin  on  the  basis  of  the  pH-activity  curve  alone.  Therefore,  in  this 
discussion,  the  term  “trypsin”  will  be  reserved  for  an  alkaline  proteinase  which  cleaves  pep- 
tide bonds  on  the  carboxyl  side  of  a basic  amino  acid  and  “chymotrypsin”  for  one  which 
cleaves  on  the  carboxyl  side  of  an  aromatic  amino  acid.  Such  designations  are  based  upon 
the  use  of  synthetic  substrates.  Proteolytic  enzymes  active  in  the  alkaline  region,  without 
adequate  demonstration  of  the  bond  specificity,  shall  be  referred  to  as  proteinases  or,  if 
necessary,  alkaline  proteinases. 

Two  proteolytic  enzymes  occur  in  whole  Aedes  aegypti  adults  (Wagner,  Tenorio  and 
Terzian,  1961).  One  of  the  enzymes  is  a trypsin  found  in  the  midgut  of  the  female  but  not 
in  the  rest  of  the  mosquito  nor  in  any  part  of  the  male.  This  enzyme  separates  into  two 
fractions  on  a DEAE-cellulose  column,  the  2 fractions  have  similar  properties.  The  other 
enzyme  hydrolyzes  denatured  hemoglobin.  It  has  maximum  activity  at  pH  7.5,  but  functions 
almost  as  well  up  to  pH  9.  The  purified  enzyme  is  quite  stable  in  acid,  losing  none  of  its 
activity  at  pH  3 when  heated  to  96  C for  ten  minutes.  Wagner  et  al  ran  2 experiments  with 
females  to  determine  whether  the  activity  was  localized  in  the  midgut;  in  one  all  of  the  ac- 
tivity was  in  the  gut;  in  the  second  53%  of  the  activity  was  in  the  gut  homogenate.  Thus, 
the  authors  concluded  that  this  proteinase  was  primarily  a digestive  enzyme. 

The  tryptic  & chymotryptic  activities  of  A.  aegypti,  Culex  pipiens  quinquefasciatus  and 
Pediculus  humanus  are  due  to  2 different  enzymes  and  for  each  species  the  chymotrypsin 
has  a higher  molecular  weight  than  the  trypsin  (Gooding,  1968,  1969).  The  major  chymo- 
trypsin fractions  from  the  midguts  of  larval  and  adult  A.  aegypti  are  approximately  the  same 
molecular  weight  (Yang  and  Davies,  1971).  Trypsin  from  adult  A.  aegypti  has  a molecular 
weight  of  21,500  (Huang,  1971a).  The  proteinases  in  Rhodnius  prolixus  and  Cimex  lectula- 
rius  have  a high  molecular  weight  (>160,000)  (Gooding,  1968,  1969). 

On  paper  electrophoresis,  using  a barbital  buffer  at  pH  8.0,  3 cationic  bands  of  proteolytic 
activity  were  found  in  Stomoxys  calcitrans  midguts  (Patterson  and  Fisk,  1958).  However, 
using  starch  gel  electrophoresis,  cationic  bands  were  never  found  but  3 bands  of  proteinase 
activity  were  found  migrating  toward  the  anode  at  pH  7.6  (tris-citrate  buffer)  and  pH  8.0 
(barbital  buffer).  By  comparing  different  electrophoretic  fractions  (from  the  starch  gel 
electrophoresis)  with  respect  to  the  relative  rates  of  hydrolysis  of  azocasein  and  azoalbumin 
Patterson  and  Fisk  concluded  that  at  least  two  “trypsin-like”  enzymes  existed  in  the  midguts 
of  S.  calcitrans. 

Crystallized  hemoglobin  is  hydrolyzed  almost  as  rapidly  by  A.  aegypti  and  C.  pipiens 
quinquefasciatus  as  denatured  hemoglobin  (Gooding,  1966a).  However  it  is  not  known 
whether  the  crude  midgut  homogenates  used  in  these  experiments  contained  substances 
which  denatured  the  hemoglobin.  If  no  denaturing  agents  were  present,  then  the  proteinases 
of  these  mosquitoes  may  differ  significantly  from  mammalian  trypsins  which  do  not  readily 
attack  native  proteins  (Sumner  and  Somers,  1947,  p.  175).  Denaturation  of  the  proteins 
within  the  mosquito’s  midgut  prior  to  digestion  has  not  been  demonstrated  and  Fisk  (1950) 
wondered  whether  the  proteinases  of  mosquitoes  normally  attacked  native  proteins  or  pro- 
teins denatured  by  some  as  yet  unknown  mechanism.  He  suggested  that  coagulation  and 
agglutination  of  blood  denature  the  proteins  sufficiently  to  permit  attack  by  mosquito 
trypsin. 


36 


Gooding 


With  proteinases  from  A.  aegypti  and  C.  pipiens  quinquefasciatus  assayed  at  pH  7.9,  the 
following  are  the  Km  values  (in  mg/ml)  for  blood  proteins:  denatured  hemoglobin,  1.51  and 
1.32;  crystallized  hemoglobin,  1.84  and  3.15;  bovine  serum  albumin  fraction  V,  19.3  and 
8.51;  and  7-globulin  fraction  II,  374,  and  6.22  (Gooding,  1966a).  At  pH  9.5  there  is  little 
hydrolysis  of  the  serum  proteins  by  either  species.  The  Km  values  for  A.  aegypti  and  C. 
pipiens  quinquefasciatus  are:  denatured  hemoglobin  2.75  and  2.10,  and  crystallized  hemo- 
globin 4.08  and  2.17.  Using  purified  A.  aegypti  midgut  trypsin,  the  Km  values  at  pH  7.9  are 
2.24  mM  for  denatured  hemoglobin  and  0.47  mM  for  benzoyl-DL-arginine-p  -nitroanilide 
(BAPNA)  (Huang,  1971a).  Davies  and  Yang  (1968)  reported  trypsin  with  a pH  optimum  of 
8.4  from  the  midguts  of  6 simuliid  species  ( Cnephia  dacotensis  (Duar  and  Shannan),  Prosi- 
mulium  decemarticulatum,  Prosimulium  fuscurn,  Simulium  rugglesi,  Simulium  venustum 
and  Simulium  vittatum).  The  Km  values  for  tosyl-L-arginine  methyl  ester  (TAME)  are  2.4 
mM  for  S.  venustum  and  3.1  mM  for  S.  rugglesi  (Yang  and  Davies,  1968b). 

The  in  vitro  temperature  optima  for  A.  aegypti  and  C.  pipiens  quinquefasciatus  protein- 
ases are  in  the  range  46  to  50  C (Gooding,  1966a,  1968),  that  for  S.  calcitrans  near  50  C 
(Patterson  and  Fisk,  1958),  and  for  C.  lectularius  and  R.  prolixus  about  45  to  50  C (Gooding, 
1968).  These  optima  are  typical  of  alkaline  proteinases.  The  temperature  optima  for  mam- 
malian trypsins  range  from  45  to  55  C (Buck,  Bier,  and  Nord,  1962).  For  the  housefly  the 
optimum  is  45  C (Lin  and  Richards,  1956),  and  for  the  larval  blowfly  it  is  44  C (Evans, 
1958).  The  A.  aegypti  proteinase  experiments  of  Fisk  (1950),  Fisk  and  Shambaugh  (1952), 
and  Shambaugh  (1954),  were  carried  out  at  approximately  40  C,  a temperature  at  which  the 
enzyme  is  functioning  at  only  half  its  maximum  rate.  Wagner  et  al  (1961)  carried  out  assays 
with  A aegypti  proteinase  at  30  C,  a temperature  at  which  the  enzyme  has  about  one  quarter 
the  activity  it  has  at  its  temperature  optimum. 

The  activity  of  the  non-trypsin  proteinase  from  A.  aegypti  is  increased  by  diisopropyl- 
fluorophosphate  (DFP),  p-chloromercuribenzoate  and  sometimes  cystine.  Crystalline  soy- 
bean trypsin  inhibitor  gives  some  inhibition,  as  do  several  cations  (ipagnesium,  calcium, 
mercury,  and  manganese)  (Wagner  et  al,  1961).  The  alkaline  proteinase  activity  of  A.  aegypti 
and  C.  pipiens  quinquefasciatus  is  inhibited  to  some  extent  by  calcium,  magnesium,  and 
manganese  when  denatured  hemoglobin  or  bovine  serum  albumin  are  used  as  the  substrates 
(Gooding,  1966a).  However  the  proteinase  from  S.  calcitrans  is  not  affected  by  several  ions 
(calcium,  magnesium,  sodium,  chloride,  or  fluoride),  penicillin  G,  or  dialysis  against  distilled 
water  (Patterson  and  Fisk,  1958). 

Cations  have  varying  effects  upon  partially  purified  A.  aegypti  trypsin  (Wagner  et  al, 
1961).  Magnesium  and  manganese  have  no  effect,  but  calcium,  mercury,  cadmium,  and  zinc 
inhibit  the  enzyme  to  varying  degrees.  The  enzyme  is  inhibited  by  p-chloromercuribenzoate 
but  cystine  has  no  effect.  This  enzyme  is  not  inhibited  by  crystalline  soybean  trypsin  inhibi- 
tor (Wagner  et  al,  1961).  However,  Akov  (1965)  stated  that  soybean  trypsin  inhibitor 
inhibited  A.  aegypti  trypsin  in  vitro  and  presented  data  to  show  that  mosquitoes  fed  on 
citrated  sheep  blood  containing  0.625  jug/ml  had  less  than  half  the  proteinase  activity  found 
in  the  controls  24  hours  after  the  meal.  Gooding  (1969)  reported  inhibition  of  trypsin  from 
A.  aegypti  and  C.  pipiens  quinquefasciatus  and  chymotrypsin  from  P.  humanus  by  soybean 
trypsin  inhibitor. 

A.  aegypti  trypsin  is  inactivated  by  DFP  (Wagner  et  al,  1961).  Phenylmethane  sulphonyl 
fluoride  (PMSF)  inhibits  trypsin  from  A.  aegypti  and  C.  pipiens  quinquefasciatus  and  chymo- 
trypsin from  P.  humanus  (Gooding,  1968,  1969).  (These  inhibitors  are  known  to  inhibit 
mamalian  trypsin  and  chymotrypsin  by  reaction  with  serine  at  the  active  center  of  the  en- 
zyme; therefore,  it  may  be  inferred  that  the  mosquito  and  louse  enzymes  studied  have  serine 
at  their  active  centers.)  Tosyl-L-lysine  chloromethyl  ketone  (TLCK)  inhibits  A.  aegypti  and 


Haematophagous  insects 


37 


C.  pipiens  quinquefasciatus  trypsin  but  not  P.  humanus  chymotrypsin,  while  tosyl-amide- 
phenylethylchloromethyl  ketone  (TPCK)  inhibits  P.  humanus  chymotrypsin  but  not  A. 
aegypti  or  C.  pipiens  quinquefasciatus  trypsin  (Gooding  1968,  1969)  (TLCK  is  a specific 
inhibitor  of  mammalian  trypsin  while  TPCK  is  a specific  inhibitor  of  mammalian  chymo- 
trypsin. Both  compounds  react  with  histidine  at  the  active  center  and  therefore  it  may  be 
inferred  that  the  insect  enzymes  studied  have  histidine  at  their  active  centers). 

Although  Fisk  (1950)  used  heparinized  whole  rabbit  blood  as  a substrate  for  most  of  his 
experiments  he  used  a 1%  solution  of  serum  albumin  for  his  studies  on  the  effects  of  pH. 
Fisk  calculated  that  during  the  assay  only  0.34%  of  the  available  protein  in  the  rabbit  blood 
was  hydrolyzed  and  concluded  that  the  presence  of  4 mg  of  blood  per  midgut  was  saturating 
the  enzyme  with  substrate.  He  noted  that  the  reaction  rate  when  serum  albumin  was  used 
was  almost  3 times  as  great  as  when  blood  was  used  as  the  substrate.  He  suggested  that  blood 
from  the  rabbit  may  contain  some  substances  which  inhibit  the  midgut  proteinase  of  A 
aegypti  and  that  some  of  the  mosquito  proteinase  may  combine  with  the  inhibitor  to  neu- 
tralize it  in  the  midgut.  Gooding  (1966a)  demonstrated  that  serum  from  both  normal  and 
Plasmodium  gallinaceum  infected  chicks  will  inhibit  the  in  vitro  activity  of  proteinases  from 
A.  aeg)?pti  and  C.  pipiens  quinquefasciatus  when  denatured  hemoglobin  is  used  as  the  sub- 
strate. To  obtain  a 50%  inhibition  of  the  proteinases,  from  1 to  7 p\  serum/ml  reaction 
mixture  was  required. 

The  sera  of  17  vertebrate  species  and  the  hemolymph  of  Periplaneta  americana  (L.) 
inhibit  A.  aegypti  midgut  trypsin  (Huang,  1971a).  The  inhibition  capacity  of  whole  serum 
varies  from  a low  of  0.01  pg  trypsin  inhibited/pig  serum  for  the  dogfish  to  a high  of  0.21 
pg  trypsin  inhibited//d  serum  for  the  chicken.  Huang  used  G-200  Sephadex  gel  filtration  to 
determine  the  minimum  number  of  inhibitors  in  serum  and  estimate  their  molecular  weights. 
P.  americana  hemolymph  has  1 inhibitor  with  a molecular  weight  of  <11,000.  All  of  the 
vertebrate  species  examined  have  an  inhibitor  with  a molecular  weight  between  3 1 ,800  and 
66,100.  This  is  the  only  inhibitor  in  4 species  (dogfish,  turkey,  chicken,  and  rat).  All  other 
vertebrate  species  have  a second  inhibitor  with  molecular  weight  >160,000.  A third  inhibi- 
tor with  a molecular  weight  between  77,600  and  107,000  occurs  in  4 species  (man;  turtle, 
frog,  and  pike). 

Huang  (1971b)  partially  purified  two  inhibitors  of  A.  aegypti  trypsin  from  bovine  serum. 
One  of  these,  inhibitor  I,  is  electrophoretically  associated  with  the  ax  -globulin  fraction  of 
serum,  has  a molecular  weight  of  about  43,500  and  combines  with  trypsin  in  the  molar  ratio 
of  3.5  inhibitor  I molecules  to  1 trypsin  molecule.  Hill  plots  indicate  that  2 molecules  of 
inhibitor  I inactivate  1 enzymic  site  of  trypsin.  Inhibition  of  trypsin  by  inhibitor  I is  com- 
petitive when  haemoglobin  is  the  substrate  at  37  C.  When  BAPNA  is  the  substrate  inhibition 
is  competitive  at  30  and  34  C and  non-competitive  at  37  and  44.5  C.  Inhibitor  II  is  electro- 
phoretically associated  with  a2  -macroglobulin,  has  a molecular  weight  of  >160,000  (possi- 
bly as  high  as  1,000,000)  and  forms  a complex  with  A.  aegypti  trypsin  in  the  ratio  1.7 
molecules  inhibitor  II/molecule  of  trypsin.  Hill  plots  indicate  that  2 molecules  of  inhibitor 
II  inactivate  1 enzymatic  site  of  trypsin.  Huang  demonstrated  the  complex  of  trypsin  and 
inhibitor  II  electrophoretically  and  by  gel  filtration.  This  complex  has  very  little  proteolytic 
activity  when  hemoglobin  is  the  substrate  but  retains  most  of  the  esterolytic  activity  when 
BAPNA  is  the  substrate.  Inhibitor  II  has  the  interesting  property  of  protecting  the  mosquito 
trypsin  against  inhibition  by  inhibitor  I,  soybean  trypsin  inhibitor  and  PMSF  but  not  against 
inactivation  by  TLCK.  Although  the  pH  optimum  is  similar  for  free  trypsin  and  trypsin- 
inhibitor  II  complex,  the  Km  for  BAPNA  is  lower  for  the  trypsin-inhibitor  II  complex 
(0.21  ± 0.007  mM)  than  for  free  trypsin  (0.47  ± 0.01  mM).  When  hemoglobin  is  used  as  a 
substrate,  inhibitor  II  is  a competitive  inhibitor  at  37  C but  when  BAPNA  is  used  as  the 


38 


Gooding 


substrate  inhibition  is  competitive  at  30  and  34  C and  non-competitive  at  37  and  44.5  C. 

The  chymotrypsin  from  A.  aegypti  larvae  is  inhibited  by  human  and  horse  sera  and  in 
both  cases  inhibition  is  associated  with  the  a-globulin  fraction  (Yang  and  Davies,  1971). 
Inhibition  by  human  a-globulin  is  competitive.  The  inhibitors  are  unaffected  by  heating  to 
60  C but  are  inactivated  at  100  C. 

In  view  of  the  presence  of  a trypsin  inhibitor  in  the  hemolymph  of  P.  americana  (Huang, 
1971a)  it  is  interesting  that  the  trypsin  activity  of  the  midgut  of  blackflies  is  approximately 
the  same  as  homogenates  of  whole  flies  (Yang  and  Davies,  1968b).  This  latter  observation 
led  Yang  and  Davies  to  conclude  that  blackfly  tissues  probably  do  not  contain  materials 
inhibiting  their  trypsin. 

The  midguts  of  several  insects  have  substances  which  influence  the  clotting  of  blood.  The 
occurrence  of  coagulins  and  anticoagulins  among  various  species  of  insects  is  summarized 
in  table  2. 

The  anterior  third  of  the  midguts  of  Glossina  tachinoides  and  Glossina  morsitans  contains 
an  anticoagulin,  but  by  removing  the  salivary  glands  it  can  be  demonstrated  that  this  anti- 
coagulin  is  derived  from  the  salivary  glands  (Lester  and  Lloyd,  1928).  However,  the  antico- 
agulin found  in  the  midgut  of  Anopheles  maculipennis  probably  does  not  originate  in  the 
salivary  glands,  since  it  is  destroyed  by  heating  to  80  C while  the  salivary  gland  anticoagulin 
is  stable  at  100  C for  35  mins  (de  Buck,  1937). 

R.  prolixus  midgut  anticoagulin  is  stable  at  60  and  80  C for  30  mins,  is  not  affected  by 
treatment  with  0.1  N HC1  or  0.1  N NaOH  at  R.T.  or  60  C for  30  mins,  but  is  destroyed  by 
heating  to  100  C for  5 mins  (Hellmann  and  Hawkins,  1964).  It  is  not  precipitated  by  centri- 
fuging at  100,000  g for  30  mins  but  is  removed  from  solution  by  dialysis  in  the  cold.  It 
prevents  clotting  of  rat,  guinea-pig,  cat,  and  human  blood.  The  gut  anticoagulin  was  desig- 
nated Prolixin-G  by  Hellmann  and  Hawkins  (1965).  Prolixin-G  inhibits  thrombin  and  has  a 
molecular  weight  between  100,000  and  200,000,  is  soluble  in  saline  but  is  insoluble  in  water 
and  is  therefore  possibly  a euglobulin.  It  is  less  stable  than  the  salivary  anticoagulin, 
Prolixin-S  (which  also  occurs  in  the  midgut)  when  tested  by  storage  at  -20  C for  24  hours, 
freeze-drying  or  dialysis.  It  is,  however,  more  stable  than  Prolixin-S  when  exposed  to  dilute 
trypsin  solutions  (Hellmann  and  Hawkins,  1965). 

An  antithrombin  in  the  midguts  of  Triatoma  maculata  is  distinct  from  the  salivary  antico- 
agulin (Hellmann  and  Hawkins,  1966).  The  gut  anticoagulin  is  not  affected  by  incubation 
•with  protamine  sulfate,  is  fairly  stable  at  60  and  80  C,  but  loses  about  half  its  activity  at 
100  C in  2 mins  and  all  its  activity  in  30  mins.  Anticoagulin  activity  is  not  lost  by  treatment 
with  0.1  N HC1  for  10  mins  at  R.T.  or  60  C,  but  is  lost  in  0.1  N NaOH  at  60  C in  10  mins. 
Freezing  and  freeze-drying  cause  a loss  of  activity  but  the  anticoagulin  is  stable  in  the  refrig- 
erator. This  antithrombin  has  a molecular  weight  between  100,000  and  200,000,  is  resistant 
to  trypsin  but  loses  some  activity  when  dialyzed  against  saline  (Hellmann  and  Hawkins, 
1966). 

Hellmann  and  Hawkins  (1966)  found  no  indication  of  fibrinolytic  activity  in  the  salivary 
glands  of  T.  maculata.  The  gut,  however,  contains  a plasminogen-activating  factor  but  no 
active  fibrinolytic  enzymes.  Guts  with  low  anticoagulin  activity  have  high  fibrinolytic  activi- 
ty and  visa  versa.  Freeze-drying  decreases  the  fibrinolytic  activity  of  the  gut  preparation  but 
storage  of  dry  preparations  under  refrigeration  and  storage  at  -20  C does  not  cause  a loss  of 
activity.  The  activity  is  not  destroyed  at  60  C but  is  in  2 mins  at  80  C or  100  C and  in  5 mins 
at  R.T.  in  0. 1 N HC1  or  0.1  N NaOH.  The  properties  of  the  gut  antithrombin  from  T.  macu- 
lata are  very  similar  to  the  gut  antithrombin  from  R.  prolixus.  (Hellmann  and  Hawkins 
1966). 

The  gut  homogenates,  but  not  the  salivary  gland  homogenates,  of  R.  prolixus  contain 


Haematophagous  insects 


39 


fibrinolytic  activity.  The  gut  contains  a fibrinolytic  activator  and  there  is  evidence  that  it 
also  contains  some  weak  fibrinolytic  enzyme  (Hellmann  and  Hawkins,  1964).  The  fibrino- 
lytic activity  in  the  gut  is  destroyed  by  heating  to  100  C for  5 mins  and  by  treating  with  0.1 
N NaOH  or  0.1  N HC1  for  30  mins  at  R.T.  A 5 minute  exposure  to  0.1  N HC1  causes  a 70% 
loss  of  activity  while  0.1  N NaOH  has  no  effect  in  5 mins.  A single  gut  contains  1 N.I.H. 
unit  of  urokinase.  A dialyzable  fibrinolytic  inhibitor  is  also  present  in  the  midguts  of  fed  R. 
prolixus ; the  source  of  this  inhibitor  (insect  or  host)  is  unknown.  The  addition  of  soybean 
trypsin  inhibitor  to  the  R.  prolixus  gut  extract  does  not  inhibit  the  lysis  of  fibrin  and  it  was 
concluded  that  “it  is  therefore  unlikely  that  the  fibrinolytic  activity  of  the  gut  extract  is  due 
to  trypsin”. 

Hawkins  and  Hellmann  (1966)  demonstrated  that  the  midgut  of  R.  prolixus  contains  a 
plasminogen  activator  which  is  detectable  by  measuring  either  fibrinolysis  or  caseinolysis 
and  they  proposed  the  name  “rhokinase”  for  this  material.  Rhokinase  activates  plasminogen 
directly  and  its  activity  in  the  assay  systems  increases  with  time.  The  source  of  rhokinase  in 
the  midgut  is  unknown  but  it  is  not  detectable  in  the  blood  of  the  guinea-pig  on  which  the 
bugs  fed,  the  salivary  glands,  the  midgut  cells  of  the  bug,  or  in  cultures  of  the  bug’s  symbi- 
ont, Nocardia  rhodnii  (Erikson). 

Lysis  of  the  blood  clot  in  the  digestive  tract  of  Glossina  austeni  is  accomplished  by  two 
agents  (Hawkins,  1966).  The  salivary  glands  and  crop  contain  a substance  presumably  from 
the  salivary  glands,  which  activates  plasminogen,  resulting  in  lysis  of  the  clot.  The  midgut 
and  hindgut  also  contain  an  enzyme  which  is  inhibited  by  soybean  trypsin  inhibitor.  This 
enzyme  was  presumed  by  Hawkins  (1966)  to  be  trypsin.  Hawkins  showed  that  this  same 
enzyme  was  responsible  for  clot  formation  of  oxalated  plasma. 

A coagulin  found  in  the  abdomen  (presumably  in  the  midgut)  of  Pthirus  pubis  can  neu- 
tralize the  anticoagulin  found  in  the  head  and  thorax  (presumably  in  the  salivary  glands) 
(Grusz,  1923).  Similarly,  the  coagulins  in  the  midguts  of  Culiseta  annulata  and  Culex  pipiens 
hasten  clotting  of  both  normal  blood  and  that  which  was  treated  with  the  salivary  glands  of 
A.  maculipennis,  C.  pipiens  quinquefasciatus,  or  C.  annulata  (de  Buck,  1937).  Presumably, 
the  coagulins  neutralize  the  salivary  anticoagulins.  The  coagulin  from  C.  annulata  is  not 
destroyed  by  drying  or  by  heating  dry  material  to  99  C for  1 hour  but  saline  solutions  are 
inactivated  by  heating  to  50  C for  15  mins. 

The  midgut  coagulin  from  G.  morsitans  is  completely  inactivated  by  treating  with  0.1  N 
KOH  for  10  mins,  or  by  heating  to  80  C for  15  mins,  while  treating  with  0.1  N HC1  for  10 
mins  destroys  about  half  the  activity  (Lester  and  Lloyd,  1928).  By  mixing  salivary  gland  and 
midgut  homogenates  in  various  ratios  and  by  adding  these  mixtures  to  sheep  blood,  the 
clotting  time  may  be  prolonged  or  shortened.  If  the  salivary  anticoagulin  is  added  to  blood  1 
min  before  adding  varying  amounts  of  coagulin,  the  clotting  time  can  be  shortened,  but  not 
to  the  same  extent  as  when  the  coagulin  is  added  to  the  anticoagulin  before  mixing  with 
the  blood.  Lester  and  Lloyd  interpreted  these  findings  as  indicating  that  the  midgut  coagu- 
lin inactivates  the  salivary  gland  anticoagulin  and  they  suggested  that  the  midgut  coagulin 
was  similar  to  vertebrate  kinase  (i.e.  the  enzyme  which  converts  prothrombin  to  thrombin). 
Lloyd  (1928),  speculating  on  the  relationship  between  the  function  of  the  midgut  coagulin 
and  clot  formation,  stated  that  the  “main  function  of  this  clot  appears  to  be  that  it  puts  a 
brake  on  to  the  fluid  meal  and  holds  it  in  the  proper  region  of  the  gut  while  digestion  be- 
gins”. The  coagulin  from  Musca  crassirostris  is  destroyed  by  heating  to  100  C for  10  mins 
and  its  concentration  in  the  midgut  reaches  a maximum  20  to  44  hours  after  a meal 
(Cornwall  and  Patton,  1914). 

In  invertases  from  P.  fuscum  and  S.  venustum  have  maximum  activity  at  pH  6.2  (Yang 
and  Davies,  1968c;  Davies  and  Yang,  1968).  The  synthesis  of  oligosaccharides  by  the  invert- 


40 


Gooding 


ase  of  S.  venustum  was  detected  by  Yang  and  Davies  but  the  products  were  not  identified  so 
it  was  not  established  whether  the  invertase  was  of  the  a-glucosidase  or  /3-fructosidase  type. 

Gasterophilus  intestinalis  midgut  amylase  has  optimal  activity  at  pH  6 and  is  activated  by 
chloride  (Tatchell,  1958).  Maltase  and  invertase  have  optimal  activity  at  pH  6.  Tatchell 
demonstrated  that  lipase  hydrolysing  tributyrin  had  maximal  activity  at  pH  7 but  no  hydrol- 
ysis of  olive  oil  or  ethyl  butyrate  was  demonstrated.  The  amylase  from  both  male  and 
female  S.  venustum  has  a pH  optimum  at  approximately  pH  6.5  (Yang  and  Davies,  1968a). 
The  Km  (starch)  is  0.65  mg/ml  for  female  S.  venustum  amylase  (Yang  and  Davies,  1968a). 

ENZYME  CONTENT  OF  THE  GUT 

Proteolytic  activity  in  the  midgut  of  Aedes  aegypti  is  significantly  higher  in  mosquitoes 
one  or  two  hours  after  a partial  blood  meal  than  in  sugar-fed  mosquitoes  (Fisk,  1950).  The 
addition  of  homogenates  of  crops  and/or  salivary  glands  does  not  markedly  increase  the 
proteolytic  activity  of  midgut  homogenates  of  unfed  mosquitoes.  Immediately  after  A. 
aegypti  feed  on  human  blood,  the  proteinase  activity  in  the  midgut  drops  below  the  level  of 
the  unfed  midgut.  When  mosquitoes  are  kept  at  26.6  C and  50%  R.H.  the  activity  rises  to  a 
maximum  about  18  hours  after  feeding  and  then  slowly  declines  (Fisk  and  Shambaugh, 
1954).  Fisk  and  Shambaugh  proposed  that  the  initial  decline  in  proteinase  activity  in  the 
midgut  was  due  either  to  depletion  of  enzyme  due  to  an  excess  of  substrate  or  to  the  pres- 
ence of  an  antitrypsin  in  the  serum.  Feeding  on  sugar  causes  a slight  increase  in  proteolytic 
activity  after  one  hour,  but  the  level  returns  to  normal  by  two  hours.  Secretion  of  the  pro- 
teolytic enzymes  is  primarily  in  response  to  serum  proteins  in  the  meal  and  there  is  a direct 
correlation  between  the  amount  of  blood  ingested  and  the  proteinase  activity  of  the  midgut 
homogenate  (Shambaugh,  1954).  Incubation  of  A.  aegypti  midgut  homogenates  with  blood 
for  1 8 hours  at  40  C does  not  result  in  production  of  detectable  quantities  of  proteinase, 
but  midguts  dissected  from  mosquitoes  18  hours  after  feeding  on  human  blood  have  a large 
quantity  of  proteinase. 

A.  aegypti  feeding  on  5-fluorouracil  (5-FU)  in  sugar  solutions  prior  to,  or  with,  the  blood 
meal  have  lower  midgut  proteinase  levels  than  controls  24  hours  after  the  blood  meal  (Akov, 
1965).  The  suppression  of  the  proteinase  level  decreases  with  time  when  the  mosquitoes  are 
taken  off  the  5— FU  diet.  Mosquito  proteinase  is  not  inhibited  in  vitro  by  5-FU. 

A.  aegypti  treated  with  metepa  (applied  topically  or  fed  a sugar  solution),  apholate  (sugar 
solution)  or  gamma  irradiation  have  a normal  amount  of  midgut  proteinase  one  day  after 
feeding  on  a rat  (Akov,  1966).  The  treated  mosquitoes,  however,  retain  blood  in  their 
midguts  longer  than  the  controls  and  two  days  after  feeding  have  much  more  midgut  protein- 
ase than  the  controls  (which,  incidently,  have  empty  midguts).  When  metepa  is  mixed  with 
citrated  sheep’s  blood  and  fed  to  A.  aegypti  through  a membrane,  the  midgut  proteinase 
activity  is  higher  than  in  the  controls  but  even  so  the  blood  meal  is  retained  longer  in  the 
metepa  treated  insects. 

Gooding  (1966a)  found  much  more  proteinase  in  the  midguts  of  A.  aegypti  and  Culex 
pipiens  quinquefasciatus  24  hours  after  feeding  on  chicks  than  in  the  midguts  of  unfed 
mosquitoes.  In  these  experiments  denatured  hemoglobin  was  used  as  the  substrate  and  the 
elevated  proteinase  levels  in  the  fed  mosquitoes  were  found  at  all  pH  values  from  4 to  1 1. 
Proteinase  activity  in  C.  pipiens  quinquefasciatus  reached  a maximum  36  hours  after  feeding 
and  in  A.  aegypti  usually  24  hours  (but  on  1 occasion  36  hours)  after  feeding  (Gooding, 
1966b).  The  time  at  which  the  maximum  concentration  of  proteinase  was  present  in  the 
midgut  was  not  influenced  by  holding  the  mosquitoes  in  continuous  light  or  continuous 
darkness.  In  a single  experiment  in  which  A.  aegypti  and  C.  pipiens  quinquefasciatus  were 


Haematophagous  insects 


41 


fed  on  normal  or  Plasmodium  gallinaceum  infected  chicks,  those  mosquitoes  which  fed  on 
the  infected  bird  had  a higher  proteinase  content  in  their  midgut.  Two  experiments  were  run 
comparing  normal  A.  aegypti  with  those  having  oocysts  of  P.  gallinaceum  on  the  midgut  at 
the  time  of  the  second  blood  meal.  In  both  experiments  the  infected  groups  of  mosquitoes 
had  higher  proteinase  activity  than  the  uninfected  groups.  The  maximum  proteinase  activity 
in  all  six  infected  groups  occurred  36  hours  after  the  blood  meal  while  in  the  two  uninfected 
groups  it  occurred  at  24  hrs  for  one  and  at  36  hours  for  the  other.  Yang  and  Davies  (1971) 
found  that  trypsin  activity  but  not  chymotrypsin  activity  in  the  midguts  of  A.  aegypti  rose 
after  a blood  meal.  Combre  et  al  (1971)  reported  that  adult  A.  aegypti  have  lower  chymo- 
trypsin activity  than  the  larvae. 

Although  the  complete  mechanism  for  the  control  of  proteinase  secretion  in  the  mosquito 
is  not  known,  it  is  clear  that  serum  proteins  stimulate  proteinase  secretion  in  A.  aegypti 
(Shambaugh,  1954).  The  greatest  secretion  results  when  mosquitoes  consume  a mixture  of 
serum  proteins,  but  the  presence  of  any  one  serum  protein  in  the  meal  also  stimulates 
secretion. 

Fisk  and  Shambaugh  (1952)  and  Shambaugh  (1954)  began  studies  to  elucidate  the  mech- 
anism which  controls  the  production  of  midgut  proteinase  in  A.  aegypti  following  a blood 
meal,  but  attempts  to  stimulate  enzyme  secretion  by  injection  of  hemolymph  from  fed  to 
unfed  mosquitoes  failed.  Detinova  (1962,  p.  59),  from  studies  of  digestion  and  ovarian 
development  in  Anopheles  maculipennis  concluded  that  “the  process  of  ovarian  develop- 
ment slows  down  the  speed  of  digestion.  Neurohormonal  regulation  of  the  duration  of  the 
digestive  process  may  therefore  be  postulated  for  mosquitoes.”  Autogenous  Aedes  atropal- 
pus  generally  do  not  take  a blood  meal  during  the  first  gonotrophic  cycle  (Hudson,  1970). 
Those  which  feed  utilize  neither  the  protein  nor  the  carbohydrate  in  the  blood  meal  during 
production  of  the  first  batch  of  eggs.  The  results  reported  by  Detinova  (1962)  and  Hudson 
(1970)  suggest  that  mosquitoes  with  mature,  or  nearly  mature,  eggs  are  incapable  of  syn- 
thesizing or  releasing  normal  quantities  of  digestive  enzymes. 

The  amount  of  midgut  proteinase  present  in  A.  aegypti  27  to  28  hours  after  engorging  on 
chicks  is  much  lower  in  mosquitoes  decapitated  within  six  hours  of  feeding  than  in  dewinged 
individuals  (Gooding,  1966b).  These  results  are  consistent  with  but  not  proof  of,  humoral 
control  of  proteinase  secretion.  However,  using  net  synthesis  of  triglycerides  as  a criterion 
for  digestion  of  the  blood-meal,  Lea  (1967)  concluded  that  ablation  of  the  median  neuro- 
secretory cells  of  mosquitoes  does  not  affect  digestion  or  absorption.  The  results  of  experi- 
ments with  5-FU  (Akov,  1965)  suggest,  but  do  not  prove,  that  the  midgut  proteinase 
secreted  by  A.  aegypti  is  formed  de  novo  after  the  ingestion  of  the  blood  meal. 

In  Simulium  venustum  the  trypsin  activity  is  higher  in  midguts  of  blood-fed  females  than 
in  females  fed  on  sucrose  only  (Yang  and  Davies,  1968b).  In  Simulium  rugglesi  fed  on  a 
duck,  the  trypsin  level  rises  steadily  for  18  hours  after  feeding,  and  remains  essentially 
unchanged  at  24  hours.  In  Prosimulium  decemarticulatum  feeding  on  a chicken,  the  trypsin 
level  rises  sharply  by  5 hours  then  drops  slightly  during  the  next  19  hours.  S.  venustum 
adults  feeding  on  a 50%  human  blood  - 0.5  M sucrose  solution,  and  kept  at  15  C nearly 
double  their  trypsin  activity  within  24  hours  and  maintain  this  level  for  about  8 days  (Yang 
and  Davies,  1968b).  At  30  C the  trypsin  level  is  nearly  triple  that  of  sugar  fed  controls  and 
is  also  maintained  at  this  elevated  level  for  8 days.  Males  feeding  on  the  blood-sucrose 
mixture  and  maintained  at  15  C have  a slightly  depressed  trypsin  level  for  4 days.  S.  venus- 
tum females  feeding  on  duck  erythrocytes  have  only  slightly  less  trypsin  than  those  feeding 
upon  whole  duck  blood;  the  reverse  is  true  when  feeding  on  material  of  bovine  origin.  The 
blood  meal  stimulates  trypsin  secretion  in  the  midgut  of  P.  decemarticulatum,  S.  rugglesi 
and  S.  venustum  (Davies  and  Yang,  1968). 


42 


Gooding 


In  Stomoxys  calcitrans  the  proteinase  activity  reaches  a maximum  about  13  hours  after  a 
blood  meal  but  remains  essentially  unchanged  after  a sucrose  meal  (Champlain  and  Fisk, 
1956).  No  depletion  of  the  midgut  proteinase  below  the  level  of  unfed  flies  occurs.  These 
authors  attributed  this  difference  in  pattern  between  A.  aegypti  and  S.  calcitrans  to  the 
different  ways  in  which  these  species  distribute  the  blood  meal  between  the  midgut  and 
crop  diverticula. 

There  is  a positive  linear  relationship  between  meal  size  and  proteinase  activity  in  the 
midgut  of  Glossina  morsitans  at  6,  18,  24,  48,  72,  and  96  hours  after  the  blood  meal 
(Langley,  1966a).  When  the  concentration  of  defibrinated  blood  in  the  meal  is  varied  from 
10%  to  100%  but  the  specific  gravity  remains  relatively  constant  (1.00-1.05),  the  amount  of 
proteinase  activity  varies  with  the  volume  of  the  meal  and  not  the  concentration  of  the 
blood  in  the  meal.  However,  since  flies  consuming  saline  alone  do  not  secrete  proteinase  it 
appears  that  some  blood  must  be  present  for  proteinase  to  be  produced.  The  stimulus  for 
proteinase  production  is  in  the  serum  rather  than  in  the  erythrocyte.  Langley  proposed  that 
stimulation  of  stretch  receptors  in  the  crop  duct  causes  impulses  to  pass  along  the  oesoph- 
ageal nerves  to  the  neuroendocrine  system,  resulting  in  the  production  and/or  release  of 
hormones  which  cause  the  middle  portion  of  the  midgut  to  produce  the  precursors  of  the 
proteinase.  He  further  suggested  that  the  enzyme  is  then  activated  by  some  component  of 
the  serum  portion  of  the  meal.  The  frequency  of  feeding  of  G.  morsitans  (every  48,  72  or  96 
hours)  does  not  affect  the  production  of  proteinase  (Langley,  1969a).  Females  have  a higher 
maximum  proteinase  level  in  the  midgut  than  males.  In  females  the  maximum  amount  of 
proteinase  occurs  24  to  48  hours  after  a meal  while  in  males  it  occurs  between  1 2 and  24 
hours  after  the  meal.  The  maximum  level  of  proteinase  in  field  caught  males  is  about  1.5 
times  the  maximum  level  in  laboratory  reared  males.  This  latter  finding  is  consistant  with 
the  fact  that  field  caught  flies  excrete  the  blood  meal  more  rapidly  than  laboratory  reared 
flies  (Langley,  1966b,  1967a). 

The  amount  of  proteinase  in  the  midgut  of  unfed  G.  morsitans  rises,  during  the  first  24 
hours  of  adult  life,  remains  constant  until  the  fly  is  96  hours  old,  and  declines  by  120  hours, 
the  flies  die  from  starvation  by  144  hours  (Langley,  1967b).  The  rise  in  proteinase  activity 
during  the  first  24  hours  is  not  caused  by  either  crawling  up  through  sand  or  flight  activity. 
Results  of  experiments  involving  puncturing  the  ptilinum,  and  injecting  material  into  the 
teneral  fly  suggest  that  distension  of  the  crop  (possibly  in  combination  with  the  presence  of 
protein  in  the  crop)  is  responsible  for  the  rise  in  proteinase  activity.  Experiments  involving 
ligaturing,  nerve  sectioning,  and  injection  of  tissue  homogenates,  demonstrated  that  the 
brain  is  involved  in  the  production  and/or  secretion  of  the  midgut  proteinase  in  the  unfed 
teneral  fly. 

During  the  first  24  hours  of  adult  life  in  Glossina  austeni  there  is  a 50%  increase  in  the 
midgut  proteinase  (Langley  and  Abasa,  1970).  This  increase  occurs  in  normal  males  and 
females  and  in  flies  irradiated  with  10  krad  as  pupae.  Twenty-four  hours  after  the  first  blood 
meal  there  is  a positive  correlation  between  meal  size  and  the  amount  of  midgut  proteinase 
activity.  The  slope  of  the  regression  line  is  not  significantly  affected  in  flies  given  10  krad  as 
pupae  or  1 5 krad  as  0 to  3 hr  old  adults.  However,  the  irradiated  flies  have  much  lower 
correlation  coefficients  (0.49)  than  the  unirradiated  controls  (0.74),  indicating  greater  varia- 
tion in  the  irradiated  flies. 

Schaefer  (1968),  largely  on  the  basis  of  work  by  Langley  (1967a)  proposed  that  the 
reactions  of  hosts  in  the  wild,  to  attack  by  blood-sucking  insects  cause  stress  in  the  latter. 
This  stress  heightens  the  activity  of  the  neurosecretory  processes  of  the  brain  and  corpora 
cardiaca  and  the  resulting  neurosecretions  stimulate  the  early  release  of  proteolytic  enzymes 
into  the  midgut. 


Haematophagous  insects 


43 


The  invertase  activity  of  A.  aegypti  midguts  increases  from  2 to  4 hours  after  the  mosqui- 
toes feed  on  blood  (Fisk  and  Shambaugh,  1954).  A depression  in  invertase  activity  in  both 
the  midgut  and  diverticula  after  a sugar  meal  persists  for  at  least  24  hours.  The  invertase 
activity  is  always  higher  in  the  midgut  than  in  the  diverticula  regardless  of  the  nature  of  the 
meal  or  the  time  after  feeding,  indicating  that  the  midgut  is  the  source  of  this  enzyme. 

There  is  no  significant  difference  in  the  invertase  content  of  sucrose-fed  and  water-fed  S. 
venustum  but  the  invertase  level  rises  immediately  after  a blood  meal  and  remains  elevated 
for  48  hours  (Davies  and  Yang,  1968;  Yang  and  Davies,  1968c). 

Some  of  the  amylase  activity  in  A.  aegypti,  Culex  pipiens,  S.  venustum  and  Simulium 
vittatum  occurs  in  the  midgut,  but  most  occurs  elsewhere  principally  in  the  hemolymph 
(Yang  and  Davies,  1968a).  With  A.  aegypti  there  is  a 3 to  4 fold  increase  in  amylase  activity 
immediately  after  feeding  on  man;  this  activity  declines  as  the  blood  is  digested.  By  com- 
paring the  amylase  activity  of  blood-fed  mosquitoes  and  human  blood  Yang  and  Davies 
concluded  that  most  of  the  amylase  activity  in  the  fed  mosquito  came  from  the  blood  meal. 

Freyvogel,  Hunter,  and  Smith  (1968)  suggested  that  esterases  demonstrated  in  the  midgut 
of  Anopheles  freeborni,  Anopheles  stephensi  and  A.  aegypti  may  have  a role  in  digestion  of 
the  blood  meal. 

Longevity  studies  indicate  that  A.  aegypti  can  utilize  the  disaccharides  sucrose,  maltose, 
trehalose,  and  melibiose,  the  trisaccharides  raffinose,  and  melizitose,  and  the  polysaccharide 
dextrin,  but  not  the  disaccharides  lactose  or  cellobiose,  the  polysaccharides  starch,  glycogen, 
or  inulin  or  the  glycosides  a-methylglucoside  or  a-methylmannoside  (Galun  and  Fraenkel, 
1957).  Homogenates  of  whole  A aegypti  hydrolyze  sucrose,  maltose,  trehalose,  raffinose, 
melezitose,  and  dextrin  but  not  melibiose,  lactose,  cellobiose,  starch,  glycogen,  inulin, 
a-methylglucoside  or  a-methylmannoside.  The  results  of  the  feeding  experiments  and  the 
enzyme  tests  are  consistent  except  in  the  case  of  melibiose,  for  which  no  hydrolytic  enzyme 
could  be  demonstrated. 

RELATIONSHIP  OF  DIGESTIVE  PROCESSES  TO  VECTORING  ABILITY. 

The  discovery  of  insect  transmission  of  vertebrate  pathogens  was  followed  within  a few 
years  by  the  demonstration  of  numerous  examples  of  vector-parasite  specificity.  An  early 
hypothesis  advanced  to  explain  this  phenomenon  was  that  the  digestive  processes  of  the 
insect  determine  which  parasites  develop.  This  hypothesis  has  been  investigated  several  times 
beginning  with  Nuttall  (1908)  and  has  remained  an  attractive  explanation.  Although  tested 
several  times  without  experimental  confirmation,  the  role  of  the  gut  in  vector-parasite 
specificity  is  still  occasionally  mentioned.  For  example  Day  and  Waterhouse  (1953a)  in  a 
review  article  stated  that  “The  physiology  of  the  mosquito  midgut  is  of  exceptional  impor- 
tance in  that  it  is  one  of  the  factors  controlling  the  establishment  of  malarial  parasites  within 
the  insect  vector”.  In  considering  the  differences  in  the  digestive  rates  of  Anopheles  sacharo- 
vi,  Anopheles  maculipennis,  and  Anopheles  superpictus,  Bates  (1949,  p.  90)  wrote  “It  has 
been  suggested  that  such  specific  differences  in  the  digestive  process  might  be  a factor  in 
determining  the  susceptibility  of  a mosquito  to  plasmodium  invasion.”  Wigglesworth  (1930) 
wrote  “there  is,  at  the  present  time,  a common  but  indefinite  impression  that  some  simple 
demonstrable  difference  in  the  chemistry  of  the  digestive  tract  (for  instance,  in  salt  content 
or  in  hydrogen-ion  concentration)  may  be  at  the  back  of  specificity  in  the  insect  host  as  a 
vector  of  pathogenic  micro-organisms.  Although  of  course  it  cannot  be  denied  that  this  may 
be  so,  I do  not  myself  see  any  a priori  reason  why  specificity  in  the  insect  host  should  be 
due  to  causes  any  less  subtle  than  say  natural  immunity  among  vertebrates.” 

The  influence  of  agglutinins  can  be  seen  by  comparing  the  distribution  of  Plasmodium 


44 


Gooding 


oocysts  in  the  midguts  of  A.  maculipennis  and  Anopheles  stephensi  (Shute,  1948).  Both 
species  have  anticoagulins  but  only  the  former  has  an  agglutinin  which  causes  the  red  blood 
cells  to  clump  and  settle  out.  As  a result  when  A.  maculipennis  is  in  its  normal,  head-up, 
vertical  position  after  a blood  meal,  the  erythrocytes  settle  to  the  posterior  part  of  the 
midgut  and  most  Plasmodium  oocysts  are  found  in  this  region.  However,  in  A.  stephensi  the 
erythrocytes  do  not  settle  out  and  the  oocysts  are  more  or  less  uniformly  distributed  over 
the  midgut. 

Lavoipierre  (1958)  reviewed  the  relationships  between  filarial  nematodes  and  their  arthro- 
pod vectors,  including  a brief  discussion  of  the  possible  role  of  digestive  physiology  of  the 
vector  in  limiting  the  intensity  of  infection. 

Chamberlain  and  Sudia  (1961),  in  a review  of  virus  transmission  by  mosquitoes,  listed 
several  hypotheses  to  explain  the  “gut  barrier”  to  infection  including  two  related  to  digestive 
physiology  (virus  inactivation  by  digestive  fluids  and  impermeability  of  the  PM).  They  em- 
phasized that  arguments  could  be  presented  to  support  or  refute  each  hypothesis  and  that 
no  mechanism  has  been  completely  proved.  For  example  the  suggestion  that  digestive  fluids 
inactivate  the  virus  can  be  argued  against  on  the  grounds  that  viable  WEE  virus  is  detectable 
in  mosquito  midgut  for  as  long  as  a day  after  the  infectious  meal.  On  the  other  hand,  since 
digestion  commences  at  the  periphery  of  the  blood  meal,  the  virus  particles  next  to  the  gut 
wall  may  be  inactivated  while  those  deeper  in  the  clot  may  be  unaffected.  Similarly,  the 
pore  sizes  of  the  peritrophic  membrane  may  explain  differences  in  susceptibility  to  viruses 
of  different  sizes  but  not  to  those  of  the  same  size. 

Young  Cimex  lectularius,  feeding  upon  mice  infected  with  Pasturella  pestis  (Lehmann  and 
Neumann)  fail  to  reduce  the  size  of  the  ingested  meal  within  a few  days  and  usually  die 
(Bacot,  1915).  Bacot  stressed  that  digestion  of  blood  is  very  rapid  in  the  midgut  of  the  flea 
and  that  with  the  destruction  of  both  the  red  blood  cells  and  the  leucocytes,  the  midgut 
becomes  very  much  like  an  artificial  culture  medium  in  which  P.  pestis  may  develop  rapidly. 
However,  in  the  crop  of  the  bug  P.  pestis  development  “differs  generally  from  that  which 
takes  place  in  the  stomach  of  the  flea  in  respect  of  its  slower  and  looser  growth,  this  limita- 
tion of  activity  being  accompanied  by  and  possibly  due  to  the  preservation  of  the  structural 
character  of  the  blood  for  many  days  after  its  ingestion  into  the  crop”. 

Duncan  (1926)  ran  tests  for  bactericidal  activity  in  the  gut  contents  and  feces  of  several 
blood-sucking  insects  ( Stomoxys  calcitrans.  Anopheles  bifurcatus  (L.),  Aedes  cinereus,  C. 
lectularius  Rhodnius  prolixus ) and  on  blood  fed  Musca  domestica  L.  Activity  was  found 
against  8 of  the  18  species  of  bacteria  used.  The  bactericidal  material  from  S.  calcitrans  was 
heat  stable  (100  C for  30  minutes)  and  was  not  destroyed  by  trypsin.  St.  John,  Simmons, 
and  Reynolds  (1930)  found  no  evidence  of  bactericidal  material  in  the  digestive  tract  of 
Aedes  aegypti. 

Packchanian  (1948a)  found  that  Leishmania  tropica  (Wright)  and  Leishmania  donovani 
fed  to  several  species  of  Triatoma  (T.  gerstaeckeri,  Triatoma  lectularia  (Stal)  (=T.  heideman- 
ni ),  T.  protracta  (Uhler),  and  T.  uhleri  Neiva)  died  in  the  gut  within  3 days.  Similarly  he 
showed  that  Trypanosoma  brucei  Plimmer  and  Bradford,  Trypanosoma  gambiense  Dutton, 
and  Trypanosoma  evansi  (Steel)  (reported  as  T.  hippicum)  died  within  10  days  of  ingestion 
by  Triatoma  spp.  (T.  gerstaeckeri,  T.  sanguisuga,  T.  protracta  and  T.  rubrofasciata ) (Packcha- 
nian, 1948b). 

The  spirochaete  Treponema  pertenue  Castellani  remains  mobile  much  longer  in  the  oe- 
sophageal diverticulum  than  in  the  stomach  of  Hippelates  pallipes  indicating  that  this  spiro- 
chaete may  be  affected  by  the  fly’s  digestive  secretions  (Kumm,  Turner,  and  Peat,  1935). 

A possible  influence  of  defecation,  during  the  act  of  feeding,  upon  intensity  of  infection 
was  shown  by  Kartman  (1953a)  who  fed  five  species  of  mosquitoes  (A.  aegypti,  Aedes 


Haematophagous  insects 


45 


albopictus,  Culex  pipiens  quinquefasciatus,  Culex  pipiens  and  Anopheles  quadrimaculatus ) 
on  a dog  infected  with  Dirofilaria  immitis  (Leidy).  As  the  mosquitoes  became  replete,  the 
only  species  observed  to  defecate  a drop  of  fluid  was  A.  quadrimaculatus.  By  counting  the 
microfilaria  in  this  drop  and  in  the  midguts  of  the  fed  mosquitoes,  it  was  estimated  that 
about  7%  of  the  microfilaria  were  lost  from  the  midgut  by  defecation. 

Kartman  (1953b)  showed  that  the  clotting  of  the  blood  in  the  mosquito  midgut  (in  for 
example  Aedes  and  Culex ) reduced  the  number  of  microfilaria  of  D.  immitis  which  could 
leave  the  midgut.  Degeneration  of  microfilaria  in  the  midgut  was  observed,  and  although  it 
was  concluded  that  this  destruction  was  due  to  the  digestive  process,  it  was  pointed  out  that 
death  may  not  have  been  caused  by  the  digestive  enzymes  but  rather  by  other  factors  in  the 
midgut  or  salivary  secretion. 

Huff  (1927)  fed  C.  pipiens  and  Aedes  sollicitans  upon  canaries  infected  with  Plasmodium 
cathemerium  and,  at  various  times  after  the  meal,  made  smears  of  the  gut  contents  and  ob- 
served the  appearance  of  the  erythrocytes  and  the  sexual  and  asexual  forms  of  the  parasites. 
In  C.  pipiens,  a susceptible  species,  the  asexual  forms  began  to  stain  abnormally  after  3 
hours,  and  after  6 hours  were  not  found.  In  A.  sollicitans  the  asexual  forms  had  not  disap- 
peared after  6 hours  and  some  were  found  until  the  end  of  the  series  20  hours  after  feeding. 
The  fate  of  the  sexual  forms  was  the  same  as  that  of  the  asexual  forms.  Huff  also  injected 
homogenates  of  the  midguts  of  both  species  into  normal  birds  at  intervals  after  the  mosqui- 
toes had  fed  on  an  infected  bird.  The  resulting  infections  in  the  birds  showed  that  the 
asexual  forms  lose  their  infectivity  after  5 or  6 hours  in  the  gut  of  both  the  susceptible  and 
the  refractory  species.  During  this  study  Huff  noted  that  some  mosquitoes  of  both  species 
differed  greatly  from  the  others  in  the  rate  of  digestion  of  blood.  Huff  (1934)  attempted  to 
determine  whether  intraspecific  variation  in  the  rate  of  digestion  was  correlated  with  the 
degree  of  susceptibility  to  malaria  parasites  in  C.  pipiens.  The  mosquitoes  were  fed  upon 
canaries  infected  with  either  P.  cathemerium  or  Plasmodium  relictum.  A second  meal  on  an 
infective  bird  was  given  5 to  8 days  later,  the  mosquitoes  then  being  dissected  and  their  mid- 
guts removed  and  examined  microscopically.  With  this  technique,  each  mosquito  served  as 
its  own  control  - those  which  had  large  oocysts  were  susceptible,  and  the  others  were  con- 
sidered refractory.  No  differences  were  observed  in  digestion  in  the  susceptible  and  refrac- 
tory individuals. 

However,  de  Buck,  Schoute,  and  Swellengrebel  (1930,  1932),  and  deBuck,  Torren,  and 
Swellengrebel  (1933),  suggested  that  refractoriness  in  A.  maculipennis  was  correlated  with 
slow  digestion  of  the  blood  meal  in  certain  varieties  during  overwintering.  Ookinetes  were 
formed  in  both  the  undigested  meal  of  the  “long  winged”  and  in  the  partially  digested  meal 
of  the  “short  winged”  mosquitoes  and  it  was  proposed  that  the  difference  in  susceptibility 
may  be  due  to  differences  in  the  ease  with  which  ookinetes  can  work  their  way  out  of  the 
blood  meal. 

Bishop  and  McConnachie  (1956)  found  no  evidence  that  exflagellation  of  Plasmodium 
gallinaceum  took  place  any  faster  in  the  stomach  of  A.  aegypti,  a susceptible  mosquito,  than 
under  a coverslip  on  a glass  slide. 

Attempts  have  been  made  to  correlate  the  development  of  oocysts  with  the  diet  of  the 
mosquito.  The  first  was  the  observation  that  Anopheles,  fed  alternately  upon  bananas  and 
gametocyte  carriers,  often  failed  to  digest  their  blood  meal  or  to  develop  oocysts  (Darling, 
1910).  Feeding  several  salts,  including  0.1  M solutions  of  CaCl2  or  MgCl2  to  A.  aegypti  de- 
creased the  number  of  P.  gallinaceum  oocysts  developing  on  the  gut  wall  (Terzian  and 
Stahler,  1960).  However,  the  same  study  showed  that  MgCl2  fed  at  a concentration  of  0.4  M 
increased  susceptibility  to  P.  gallinaceum.  The  presence  of  CaCl2  and  MgCl2  in  the  diet  of  A. 
aegypti  and  A.  quadrimaculatus  inhibited  the  in  vivo  digestion  of  blood  (Terzian,  1958, 


46 


Gooding 


1963),  and  these  same  salts  had  an  inhibitory  effect  upon  the  in  vitro  activity  of  A aegypti 
proteinase  (Wagner,  Tenorio  and  Terzian,  1961;  Gooding,  1966a).  On  the  other  hand, 
feeding  a chloramphenicol-dihydrostreptomycin-sugar  solution  to  C.  pipiens  quinquefascia- 
tus  inhibited  digestion  of  blood  by  this  species,  but  increased  its  susceptibility  to  P.  relictum 
(Micks  and  Ferguson,  1961).  There  was  a slight  increase  in  the  per  cent  of  the  mosquitoes 
infected  (74%  in  controls,  85%,  in  treated  mosquitoes)  and  a considerable  increase  in  the 
intensity  of  the  infection  (56  oocysts/midgut  in  controls  and  110  oocysts/midgut  in  the 
antibiotic  treated  mosquitoes). 

Attempts  have  been  made  to  influence  the  level  of  infection  by  decapitating  the  mosqui- 
toes after  a blood  meal.  Rozeboom  (1961)  found  a decrease  in  the  percentage  of  mosquitoes 
infected,  in  the  number  of  oocysts  per  midgut,  and  in  the  size  of  the  oocysts  (P.  gallinaceum ) 
when  A.  aegypti  were  decapitated  within  6 hours  of  feeding.  These  results  may  indicate  a 
lower  nutritive  environment  for  the  P.  gallinaceum  oocyst  in  the  decapitated  mosquitoes, 
since  decapitation  was  subsequently  shown  to  influence  the  amount  of  proteinase  in  the 
midgut  of  A.  aegypti  (Gooding,  1966b).  However,  Rozeboom  (1961)  stated  that  “blood 
digestion  does  proceed  in  the  decapitated  females  which  survive  3 to  5 days.  In  many  spec- 
imens a residue  of  the  blood  meal  may  remain  in  the  gut;  in  others  the  gut  becomes  com- 
pletely empty  as  in  normal  mosquitoes.  Thus  a sufficient  degree  of  normal  digestive  pro- 
cesses, including  changes  in  the  epithelium,  continued  to  take  place  in  decapitated  mosqui- 
toes to  permit  a somewhat  reduced  number  of  zygotes  to  be  taken  up  by  the  epithelium”. 
Yoeli,  Upmanis,  and  Most  (1962)  found  that  decapitation  of  A.  quadrimaculatus  after  a 
blood  meal  did  not  interfere  with  the  normal  development  of  the  larvae  of  D.  immitis. 

Stohler  (1961)  considered  in  some  detail  the  relationship  between  the  PM  of  Diptera  and 
the  role  of  the  latter  as  vectors  of  blood  parasites.  He  concluded  that  the  PM  could  influence 
the  intensity  of  an  infection  by  imprisoning  a portion  of  the  parasites  within  the  blood  meal 
but  that  the  parasites  can  usually  escape  from  the  meal  through  the  viscous  portion  of  the 
PM,  through  its  open,  posterior  end  or  at  the  time  the  PM  breaks  up  if  the  parasites  have  not 
already  been  killed.  He  stated,  however,  that  the  full  role  of  the  PM  in  influencing  vectoring 
ability  of  flies  was  not  completely  resolved,  and  that  a study  of  closely  related  species  which 
differed  widely  in  their  vectoring  abilities  could  be  very  useful. 

Bates  (1949,  p.  229)  considered  the  critical  stage  in  determining  the  susceptibility  of  the 
mosquito  to  be  the  penetration  of  the  gut  wall  by  the  ookinete.  Mariani  (1961)  suggested 
that  the  PM  in  Anopheles  labranchiae  Falleroni  may  hinder  the  passage  of  malaria  zygotes 
from  the  blood  meal  into  the  mosquito.  He  discussed  the  importance  of  the  PM  in  vectoring 
ability  of  mosquitoes. 

Ookinetes  of  P.  gallinaceum  that  fail  to  penetrate  the  PM  between  20  and  30  hours  after 
A.  aegypti  feeds  will  perish.  After  considering  the  frequency  with  which  ookinetes  are  found 
near  the  PM,  Stohler  (1957)  suggested  that  the  PM  constitutes  a physical  barrier  to  the  pen- 
etration of  the  ookinetes  and  that  penetration  of  the  gut  epithelium  does  not  constitute  a 
significant  barrier  to  infection.  During  the  early  stages  of  its  formation,  the  PM  does  not 
constitute  a barrier  to  ookinete  penetration,  but  with  subsequent  hardening  it  becomes  pro- 
gressively more  impenetrable. 

Interpretation  of  the  role  of  digestion  in  vector-plasmodia  specificity  is  complicated  by 
our  uncertainty  about  the  course  of  development  of  the  malaria  parasite  within  the  gut  of 
the  mosquito  and  by  the  paucity  of  information  on  the  stage  at  which  development  of  the 
parasite  stops.  Most  of  the  evidence  suggests  that  decreased  susceptibility  is  correlated  with 
a decreased  rate  of  digestion.  It  thus  appears  that  if  the  digestive  processes  of  the  mosquito 
play  a role  in  the  infection  of  the  mosquito  by  plasmodia,  it  would  be  by  providing  nutrients 
to  the  developing  parasite.  The  possibility  that  the  digestive  enzymes  of  a refractory  species 


Haematophagous  insects 


47 


act  directly  on  the  parasite  has,  however,  not  been  adequately  investigated  and  thus  cannot 
be  eliminated  at  this  time. 

The  influence  of  the  PM  upon  the  establishment  of  Trypanosoma  grayi  (Novy)  in  Glossina 
palpalis  has  been  studied  by  Hoare  (1931).  T.  grayi  are  confined  to  the  midgut  lumen  by  the 
PM  for  2 or  3 days  after  an  infective  meal.  They  then  migrate  back  to  the  hind  gut  and 
escape  from  the  PM  taking  up  residence  between  the  PM  and  the  gut  wall  by  6 to  8 days  after 
the  infective  meal  and  eventually  migrating  foreward  in  this  space.  The  trypanosomes  con- 
tinue to  occupy  this  space  for  the  remainder  of  their  residence  in  the  tsetse  fly.  Lewis  (1950) 
observed  that  the  blackfly  PM  prevented  many  microfilaria  from  entering  the  body  cavity 
and  he  concluded  that  “Frequently  therefore,  the  membrane  protects  the  fly  itself  from 
heavy  infection  without  preventing  it  from  transmitting  the  parasite”. 

CONCLUDING  REMARKS 

The  blood-sucking  insects  are  parasites  ingesting  a meal  which  is  well  defined  both  in 
respect  of  its  composition  and  the  time  it  is  consumed.  As  such  these  insects  should  be  ideal 
for  studies  of  digestive  physiology  and  nutrition  as  they  relate  to  parasitism  in  general  and 
host-parasite  relationships  in  particular. 

The  size  of  a blood-sucking  insect  ultimately  determines  the  size  of  the  blood  meal  it  can 
ingest.  In  general  the  blood  meal  is  rather  large  compared  with  the  size  of  the  insect,  and  all 
the  greater  when  one  considers  it  in  relation  to  the  amount  of  tissue  available  for  synthesis 
of  the  digestive  enzymes  and  absorption  of  the  products  of  digestion.  It  then  appears  that 
blood-sucking  insects  can  digest  relatively  larger  quantities  of  blood  at  a time.  This  however 
is  more  apparent  than  real  for  most  of  them  have  some  mechanism  which  limits  the  amount 
of  blood  being  digested  at  any  time.  In  some  there  are  no  anticoagulins  and  the  blood  clots 
in  the  midgut,  while  many  of  those  which  do  have  anticoagulins  also  have  agglutinins.  The 
meal  is  further  concentrated  by  removal  of  water  during  or  just  after  feeding.  Enzymes  are 
then  secreted  onto  the  surface  of  the  meal.  In  bugs  and  many  of  the  higher  flies  most  of  the 
meal  is  stored  (and  in  some  species  concentrated)  in  the  anterior  part  of  the  midgut  without 
any  digestion  taking  place  and  then  passes  to  the  posterior  part  of  the  midgut  in  small  quan- 
tities for  digestion.  The  net  result  of  both  of  these  methods  is  that  only  a small  portion  of 
the  meal  is  digested  at  any  one  time  and  that  the  digestive  enzymes  are  never  mixed  with 
the  total,  freshly  ingested,  meal. 

Digestion  of  only  a small  fraction  of  the  meal  at  a time  has  several  advantages  to  the 
insect.  One  advantage  is  that  if  the  enzymes  were  mixed  thoroughly  with  the  entire  meal 
there  may  be  such  an  excess  of  substrate  that  substrate  inhibition  would  significantly  reduce 
the  rate  of  digestion.  The  proteinase  inhibitors  in  serum  may  also  inhibit  most  of  the  diges- 
tive proteinases  thus  reducing  the  rate  of  digestion  or  necessitating  secretion  of  increased 
amounts  of  enzyme.  Thus  another  advantage  of  having  only  a small  portion  of  the  meal 
exposed  to  the  digestive  enzymes  is  that  the  serum  proteinase  inhibitors  may  be  titrated  out 
by  the  digestive  enzymes  or  destroyed  by  a concerted  attack  by  the  midgut  proteinases. 
A third  advantage  to  digesting  small  quantities  of  blood  close  to  the  midgut  epithelium  is 
that  the  products  of  digestion  are  readily  available  for  absorption  rather  than  having  to 
move  from  the  center  of  the  midgut. 

Salivary  gland  anticoagulins  and  agglutinins  are  widespread,  but  not  universal,  charac- 
teristics of  blood-sucking  insects.  Whether  these  substances  indicate  a degree  of  conver- 
gence selected  for  by  the  nature  of  the  blood  meal  or  the  retention  of  primitive  char- 
acters is  unknown.  It  would  indeed  be  interesting  to  examine  the  saliva  of  several  non- 
haematophagous  insects  for  anticoagulins  and  agglutinins.  The  specific  contribution  of  sali- 


48 


Gooding 


vary  agglutinins  and  anticoagulins  to  the  denaturation  and  digestion  of  the  blood  meal  has 
not  been  investigated.  There  are  however  reports  that  mosquitoes  and  tsetse  flies  can  digest 
blood  without  saliva.  Whether  the  efficiency  of  the  process  is  unaltered  in  surgically  modi- 
fied insects  is  not  definitely  established.  One  might  expect  some  effect  on  digestion  in 
Glossina  austeni  whose  salivary  ducts  have  been  cut  since  a plasminogen  activator  from  the 
salivary  glands  probably  contributes  to  clot  lysis  in  the  midgut. 

The  role  of  digestive  physiology  in  host  parasite  relations  has  not  been  systematically 
examined.  The  work  on  digestion  by  Pediculus  humanus  indicates  that  this  highly  host 
specific  ectoparasite  encounters  difficulties  in  digesting  guinea  pig  blood.  To  what  extent 
these  difficulties  are  peculiar  to  the  P.  humanus  — guinea  pig  system  is  unknown.  Insuffi- 
cient work  has  been  done  on  comparison  of  digestive  physiology  of  insects  varying  in  the 
degree  of  host  specificity.  Vertebrate  serum  contains  some  proteins  necessary  for  the  secre- 
tion or  activation  of  digestive  proteinases  and  others  which  inhibit  proteinases.  The  relative 
concentrations  or  activities  of  these  two  kinds  of  proteins  in  various  vertebrate  sera  are 
unknown,  as  are  the  responses  of  various  insects  to  these.  It  is  conceivable  that  certain 
highly  host  specific  blood-sucking  insects  could  have  very  precise  requirements  with  respect 
to  both  the  proteinase  stimulators  and  the  inhibitors.  The  fate  of  these  stimulators  and 
inhibitors  in  the  digestive  tracts  of  either  specific  or  non-specific  blood-sucking  insects  is 
unknown  but  worthy  of  investigation. 

On  the  basis  of  work  with  synthetic  substrates  as  well  as  specific  inhibitors  it  appears 
that  most  of  the  proteinase  activity  in  the  mosquito  midgut  is  due  to  a trypsin,  with  much 
smaller  amounts  of  chymotrypsin  also  being  present.  However,  immunological  studies  indi- 
cate that  digestion  products  from  at  least  some  blood  proteins  in  the  midgut  of  Culex 
pipiens  quinquefasciatus  are  different  from  those  produced  by  mammalian  trypsin.  These 
findings  indicate  that  either  the  small  amount  of  chymotrypsin  may  have  a marked  qualita- 
tive influence  upon  digestion  in  the  mosquito  or  that  the  mosquito  trypsin  has  a different 
bond  specificity  than  mammalian  trypsin  when  whole  proteins  are  used  as  the  substrate. 

In  this  article  I have  summarized  a substantial  portion  of  the  literature  on  digestion  in 
blood-sucking  insects  and  indicated  some  areas  in  which  further  research  would  be  profit- 
able. In  subsequent  articles  in  this  series  I propose  to  report  on  digestion  in  a variety  of 
blood-sucking  insects  and  on  contributions  to  the  solution  of  some  of  the  problems  indi- 
cated in  this  article. 


ACKNOWLEDGEMENTS 


I am  happy  to  acknowledge  the  assistance  of  J.  Belicek  and  J.  Rickert  in  translation 
of  Russian  and  German  papers,  and  to  thank  B.  S.  Heming,  Department  of  Entomology, 
University  of  Alberta,  for  his  many  helpful  comments  on  the  manuscript.  The  research  for 
this  paper  was  supported  (in  part)  by  the  Defence  Research  Board  of  Canada,  Grant  Number 
6801-41. 


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biological  notes.  Int.  J.  publ.  Hlth  1 :3 16-346. 

Service,  M.W.  1968a.  Observations  on  feeding  and  oviposition  in  some  British  mosquitoes. 
Entomologia  exp.  appl.  11:277-285. 

Service,  M.W.  1968b.  Some  environmental  effects  on  blood-fed  hibernating  Culiseta  annulata 
(Diptera:  Culicidae).  Entomologia  exp.  appl.  1 1:286-290. 

Service,  M.W.  1968c.  Blood  digestion  and  oviposition  in  Culicoides  impunctatus  Goetghebuer 
and  C.  obsoletus  (Meigen)  (Diptera,  Ceratopogonidae).  Ann.  trop.  Med.  Parasit.  62:325- 
330. 

Shambaugh,  G.F.  1954.  Protease  stimulation  by  foods  in  adult  Aedes  aegypti  Linn.  Ohio  J. 
Sci.  54:151-160. 

Shute,  P.G.  1935.  Agglutination  of  the  red  blood  corpuscles  of  man,  animals  and  birds  by 
the  salivary  glands  of  Anopheles  maculipennis.  J.  trop.  Med.  Hyg.  38:277-278. 

Shute,  P.G.  1948.  The  comparative  distribution  of  oocysts  of  human  malaria  parasites  on 
the  stomach  wall  of  Anopheles  mosquitoes.  Trans.  R.  Soc.  trop.  Med.  Hyg.  42:324. 

Snodgrass,  R.E.  1935.  Principles  of  Insect  Morphology.  McGraw-Hill  Book  Co.  Inc.  N.Y., 
London,  ix  + 667  pp. 

Stage,  H.H.  and  W.W.  Yates.  1936.  Some  observations  on  the  amount  of  blood  engorged  by 
mosquitoes.  J.  Parasit.  22:298-300. 

Staubli,  W.,  T.A.  Freyvogel  and  J.  Suter.  1966.  Structural  modification  of  the  endoplasmic 
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Stohler,  H.  1957.  Analyse  des  Infektionsverlaufes  von  Plasmodium  gallinaceum  im  Darme 
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Stohler,  H.R.  1961.  The  peritrophic  membrane  of  blood  sucking  Diptera  in  relation  to  their 
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Suenaga,  O.  1965.  A rearing  method  of  stable  fly  and  quantity  of  blood  taken  up  by  a fly. 
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Sumner,  J.B.  and  G.F.  Somers.  1947.  Chemistry  and  methods  of  enzymes.  (Second  Edition). 
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Tashiro,  H.  and  H.H.  Schwardt.  1949.  Biology  of  the  major  species  of  Horse  Flies  of  central 
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Tatchell,  R.J.  1958.  The  physiology  of  digestion  in  the  larvae  of  the  horse  bot-fly,  Gastero- 
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Templis,  C.F.  and  M.G.  Lofy.  1963.  A modified  precipitin  method  for  identification  of  mos- 
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Terzian,  L.A.  1963.  The  effects  of  certain  cations  and  antibiotics  on  blood  digestion  in  mos- 
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Terzian,  L.A.  and  N.  Stahler.  1960.  Some  inorganic  acids,  bases  and  salts  as  determinants  of 
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60 


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RAPID  ORIENTATION  OF  WAX  EMBEDDED  SPECIMENS 


D A.  CRAIG 

Department  of  Entomology 
University  of  Alberta 
Edmonton  7,  Alberta 


Quaestiones  entomologicae 
8 : 61-62  1972 


A rapid  method  for  orientation  of  wax  embedded  specimens  in  precast  wax  blocks  with 
an  electrical  heat  probe  is  given. 

Pour  V orientation  de  specimens  imbibes  au  prealable  de  cire  dans  des  blocs  de  cire  moules 
d’avance,  une  methode  simple  est  decrite  ne  necessitant  qu’un  simple  fil  metallique  chauffe 
electriquement. 

A procedure  for  the  rapid,  accurate  embedding  and  orientation  of  simuliid  (Diptera) 
embryos  and  larval  heads  in  wax  has  been  developed  which  should  prove  useful  for  other 
small  animals. 

The  specimens  are  impregnated  with  wax  in  a small  container  on  a hot  plate  adjacent  to 
a stereoscopic  microscope.  Then  a previously  cast  squared  wax  block  is  positioned  in  the 
optical  axis  of  the  microscope.  Next,  each  specimen  is  lifted  from  the  molten  wax  with  a 
fine  needle.  On  removal  the  wax  solidifies  rapidly  around  the  needle  and  specimen.  Then  the 
specimen  is  carried  to  the  top  of  the  wax  block  and  held  there  while  a small  pool  of  molten 
wax  is  formed  in  the  top  of  the  block  with  an  electrically  heated  probe.  Then  the  probe  is 
touched  to  the  needle  above  the  specimen.  This  melts  the  wax  and  the  specimen  slides  off 
into  the  pool. 

The  probe  consists  of  part  of  a straightened  wire  paper-clip  with  one  end  inserted  in  a 
glass  rod.  Three  inches  of  oxide  coated,  0.008  inches  diameter  resistance  wire  is  twisted 
around  the  clip  near  the  tip  and  the  ends  of  the  wire  are  connected  to  the  secondary  winding 
of  a variable  transformer  for  a microscope  lamp  which  allows  the  temperature  of  the  probe 
to  be  controlled. 

The  specimen  is  oriented  with  the  needle,  while  the  probe  prevents  the  pool  of  wax  from 
solidifying  and  determines  the  depth  of  the  specimen  in  it  (Fig.  1).  During  final  cooling  the 
wax  solidifies  from  the  bottom  and  holds  the  specimen  steady.  If  the  specimen  has  been 
maintained  in  the  microscope’s  optical  axis,  the  orientation  of  the  specimen  is  known  in 
relation  to  the  sides  of  the  wax  block  and  any  required  orientation  can  be  repeated.  The 
block  is  sectioned  in  the  normal  manner  with  particular  care  given  to  its  orientation  to  the 
microtome  knife. 

With  this  technique  it  is  possible  to  get  perfect  transverse,  sagittal,  and  frontal  sections. 
Fig.  2 shows  a sagittal  section  of  the  recurrent  nerve  (r.n)  in  Cnephia  dacotensis  and  Fig.  3, 
the  stomodaeum  (st.)  of  Gymnopais  sp. 


I wish  to  thank  B.S.  Heming  for  constructive  criticism  and  J.S.  Scott  for  photographic 
assistance.  The  work  was  supported  by  the  National  Research  Council  of  Canada. 


ACKNOWLEDGEMENTS 


62 


Craig 


10  mm 

i 1 


Figure  1.  Larval  head  of  Cnephia  dacotensis  during  orientation,  pr  = probe,  nd  = needle. 


Figure  2.  Sagittal  section  of  larval  head  of  Cnephia  dacotensis  showing  recurrent  nerve  (m).  Figure  3.  Sagittal  section  of 
embryo  of  Gymnopais  sp.  showing  stomodaeum  (st.). 


h* 


63 


Book  Review 

HODGES,  R.W.  1971.  Sphingoidea.  Fascicle  21.  In  The  moths  of  America  north  of  Mexico, 
including  Greenland.  FERGUSON,  R.B.  FRANCLEMONT,  J.G.  HODGES,  R.W.  MUNROE, 
E.G.  DOMINICK,  R.B.  EDWARDS,  C.R.  Editors.  E.W.  Classey  Ltd.  & R.B.D.  Publications 
Inc.,  London,  xii  + 158  pp.,  4 black  & white,  14  color  plates;  2 pages  of  line  drawings  at  the 
end,  19  groups  of  line  drawings  in  the  text.  Size  8-7/8"  x 1 lW,  wrap  cover,  103  references. 
Price:  £-10.00;  $24.00. 

Printed  in  England,  “The  Sphingoidea”  section  of  this  encyclopaedic  work  on  North 
American  moths  is  the  first  of  41  planned.  The  work  is  scheduled  for  completion  in  about 
twelve  years  and  the  “Announcement”  of  publication  states:  “it  is  intended  that  a similar 
work  on  skippers  and  butterflies  will  follow.” 

“The  moths  of  America  north  of  Mexico,  including  Greenland”  will  be  the  first  compre- 
hensive treatise  on  more  than  10,000  moth  species  known  from  that  region.  It  is  superbly 
illustrated  and  the  14  color  plates  contained  in  fascicle  21  deserve  special  mention.  The 
plates  were  reproduced  from  4"  x 5"  transparencies  taken  by  R.B.  Dominick  and  C.R. 
Edwards  and  printed  by  offset  lithography  in  four  colors.  Credit  must  be  given  to  them  as 
well  as  to  The  Curwen  Press  of  London. 

Dr.  R.W.  Hodges,  of  the  U.S.D.A.,  Systematic  Entomology  Laboratory  in  Washington, 
D.C.,  the  author  of  “Sphingoidea”,  earned  his  doctorate  at  Cornell  University  in  insect 
taxonomy.  “Sphingoidea”  is  a synthesis  of  past  revisionary  studies  of  the  group.  One  species 
is  described  as  new  and  one  new  genus  is  proposed,  along  with  13  new  combinations. 

The  work  is  intended  for  use  by  both  the  professional  and  the  amateur  entomologist. 
Two  pages  of  line  drawings  at  the  end  together  with  fig.  1 , provide  a good  introduction  to 
structural  characteristics.  The  text  figures  by  Dr.  Hodges’  wife  Elaine  R.  Hodges  are  fully 
labelled  and  self  explanatory;  scales  apparently  vary  but  are  not  indicated.  It  is  unfortunate 
that  genital  armatures  are  not  pictured  for  all  the  species  described. 

The  book  begins  with  an  introductory  note  followed  by  the  introduction  to  and  supra- 
specific  classification  of  North  American  Sphingidae.  It  contains  a key  to  genera  based  on 
adults,  a partial  key  to  genera  based  on  pupae  (after  Mosher,  1918),  and  a partial  key  to 
genera  based  on  mature  larvae  (after  Forbes,  1911).  For  each  genus,  a complete  citation  of 
its  original  description,  type  species  designation,  synonymy,  generic  description  and  key  to 
its  species  are  given.  For  each  species,  a complete  citation  of  its  original  description,  syno- 
nymy, type  locality,  and  where  applicable  an  official  common  name  are  given.  Species  are 
briefly  and  unevenly  discussed.  A review  of  important  literature  on  the  group  concludes  the 
text.  The  color  plates  portray  life  size  199  specimens  of  all  species  described  and  the  major 
polymorphs.  Each  plate  is  faced  by  legend  and  followed  by  explanatory  notes.  The  book  is 
concluded  by  indices  to  animal  names  and  to  plant  names. 

According  to  the  “Announcement”,  the  completed  work  will  include  an  introduction,  to 
be  published  last.  This  part  is  intended  to  include  sections  on  morphology,  phylogeny, 
ecology,  faunal  history,  distribution,  variation,  migration,  and  dispersal.  These  are  aspects 
either  not  covered  in  “Sphingoidea”  or  discussed  only  briefly.  I fear  that  when  published, 
years  from  now,  they  may  not  be  adequate  for  the  whole  work.  For  example,  species 
zoogeography  is  briefly  discussed  here  and  there  in  “Sphingoidea”  but  no  distribution  maps 
are  given.  For  zoogeographic  or  dispersal  studies,  all  specimens  must  be  examined  again.  The 
omission  of  illustrations  of  immature  stages  is  another  of  the  few  weaknesses.  Minor  errors 
include,  on  p.  47  penultimate  line:  “ Isoparce  is  a monotypic  genus.”  A genus  proposed  for 
a single  species  is  better  described  as  monobasic  since  all  genera  have,  by  rule,  only  one  type 
species.  A little  more  serious  is  incorrect  binominal  nomenclature,  as  on  p.  149,  “ Pluto  has 


l 


64 


been  taken  in  Southern  Texas....”.  I believe  the  author  meant:  “Representatives  of  Xylopha- 
nes  pluto  have  been  taken  in  Southern  Texas....”. 

Further  sections  of  “The  moths  of  America  north  of  Mexico,  including  Greenland”  are 
impatiently  awaited.  They  will  become  indispensable  for  any  lepidopterist  interested  in  the 
North  American  fauna.  But  it  is  to  be  hoped  that  the  text  in  succeeding  volumes  might  be 
fuller.  The  cutting  of  corners  in  such  an  important  part  of  such  an  important  work  cannot 
be  justified. 


Josef  Belicek 
Department  of  Entomology 
University  of  Alberta 
Edmonton  7,  Alberta 


ANNOUNCEMENT 

In  connection  with  the  50th  Anniversary  of  the  Department  of  Entomology  at  the 
University  of  Alberta  two  symposia  are  being  organised  for  the  third  week  in  May  1972. 
Readers  who  wish  to  be  kept  advised  of  developments  should  write  to  the  person  named. 

1)  Biting  fly  control  and  environmental  quality  16-18  May:  Dr.  Susan  B.  Mclver, 
Department  of  Parasitology,  School  of  Hygiene,  University  of  Toronto,  Toronto  181, 
Ontario,  Canada. 

2)  Entomology  in  education  and  education  in  entomology  19  May:  Dr.  Brian  Hock- 
ing, Department  of  Entomology,  University  of  Alberta,  Edmonton  7,  Alberta. 


Publication  of  Quaestiones  Entomologicae  was  started  in  1965  as  part 
of  a memorial  project  for  Professor  E.  H.  Strickland,  the  founder  of  the 
Department  of  Entomology  at  the  University  of  Alberta  in  Edmonton 
in  1922. 

It  is  intended  to  provide  prompt  low-cost  publication  for  accounts  of 
entomological  research  of  greater  than  average  length,  with  priority 
given  to  work  in  Professor  Strickland’s  special  fields  of  interest  including 
entomology  in  Alberta,  systematic  work,  and  other  papers  based  on  work 
done  at  the  University  of  Alberta. 

Copy  should  conform  to  the  Style  Manual  for  Biological  Journals 
published  by  the  American  Institute  of  Biological  Sciences,  Second 
Edition,  1964,  except  as  regards  the  abbreviations  of  titles  of  periodicals 
which  should  be  those  given  in  the  World  List  of  Scientific  Periodicals, 
1964  Edition.  The  appropriate  abbreviation  for  this  journal  is  Quaest.  ent. 
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Quaestiones 


entomologicae 


MU3.  COMP.  ZOOl — 
LIBRARY 

my  1 2 1972 

HARVARD 

UNIVERSITY 


A periodical  record  of  entomological  investigations. 


published  at  the  Department  of  Entomology, 
University  of  Alberta,  Edmonton,  Canada. 


VOLUME  VIII 


NUMBER  2 


APRIL  1972 


QUAESTIONES  ENTOMOLOGICAE 


A periodical  record  of  entomological  investigations  published  at  the  Department  of 
Entomology,  University  of  Alberta,  Edmonton,  Alberta. 

Volume  8 Number  2 1 April  1972 


CONTENTS 


Editorial  - Dissection  of  Science  65 

Griffiths  — Studies  on  boreal  Agromyzidae  (Diptera).  I. 

Phy  to  my  za  miners  on  Saxifragaceae 67 

Schaaf  - The  parasitoid  complex  of  Euxoa  ochrogaster  (Guenee) 

(Lepidoptera:  Noctuidae) 81 

Kevan  - Collembola  on  flowers  on  Banks  Island,  N.W.T 121 

Book  review  123 

Book  review  124 

Book  review  125 

Announcement 127 


Editorial  — Dissection  of  Science 

Politicians  — in  the  sense  of  those  who  would  make  policy  — seem  to  have  it  in  for  science. 
Commissions,  councils,  and  advisory  bodies,  in  Canada  as  in  many  other  countries,  have 
come  into  being  in  the  last  few  years,  with  instructions  to  take  a pragmatic  look  at  scientists 
and  their  activities.  Such  groups  have  been  busy  dissecting  “science”  in  every  conceivable 
way  and  attempting  to  fit  the  pieces  into  categories  of  likenesses.  But  no  matter  how  they 
dissect  it  they  seem  to  wind  up  with  a number  of  bits  such  as  technology,  and  industrial 
research  -which  are  hardly  science  at  all-  and  to  be  left  with  an  amorphous  mass  of  true 
science  which  they  can  divide  no  further,  a sort  of  Lucretius’  atom  of  knowledge.  This 
should  not  surprise  us,  for  science  —true  science—  is  but  weakly  represented,  if  at  all,  among 
the  membership  of  such  groups:  a true  scientist  and  a politician  are  a world  apart. 

Science  once  meant  knowledge;  scientific,  making  knowledge.  It  has  now  come  to  mean 
new  knowledge  ‘made’  by  the  scientific  method  of  hypothesis  and  experiment;  or  at  most 
knowledge  susceptible  of  verification  by  experiment.  Your  narrow-minded  scientist  may 
consider  this  the  only  true  knowledge.  Research,  often  synonymized  with  scientific  method, 
by  derivation  means  having  another  look  —a  small  step  from  verification  by  experiment. 
While  repetition  resulting  from  inadequate  literature  (re)search  is  justly  frowned  upon, 
usually  for  sound  economic  reasons,  it  can  cost  more  today  to  retrieve  published  informa- 
tion than  to  repeat  the  work.  Repetition  for  confirmation  of  results  which  have  been  called 
in  question  is  another  matter  and  is  often  necessary. 

Most  research  is  an  extension  from  previous  research;  if  it  has  an  avowed  purpose  this  is 
sometimes  referred  to  as  applied  research.  The  application  of  science  or  research  is  technolo- 
gy, which  uses  existing  knowledge  rather  than  making  new.  Research  which  breaks  new 
ground  and  is  not  an  extension  from  previous  research  cannot,  by  its  nature,  have  a purpose 
beyond  the  creation  of  new  knowledge.  Indeed  the  term  research  is  clearly  inappropriate 
here  and  such  work  is  often  referred  to  as  pure,  basic,  or  fundamental  science.  None  of 
these  terms  is  without  objection.  Pure,  because  although  most  other  work  may  be  biassed 
by  economic  purpose  it  is  unjust  to  imply  that  it  is  all  impure.  Basic,  because  to  a physical 
scientist  other  work  may  then  become  acid.  Fundamental,  because  of  its  implications  to  a 
biologist.  The  description  ‘free  basic  science’  has  recently  become  current  for  this  type  of 


66 


study,  and  free  is  certainly  apposite,  in  both  its  meanings.  For  who  can  regulate  where  all 
are  ignorant?  And  contrary  to  popular  belief,  basic  discoveries  are  not  usually  expensive, 
though  the  minds  that  can  make  them  are  rare,  and  the  rewards  often  wanting  or  post- 
humous. We  might  perhaps  call  this  kind  of  study  foundation  science. 

In  the  field  of  entomology,  taxonomy,  despite  the  slender  support  it  gets,  is  close  to 
foundation  science,  for  although  the  structure,  functions,  and  even  relationships  of  undes- 
cribed species  have  been  worked  on,  and  molecular  studies  are  sometimes  reported  without 
reference  to  a species,  it  is  usually  otherwise.  The  discovery  and  description  of  a “new” 
species  is  new  knowledge,  allowing  the  further  extension  of  knowledge  by  the  morpholo- 
gist, physiologist,  —and  even  the  molecular  biologist.  If,  as  sometimes  happens,  a new  species 
is  discovered  by  chance,  however,  though  foundation  it  is  not  science,  in  the  derived  sense. 
Often,  of  course,  the  existence  of  a new  species  can  be  hypothesized  from  known  species; 
its  discovery  is  then  truly  scientific  —but  no  longer  quite  foundation. 

Though  technological  virtuosity  now  allows  us  to  erect  a large  building  on  a small  base,  if 
the  edifice  of  science  is  to  serve  us  well  its  foundations  must  outspan  its  superstructure. 
This  is  not  now  so.  The  economic  rewards  of  technology  have  tempted  it  too  far  beyond  its 
foundation  science  in  too  many  directions  —dust  bowls,  pollution,  oil  spills,  thalidomide 
babies,  the  drug  problem—  are  some  of  the  consequences. 

Politicians,  policy,  and  police  are  words  of  common  origin,  collectively  implying  regula- 
tion, and  a settled  course  of  action.  But  foundation  science  is  born  of  curiosity  in  the  face 
of  ignorance  and  leads  none  knows  where.  To  regulate  it  is  to  destroy  it.  Direct  policy  for 
foundation  science  is  thus  a contradiction  in  terms.  But  in  a favorable  intellectual  climate  it 
thrives.  And  such  an  intellectual  climate  can  be  created.  If  policy  would  encourage  techno- 
logy it  must  first  provide  the  climate  for  foundation  science.  Failing  this  it  must  restrain 
technology  within  the  existing  foundations  of  science.  Advisory  bodies  concerned  with 
science  policy  should  devote  less  of  their  time  to  shuffling  funds  around  among  the  varyingly 
squeaky  wheels  at  various  levels  of  technology  and  applied  and  industrial  research,  and  more 
of  it  seeking  ways  and  means  of  creating  a climate  for  foundation  science.  Industry  can 
afford  its  own  research.  Never  since  the  Renaissance  has  the  world  been  in  such  dire  need 
of  new  thought  at  a basic  level  —of  enlarged  foundations  of  knowledge. 


Brian  Hocking 


STUDIES  ON  BOREAL  AGROMYZIDAE  (DIPTERA).  I. 

PHYTOMYZA  MINERS  ON  SAXIFRAGACEAE 

GRAHAM  C.  D.  GRIFFITHS 
Department  of  Entomology 

University  of  Alberta  Quaestiones  entomologicae 

Edmonton  7,  Alberta  8 : 67-80  1972 


Five  species  of  Phytomyza  are  known  as  leaf-miners  of  Saxifragaceae.  Three  of  these  are 
confined  to  Saxifraga,  as  follows:  Phytomyza  deirdreae  n.  sp.  (Western  Canada,  Alaska  and 
Japan,  type-locality  Sitka),  P.  saxifragae  Hering  (Central  Europe  and  Balkans)  and  P.  aizoon 
Hering  (Central  Europe).  On  other  genera  of  Saxifragaceae  two  new  species  are  recorded: 
Phytomyza  tiarellae  n.  sp.  on  Tiarella  and  Tolmiea  (type-locality  Sitka,  Alaska)  and  P. 
mitellae  n.  sp.  on  Mitella  (type-locality  Edmonton,  Alberta). 

Cinq  especes  de  Phytomyza  sont  connues  comme  mineuses  dans  les  feuilles  des  Saxi- 
fragacees.  Trois  de  ces  especes  sont  limitees  a la  Saxifraga,  tel  que:  Phytomyza  deirdreae 
n.  sp.  (L’ouest  du  Canada,  Alaska  etJapon,  localite-type  Sitka),  P.  saxifragae  Hering  ( Europe 
centrale  et  Balkans)  et  P.  aizoon  Hering  (Europe  centrale).  Sur  les  autres  genres  de  Saxi- 
fragacees  deux  especes  nouvelles  sont  rapportees:  Phytomyza  tiarellae  n.  sp.  sur  Tiarella  et 
sur  Tolmiea  (localite-type  Sitka,  Alaska)  et  P.  mitellae  n.  sp.  sur  Mitella  (localite-type 
Edmonton,  Alberta). 

Fiinf  Phytomyza -Arten  sind  als  Blattminierer  von  Saxifragaceae  bekannt.  Drei  von  diesen 
sind  auf  Saxifraga  beschrankt,  wie  folgt:  Phytomyza  deirdreae  n.  sp.  (Westlich  Kanada, 
Alaska  und  Japan,  Fundort  vom  Typus  Sitka),  P.  saxifragae  Hering  (Mitteleuropa  und 
Balkanhalbinsel)  und  P.  aizoon  Hering  (Mitteleuropa).  An  anderen  Saxifragaceen-Gattungen 
werden  zwei  neue  Arten  besprochen:  Phytomyza  tiarellae  n.  sp.  an  Tiarella  und  an  Tolmiea 
(Fundort  vom  Typus  Sitka,  Alaska)  und  P.  mitellae  n.  sp.  an  Mitella  (Fundort  vom  Typus 
Edmonton,  Alberta). 


The  present  paper  is  the  first  of  a series  dealing  with  boreal  and  arctic  Agromyzidae,  both 
from  the  Palaearctic  and  Nearctic  regions.  The  distinction  between  these  regions  becomes 
unnatural  at  the  level  of  the  boreal  forest,  because  many  of  the  species  found  here  are  dis- 
tributed in  both  regions  or  have  their  closest  relatives  in  the  other  region.  I will  be  particu- 
larly concerned  in  this  series  with  making  critical  comparisons  between  European  and  North 
American  material,  in  order  to  establish  which  species  are  holarctic.  The  Agromyzidae  are 
well  suited  for  studies  of  historical  biogeography,  because  their  restricted  choice  of  larval 
host-plants  allows  hypotheses  about  their  dispersal  to  be  correlated  with  the  likely  dispersal 
of  their  host-plants. 

The  references  listed  in  this  series  with  the  synonyms  of  each  species  will  refer  only  to 
works  which  contain  nomenclatural  proposals  or  present  substantial  new  information  on 
the  species.  References  in  catalogues,  faunal  lists  and  summarizing  works  will  not  be  listed 
in  synonymies  unless  meeting  the  above  criteria. 

In  the  present  paper  I deal  with  the  miners  of  Saxifragaceae  sensu  stricto.  I do  not  treat 
the  miners  of  the  Hydrangeaceae  (including  Philadelphus,  Deutzia  and  Hydrangea),  which 
l are  included  in  Saxifragaceae  in  some  botanical  classifications.  Names  of  plants  are  used  in 
the  sense  of  Webb  (1964)  for  European  species,  and  of  Hulten  (1968)  for  North  American 
species. 


68 


Griffiths 


The  holotypes  of  the  new  species  described  in  this  paper  will  be  deposited  in  the  Canadian 
National  Collection  (Ottawa).  Other  North  American  material  is  in  the  University  of  Alberta 
collections  and  in  my  personal  collection.  Leaf  mines  of  the  North  American  species  are 
preserved  in  my  herbarium  of  mines. 

TERMS  APPLIED  TO  MALE  GENITALIA 

I have  discussed  elsewhere  (Griffiths,  in  press)  the  terms  applied  to  the  male  postabdomen 
and  genitalia  of  cyclorrhaphous  Diptera  in  general.  The  proposals  in  that  book  entail  modi- 
fications of  the  terms  in  use  for  some  parts  of  the  male  genitalia  of  Agromyzidae.  Table  1 
sets  out  the  terms  used  in  the  present  series  of  papers,  with  the  equivalent  term  or  terms 
used  in  recent  literature  on  Agromyzidae. 


Table  1 . Equivalence  of  terminology. 


Equivalent  previous  terminology 

Revised  terms 

Areas  of  aedeagus 

basal  section  (basiphallus)  and  phallophore 

basal  section 

distal  and  median  sections 

distal  section 

hypophallus 

medial  lobe(s) 

Sclerite s of  aedeagus 

arms  of  basiphallus,  or  sclerites  of  basiphallus 

basal  sclerites  or  paraphalli 

paraphalli 

paramesophalli 

sclerites  of  hypophallus 

sclerites  of  medial  lobe(s) 

(The  application  of  the  terms  phallophore,  distiphallus  and  mesophallus  is  unchanged.) 

Other  parts  of  genital  segment 

epandrium 

periandrium 

surstyli 

telomeres 

(The  application  of  the  terms  epiphallus,  aedeagal  hood,  hypandrium,  pregonites  and 
postgonites  is  unchanged.) 


I now  consider  the  narrow  dorsal  band  of  sclerotization  found  after  the  6th  tergum  in 
some  agromyzid  species  as  a remnant  of  the  inverted  8th  sternum  (a  large  sclerite  in  many 
other  families  of  Schizophora). 

A special  difficulty  already  recognized  by  other  authors  (Nowakowski,  1964;  von  Tschirn- 
haus,  1969)  involves  the  application  of  the  terms  “dorsal”  and  “ventral”  to  the  aedeagus.  In 
Agromyzidae  and  many  other  families  of  Schizophora,  the  aedeagus  is  swung  by  muscular 
action  through  a wide  arc  from  a posteriorly  directed  copulatory  position  to  an  anteriorly 
directed  rest  position  (Griffiths,  in  press).  Which  side  of  the  aedeagus  is  dorsal  and  which 
ventral  thus  depends  on  the  position  of  the  organ.  The  convention  in  descriptions  of  Agro- 


Boreal  Agromyzidae 


69 


myzidae  is  to  apply  these  terms  with  reference  to  the  rest  position  of  the  aedeagus.  Probably 
little  would  be  gained  by  attempting  to  change  this  convention.  But  the  ambiguity  of  these 
terms  should  be  appreciated.  In  discussions  where  an  equivalent  application  of  such  terms  as 
“dorsal”  and  “ventral”  throughout  the  Diptera  is  needed  these  terms  should  be  applied  to 
the  copulatory  position  of  the  aedeagus  in  those  groups  of  Schizophora  which  show  the 
swinging  mechanism. 


ABBREVIATIONS 


The  following  conventional  abbreviations  are  used  in  descriptions: 


acr 

dc 

ia 

mg2 , mg3 , mg4 
ori 
ors 
pa 


acrostichal  setulae 
dorsocentral  bristle(s) 
intra-alar  setulae 

second,  third  and  fourth  costal  sections 
lower  orbital  bristle(s) 
upper  orbital  bristle(s) 
postalar  bristle(s) 


RELATIONSHIPS  OF  SPECIES  TREATED 

In  my  discussion  of  the  Phytomyza  syngenesiae  group  (Griffiths,  1967)  I alluded  to  the 
possibility  of  defining  as  one  of  the  segregates  of  Phytomyza  in  the  present  sense  a group 
containing  the  syngenesiae  group,  the  milii  group  and  P.  nigra  Meigen.  The  species  now 
treated  in  this  paper,  as  well  as  some  of  the  Phytomyza  miners  of  Gentianaceae  and  Capri- 
foliaceae,  may  be  added  to  this  list.  Hardy’s  (1849)  name  Chromatomyia  may  be  applied  to 
this  group  (whether  as  genus  or  subgenus),  when  a division  of  Phytomyza  in  the  present 
wide  sense  is  proposed  (see  Griffiths,  1967).  But  such  a formal  proposal  would  be  premature 
at  the  present  time,  as  the  male  genitalia  of  many  European  species  have  still  not  been  de- 
scribed. The  structure  of  the  distal  section  of  the  aedeagus  in  this  group  is  strongly  modified 
(apomorphous).  Most  characteristic  is  the  presence  of  a pair  of  dorsal  “supporting  sclerites”, 
arising  from  the  base  of  the  distal  section  (Fig.  8).  I accept  von  Tschirnhaus’  (1969)  opinion 
that  these  sclerites  should  not  be  called  the  “distiphallus”  (as  in  my  1 967  paper),  and  follow 
him  in  calling  them  supporting  sclerites  (“Stiitzsklerite”).  The  medial  lobe  (“hypophallus”) 
is  poorly  or  not  at  all  differentiated.  And  it  is  doubtful  whether  a true  distiphallus  (contain- 
ing a bifid  terminal  portion  of  the  ejaculatory  duct)  is  retained  in  any  members  of  this 
group.  Von  Tschirnhaus  uses  the  term  distiphallus  for  the  distal  tubule  containing  the  ejacu- 
latory duct  in  the  syngenesiae  group;  but  since  this  is  unpaired  it  more  probably  represents 
the  mesophallus  (as  assumed  in  my  1967  paper)  or  a secondary  sclerotization. 

All  species  with  the  type  of  aedeagus  described  above  also  show  a characteristic  apomor- 
phous type  of  puparium.  The  puparium  remains  within  the  host  plant,  with  its  anterior 
spiracles  bent  downwards  so  that  they  project  through  the  epidermis.  Hardy  ( 1 849)  charac- 
terized his  proposed  genus  Chromatomyia  on  the  basis  of  this  puparium  type.  However  this 
puparium  type  has  a wider  distribution  than  the  type  of  aedeagus  described  above.  Either 
* the  apomorphous  puparium  type  indicates  a wider  monophyletic  group  inclusive  of  the 
group  characterized  by  the  apomorphous  type  of  aedeagus;  or  the  puparium  type  has 
evolved  more  than  once.  The  latter  possibility  cannot  be  evaluated  without  studies  of  addi- 
tional groups  of  species.  But  I am  confident  that  the  species  which  show  both  the  modified 
form  of  aedeagus  and  the  Chromatomyia- type  of  puparium  form  a monophyletic  group, 
deserving  eventually  of  nomenclatural  recognition. 


70 


Griffiths 


DIAGNOSIS 

The  species  treated  in  this  paper  can  be  identified  most  readily  as  larvae  or  puparia.  The 
three  species  on  Saxifraga  show  obvious  differences  in  the  form  of  the  posterior  larval  (and 
puparial)  spiracles  (Fig.  3-5).  In  the  new  species  tiarellae  (on  Tiarella  and  Tolmiea)  and 
mitellae  (on  Mitella)  these  spiracles  have  a characteristic  crescentic  form  (Fig.  6-7).  No  other 
agromyzid  larvae  are  known  to  mine  the  leaves  of  Saxifragaceae. 

Caught  males  of  these  species  can  be  identified  by  study  of  their  genitalia,  particularly 
the  form  of  the  distal  section  of  the  aedeagus.  I am  doubtful  whether  reliable  diagnosis  is 
possible  on  the  basis  of  the  external  form  of  the  adult. 

The  three  new  species  may  be  included  in  Spencer’s  (1969:219)  key  to  Phytomyza 
species  of  Canada  and  Alaska  by  the  following  extensions. 

56.  Sides  of  thorax  bright  yellow notopleuralis  Spencer 

Sides  of  thorax  predominantly  dark 56a 

56a.  Aedeagus  as  in  Spencer’s  Fig.  395,  with  membranous  distal  section 

agromyzina  Meigen 

Aedeagus  as  in  Figs.  13  and  15-17 56b 

56b.  Mesonotum  strongly  shining;  aedeagus  as  in  Fig.  13 mitellae  n.  sp. 

Mesonotum  weakly  shining;  aedeagus  as  in  Fig.  17 tiarellae  n.  sp. 


58.  Distal  section  of  aedeagus  with  cylindrical  mesophallus  and  distiphallus  consisting  of 


divergent  tubules  (Spencer’s  Fig.  442) ilicis  Curtis 

Aedeagus  not  of  this  type 59 

59.  Aedeagus  as  in  Spencer’s  Figs.  447-448  involucratae  Spencer 

Aedeagus  as  in  Spencer’s  Fig.  460  milii  Kaltenbach 

Aedeagus  as  in  Figs.  8-9 deirdreae  n.  sp. 


TREATMENT  OF  SPECIES 
Phytomyza  deirdreae  new  species 

"Phytomyza  saxifragae  Hering”.  Sasakawa,  1956:105.  —1961:467. 

Adult.  — Head  (Fig.  2)  with  orbits  not  or  only  very  narrowly  projecting  above  eyes  in 
lateral  view;  genae  in  middle  about  XA  of  eye  height;  eye  pubescence  fine  and  inconspicuous. 
Frons  at  level  of  front  ocellus  about  twice  width  of  eye.  Two  ors,  of  equal  length,  posteri- 
orly directed;  two  ori,  inwardly  directed,  anterior  at  least  half  as  long  as  posterior;  orbital 
setulae  one-rowed.  Peristomal  margin  with  vibrissa  and  6-8  upcurved  peristomal  setulae. 
Third  antennal  article  rounded  distally,  with  only  short  pubescence. 

3 + 1 dc;  acr  numerous,  in  5-7  rows  anteriorly,  4-5  rowed  posteriorly;  presutural  ia  numer- 
ous; 11-16  postsutural  ia;  inner  pa  long,  over  half  as  long  as  outer  pa. 

Second  cross-vein  (m-m)  absent.  Costal  ratio  mg2/mg4  2.9-3. 1 in  type  series  (about  3.5 
in  Japanese  material  according  to  Sasakawa,  1956).  Wing  length  about  2.5  mm  (both  sexes). 

Colour  largely  dark.  Centre  of  frons  dark  brown,  only  slightly  paler  than  black  orbits  and 
ocellar  plate;  genae  dark  brown.  Antennae  black.  Palpi  black;  labella  brown  or  yellow- 
brown.  Thorax  finely  grey-dusted,  only  weakly  shining,  completely  black  except  whitish 
seams  of  notopleural  and  mesopleural  sutures;  squamal  margin  and  fringe  infuscated;  wing 
base  infuscated.  Legs  dark,  with  tips  of  femora  yellow-brown  (but  only  those  of  front  legs 


Boreal  Agromyzidae 


71 


distinctly  so).  Basal  cone  of  ovipositor  (9)  dusted  on  about  basal  two-thirds. 

Male  postabdomen  with  8th  sternum  fused  with  6th  tergum.  Telomeres  not  delimited 
from  periandrium,  indicated  by  dense  group  of  short  setulae.  Aedeagus  as  in  Fig.  8-9,  with 
large  ventral  area  enclosed  by  membrane,  without  medial  lobe;  supporting  sclerites  fused 
basally,  in  form  of  Y-shaped  structure  with  base  confluent  with  short  stretch  of  sclerotiza- 
tion  of  ejaculatory  duct;  other  distal  sclerites  (?  mesophallus  or  paramesophalli)  better 
developed  than  in  saxifragae  and  aizoon,  extending  anteriorly  below  supporting  sclerites. 
Ejaculatory  apodeme  (Fig.  10)  very  small. 

Additional  figures  and  information  on  the  female  genitalia  (not  considered  here)  are 
given  by  Sasakawa  (1961). 

Puparium  and  third  instar  larva.  - Mandibles  with  two  alternating  teeth;  right  mandible 
longer  than  left.  Anterior  spiracles  two-horned,  with  at  least  25  bulbs.  Posterior  spiracles 
(Fig.  3)  with  about  40-45  bulbs,  with  two  very  long  and  slender  horns  which  are  directed 
more  or  less  vertically  on  puparium.  Colour  of  puparium  variable  (white,  brown  or  blackish). 
Length  of  puparium  2. 3-2. 5 mm. 

Other  figures  are  given  by  Sasakawa  (1961). 

Mine.  — Larvae  leaf-miners  on  certain  Saxifraga  species  (see  records  below).  Mine  (Fig. 
22)  at  origin  with  short  linear  channel,  but  soon  broadened  into  irregular  blotch  (the  latter 
in  some  cases  enclosing  the  initial  linear  channel),  appearing  white  or  greenish  white  in  in- 
cident light;  faeces  scattered  as  discrete  particles  throughout  mine;  main  part  of  mine  nor- 
mally formed  on  upper  surface  of  leaf,  with  pupation  following  at  end  of  short  channel 
without  faeces  on  lower  surface  (but  a few  mines  formed  entirely  on  the  lower  surface  were 
also  found).  Puparium  with  its  ventral  surface  adjacent  to  surface  of  leaf,  with  its  anterior 
spiracles  projecting  ventrally  through  epidermis. 

Types.  — Holotype  6,  3 99  paratypes  from  larvae  and  puparia  19.viii.69  on  Saxifraga 
ferruginea  Graham,  Harbour  Mountain  (1900  feet  elevation),  Sitka,  Alaska,  emerged  l.ix.69, 
2.ix.69,  5.V.70  (holotype)  and  6.V.70,  leg.  D.  E.  and  G.  C.  D.  Griffiths. 

Additional  records.  — I hope  to  obtain  further  material  from  larvae  and  puparia  collected 
1 5-23 .viii.7 1 on  Saxifraga  lyallii  Engler,  S.  nivalis  L.  and  S.  punctata  L.  on  the  slopes  above 
the  Mount  Cavell  chalet,  Jasper  National  Park,  Alberta,  at  elevations  between  5900  and 
7900  feet. 

Additional  records  for  North  America,  based  on  my  own  collections  of  larvae  and  puparia 
which  yielded  parasites,  are  as  follows: 

Puparia  20.viii.69  on  Saxifraga  punctata  L.,  same  locality  as  type  series  (1000  feet  eleva- 
tion); puparia  27.viii.69  on  Saxifraga  punctata  L.,  Chilkat  Pass  (3000  feet  elevation), 
Haines  highway,  British  Columbia;  larvae  and  puparia  17-19.vii.68  on  Saxifraga  hieraci- 
folia  Waldst.  and  Kit.  and  S.  punctata  L.,  Eagle  Summit  (3900  feet  elevation),  Steese 
highway,  Alaska. 

Sasakawa  (1956)  described  material  bred  from  Saxifraga  sachalinensis  Fr.  Schm.,  Jyo- 
zankei,  Hokkaido,  Japan  (leg.  Y.  Nishijima).  In  his  1961  work  he  also  records  this  species 
on  S.  fusca  Maxim.,  Mount  Hakusan,  Toyama  Prefecture  (Japan). 

Dedication.  — I am  pleased  to  dedicate  this  species  to  my  wife  Deirdre,  who  has  assisted 
me  ably  on  field  work. 

Discussion.  — The  description  of  this  species  brings  the  total  of  known  Phytomyza  miners 
of  Saxifraga  to  three.  The  other  two  species  {saxifragae  and  aizoon ) are  known  only  from 
Europe.  The  three  species  are  probably  monophyletic,  as  evidenced  by  the  similar  form  of 
the  aedeagus.  The  most  obvious  differences  between  them  are  in  the  form  of  the  posterior 
larval  (and  puparial)  spiracles  (Fig.  3-5).  There  are  also  slight  differences  in  the  form  of  the 
distal  section  of  the  aedeagus.  There  are  probably  some  statistical  differences  in  the  external 


72 


Griffiths 


form  of  the  adult,  for  instance  in  the  costal  ratio  and  numbers  of  thoracic  setulae;  but  the 
available  material  of  all  species  is  too  limited  for  reliable  statistical  treatment. 

The  occurrence  of  Phytomyza  miners  on  Saxifraga  in  Finland  is  indicated  by  Linnaniemi’s 
(1913,  Fig.  29)  photograph  of  mines  on  Saxifraga  nivalis  L.  I think  these  mines  may  well  be 
those  of  deirdreae,  but  no  firm  opinion  can  be  given  in  the  absence  of  information  on  the 
form  of  the  puparia.  Hering  (1957)  includes  Linnaniemi’s  record  as  no.  4648  in  his  key  to 
miners  of  Saxifraga. 

I regard  the  Japanese  material  described  by  Sasakawa  (1956,  1961)  as  probably  conspe- 
cific  with  my  North  American  material,  not  with  the  Central  European  species  Phytomyza 
saxifragae  Hering.  Sasakawa’s  (1961)  Fig.  143n  indicates  long  and  slender  horns  on  the  pos- 
terior larval  spiracles,  and  his  figure  of  the  aedeagus  (143d)  also  agrees  substantially  with 
that  of  the  holotype  of  deirdreae.  I detect  a discrepancy  only  in  his  figure  (143c)  of  the 
telomeres  (“surstyli”).  The  group  of  short  spines  indicated  by  Sasakawa  are  represented  by 
rather  longer  setulae  in  the  holotype. 

The  known  distribution  of  Phytomyza  deirdreae  is  indicated  on  Fig.  19. 

Phytomyza  saxifragae  Hering  1924 

Phytomyza  saxifragae  Hering.  Hering,  1924:38.  —1927:135.  De  Meijere,  1926:289.  —1941: 

25.  Hendel,  1928:99.  —1935:473.  Holotype  9,  Herculesbad  (Roumania),  in  the  Zoolo- 

gisches  Museum,  Humboldt  Universitat,  Berlin. 

Adult.  — Hendel  (1935)  has  described  the  external  form  of  the  adult  in  detail.  I am  unable 
to  separate  this  species  from  deirdreae  on  external  characters.  The  costal  ratio  mg2/mg4  is 
3. 3-3. 5 in  the  specimens  I examined.  The  sclerites  of  the  wing  base  are  paler  than  in  deir- 
dreae, but  Hendel’s  (1935)  description  “Fliigelwurzel  weisslichgelb,  kontrastierend”  seems 
exaggerated.  The  colour  difference  is  not  so  great  that  I would  rely  on  it  for  identification. 

Male  postabdomen  and  genitalia  similar  to  those  of  deirdreae,  but  with  some  difference 
in  form  of  distal  section  of  aedeagus  (Fig.  11);  base  of  supporting  sclerites  not  confluent 
with  short  stretch  of  sclerotization  of  ejaculatory  duct;  area  below  supporting  sclerites 
membranous. 

Puparium  and  third  instar  larva.  — Differing  from  deirdreae  in  form  of  posterior  spiracles, 
which  have  22-25  bulbs  in  a widely  open  bow  (Fig.  5),  with  only  one  prominent  horn  which 
is‘directed  more  or  less  horizontally  on  puparium.  See  further  the  descriptions  and  figures 
of  de  Meijere  (1926,  1941).  Puparium  black  ventrally,  red  dorsally  (Hering,  1927). 

Mine.  — Larvae  leaf-miners  on  Saxifraga  rotundifolia  L.  Mine  (Fig.  23)  primarily  linear 
according  to  Hering  (1924,  1927)  and  Hendel  (1928,  1935),  but  seldom  extended,  usually 
crossing  itself  or  blending  to  form  secondary  blotch,  appearing  whitish  in  incident  light. 
Hendel  gives  the  length  of  the  mine  as  about  14  cm,  and  its  greatest  terminal  width  as 
2.75-3.0  mm.  Faeces  scattered  as  discrete  particles  on  either  side  of  mine  channel  (separated 
by  about  5 mm  in  terminal  part  of  mine).  Main  part  of  mine  normally  formed  on  upper 
surface  of  leaf  (but  sometimes  on  lower  surface  according  to  Hering),  with  pupation  nor- 
mally following  on  lower  surface.  Puparium  (when  internal)  with  its  ventral  surface  adjacent 
to  surface  of  leaf,  with  its  anterior  spiracles  projecting  ventrally  through  epidermis. 

Hering  (1924,  1927)  stated  that  puparia  were  found  inside  the  leaf,  which  I think  must 
be  their  normal  location  in  view  of  their  morphological  adaptation  to  this  end  (anterior 
spiracles  turned  downwards).  However  Hendel  (1928)  reported  that  larvae  may  also  leave 
the  leaf  to  pupate. 

A photograph  of  the  leaf  mine  is  given  by  Hendel  (1928,  Tafel  V). 

Material  examined.  — 1 9 from  mine  on  Saxifraga  rotundifolia  L.,  West  Rila  mountains, 


Boreal  Agromyzidae 


73 


Bulgaria,  emerged  31.viii.39,  leg.  H.  Buhr.  1 6 from  mine  on  Sax  if raga  rotundifolia  L.,  Vais, 
Switzerland,  emerged  8.vii.29,  leg.  W.  Hopp.  1 6 from  mine  on  Saxifraga  rotundifolia  L., 
Rigi,  Switzerland,  emerged  2.viii.25,  leg.  M.  Hering. 

Additional  records.  — This  species  was  originally  described  from  Herculesbad  in  the  Banat 
region  of  Roumania  (Hering,  1924)  (holotype  9 emerged  30.V.22  from  puparium  collected  13. 
v.22).  Hering  also  refers  in  that  paper  to  the  finding  of  mines  at  Konigssee,  near  Berchtesga- 
den  in  Bavaria  (Germany).  There  are  also  sheets  in  Hering ’s  mine  herbarium  (now  in  the  Brit- 
ish Museum)  for  the  Plockenpass,  Carinthia  (Austria),  27.vi.29,  leg.  Hedicke;  and  for  Brunn- 
steinsee,  Warscheneck-Gebirge,  Austria  (1600  metres  elevation),  28.viii.60,  leg.  E.  M.  Hering. 

Discussion.  — The  above  records  indicate  that  this  species  is  widely  distributed  at  high 
elevations  in  the  mountains  of  central  Europe  and  the  Balkans  (Fig.  20).  Buhr  (reported  by 
de  Meijere,  1941)  gives  its  altitudinal  range  in  the  West  Rila  mountains  as  1600  to  2200 
metres.  Webb  (1964)  indicates  that  the  host-plant  is  widely  distributed  in  the  mountains  of 
central  and  southern  Europe,  but  does  not  occur  in  northern  Europe. 

Phytomyza  aizoon  Hering  1932 

Phytomyza  aizoon  Hering.  Hering,  1932:162.  Hendel,  1934:337.  De  Meijere,  1938:87.  Syn- 
types  6 9,  Mauthen  (Carinthia,  Austria),  in  the  Zoologisches  Museum,  Humboldt  Univer- 
sitat,  Berlin. 

Adult.  — Hering  (1932)  and  Hendel  (1934)  have  described  the  external  form  of  the  adult 
in  detail.  Adults  of  this  species  are  substantially  similar  on  external  characters  to  those  of 
the  previous  two  species  (. saxifragae  and  deirdreae ),  but  I note  the  following  points.  Accord- 
ing to  Hendel  the  orbits  in  aizoon  are  distinctly  projecting  above  the  eye  in  lateral  view 
(his  Fig.  345),  and  the  arista  is  thickened  to  about  its  middle  (only  on  about  its  basal  third 
in  the  other  two  species).  8-1 1 postsutural  ia.  The  costal  ratio  mg2/mg4  is  only  2.4  in  the 
paratype  examined  by  me;  the  value  3.0  in  the  original  description  (Hering,  1932)  is  prob- 
ably an  overestimate,  as  already  implied  by  Hendel’s  (1935)  placement  of  aizoon  in  his  key 
(p.  511).  Size  smaller  (wing  length  about  1.75  mm). 

Male  postabdomen  and  genitalia  similar  to  those  of  deirdreae  and  saxifragae,  but  with 
some  difference  in  form  of  distal  section  of  aedeagus  (Fig.  12);  base  of  supporting  sclerites 
not  confluent  with  short  stretch  of  sclerotization  of  ejaculatory  duct;  area  below  supporting 
sclerites  membranous,  not  extending  so  far  anteriorly  as  in  saxifragae. 

Puparium  and  third  instar  larva.  — Differing  very  obviously  from  saxifragae  and  deirdreae 
in  form  of  spiracles.  Anterior  spiracles  knob-shaped,  with  only  9-10  bulbs  (Hering,  1932). 
Posterior  spiracles  (Fig.  4)  small,  knob-shaped,  with  only  9-12  bulbs.  Puparium  white,  2.3 
mm  long. 

Mine.  — Larvae  leaf-miners  on  Saxifraga  paniculata  Miller  (=  aizoon  Jacq.).  Hering  (1932, 
1957)  describes  the  mine  as  a gradually  widening  upper-surface  channel,  sometimes  branch- 
ing, often  becoming  blotch-like  terminally;  appearing  greenish  or  brownish  in  incident  light; 
with  mine  channel  sometimes  becoming  swollen  subsequently  due  to  formation  of  callus 
tissue;  faecal  particles  present.  Puparium  remaining  in  mine,  with  its  ventral  surface  adjacent 
to  surface  of  leaf,  with  its  anterior  spiracles  projecting  ventrally  through  epidermis. 

Material  examined.  — 1 6 paratype  from  mine  24.vii.29  on  Saxifraga  paniculata  Miller, 
Mauthen,  Carinthia,  Austria,  emerged  3.viii.29,  leg.  O.  Hering. 

Additional  records.  - Hering  (1932)  records  this  species  for  Mauthen  (Carinthia,  Austria) 
and  Zernez,  Switzerland  (adult  emerged  16.viii.29  from  mines  collected  12.vii.  29,  leg. 
Hopp).  The  only  additional  collection  which  I have  traced  is  by  Zavrel  on  12.ix.52  at  Berg 
Kotouc,  Stramberg,  Eastern  Moravia  (Czechoslovakia)  (sheet  in  Hering’s  mine  herbarium). 


74 


Griffiths 


Discussion.  — The  above  records  suggest  a restricted  distribution  for  this  species  in  the 
mountains  of  central  Europe  (Fig.  21),  where  it  is  sympatric  with  saxifragae.  But  the  real 
distribution  may  well  be  much  wider,  for  Webb  (1964)  indicates  that  the  host-plant  is  wide- 
ly distributed  also  in  southern  Europe,  Asia  Minor  and  the  Caucasus,  and  occurs  locally  in 
Norway.  A “variety”  of  the  host-plant  occurs  in  North  America  (mainly  in  the  East),  but 
has  not  yet  been  examined  for  leaf  miners. 

Phytomyza  tiarellae  new  species 

Adult.  — Head  (compare  Fig.  1)  with  proportionately  large  eyes;  orbits  not  projecting 
above  eyes  in  lateral  view;  genae  in  middle  less  than  V*  of  eye  height;  eye  pubescence  fine 
and  inconspicuous.  Frons  at  level  of  front  ocellus  about  twice  width  of  eye.  Two  ors,  of 
equal  length,  posteriorly  directed;  two  ori,  inwardly  directed,  anterior  pair  variable  in  length 
(only  slightly  shorter  than  posterior  pair  in  holotype,  but  less  than  half  as  long  in  paratype); 
orbital  setulae  one-rowed.  Peristomal  margin  with  vibrissa  and  5-6  upcurved  peristomal  setu- 
lae.  Third  antennal  article  rounded  distally,  slightly  longer  than  high,  with  fairly  long  pale 
pubescence. 

3 + 1 dc;  acr  numerous,  in  5-6  rows  anteriorly,  becoming  4-5  rowed  posteriorly;  pre- 
sutural  ia  numerous;  6-10  postsutural  ia;  inner  pa  about  half  as  long  as  outer  pa. 

Second  cross-vein  (m-m)  absent.  Costal  ratio  mg2/mg4  3.0  in  male  holotype,  3.5  in  female 
paratype.  Wing  length  2.1  mm  (holotype),  2.5  mm  (paratype). 

Colour  largely  dark.  Centre  of  frons  partly  brown  (paler  than  black  orbits  and  ocellar 
plate);  genae  brown.  Antennae  black.  Palpi  black;  labella  yellow.  Thorax  finely  grey-dusted, 
only  weakly  shining,  completely  black  except  whitish  seams  of  sutures  (especially  noto- 
pleural  and  mesopleural  sutures);  squamal  margin  and  fringe  infuscated,  but  wing  base  con- 
trastingly whitish.  Legs  with  coxae,  trochanters  and  femora  largely  dark,  but  with  tips  of 
femora  and  whole  of  tibiae  and  tarsi  contrastingly  deep  yellow  or  yellow-brown.  Abdomen 
largely  dark  brown.  Basal  cone  of  ovipositor  (9)  dusted  on  about  basal  two-thirds. 

Male  postabdomen  with  8th  sternum  fused  with  6th  tergum.  Telomeres  not  delimited 
from  periandrium,  indicated  by  dense  group  of  short  setulae.  Aedeagus  as  in  Fig.  15-17, 
with  medial  lobe  weakly  differentiated;  supporting  sclerites  closely  approximated,  parallel; 
small  membranous  lobe  present  distal  to  supporting  sclerites;  sclerites  below  supporting 
sclerites  (?  paramesophalli)  appearing  broad  basally  in  lateral  view,  extending  distally  almost 
as  far  as  supporting  sclerites.  Ejaculatory  apodeme  (Fig.  18)  very  small. 

Puparium  and  third  instar  larva.  - Mandibles  with  two  alternating  teeth;  right  mandible 
longer  than  left.  Anterior  spiracles  two-horned,  with  about  20  bulbs.  Posterior  spiracles 
(Fig.  6)  one-horned,  with  18-23  bulbs  arranged  more  or  less  in  crescent.  Puparium  brown 
or  white,  with  darker  strip  on  ventral  surface.  Length  of  puparium  1. 9-2.1  mm. 

Mine.  - Larvae  leaf-miners  on  Tiarella  trifoliata  L.  and  Tolmiea  menziesii  (Pursh).  Mine 
(Fig.  24)  entirely  linear,  appearing  white  in  incident  light,  up  to  20-25  cm  long,  about  2 mm 
wide  terminally;  faeces  scattered  as  discrete  particles  (mostly  separated  by  over  1 mm),  or 
forming  short  “threads”  (Fadenstiicke)  in  terminal  part  of  mine;  mine  formed  entirely  on 
upper  surface  of  leaf,  but  with  puparium  formation  following  on  lower  surface  at  end  of 
mine  channel.  Puparium  with  its  ventral  surface  adjacent  to  surface  of  leaf,  with  its  anterior 
spiracles  projecting  ventrally  through  epidermis. 

Types.  — Holotype  6,  1 9 paratype  from  larvae  and  puparia  22-24.viii.69  on  Tiarella 
trifoliata  L.,  Starrigavan,  Sitka,  Alaska  (near  sea  level),  emerged  9.ix.69  and  8.V.70  (holo- 
type), leg.  G.  C.  D.  Griffiths. 

Discussion.  — Puparia  were  also  collected  at  the  type  locality  on  Tolmiea  menziesii 


Boreal  Agromyzidae 


75 


(Pursh),  but  only  parasites  obtained  from  this  sample. 

The  known  host-plants  of  tiarellae  are  both  distributed  mainly  in  the  rain  forest  of  the 
Pacific  coast  of  North  America,  with  ranges  from  Alaska  to  northern  California  (Hulten, 
1968).  A similarly  restricted  distribution  may  also  be  expected  for  the  fly.  The  mean  annual 
rainfall  at  the  type  locality  is  probably  about  100  inches,  on  the  basis  of  data  for  the  Sitka 
Magnetic  weather  station. 

The  emergence  of  one  of  the  adult  flies  soon  after  collection  of  the  puparia  indicates  that 
this  species  is  at  least  partly  multivoltine,  unlike  the  species  next  to  be  described. 

Phytomyza  mitellae  new  species 

Adult.  - External  form  of  adult  as  described  for  tiarellae,  except  as  follows.  Frons  about 
1.5  times  eye  width  at  level  of  front  ocellus.  Anterior  pair  of  ori  well  developed  in  all  speci- 
mens, at  least  half  as  long  as  posterior  pair  (Fig.  1).  Costal  ratio  mg2/mg4  2.4  in  male  holo- 
type,  2. 7-3.0  in  female  paratypes.  Wing  length  1.9-2. 2 mm.  Ocelli  bright  red  in  most  speci- 
mens, but  yellow  in  two  females  (as  normally  in  Phytomyza,  including  all  other  species 
treated  in  this  paper).  Mesonotum  strongly  shining,  with  only  very  fine  dusting;  sides  of 
mesonotum  brown;  mesopleuron  with  dorsal  and  posterior  margins  narrowly  white;  squamae 
pale,  with  only  their  fringe  infuscated.  Abdomen  brown,  in  some  specimens  yellowish  on 
sides  at  base. 

Male  postabdomen  and  genitalia  very  similar  to  those  of  tiarellae,  but  with  some  differ- 
ence in  form  of  distal  section  of  aedeagus  (Fig.  13);  sclerites  below  supporting  sclerites 
(?  paramesophalli)  appearing  narrower  in  lateral  view,  not  extending  so  far  distally  (with 
their  apices  well  short  of  apices  of  supporting  sclerites). 

Puparium  and  third  instar  larva.  — Very  similar  to  tiarellae ; posterior  spiracles  (Fig.  7) 
one-horned,  with  14-17  bulbs  arranged  more  or  less  in  crescent.  Puparium  uniformly  brown 
or  yellow-brown.  Length  of  puparium  1. 7-2.1  mm. 

Mine.  — Larvae  leaf-miners  on  Mitella  nuda  L.  Mine  (Fig.  25)  entirely  linear,  appearing 
white  in  incident  light,  10-1 1 cm  long,  1 .5-2.0  mm  wide  terminally;  faeces  deposited  as  fine 
particles,  forming  more  or  less  continuous  strip  in  early  part  of  mine,  separated  (but  mostly 
by  less  than  1 mm)  in  terminal  part  of  mine;  mine  formed  entirely  on  upper  surface  of  leaf, 
but  with  puparium  formation  following  on  lower  surface  at  end  of  mine  channel.  Puparium 
with  its  ventral  surface  adjacent  to  surface  of  leaf,  with  its  anterior  spiracles  projecting  ven- 
trally  through  epidermis. 

Types.  — Holotype  <3,  6 99  paratypes  from  larvae  and  puparia  21  .viii-27.ix.70  on  Mitella 
nuda  L.,  Edmonton  (White  Mud  Creek  and  north-facing  slopes  of  river  valley),  Alberta, 
emerged  14-26.V.71  (holotype  14.V.71),  leg.  D.  C.  Christophel,  V.  K.  Sehgal,  D.  E.  and 
G.  C.  D.  Griffiths. 

Additional  records.  — This  species  also  occurs  at  Elk  Island  National  Park,  Alberta  (mines 
with  larvae  noted  on  2 1 .ix.7 1 ). 

Discussion.  — The  host  plant  is  common  in  the  ground  layer  of  forest  in  the  Edmonton 
district.  It  is  one  of  the  few  herbs  whose  leaves  remain  green  through  the  winter  beneath 
the  snow  cover.  The  fly  seems  to  be  univoltine,  since  no  mines  have  been  found  before  late 
August.  Feeding  larvae  continued  to  be  found  up  to  September  27th,  the  last  pupating  in 
the  insectary  on  October  1st.  This  is  well  after  leaf  fall  and  the  onset  of  frost. 

I have  no  doubt  that  mitellae  and  tiarellae  are  monophyletic;  for  instance,  the  crescentic 
form  of  the  hind  spiracles  of  the  puparium  and  the  presence  on  the  male  aedeagus  of  a 
membranous  lobe  distal  to  the  supporting  sclerites,  are  both  synapomorphous  characters 
of  these  two  species. 


76 


Griffiths 


ACKNOWLEDGEMENTS 

I am  grateful  to  acknowledge  the  financial  support  of  the  Boreal  Institute  of  the  Univer- 
sity of  Alberta  for  my  field  work  in  Alaska,  the  Yukon  and  neighbouring  areas.  Material  of 
European  species  was  kindly  lent  by  K.  A.  Spencer  (London,  England)  and  H.  J.  Hannemann 
(Zoologisches  Museum,  Humboldt  University,  Berlin).  My  wife  Deirdre  prepared  the  illustra- 
tions of  leaf  mines  (Fig.  22-25). 


REFERENCES 

Griffiths,  G.  C.  D.  1967.  Revision  of  the  Phytomyza  syngenesiae  group  (Diptera,  Agromy- 
zidae),  including  species  hitherto  known  as  “ Phytomyza  atricornis  Meigen”.  Stuttg.  Beitr. 
Naturk.  no.  177.  28  pp. 

Griffiths,  G.  C.  D.  in  press.  Studies  on  the  phylogenetic  classification  of  Diptera  Cyclor- 
rhapha,  with  special  reference  to  the  structure  of  the  male  postabdomen.  Uitgeverij  Dr. 
W.  Junk,  The  Hague. 

Hardy,  J.  1849.  On  the  primrose-leaf  miner,  with  notice  of  a proposed  new  genus,  and 
characters  of  three  species  of  Diptera.  Ann.  Mag.  nat.  Hist.  4:385-392. 

Hendel,  F.  1928.  Blattminenkunde  Europas.  I.  Die  Dipterenminen  2.  Fritz  Wagner,  Vienna: 
65-100. 

Hendel,  F.  1931-1936.  Agromyzidae.  Fliegen  palaearkt.  Reg.  6(2),  Teil  59.  570  pp. 

Hering,  M.  1924.  Zur  Kenntnis  der  Blattminenfauna  des  Banats.  I.  Z.  wiss.  InsektBiol. 
19:1-15,  31-41. 

Hering,  M.  1927.  Agromyzidae.  Tierwelt  Dtl.  6.  172  pp. 

Hering,  M.  1932.  Minenstudien  11.  Z.  wiss.  InsektBiol.  26:157-182. 

Hering,  E.  M.  1957.  Bestimmungstabellen  der  Blattminen  von  Europa  einschliesslich  des 
Mittelmeerbeckens  und  der  Kanarischen  Inseln.  Uitgeverij  Dr.  W.  Junk,  The  Hague.  1 185 
+ 86  pp.  (3  vols.). 

Hulten,  E.  1968.  Flora  of  Alaska  and  neighbouring  territories.  Stanford  University  Press, 
Stanford,  California,  xxii  + 1008  pp. 

Linnaniemi,  W.  M.  1913.  Zur  Kenntnis  der  Blattminierer  speziell  derjenigen  Finnlands.  I. 
Acta  Soc.  Fauna  FI.  fenn.  37,  no.  4.  137  pp. 

Meijere,  J.  C.  H.  de.  1926.  Die  Larven  der  Agromyzinen  (Fortsetzung  und  Schluss).  Tijdschr. 
Ent.  69:227-317. 

Meijere,  J.  C.  H.  de.  1938.  Die  Larven  der  Agromyzinen.  Vierter  Nachtrag.  Tijdschr.  Ent. 
81:61-116. 

Meijere,  J.  C.  H.  de.  1941.  Die  Larven  der  Agromyzinen.  Sechster  Nachtrag.  Tijdschr.  Ent. 
84:13-30. 

Nowakowski,  J.  T.  1964.  Studien  iiber  Minierfliegen  (Dipt.  Agromyzidae).  9.  Revision  der 
Artengruppe  Agromyza  reptans  Fall. -A  rufipes  Meig.  Dt.  ent.  Z.  11:175-213. 

Sasakawa,  M.  1956.  A new  species  related  to  Phytomyza  saxifragae  Hering  (Diptera,  Agro- 
myzidae). Insecta  matsum.  19:105-108. 

Sasakawa,  M.  1961.  A study  of  the  Japanese  Agromyzidae  (Diptera).  Part  2.  Pacif.  Insects 
3:307-472. 

Spencer,  K.  A.  1969.  The  Agromyzidae  of  Canada  and  Alaska.  Mem.  ent.  Soc.  Can.  no.  64. 
3 1 1 pp. 

Tschirnhaus,  M.  von.  1969.  Zur  Kenntnis  der  Variability,  Eidonomie  und  Verwandtschaft 
bemerkenswerter  Agromyzidae  (Diptera).  Senckenberg.  biol.  50:143-157. 

Webb,  D.  A.  1964.  Saxifragaceae.  Flora  eur.  1:364-381. 


Boreal  Agromyzidae 


77 


Fig.  1.  Phytomyza  mitellae  n.  sp.,  head  in  left  lateral  view.  Fig.  2.  Phytomyza  deirdreae  n.  sp.,  head  in  left  lateral  view. 
Fig.  3.  Phytomyza  deirdreae  n.  sp.,  posterior  spiracles  of  puparium  in  left  lateral  view.  Fig.  4.  Phytomyza  aizoon  Hering, 
posterior  spiracles  of  puparium  in  caudal  view.  Fig.  5.  Phytomyza  saxifragae  Hering,  posterior  spiracles  of  puparium  in 
left  lateral  view.  Fig.  6.  Phytomyza  tiarellae  n.  sp.,  posterior  spiracles  of  puparium  (+  dorsal  view).  Fig.  7.  Phytomyza 
mitellae  n.  sp.,  posterior  spiracles  of  puparium  (±  dorsal  view). 


78 


Griffiths 


Fig.  8-10.  Phytomyza  deirdreae  n.  sp.,  holotype  <5:  8,  aedeagus  and  associated  structures  in  lateral  view  (AED  aedeagus, 
AEDAD  aedeagal  apodeme,  AEDH  aedeagal  hood,  PHPH  phallophore,  POG  postgonite,  SSC  supporting  sclerite); 
9,  aedeagus  in  ventral  view;  10,  ejaculatory  apodeme.  Fig.  11.  Phytomyza  saxifragae  Hering,  Vais  (Switzerland),  aedea- 
gus (d)  in  lateral  view.  Fig.  12.  Phytomyza  aizoon  Hering,  paratype  <5,  Mauthen  (Austria),  aedeagus  in  lateral  view. 
Fig.  13-14.  Phytomyza  mitellae  n.  sp.,  holotype  6:  13,  aedeagus  in  lateral  view;  14,  ejaculatory  apodeme.  Fig.  15-18. 
Phytomyza  tiarellae  n.  sp.,  holotype  <5:  15,  distal  section  of  aedeagus  in  dorsal  view;  16,  distal  section  of  aedeagus  in 
ventral  view;  17,  aedeagus  in  lateral  view;  18,  ejaculatory  apodeme. 


Boreal  Agromyzidae 


79 


Fig.  19.  Collection  sites  for  Phytomyza  deirdreae  n.  sp.  Fig.  20.  Collection  sites  for  Phytomyza  saxifragae  Hering. 
Fig.  21.  Collection  sites  for  Phytomyza  aizoon  Hering. 


80 


Griffiths 


Fig.  22.  Leaf-mine  of  Phytomyza  deirdreae  n.  sp.  on  Saxifraga  punctata  L.  Fig.  23.  Leaf-mine  of  Phytomyza  saxifragae 
Hering  on  Saxifraga  rotundifolia  L.  (after  Hering,  1927).  Fig.  24.  Leaf-mine  of  Phytomyza  tiarellae  n.  sp.  on  Tiarella 
trifoliata  L.  Fig.  25.  Leaf-mine  of  Phytomyza  mitellae  n.  sp.  on  Mitella  nuda  L. 


THE  PARASITOID  COMPLEX  OF  EUXOA  OCHROGASTER  (GUENEE) 
(LEPIDOPTERA:  NOCTUIDAE) 


A.  C.  SCHAAF 

Sugar  Research  Department 

Sugar  Manufacturers  ’ Association  Quaes tiones  en  tomologicae 

Mandeville  P.  0.,  Jamaica,  W.  1.  8 : 81-120  1972 

Twenty-seven  species  of  parasitoids  have  been  recorded  as  being  reared  from  Euxoa 
ochrogaster  (Guenee).  Three  of  these  records  are  not  valid  because  they  are  based  on  mis- 
identifications.  Descriptions  of  the  available  immature  stages  of  15  species  are  provided. 
Of  the  remaining  species,  four  had  been  previously  described,  two  could  not  be  studied 
because  of  taxonomic  difficulties,  and  no  material  was  available  of  three  more.  A brief 
discussion  of  the  host  specificity  and  the  role  of  the  parasitoids  in  regulating  E.  ochrogaster 
populations  is  given. 

Vingt-sept  especes  de  parasitoids  ont  ete  specifiees  comme  faisant  par  tie  de  /’Euxoa 
ochrogaster  (Guenee).  Cependant,  trois  de  ces  specifications  se  sont  relevees  fausses  a cause 
de  certaines  erreurs  d’identification.  Des  descriptions  de  15  especes  a Tetat  precoce  sont 
publiees.  Concernant  le  reste,  quatre  ont  ete  anterieurement  decrites,  deux  de  ces  especes 
n’ont  pas  pu  etre  observees  encore  de  fagon  specifique  a cause  de  difficultes  d’ordre  taxono- 
mique  et  les  trois autres  demeurent  inconnues  par  manque  de  materiels.  Une  breve  discussion 
sur  la  specificite  et  le  role  des  parasitoids  en  relation  avec  le  controle  des  groupes  d’E. 
ochrogaster  a ete  pourvue. 

The  red-backed  cutworm,  Euxoa  ochrogaster  (Guenee)  is  a well-known,  destructive 
ground  cutworm  native  to  the  prairie  provinces  of  Canada.  The  purpose  of  this  study  is  to 
provide  a method  of  identifying  some  of  the  immature  stages  of  the  parasitoids  of  the  red- 
backed  cutworm.  Some  groups  of  parasitoids  are  difficult  to  identify  to  species  as  adults 
but  may  be  identified  using  morphological  or  behavioural  characters  of  immature  stages. 
The  advantages  of  recognizing  the  immature  stages  of  the  parasitoids  of  any  host  species 
are:  1.  If  research  with  live  adult  parasitoids  is  necessary,  identification  using  the  remains  of 
the  immature  stages  prevents  damage  to  the  living  specimens.  2.  It  is  not  necessary  to  rear 
either  the  host  or  the  parasitoids  to  maturity  to  obtain  data  on  the  host-parasitoid  relations. 
This  allows  analysis  of  host  specimens  which  died,  and  of  hosts  killed  to  enlarge  a sample 
size  when  there  was  no  time  to  rear  all  the  hosts  or  parasitoids  to. maturity.  3.  Super-, 
hyper-,  or  multiple  parasitoid  attack  can  usually  be  recognized  only  by  the  dissection  of 
host  material  before  any  of  the  parasitoids  can  mature  and  emerge.  Analysis  of  inter-specific 
and  intra-specific  competition  of  parasitoids  as  well  as  the  interactions  with  predators  and 
disease  is  possible  from  the  results  of  such  dissections. 

The  term  parasitoid  is  used  in  this  paper  rather  than  parasite  which  is  usually  used  when 
referring  to  entomophagous  groups  which  attack  single  host  units.  The  need  for  the  term 
parasitoid  arises  from  the  fact  that  ecologically  the  action  of  such  entomophagous  species 
is  different  from  that  of  either  predators  or  true  parasites.  Doutt  (1964)  outlined  the  ways 
in  which  parasitoids  differ  from  parasites. 

PROCEDURES  AND  REVIEW 


Materials  and  methods 

This  research  was  started  during  the  summer  of  1967  in  conjunction  with  the  work  of 
Dr.  J.  H.  Frank  (1971a,  b)  on  the  predator  complex  attacking  E.  ochrogaster.  The  field 
research  was  carried  out  at  Calahoo,  Alberta  on  the  farm  of  Mr.  C.  Bergstreiser.  The  location 


82 


Schaaf 


of  the  field  site  was  South-west  8,  Township  55  - Range  27:  West  4th  Meridian.  This  site 
was  first  investigated  June  22,  1967  following  a report  of  a cutworm  outbreak  to  the 
Alberta  Department  of  Agriculture.  The  outbreak  was  limited  in  both  cutworm  numbers 
and  in  area  attacked  and  little  damage  was  caused.  Most  of  the  cutworms  were  in  the  final 
instar.  The  center  of  the  outbreak  was  used  as  a test  area  and  the  balance  of  the  field  was 
sprayed  with  insecticide.  During  the  spring  and  summer  of  1968,  plots  were  established  in 
an  oat  field  adjacent  to  the  barley  field  in  which  the  outbreak  had  occurred  the  year  before. 
The  oat  field  was  summer-fallowed  the  year  before.  Cutworms  were  collected  in  two  types 
of  pitfall  traps:  3.05  m x 0.12  m eavestroughs  and  plastic  containers  8.7  cm  diameter  at 
the  top  x 10.2  cm  depth,  by  hand  collecting,  and  by  systematic  sieving  of  quadrat  samples. 
The  numbers  of  cutworms  captured  were  higher  than  the  season  before  but  inadequate  for 
a meaningful  population  study  of  either  the  host  or  its  parasitoids.  All  live  hosts  were  reared 
in  both  1967  and  1968.  A small  number  of  cutworms  was  collected  in  1969  at  Calahoo 
using  3.05  m eavestroughs  as  pitfall  traps  and  preserved  for  further  study. 

Captured  cutworms  were  reared  in  the  laboratory  in  individual  plastic  petri  dishes.  Each 
dish  was  provided  with  moist  filter  paper  and  fresh  dandelion  ( Taraxacum  officinale  Weber) 
leaves  on  at  least  every  second  day.  The  dishes  were  changed  when  feces  had  badly  contam- 
inated the  filter  paper.  To  prevent  or  reduce  the  spread  of  disease  in  the  laboratory  new 
dishes  were  used  as  much  as  possible  or  the  old  dishes  were  sterilized  with  a KOH  solution. 
Forceps  used  to  handle  specimens  were  rinsed  in  95%  ethyl  alcohol  after  each  specimen  was 
handled.  Dishes  infested  with  mites  were  changed  but  mites  were  still  a serious  source  of 
mortality  in  laboratory  reared  cutworms.  Specimens  were  reared  in  a controlled  temperature 
cabinet  at  between  18  and  21  C.  All  specimens  which  were  found  dead  in  the  field  (usually 
drowned)  and  most  of  those  which  died  in  the  laboratory  were  preserved  in  70%  ethyl 
alcohol.  Some  of  the  hosts  which  showed  obvious  stages  of  parasitoid  attack  were  killed  and 
stored  in  alcohol  for  further  study.  All  hosts  which  failed  to  produce  either  adults  or 
parasitoids  were  checked  by  dissection  for  evidence  of  parasitoid  attack. 

The  basic  parasitoid  list  was  formed  using  three  sources  of  information.  A literature 
review  was  carried  out  to  establish  all  the  recorded  parasitoids.  The  best  sources  of  informa- 
tion were  King  and  Atkinson  (1928),  Thompson  (1945),  and  Graham  (1965).  Specimens  of 
adult  parasitoids  which  had  been  reared  in  my  study  were  compared  with  known  specimens 
or  were  identified  by  Dr.  Mason  or  Dr.  Peck  of  the  Entomology  Research  Institute,  Ottawa. 
The  final  source  of  information  was  host  labels  on  reared  specimens  in  the  Canadian  National 
Collection  (C.N.C.).  The  host  labels  aided  by  allowing  me  to  find  specimens  which  had 
originally  been  recorded  at  the  generic  level  only,  or  which  were  not  previously  recorded  at 
all. 

Specimens  of  most  of  the  species  studied  were  borrowed  from  the  C.N.C.  The  only 
bombyliid  personally  examined  was  Poecilanthrax  alcyon  (Say).  The  only  other  parasitoid 
species  which  I did  not  borrow  specimens  of  was  Copidodoma  bakeri  (Howard). 

The  methods  of  differentiating  species  and  the  preparation  of  the  specimens  for  study 
will  be  discussed  in  the  sections  dealing  with  each  family  of  parasitoids. 

Biology  of  E.  ochrogaster 

The  biology  of  the  red-backed  cutworm  has  been  discussed  by  King  (1926),  McMillan 
(1930),  and  Strickland  (1923).  Jacobson  (1970)  gives  details  of  the  laboratory  ecology. 
Hardwick  (1965)  reviews  the  taxonomy  and  the  geographical  range  of  E.  ochrogaster. 

One  useful  character  is  the  appearance  of  the  pupa  before  and  after  the  parasitoids 
emerge.  Fig.  60  shows  a typical  pupa  which  would  produce  an  E.  ochrogaster  adult.  Directly 
after  the  prepupa  has  molted  to  the  pupa,  the  cuticle  is  a pale  off-white.  The  cuticle  darkens 


Parasitoids  of  Euxoa 


83 


quickly  to  a light  brown  and  remains  this  color.  As  the  normal  pupa  develops,  it  gradually 
darkens  and  shortens  until  just  before  emergence,  when  it  is  very  black  and  the  surface  is 
distorted.  The  adult  emerges  through  the  dorsal  side  and  many  of  the  sutures  release, 
leaving  the  pupal  remains  badly  damaged.  The  abdominal  segments  of  the  pupa  are  often 
telescoped  anteriorly  at  emergence  (Fig.  61).  After  emergence,  the  pupa  returns  to  a light 
brown  color. 

Predators,  diseases,  and  non-insect  parasitoids 

King  and  Atkinson  (1928)  list  several  predators  of  E.  ochrogaster  immature  stages.  Frank 
(1971a)  studied  the  carabid  predators  of  E.  ochrogaster  extensively.  I found  that  the  lycosid 
spider,  Trichosa  terricola  Thorell,  killed  many  cutworms  in  pitfall  traps.  From  its  ability  to 
kill  even  the  largest  cutworms,  it  is  probably  an  important  predator  at  Calahoo. 

The  role  of  diseases  in  the  control  of  the  red-backed  cutworm  is  in  need  of  study.  King 
and  Atkinson  (1928)  carried  out  a preliminary  study  but  did  not  have  the  pathogen  identi- 
fied. I also  found  extensive  mortality  from  an  unidentified  disease  in  my  laboratory  colonies. 

The  only  non-insect  parasitoid  found  attacking  the  red-backed  cutworm  was  a single 
nematode.  It  was  reared  from  a fifth  instar  red-backed  cutworm  in  late  June,  1969  from 
material  collected  at  Calahoo.  This  specimen  was  examined  by  Dr.  H.  E.  Welch,  who  stated 
that  it  probably  belongs  to  the  genus  Agamermis  Cobb,  Steiner,  and  Cristie.  Positive  identi- 
fication was  not  possible  because  the  specimen  was  immature. 

TACHINIDAE 


The  morphology  of  immature  tachinids 

The  morphology  of  the  final  instar  larvae  and  puparia  of  tachinids,  as  well  as  of  other 
higher  Diptera,  is  poorly  understood.  A recent  work  by  Menees  (1962)  offers  an  explanation 
of  the  origins  of  the  cephalopharyngeal  structures  of  the  various  larval  instars.  In  his  work 
he  shows  that  the  mouth  hooks  are  chiefly  maxillary  in  origin,  and  that  beyond  the  first 
instar  there  is  no  evidence  of  vestigial  mandibular  structures.  Various  authors  describing 
these  structures  use  different  terminologies  which  assume  different  origins  of  the  structures 
(Zuska,  1963;  Sanjean,  1957).  Others  based  their  terminologies  on  convenient  names  (Fin- 
layson,  1960).  The  system  which  I use  is  outlined  in  Fig.  1-4  and  includes  arbitrary  terms 
not  based  on  any  morphological  assumptions. 

The  final  instar  tachinid  larva  has  12  segments,  but  the  puparium  has  only  11  due  to 
the  invagination  of  the  pseudocephalon  and  part  of  the  first  thoracic  segment  when  the 
larval  skin  becomes  the  puparium  (Zuska,  1963).  This  leaves  the  cephalopharyngeal  struc- 
tures lying  in  the  immediate  anterior  end  of  the  puparium,  attached  to  part  of  the  un- 
sclerotized  final  instar  larval  skin.  Horizontal  and  vertical  sutures  in  the  puparium  release 
when  the  adult  emerges.  The  flaps  which  are  formed  at  emergence  are  connected  at  their 
midpoints  to  the  rest  of  the  puparium.  The  dorsal  flap,  carrying  the  anteiror  spiracles,  often 
is  lost.  The  ventral  flap  is  less  often  lost  and  contains  the  cephalopharyngeal  structures. 

The  puparium  retains  many  of  the  characters  of  the  final  instar  larva.  One  of  these  charac- 
ters is  the  pattern  of  spinules.  Zuska  (1963)  states  that  these  patterns  may  vary  due  to 
different  hosts  and  other  factors,  and  that  thus  is  not  a good  taxonomic  character.  Colour  is 
often  used  as  a character  in  the  description  of  the  puparium  (Greene,  1921;  Strickland, 
1923),  but  as  pointed  out  by  Zuska  (1963)  and  from  my  observation,  the  variation  is  too 
great  for  it  to  be  of  much  use.  The  most  reliable  puparium  characters  to  work  with  are  the 
posterior  spiracles  (Fig.  3).  Unfortunately,  the  difference  between  closely  related  species  is 
not  always  sufficient.  While  all  of  the  species  in  this  study  had  three  orificia,  some  groups 


84 


Schaaf 


of  tachinids  have  four  or  more.  The  cicatrix,  remnants  of  the  second  instar  spiracle,  was 
evident  in  all  specimens  studied.  As  the  anterior  spiracles  are  often  lost,  they  are  not  a good 
character  to  base  general  classifications  on.  In  addition,  the  number  of  openings  or  pori 
varies  intraspecifically. 

The  cephalopharyngeal  structures  (Fig.  2)  found  inside  the  puparium  are  generally  good 
characters,  but  the  variability  of  some  of  the  parts  must  be  considered.  Basically,  the 
cephalopharyngeal  structures  are  formed  of  three  sclerites:  the  anterior,  median,  and  poste- 
rior. These  may  be  fused  to  their  adjacent  members  so  that  only  one  or  two  sclerites  are 
apparent  and  functional.  The  anterior  parts  appear  to  be  very  constant  whereas  the  posterior 
portion  may  vary  a great  deal  in  shape  or  degree  of  sclerotization.  As  the  posterior  sclerite 
is  the  least  sclerotized  portion  of  the  structure,  it  may  be  twisted  or  bent  in  such  a way  as 
to  obscure  its  true  appearance. 

Sclerites  in  addition  to  the  basic  three,  occur  in  some  species.  Sanjean  (1957)  offers 
names  for  three  such  sclerites  in  sarcophagid  larvae,  but  there  is  no  evidence  to  show  which 
are  present  in  my  specimens.  As  the  true  origin  of  these  sclerites  is  not  known,  I have  called 
them  auxiliary  sclerites.  While  dissecting  the  host,  the  cast  cephalopharyngeal  structures  of 
earlier  instars  may  be  found  but  not  often  enough  to  be  of  use  in  identifying  a species. 

Gonia  Meigen 

The  concept  of  the  genus  Gonia  has  been  reviewed  by  Tothill  (1924),  Morrison  (1940), 
and  Brooks  (1943).  Brooks  regarded  Gonia  as  a composite  of  several  genera  which  he 
separated  and  described.  His  work  separated  the  species  which  I am  considering  into  three 
genera:  Gonia  (capitata,  sequax);  Reaumuria  ( aldrichi ),  and  Fuscigonia  (fuscicollis).  Sabrosky 
and  Arrtaud  (1965)  restored  Gonia  to  its  original  concept,  which  will  be  used  in  this  paper. 
As  will  be  shown  in  the  discussion  of  the  species  of  this  genus,  more  work  is  needed  on  their 
taxonomy. 

The  characteristics  of  Gonia  puparia  are  as  follows.  The  puparia  are  robust,  larger  than 
9 mm  in  length,  and  are  patterned  or  completely  covered  with  spinules.  The  posterior 
spiracles  are  large,  protruding,  and  heavily  sclerotized  while  the  anterior  spiracles  are  diverse 
in  character.  The  anal  protuberance  is  small  and  insignificant.  The  cephalopharyngeal  struc- 
ture is  two-articled  with  the  anterior  and  median  sclerites  being  fused.  The  dorsal  anterior 
portion  of  the  posterior  sclerite  forms  an  arm  which  projects  forward  to  the  anterior 
sclerite.  A sclerotized  band  of  different  widths  surrounds  the  inner  angle  between  the  dorsal 
and  ventral  processes  of  the  posterior  sclerite.  The  entire  structure  of  the  cephalopharyngeal 
apparatus  has  a triangular  form  with  the  lines  of  the  anterior-median  and  posterior  sclerites 
being  nearly  straight. 

The  females  of  this  genus  typically  lay  their  eggs  on  vegetation  which  may  be  eaten  by 
host  larvae.  The  eggs  hatch  in  the  host  gut  and  the  larvae  penetrate  into  the  body  cavity. 
The  larvae  develop  to  the  second  instar  in  the  host  larva  and  complete  development  after 
the  host  pupates.  The  puparium  is  found  in  the  host  pupa.  Strickland  (1923)  provides  a 
detailed  account  of  the  life-cycle  of  a species  he  called  Gonia  capitata.  King  and  Atkinson 
(1928)  did  not  differentiate  between  three  species  of  Gonia  which  they  found  but  stated 
that  as  a group,  they  tended  to  select  plants  which  were  most  likely  to  be  eaten  by  host 
cutworms.  None  of  these  authors  believed  that  Gonia  species  would  ever  show  a high 
effective  rate  of  parasitism  in  E.  ochrogaster  populations  as  has  been  found  in  Agrotis 
orthogonia  Morrison  populations. 

When  the  cutworm  is  attacked  by  Gonia  sp.,  the  pupa  darkens  to  a deep  brown  because  of 
the  presence  of  the  puparium.  The  emerging  adult  causes  a transverse  break  across  the  head 


Parasitoids  of  Euxoa 


85 


of  the  host  pupa.  (Fig.  62).  The  break  usually  closes  and  the  host  pupa  remains  intact.  The 
abdominal  segments  remain  very  much  like  those  of  a normal  pupa.  In  some  specimens,  the 
host  pupa  is  expanded  around  the  puparium  and  is  slightly  collapsed  directly  behind  it. 

Gonia  aldrichi  Tothill 

King  and  Atkinson  (1928)  recorded  Gonia  aldrichi  as  reared  from  E.  ochrogaster.  They 
stated  that  aldrichi  is  the  most  important  parasitoid  of  E.  ochrogaster  in  the  genus  Gonia 
and  is  widely  distributed  in  Saskatchewan.  It  appears  that  at  least  two  species  are  currently 
included  in  the  concept  of  G.  aldrichi  and  both  have  been  reared  from  E.  ochrogaster.  These 
species  will  be  designated  here  as  G.  aldrichi  No.  1 and  G.  aldrichi  No.  2. 

Description  of  puparia.  — The  puparia  are  similar  in  both  of  these  species.  The  exit  hole 
from  the  host  pupa  is  a transverse,  irregular  break  across  the  head  of  the  pupa  (Fig.  62). 
After  the  adult  has  emerged,  the  break  is  usually  only  slightly  open  except  in  the  unusual 
cases  where  the  anterior  region  of  the  host  pupa  is  broken  off.  The  posterior  spiracles  (Fig. 
9,  10)  are  large  and  the  orificial  ridges  are  high  and  prominent.  The  orificial  ridges  occur  very 
close  to  the  edge  of  the  spiracular  plate  and  the  ventral  ridge  often  appears  continuous  with 
the  edge.  The  cicatrix  is  usually  poorly  developed  but  varies  in  size  and  prominence  from 
specimen  to  specimen.  Spinules  cover  most  of  the  puparium  in  indistinct  bands.  The  spinules 
occur  singly,  and  are  randomly  distributed  (Fig.  1 1).  The  anterior  spiracles  have  two  very 
different  shapes,  one  of  which  consists  of  two  or  three  pori  on  a distinct  pedicel  (Fig.  8), 
and  the  other  with  12  or  more  pori  (Fig.  7)  surrounding  and  partially  obscuring  the  pedicel. 
The  difference  in  anterior  spiracle  shape  could  not  be  correlated  to  other  characters.  Also, 
the  number  of  pori  vary  on  the  same  specimen  though  never  from  one  type  to  the  other. 

Description  of  larvae.  — The  main  difference  between  the  two  species  lies  in  the  shape  of 
the  cephalopharyngeal  structures.  The  entire  structure  of  G.  aldrichi  No.  1 (Fig.  5)  is  a 
wider  triangle  than  that  of  G.  aldrichi  No.  2 (Fig.  6).  The  angle  between  the  dorsal  and 
ventral  arms  of  the  posterior  sclerite  is  greater  in  No.  1 than  in  No.  2.  The  inner  angle  of 
No.  1 is  only  lightly,  if  at  all  sclerotized,  whereas  in  No.  2 a definite  band  of  up  to  one- 
quarter  the  width  of  the  dorsal  arm  extends  around  the  inner  angle  from  near  the  tip  of  the 
dorsal  arm  to  past  the  widened  area  of  the  lower  arm.  The  lower  arm  of  No.  1 lacks  any 
definite  widening  along  its  length.  The  fused  anterior  and  median  sclerite  of  No.  1 is  shorter 
than  that  of  No.  2.  The  blade  of  the  mouth  hooks  of  No.  2 has  a definite  S-curve  shape 
whereas  in  No.  1 the  curve  is  a simple  arc.  The  overall  sizes  of  No.  1 and  2 are  similar. 

The  second  instar  larvae  of  Gonia  sp.  (Fig.  12)  were  found  in  my  dissections  of  dead 
host  larvae.  As  the  puparia  and  cephalopharyngeal  structures  of  all  the  reared  specimens 
were  similar  to  the  two  G.  aldrichi  species,  it  is  safe  to  call  these  second  instar  larvae  G. 
aldrichi  also.  The  cephalopharyngeal  structures  of  these  larvae  are  fused  into  a single  sclerite 
(Fig.  13).  The  larva  is  5 to  6 mm  long,  curved  ventrally  and  patterned  by  black  papillae. 
The  patterns  of  papillae  were  not  constant. 

Biology.  — While  no  adults  were  obtained  from  reared  specimens  in  my  study,  several 
puparia  were  found  in  reared  host  pupae.  The  reared  Gonia  puparia  compared  favorably 
with  borrowed  specimens.  It  is  likely  that  faulty  rearing  conditions,  caused  the  failure  of 
adult  emergence.  During  the  dissection  of  dead  host  larvae,  five  specimens  were  found  to 
contain  from  one  to  five  second  instar  Gonia  larvae.  In  one  of  these  host  larvae,  two  of  the 
five  Gonia  larvae  were  damaged  and  partially  disintegrated,  while  in  another  only  the  cepha- 
lopharyngeal structures  of  a larva  were  found,  as  well  as  a healthy  Gonia  larva. 

Hosts.  — Specimens  of  G.  aldrichi  examined  were  reared  from  E.  ochrogaster  and  A. 
orthogonia. 


£J_ 


6 


0.1 


Fig.  1-4.  Immature  tachinids.  1.  posterior  view  of  puparium.  a,  b,  maximum  and  minimum  distance  between  posterior 
spiracles;  c,  orificium;  d,  spiracular  plate;  e,  supra-anal  protuberance;  f,  distance  of  anus  from  posterior  spiracles;  g,  anus. 
2.  cephalopharyngeal  structures  of  3rd  instar  larva,  h,  anterior  sclerite;  i,  dorsal  process  of  posterior  sclerite;  j,  median 
sclerite;  k,  posterior  sclerite;  1,  ventral  arm  of  posterior  sclerite.  3.  side  view  of  posterior  spiracle,  m,  orificial  ridge; 
n,  stigmatophore;  (from  Zuska,  1963).  4.  dorsal  view  of  puparium.  o,  p,  width  and  length  of  puparium.  Fig.  5-8. 
Gonia  aldrichi.  5.  cephalopharyngeal  structures  of  G.  aldrichi  No.  1.  6.  G.  aldrichi  No.  2.  7,  8.  anterior  spiracles.  Scales 
in  millimeters. 


Parasitoids  of  Euxoa 


87 


Gonia  capitata  (De  Geer) 

Gonia  capitata  was  recorded  as  being  reared  from  E.  ochrogaster  by  Strickland  (1923). 
Strickland  noted  that  this  could  be  a mistaken  identification  and  that  the  species  studied 
could  be  divided  into  five  groups.  It  is  now  recognized  that  G.  capitata  is  exclusively  a 
European  species  (Brooks,  1943;  Sabrosky  and  Arnaud,  1965).  This  invalidates  the  figures 
and  descriptions  of  Greene  (1921).  The  biological  work  of  Strickland  must  now  be  regarded 
as  being  of  Gonia  sp.  but  is  nevertheless  a valuable  source  of  information. 

Gonia  fuscicollis  Tothill 

Gonia  fuscicollis  was  recorded  as  being  reared  from  E.  ochrogaster  by  King  and  Atkinson 
(1928).  Brooks  (1943)  regarded  this  species  as  being  so  different  from  the  other  species  of 
Gonia  that  he  created  the  genus  Fuscigonia  for  it.  Unfortunately,  no  specimens  were  availa- 
ble for  study. 

Nothing  is  known  of  the  biology  of  this  species  and  it  is  likely  that  it  is  not  an  important 
parasitoid  of  any  of  the  economic  cutworms.  The  description  of  the  immature  stages  is 
necessary  in  the  future,  however,  to  help  separate  the  large  number  of  species  in  this  genus. 

Gonia  sequax  Williston 

Gonia  sequax  has  not  been  recorded  as  being  reared  from  E.  ochrogaster . I am  including 
G.  sequax  in  this  study  as  I am  sure  that  it  is  a potential,  if  not  actual  parasitoid  of  E. 
ochrogaster,  as  well  as  to  represent  the  capitata  species  group  (Brooks,  1943). 

Description  of  puparia.  — The  puparium  of  G.  sequax  is  very  similar  to  that  of  G.  aldrichi 
in  size  and  shape.  The  posterior  spiracles  (Fig.  15)  differ  in  that  the  orificial  ridges  are  set 
further  in  from  the  edge  of  the  spiracular  plate  than  those  of  G.  aldrichi.  The  orificial  ridges 
enclose  the  cicatrix  which  is  fairly  prominent.  The  anterior  spiracles  (Fig.  16)  have  only  one 
porus  on  the  end  of  a long  pedicel.  Only  one  specimen  was  examined  so  that  this  may  not 
be  characteristic  of  all  members  of  this  species.  The  spination  of  the  puparia  differs  sharply 
from  that  of  G.  aldrichi.  The  spines  (Fig.  17)  tend  to  be  in  groups  or  series  of  three  or  more 
and  often  form  long,  irregular  rows. 

Description  of  larvae.  — The  cephalopharyngeal  structures  (Fig.  14)  are  similar  to  both 
G.  aldrichi  No.  1 and  No.  2 but  differ  from  both  enough  to  be  separated.  The  blade  of  the 
mouth  hooks  is  a simple  arc  as  in  G.  aldrichi  No.  1 but  the  overall  shape  is  closer  to  that  of 
No.  2.  Distinct  from  both  G.  aldrichi  No.  1 and  No.  2 is  the  anterior  projection  of  the  dorsal 
arm  of  the  posterior  sclerite.  It  is  long,  distinct,  and  well  developed,  and  usually  has  a large 
open  space  between  it  and  the  anterior-median  sclerite. 

Biology.  — Little  is  known  of  the  biology  of  this  species  and  how  it  differs  from  other 
members  of  the  genus. 

Hosts.  — This  species  has  been  reared  from  Agrotis  orthogonia  from  Alberta  and  Sas- 
katchewan. 


Bonnetia  comta  (Fallen) 

The  puparia  and  posterior  spiracles  of  Bonnetia  comta  were  described  by  Greene  (1921) 
and  all  the  life  stages  including  the  above  structures  were  described  by  Strickland  (1923). 
Allen  (1926)  described  the  first  instar  larvae.  Greene’s  drawings  either  are  not  clear  enough 
or  are  of  a different  species  than  what  is  now  called  B.  comta. 


88 


Schaaf 


Fig.  9-13.  Gonia  aldrichi.  9.  posterior  spiracle.  10.  posterior  view  of  puparium.  11.  spine  pattern  of  puparium.  12. 
second  instar  larva.  13.  cephalopharyngeal  structures  of  second  instar  larva.  Fig.  14-17.  Gonia  sequax.  14.  cephalo- 
pharyngeal  structures.  15.  posterior  spiracle.  16.  anterior  spiracle  (side  view).  17.  spine  pattern  of  puparium.  Scale  in 
millimeters. 


Parasitoids  of  Euxoa 


89 


Description  of  puparia.  - The  puparia  are  large,  sub-elliptical,  smooth-surfaced  without 
any  trace  of  spination,  and  show  only  vague  segmentation  marks  (Fig.  22).  The  anterior 
stigmata  (Fig.  20)  are  raised  and  show  either  five  or  six  pori  which  are  arranged  in  a 
curvilinear  pattern  with  the  axes  of  the  individual  porus  pointing  towards  the  center  of  the 
curve.  Both  the  stigmatal  plates  and  the  supraanal  protuberance  are  widely  separated  from 
the  true  anus  and  occur  high  on  the  dorsal  surface  of  the  puparium  (Fig.  21).  The  stigmatal 
plates  of  the  posterior  spiracles  (Fig.  19)  are  low  and  flat  with  the  orificial  ridges  being  only 
slightly  raised  but  quite  wide.  The  cicatrix  is  also  low  but  is  large  and  distinct.  Occasionally, 
especially  with  transmitted  light,  a weakly  sclerotized  region  can  be  seen  between  the  inner 
and  medial  orificial  ridges.  The  supraanal  protuberance  is  pronounced  and  in  some  speci- 
mens is  higher  than  are  the  stigmatal  plates.  From  it  runs  a distinct  ridge  which  separates 
the  stigmatal  plates. 

Description  of  larvae.  - Strickland  (1923)  describes  the  life  stages  and  gives  figures  for 
them.  The  following  passage  is  taken  from  his  paper  to  describe  the  final  instar  larvae,  which 
I have  not  examined. 

“The  smallest  specimen  seen  measures  12  mm.  long  and  4 mm.  wide;  the  largest,  which 
was  almost  mature,  was  15.75  mm.  by  4.25  mm.  While  living,  the  larvae  that  were  dis- 
sected from  their  host  constantly  changed  their  shape  by  violent  muscular  contractions. 
The  transparent  cuticle  revealed  the  yellow  and  brown  viscera  in  strong  contrast  to  the 
voluminous  white  fat-body.  When  killed  . . . the  larva  is  arcuate,  dorsum  concave,  tapering 
cephalad  and  slightly  so  to  the  bluntly  rounded  caudal  extremity.  Eleven  segments,  only, 
were  seen.  Dorso-laterally  between  segments  I and  II  there  are  a pair  of  blackened 
spiracles  . . . each  of  which  possesses  six  respiratory  papillae  that  open  into  a short  con- 
stricted felt-chamber,  behind  which  are  a pair  of  stout  trachea  which  run  the  length  of  the 
body  and  connect  with  the  caudal  spiracles.  The  cuticle  is  almost  destitute  of  armature, 
though  minute  simple  spines  are  present  on  all  of  the  intersegmental  areas.  These  are 
most  numerous  in  the  anterior  and  posterior  segments  where  they  form  a fine  network 
of  rows  that  encircle  the  body.  In  addition  there  are  traces  of  intersegmental  hooks  be- 
tween the  four  anterior  segments.  The  buccal-pharyngeal  armature  . . . differs  little  from 
that  of  the  preceding  stage  except  in  size.  The  over-all  measurement  is  1.0  mm.  to  1.1 
mm.,  the  mandibular  hooks  being  0.17  mm.  to  0.18  mm.  long.” 

Strickland’s  drawings  of  the  cephalopharyngeal  structures  provide  the  appropriate  general 
impressions  but  are  inaccurate  in  one  aspect.  Ventral  to  the  anterior  sclerite  lies  an  auxiliary 
sclerite  as  shown  in  Fig.  18.  This  sclerite  is  always  shown  as  being  solidly  fused  to  the  ante- 
rior sclerite  in  Strickland’s  paper.  I have  examined  12  borrowed  specimens  and  three  slides 
which  were  part  of  Strickland’s  study  and  my  interpretation  is  as  follows:  The  posterior 
sclerite  is  very  slightly  sclerotized  and  while  it  has  a characteristic  shape,  it  is  often  twisted 
or  bent  in  the  puparium.  The  anterior  process  of  this  structure  becomes  progressively  less 
sclerotized  until  it  terminates  in  a ligament-like  structure  which  connects  with  the  posterior 
process  of  the  anterior  sclerite.  The  articulation  between  the  median  and  posterior  sclerite 
is  very  weak  and  usually  releases  when  the  entire  structure  is  being  removed  from  the  pu- 
parium. In  contrast,  the  anterior  articulation  is  very  solid  and  it  is  often  difficult  to  find  the 
division  between  the  two  sclerites  except  at  the  tip  of  the  posterior  process  of  the  anterior 
sclerite. 

Biology.  — The  life  history  and  behavior  of  B.  comta  is  well  outlined  by  Strickland 
(1923).  This  species  belongs  to  the  tachinids  which  either  larviposit  or  oviposit  in  an  area 
where  the  first  instar  larvae  can  actively  attach  themselves  to  a host  and  enter  from  the  out- 
side of  the  body.  Strickland  found  that  the  cutworms  were  attacked  in  the  third  or  fourth 
instar  and  death  resulted  during  the  prepupal  stage  when  the  mature  larva  emerged  to  enter 


90 


Schaaf 


the  soil  and  pupate. 

Hosts.  — Two  generations  occur  per  year  with  the  larvae  of  the  second  generation  over- 
wintering in  their  hosts.  The  hosts  recorded  for  this  species  in  Strickland’s  (1923)  paper  are 
as  follows:  Euxoa  tristicula  Morrison,  Montana;  E.  messoria  Harris,  Washington;  Agrotis 
ypsilon  L.,  California;  A.  orthogonia  Morrison,  Alberta;  Copablepharon  viridisparsa  Dod., 
Alberta;  Polia  acutermina  Sm.,  Alberta  (induced  parasitism).  King  and  Atkinson  (1928)  re- 
corded B.  comta  as  reared  from  E.  ochrogaster  but  found  it  to  be  an  insignificant  parasite 
during  their  studies  in  Saskatchewan. 


Periscepsia  Gistel 

This  genus  has  been  handled  in  several  different  ways  by  various  authors.  Sabrosky  and 
Arnaud  (1965)  list  six  different  synonyms  of  Periscepsia.  The  confusion  with  regard  to  P. 
helymus  and  P.  laevigata  lies  in  the  fact  that  the  former  has  been  placed  in  seven  different 
name  combinations  and  the  latter  in  eight.  In  addition,  many  authors  confused  P.  laevigata 
with  P.  helymus  and  named  it  as  the  latter.  The  major  generic  names  in  which  both  species 
have  been  placed  and  which  have  been  used  in  the  literature  are  Phorichaeta,  Metachaeta, 
and  Wagneria.  P.  helymus  has  also  had  the  specific  names  helyma  and  sequax,  while  P. 
laevigata  has  been  known  as  helymus,  atra,  and  carbonaria.  The  character  recognized  by 
Reinhard  (1955)  and  Wood  (pers.  comm.)  to  separate  these  species  is  the  presence  of  setae 
on  the  median  portion  of  the  first  wing  vein  in  P.  laevigata  as  opposed  to  a bare  first  wing 
vein  in  P.  helymus.  Because  of  the  confusion  which  has  existed  about  these  two  species,  one 
must  regard  the  host  lists  from  the  literature  with  caution. 

Periscepsia  helymus  (Walker) 

The  puparia  of  P.  helymus  were  described  by  Greene  (1921)  under  two  synonymous 
names,  Phorichaeta  sequax  (Williston)  and  Metachaeta  helymus,  but  the  puparia  were  differ- 
ent from  each  other. 

Description  of  puparia.  — Greene’s  (1921)  descriptions  and  figures  of  the  puparia  under 
the  name  Metachaeta  helymus  closely  resemble  P.  helymus , while  those  of  Phorichaeta  se- 
quax differ  sufficiently  from  the  other  two  to  be  considered  different.  Also,  the  puparia 
which  I measured  were  significantly  larger  (4.5  x 2.1  mm)  than  the  dimensions  given  by 
Greene  for  Phorichaeta  sequax  (3.5  x 1.5  mm).  Even  the  ranges  of  the  specimens  I studied 
(4.08  mm  - 4.88  mm  x 1.92  mm  - 2.16  mm)  did  not  include  his  dimensions.  M.  helymus 
dimensions  were  4.75  mm  x 1.75  mm  and  hence  are  similar  to  the  measurements  taken 
from  my  study  series. 

From  a lateral  view  of  the  puparium,  the  posterior  regions  carrying  the  stigmatal  plates 
and  anus  appear  extended  (Fig.  24).  From  a posterior  view  the  puparia  appear  nearly  circu- 
lar with  a raised  central  region  (Fig.  25).  The  spiracular  plates  are  terminal  in  position.  The 
spiracular  plates  of  the  posterior  spiracles  (Fig.  26)  are  narrowly  separated,  only  slightly 
raised  and  are  quite  flat.  The  cicatrix  is  large,  round,  and  slightly  concave  in  the  central 
region.  The  entire  shape  of  the  spiracular  plate  varies  from  oblong  to  slightly  curved.  The 
anterior  spiracles  are  especially  poor  characters  in  this  group  because  they  are  often  lost  at 
emergence.  The  stigmata  are  small  and  tuberculate  with  a cluster  of  pori  in  a crescent  (Fig. 
28).  The  number  of  pori  is  not  constant. 

Description  of  larvae.  - No  specimens  of  the  final  instar  larvae  were  available  for  study, 
therefore  I am  relying  on  the  cephalopharyngeal  structures  recovered  from  the  puparia  for 
characters.  The  cephalopharyngeal  structure  (Fig.  23)  is  very  distinctive  but  the  variations 


Parasitoids  of  Euxoa 


91 


which  occur  tend  to  confuse  its  appearance.  The  anterior  sclerite  is  heavily  sclerotized  with 
the  ventral  process  being  almost  as  long  as  the  actual  mouth  hooks.  Behind  the  ventral 
process  lies  an  auxiliary  sclerite  which  may  be  concealed  or  lie  in  several  different  positions 
in  the  same  general  region.  The  median  and  posterior  sclerites  are  fused  but  with  adequate 
lighting,  the  suture  can  be  seen.  Projecting  dorsally  from  the  median  sclerite  is  a hook-like 
process  which  curves  anteriorly.  The  posterior  sclerite  varies  considerably  in  the  degree  of 
sclerotization  from  a dark  colour  which  nearly  obscures  the  hook,  to  a very  weakly  sclero- 
tized, clear  structure.  In  the  latter  case,  many  structural  differences  appear  but  are  probably 
insignificant.  In  this  species  the  anterior  process  of  the  posterior  sclerite  is  very  weak  or 
lacking.  In  some  specimens  a membranous  connection  may  be  seen  between  the  anterior 
and  posterior  sclerites  in  the  dorsal  regions. 

Biology.  - Little  is  known  of  the  biology  of  this  species  except  that  it  attacks  cutworm 
larvae  and  that  more  than  one  may  emerge  from  a single  host  (Reinhard,  1955).  Guppy 
(1967)  records  three  or  fourP.  helymus  as  being  reared  from  a single  sixth  instar  P.  unipunc- 
ta  Haworth  larva. 

Hosts.  - In  the  literature,  P.  helymus  has  been  recorded  as  reared  from  the  following 
hosts:  Heliophila  commoides  Guenee  (Tothill,  1913),  Ontario;  Pseudaletia  unipuncta  Ha- 
worth (Baker,  1914),  Ontario;  Euxoa  ochrogaster  (King  and  Atkinson,  1928),  Saskatchewan. 
Reinhard  (1955)  lists  the  following  hosts  not  recorded  above:  black  army  cutworm,  Actebia 
fennica  Tausch,  Michigan;  Peridroma  saucia  Hiibner,  California;  Cirphis  sp.  Hampson,  Wash- 
ington; Euxoa  auxiliaris,  Alberta;  Polia  adjuncta;  Grapholitha  sp.  Conistra  devia  Grote; 
Lithophane  innominata  Smith,  Maine.  Specimens  which  I examined  were  reared  from:  H. 
commoides,  Ontario;  A eliminata  Gn.,  New  Brunswick;  R hynchagrotis  cupida  Grote,  New 
Brunswick;  Andropolia  vancouvera  Strand,  British  Columbia;  Andropolia  sp.  Grote,  Alberta; 
A.  contacta  Walker,  Alberta. 

Periscepsia  laevigata  (Van  der  Wulp) 

There  appears  to  be  no  previous  description  of  the  immature  stages  of  this  species  in  the 
literature. 

Description  of  puparium.  — Only  the  puparium  of  one  specimen  was  studied  and  it  lacked 
the  anterior  flaps  so  that  neither  the  cephalopharyngeal  structures  nor  the  anterior  spiracles 
were  available  for  study.  The  posterior  spiracles  are  similar  to  those  of  P.  helymus  but  the 
orificial  ridges  tend  to  be  higher,  wider,  and  more  rounded  (Fig.  27).  The  orificia  follow  the 
top  of  the  ridges  almost  to  the  level  of  the  spiracular  plates.  While  the  cicatrix  is  large  and 
round  it  is  not  as  distinct  as  that  of  P.  helymus.  The  best  differentiating  character  is  the 
median  orificium  which  is  very  curved  in  P.  laevigata  but  nearly  straight  in  P.  helymus.  The 
rounded  shape  of  the  orificial  ridge  of  P.  laevigata  contrasts  well  with  the  long,  narrow  shape 
ini5,  helymus. 

Biology.  - Little  is  known  of  the  biology  of  this  species  except  that  it  attacks  fifth  instar 
cutworm  larvae  (Guppy,  1967).  The  range  of  the  species  is  from  Guatemala  to  Canada 
(Reinhard,  1955). 

Hosts.  — The  problem  of  host  records  is  important  in  regards  to  this  species.  As  one 
cannot  be  sure  of  the  accuracy  of  earlier  identifications,  it  is  possible  that  the  following 
list  is  either  incomplete  or  inaccurate.  Reinhard  (1955)  lists  the  following  hosts:  Pseudaletia 
unipuncta,  Euxoa  auxiliaris,  Grapholitha  sp.  Hiibner;  Lascoria  ambigualis  Walker,  Elaphria 
nucicolora  Grote,  and  unidentified  cutworms.  Guppy  (1967)  records  P.  unipuncta  from 
Ontario  as  a host.  The  specimen  I examined  was  reared  from  an  unidentified  ‘phalaenid’ 
from  Big  Beaver,  British  Columbia. 


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Fig.  18-22.  Bonnetia  comta.  18.  cephalopharyngeal  structures.  19.  posterior  spiracle.  20.  anterior  spiracle.  21.  posterior 
view  of  puparium.  22.  puparium.  Fig.  23-26,  28.  Periscepsia  helymus.  23.  cephalopharyngeal  structures.  24.  puparium. 
25.  posterior  view  of  puparium.  26.  posterior  spiracle.  28.  anterior  spiracle.  Fig.  27.  Posterior  spiracle  of  P.  laevigata. 
Scale  in  millimeters. 


Parasitoids  of  Euxoa 


93 


BOMBYLIIDAE 

Biology  and  morphology  of  immature  bombyliids 

Five  species  in  two  genera  of  Bombyliidae  have  been  recorded  as  primary  parasitoids,  and 
one  other  species  as  a hyperparasitoid  of  Euxoa  ochrogaster . Brooks  (1952)  illustrated  the 
final  instar  larvae  and  the  pupae  of  five  of  these  six  species  and  provided  keys  for  their 
identification.  Painter  and  Hall  (1960)  provide  keys  for  adults  to  the  genera  of  Bombyliidae, 
to  the  subgenera  of  Villa,  and  to  the  known  species  of  Poecilanthrax . Generally,  little  is 
known  of  the  biology  of  bombyliids.  No  comprehensive  work  has  been  done  on  the  biology 
of  any  species  attacking  Lepidoptera.  The  terminology  used  for  the  morphology  of  the 
immature  stages  is  based  on  Berg  (1940)  in  his  work  on  the  immature  stages  of  Systoechus 
vulgaris  Loew. 

While  Bohart  et  al.  (1960)  suggest  that  there  may  be  four  larval  instars,  other  authors 
have  only  described  three  (Hynes,  1947;  Berg,  1 940;  Clausen,  1928).  The  first  instar  larva  is 
active,  vermiform,  and  adapted  to  move  through  the  soil  and  seek  out  its  host  (Clausen, 
1928;  Bohart  et  al.,  1960;  Berg,  1940).  In  parasitoid  species  the  first  instar  larvae  penetrate 
their  host  and  develop  internally.  The  second  instar  larva  is  far  less  mobile  than  the  first  and 
is  more  maggot-like  (Clausen,  1940;  Bohart  et  al.,  1960).  It  has  lost  its  adaptations  for 
moving  through  the  soil  and  likely  can  only  move  in  open  areas.  In  species  which  are  internal 
parasitoids,  the  larva  likely  molts  immediately  after  attacking  the  host  due  to  the  radically 
different  environment  in  which  it  then  lives.  Because  the  cast  skin  of  the  final  instar  larva 
can  be  found  in  the  pupal  case  of  the  host,  descriptions  of  this  stage  are  available  for  several 
species.  Brooks  (1952)  reviews  the  taxonomically  important  characters  of  both  the  final 
instar  larvae  and  the  pupae  of  noctuid-attacking  bombyliids.  In  the  final  instar  larvae  he 
found  the  structure  of  the  mouth  parts  and  the  head  sclerites  to  be  important.  In  the  pupae 
he  found  that  the  head  tubercles  and  mouth  part  sizes  were  of  specific  use,  whereas  the 
apical  segments  were  of  little  use  due  to  the  general  uniformity  in  the  group  and  the  varia- 
bility within  the  species  involved. 

Bombyliids  have  been  observed  apparently  ovipositing  in  loose  sand  or  dust,  but  this  has 
not  been  confirmed  (Painter  and  Hall,  1960).  Brooks  (1952)  noted  that  the  species  attacking 
noctuids  fly  from  the  latter  half  of  July  through  to  September,  but  the  hosts  are  not 
attacked  until  their  fourth,  fifth  or  sixth  instars  the  next  season.  He  felt  that  the  species 
overwinters  as  eggs  in  the  soil  or  old  vegetation.  Painter  and  Hall  feel  that  the  first  instar 
larvae  seek  out  small  caterpillars  as  hosts  and  remain  inactive  until  the  host  pupates,  when 
they  rapidly  develop  and  kill  the  host.  The  bombyliid  pupa  breaks  out  of  the  host  pupal 
remains,  and  with  the  use  of  directed  spines  and  bristles  moves  to  the  soil  surface  where  the 
adult  emerges  quickly  (Painter  and  Hall,  1960;  Allen,  1921;  Snow,  1925).  It  is  likely  that 
the  abrasion  from  moving  through  the  soil  accounts  for  much  of  the  intraspecific  dif- 
ferences which  have  been  observed  in  the  shape  of  the  head  tubercles,  apical  segments,  and 
spines. 

Other  than  Allen’s  (1921)  report  that  Poecilanthrax  lucifer  (Fabr.)  attacked  25%  of  the 
fall  army  worm,  Spodoptera  frugiperda  (Smith)  in  Mississippi,  bombyliids  have  been  re- 
garded as  minor  parasitoids  of  noctuids  (Brooks,  1952;  King  and  Atkinson,  1928). 

Villa  Lioy 

This  genus  has  a wide  host  range  as  internal  or  external  parasites  of  Diptera,  Lepidoptera, 
Hymenoptera,  and  Coleoptera.  Painter  and  Hall  (1960)  provide  a key  to  the  adult  sub- 
genera. Two  of  these,  Villa  and  Hemipenthes,  are  of  interest  to  us. 


94 


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Villa  (Villa)  alternata  (Say) 

This  species  was  recorded  by  Brooks  (1952)  as  a parasite  of  E.  ochrogaster  which  was 
collected  as  fourth,  fifth,  and  sixth  instar  larvae  in  May  and  June.  Records  show  this  species 
to  range  across  the  prairies  of  Canada.  Brooks  provides  illustrations  of  the  mouth  hooks, 
head  capsule,  and  apical  platelets  of  the  mature  larvae  as  well  as  of  the  entire  pupa,  its  head, 
and  apical  structures. 

Hosts.  — Euxoa  flavicollis  Sm.,  E.  tesselata  Harris,  Agrotis  orthogonia,  Feltia  ducens 
Walker,  E.  ochrogaster,  (Brooks,  1952);  tenebrionid  larvae  (Clausen,  1940). 

Villa  (Villa)  fulviana  (Say) 

This  species  was  recorded  by  King  and  Atkinson  (1928)  as  reared  from  E.  ochrogaster, 
but  only  from  Euxoa  sp.  Hiibner  by  Brooks  (1952).  It  has  been  collected  in  host  pupae  from 
June  30  to  July  12  and  found  to  emerge  as  adults  in  the  autumn.  King  and  Atkinson  felt 
that  it  overwinters  in  alternative  hosts.  Illustrations  of  the  larvae  and  pupae  are  provided  by 
Brooks. 

Hosts.  - E.  ochrogaster  (King  and  Atkinson,  1928),  Euxoa  sp.  Hiibner  (Brooks,  1952). 

Villa  (Villa)  lateralis  (Say) 

King  and  Atkinson  (1928)  record  this  species  as  having  attacked  two  specimens  of  E. 
ochrogaster  and  emerging  in  the  autumn.  No  reference  is  made  to  this  species  by  Brooks 
(1952).  No  drawings  of  any  of  the  life  stages  are  known  and  no  material  appears  to  be 
available  at  the  present  time. 

Villa  (Hemipenthes)  moroides  (Say) 

Species  of  the  subgenus  Hemipenthes  are  hyperparasitic  upon  the  primary  parasites  of 
Lepidoptera  (Clausen,  1940).  Clausen  refers  to  members  of  this  subgenus  as  attacking 
ichneumonid  parasitoids,  but  Brooks  (1952)  records  three  species  as  reared  from  tachinid 
hosts.  V.  moroides  was  reared  from  Gonia  spp.  and  Bonnetia  comta , which  were  reared  from 
noctuid  hosts.  Clausen  stated  that  the  exact  relationship  between  the  host,  parasitoid,  and 
hyperparasitoid  was  not  established  and  that  it  was  possible  that  the  ichneumonids  were 
attacked  in  the  pupal  stage  independently  of  the  host.  It  appears  that  a true  hyperparasitoid 
role  has  been  established  for  V.  moroides. 

Poecilanthrax  Osten  Sacken 

This  is  a widespread  genus  which  attacks  chiefly  noctuid  larvae.  Painter  and  Hall  (1960) 
list  1 5 species  of  cutworms  and  army  worms  which  are  attacked  by  eight  species  of  Poeci- 
lanthrax. 


Poecilanthrax  alcyon  (Say) 

This  species  was  recorded  as  a parasitoid  of  E.  ochrogaster  by  King  and  Atkinson  (1928) 
under  the  name  of  halcyon.  Painter  and  Hall  (1960)  give  the  distribution  of  the  species  in 
southern  Canada  and  the  United  States,  but  fail  to  show  the  correct  boundaries  of  northern 
distribution.  The  range  extends  from  Texas  to  the  Northwest  Territories,  and  with  the  ex- 


Parasitoids  of  Euxoa 


95 


ception  of  Southern  California,  east  of  the  Rocky  Mountains  to  the  Atlantic  Ocean. 

Brooks  (1952)  illustrated  the  final  instar  larva  (Fig.  32)  and  pupa.  Painter  and  Hall  (1960) 
discuss  the  entire  species  at  length,  as  P.  alcyon  is  the  type  of  the  genus,  and  give  a detailed 
description  of  the  adult.  Fig.  29,  30,  31  and  33  are  original  drawings  of  a specimen  of 
P.  alcyon  reared  from  E.  ochrogaster . 

Only  one  specimen  of  a pupa  attacked  by  Poecilanthrax  alcyon  (Fig.  63)  was  examined. 
This  specimen  was  a light  brown  similar  to  one  from  which  a moth  had  emerged.  The  adult 
P.  alcyon  emerges  from  the  dorsal  surface  behind  the  head,  leaving  the  pupa  intact.  The 
abdominal  segments  are  fully  extended  after  emergence. 

Hosts.  - E.  ochrogaster,  E.  flavicollis,  Chorizagrotis  thanatologia,  Pseudaletia  unipuncta, 
(Brooks,  1 95 2) ; peridroma  margaritosa  (Walkden,  1950). 

Poecilanthrax  mllistonii  (Coquillet) 

The  adults  of  this  species  were  reported  by  King  and  Atkinson  (1928)  to  emerge  either  in 
the  autumn  or  the  following  June.  They  felt  that  the  species  overwinters  as  larvae  in  noctuid 
hosts  which  overwinter  in  the  larval  stage.  Painter  and  Hall  (1960)  show  the  distribution  as 
from  south  of  the  United  States  to  the  middle  of  the  prairie  provinces  of  Canada,  and  from 
the  west  coast  through  to  the  central  great  plains.  Figures  of  the  larvae  and  pupae  are  given 
by  Brooks  (1952)  and  those  of  the  pupae  are  reprinted  by  Painter  and  Hall  (1960). 

Hosts.  - Agroperina  dubitans  Walker,  Chorizagrotis  thanatologia  Dyar,  S.  devestator, 
Euxoa  flavicollis,  E.  ochrogaster,  E.  tessellata,  Feltia  ducens,  (Brooks,  19 52);  Chorizagrotis 
auxiliaris  Grote,  Euxoa  scandens  Riley,  (Walkden,  1950). 

ICHNEUMONIDAE 
Morphology  of  immature  ichneumonids 

The  ichneumonid  parasitoids  have  many  taxonomic  characters  which  are  useful  to  sepa- 
rate their  immature  stages.  The  terminology  and  approach  to  description  follows  that  of 
Finlayson  (1960).  The  cocoons,  if  present,  can  be  separated  using:  size,  color  and  shape, 
and  location  of  the  adult  emergence  hole  from  the  cocoon.  The  spiracles  were  of  limited 
use  when  studying  these  species  as  the  differences  I found  were  insignificant.  The  best  sepa- 
rating characters  for  the  final  instar  larvae  were  found  in  the  cephalic  structures  (Fig.  34). 
To  differentiate  between  species,  the  presence  or  absence  of  sclerites,  and  the  shape  of  man- 
dibles were  the  best  characters  found. 

Preparation  of  specimens.  — After  having  been  soaked  in  water  for  at  least  24  hours,  the 
final  instar  larval  skins  were  removed  from  the  cocoons  by  means  of  fine  forceps  or  a 
hooked  pin.  The  larval  skin  was  gently  unfolded  and  removed  from  the  adult  meconium 
which  covered  many  of  the  specimens.  The  skin  was  then  placed  in  10%  KOH  for  24  hours 
at  room  temperature,  or  longer  if  it  was  not  cleared  enough.  Boiling  in  KOH  was  found  to 
disarticulate  the  cephalic  structures  resulting  in  the  loss  of  sclerites,  and  so  was  not  used. 
The  cleared  larval  skin  was  mounted  in  polyvinyl  lactophenol  on  microscope  slides  for 
further  examination.  Polyvinyl  lactophenol  was  chosen  because  it  is  a mild  clearing  agent 
and  thus  aids  in  the  ease  of  sclerite  recognition. 

Four  species  of  the  tribe  Ichneumonini  have  been  recorded  as  parasitoids  of  E.  ochro- 
gaster. Three  species,  Eutanyacra  suturalis,  Diphyus  No.  1,  and  Spilichneumon  superbus 
were  examined  and  found  to  be  similar  in  the  following  ways.  The  females  lay  their  eggs  in 
the  cutworm  larvae  but  the  host  is  not  killed  until  it  reaches  the  pupal  stage.  None  spin  any 
apparent  cocoon  and  all  use  the  host  pupa  for  protection  during  their  pupal  stage.  All  have 


96 


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Fig.  29-33.  Poecilanthrax  alcyon.  29.  pupa.  30.  ventral  view  of  pupal  head.  31.  ventro-lateral  view  of  posterior  end  of 
pupa.  32.  final  instar  larva  (Brooks,  1952).  33.  spine  and  seta!  pattern  of  third  abdominal  segment  of  pupa.  Scale  in 
millimeters. 


Parasitoids  of  Euxoa 


97 


greatly  reduced  larval  mouth  parts  which  differ  between  species  mainly  in  the  mandible 
shape.  One  pair  of  sclerites  located  behind  the  mandibles  do  not  conveniently  fit  into  the 
Finlayson  (1960)  sclerite  classification.  It  is  possible  that  these  are  modified  from  the 
suspensorial  sclerite.  In  all  these  species,  the  stipital  and  labial  sclerites  are  lost.  The  shape 
of  the  spiracles  is  quite  constant  (Fig.  37),  and  is  of  no  use  to  separate  species.  Several 
ichneumonine  larvae  were  dissected  from  final  instar  cutworm  larvae  and  one  was  removed 
from  a host  pupa.  Those  removed  from  the  cutworms  were  an  early  instar  (Fig.  36)  and 
could  be  either  Diphyus  No.  1 or  S.  superbus.  The  one  removed  from  the  pupa  had  cephalic 
structures  which  were  clearly  those  of  Diphyus  No.  1.  When  attacked  by  any  of  these  spe- 
cies, the  host  pupa  does  not  darken  as  it  normally  does,  but  remains  the  shiny  brown  of  a 
healthy  3 to  7 day  old  pupa.  The  anterior  portion  of  the  host  pupa  is  chewed  and  broken 
by  the  emerging  ichneumonine  adult  (Fig.  64).  The  pupal  remains  are  distinct  from  those 
from  which  a moth  has  emerged  (Fig.  61).  The  unexamined  species,  Pseudambly  teles  sub- 
fuscus,  is  likely  similar  to  the  examined  species. 

Eutanyacra  suturalis  (Say) 

Description  of  larvae.  - The  cephalic  structures  (Fig.  35)  of  the  final  instar  larva  with 
incomplete  epistoma.  Pleurostoma  wide,  heavily  sclerotized,  and  with  a very  irregular,  poor- 
ly defined  edge.  Superior  mandibular  process  wide  and  short,  inferior  mandibular  process 
reduced  to  one.  Pleurostoma  and  hypostoma  closely  connected  but  limited  movement 
between  them  possible.  Hypostomal  arms  heavy  and  of  irregular  width.  Stipital  and  labial 
sclerites  missing.  Labial  and  maxillary  palpi  small  and  difficult  to  see.  Silk  press  small  and 
very  lightly  sclerotized.  Mandibles  large,  robust,  heavily  sclerotized  with  slightly  curved 
blades  which  continued  into  base  without  clear  marking.  Behind  or  above  mandibles,  the 
two  sclerites  heavily  sclerotized  and  irregularly  larger  dorsally  than  ventrally. 

Biology.  — E.  suturalis  females  were  observed,  collected,  and  reared  during  the  Actebia 
fennica  outbreak  at  Worsley,  Alberta  in  1967.  The  females  were  seen  to  fly  and  hunt  during 
the  late  afternoon  and  early  evening.  They  were  found  hunting  in  fields  which  had  been 
defoliated  several  days  before  by  the  passing  cutworm  army.  The  hunting  females  ran 
rapidly  on  the  soil  surface  and  searched  around  soil  clods  and  in  large  cracks  in  the  soil. 
While  hunting,  they  could  be  easily  approached  and  could  be  captured  by  hand.  Captured 
females  would  attack  field-caught  A.  fennica  larvae,  but  oviposition  was  never  observed.  The 
attack  consisted  of  mounting  the  cutworm  lengthwise  and  curling  the  tip  of  the  abdomen 
under  so  that  the  ovipositor  touched  the  cutworm.  At  this  point,  every  attacking  female 
was  flipped  off  the  cutworm  by  a violent  twisting  movement  of  the  cutworm.  In  no 
instance  was  such  a cutworm  reattacked,  and  I never  succeeded  in  rearing  an  E.  suturalis 
adult  from  an  offered  cutworm.  In  the  laboratory,  adult/:,  suturalis  emerged  10  to  14  days 
after  host  pupation. 

Hosts.  — Whitehouse  (1922)  recorded  E.  suturalis  as  reared  from  E.  ochrogaster  in  Alber- 
ta. The  following  hosts  were  found  by  examining  host  labels  in  the  C.N.C.:  Actebia  fennica, 
British  Columbia;  E.  ochrogaster,  Saskatchewan;  E.  flavicollis,  Saskatchewan;  Scotogramma 
trifolii  Rottenburg,  Saskatchewan. 

Spilichneumon  superbus  (Provancher) 

Description  of  larvae.  — The  cephalic  structures  of  the  final  instar  larva  (Fig.  39)  are 
similar  to  those  of  E.  suturalis  and  Diphyus  No.  1.  The  size  is  closer  to  that  of  Diphyus 
No.  1 than  to  that  of  E.  suturalis.  The  mandibles  are  short  and  broad  with  a wide,  straight 


98 


Schaaf 


blade  which  is  continuous  in  appearance  with  the  base.  The  edge  of  the  pleurostoma  is  well 
defined  as  in  Diphyus  No.  1 . 

Biology.  — This  species  was  reared  from  A.  fennica,  E.  ochrogaster  and  F.  ducens  during 
the  current  study.  Because  it  was  not  recognized  as  being  different  from  Diphyus  No.  1 
while  live  adults  were  available,  the  adults  were  mixed  and  little  data  on  either  behavior  or 
biology  was  obtained.  The  adults  emerged  from  the  host  about  3 weeks  after  host  pupation. 

Hosts.  — The  following  hosts  of  S.  superbus  have  been  recorded  (Heinrich,  1960): 
Chorizagrotis  auxiliaris,  Alberta;  E.  ochrogaster , Manitoba;  E.  scandens,  E.  flavicollis,  E. 
messoria,  Feltia  ducens,  Saskatchewan;  Pseudaletia  unipuncta,  Hawaii  (introduced). 

Pseudambly  teles  subfuscus  (Cresson) 

Strickland  (1923)  recorded  Ambly teles  subfuscus  as  reared  from  E.  ochrogaster.  This 
species  was  subsequently  transferred  to  Pseudambly  teles  which  is  now  considered  to  be 
congeneric  with  Diphyus  Kriechbaumer  (Heinrich,  1961).  The  genus  Diphyus  is  now  being 
reworked  and  the  different  species  are  currently  denoted  by  numbers  in  the  C.N.C.  In  the 
C.N.C.  only  Diphyus  No.  1 of  the  genus  Diphyus  had  host  labels  associating  it  with  E. 
ochrogaster.  Dr.  Mason  (pers.  comm.)  states  that  Diphyus  No.  1 does  not  include  what  was 
P.  subfuscus.  There  is  no  other  evidence  currently  available  to  determine  which  of  the 
Diphyus  species  was  P.  subfuscus. 

Strickland  (1923)  described  the  biology  of  P.  subfuscus  from  his  research.  Unfortunately 
little  of  the  data  which  he  presents  is  of  use  to  separate  P.  subfuscus  from  any  of  the  other 
ichneumonines  which  attack  E.  ochrogaster.  The  major  biological  fact  which  he  describes 
is  that  the  eggs  are  laid  in  the  salivary  glands  of  the  host  cutworm.  During  my  dissections  of 
E.  ochrogaster  larvae,  I have  seen  neither  the  eggs  which  he  describes  nor  the  resulting  scar 
on  the  salivary  glands.  Until  the  taxonomy  of  Diphyus  is  better  understood,  the  status  of 
P.  subfuscus  as  a parasitoid  of  E.  ochrogaster  will  be  unclear. 

Hosts.  — Strickland  (1923)  reared  P.  subfuscus  from  Chorizagrotis  auxiliaris  and  E.  ochro- 
gaster in  Alberta,  while  Gibson  (1917)  reared  it  from  Euxoa  excellans  Grote  in  British 
Columbia. 


Diphyus  No.  1 

Description  of  larvae.  — The  cephalic  structures  of  the  final  instar  larvae  (Fig.  38)  of 
Diphyus  No.  1 are  similar  to  those  of  E.  suturalis.  The  mandibles  differ  from  those  of  E. 
suturalis  in  that  they  have  a long  narrow  blade  which  is  well  marked  off  from  the  base. 
When  rotated  the  blades  appear  to  be  curved  posteriorly.  The  sclerites  above  the  mandibles 
tend  to  be  more  squared  than  those  of  E.  suturalis.  The  edge  of  the  epistoma  in  Diphyus  No. 
1 is  well  defined. 

Biology.  — Using  the  final  instar  cephalic  structures,  it  was  found  that  this  species  was  a 
parasitoid  of  E.  ochrogaster  at  Calahoo.  Adults  emerged  from  the  host  pupae  about  3 weeks 
after  host  pupation. 

Hosts.  — The  following  hosts  were  found  by  examining  Diphyus  No.  1 specimens  in  the 
C.N.C.:  E.  ochrogaster,  Alberta;  C.  auxiliaris,  Alberta;  unidentified  cutworm,  British  Colum- 
bia. 


Camp  ole  tis  atkinsoni  (Viereck) 


King  and  Atkinson  (1928)  reported  C.  atkinsoni  as  reared  from  E.  ochrogaster.  This 


Fig.  34.  Generalized  cephalic  structures  of  final  instar  ichneumonoid  larva  (from  Finlayson,  1960).  a,  vacuole;  b,  sus- 
pensorial  sclerite;  c,  antennal  socket;  d,  antenna;  e,  mandible;  f,  lacinial  sclerite;  g,  sensorium;  h,  maxillary  palp;  i,  silk 
press;  j,  prelabial  sclerite;  k,  labial  palp;  1,  labial  sclerite;  m,  labral  sclerite;  n,  epistoma  (incomplete);  o,  superior  mandi- 
bular process;  p,  pleurostoma;  q,  teeth;  r,  inferior  mandibular  process;  s,  hypostoma;  t,  hypostomal  spur;  u,  blade  of 
mandible;  v,  dorsal  arm  of  labial  sclerite;  w,  stipital  sclerite.  Fig.  35.  Cephalic  structures  of  final  instar  larva  of  Eutanyacra 
suturalis.  Fig.  36.  Ichneumonine  larva.  Fig.  37.  Larval  spiracle  of  Spilichneumon  superbus.  Fig.  38.  Cephalic  structures 
of  final  instar  larva  of  Diphyus  No.  1.  Fig.  39.  Cephalic  structures  of  final  instar  larva  of  S.  superbus.  Scale  in  millimeters. 


100 


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species  appears  to  be  one  of  the  more  important  parasitoids  of  E.  ochrogaster.  This  will  be 
discussed  later. 

Description  of  cocoon.  - The  cocoon  (Fig.  40)  (2. 1-2.4  x 5. 4-6. 2 mm)  consists  of  two 
layers:  a very  thin  outer  layer  of  silk  which  is  also  used  to  secure  the  cocoon  to  the  sub- 
strate, and  a tough  inner  parchment-like  layer.  The  freshly  spun  cocoon  is  a light  yellowish 
white  but  the  mature  cocoon  varies  from  a dark  tan  to  a dull  brown.  Some  of  the  specimens 
reared  in  this  study  were  observed  to  spin  their  cocoons  with  the  host  remains  attached  to 
the  posterior  end.  In  other  cases,  the  larva  would  crawl  a few  centimeters  before  spinning  a 
cocoon.  In  all  cases  the  cocoon  took  12  to  18  hours  to  complete.  The  exit  hole  is  on  the 
dorsal  edge  of  the  anterior  end  of  the  cocoon.  It  is  irregular  in  outline  as  the  adult  chews  its 
way  out.  The  remains  of  the  final  instar  larva  are  stuck  to  one  side  near  the  end  of  the 
cocoon.  These  remains  are  often  difficult  to  find  and  remove.  The  remains  of  the  pupa  and 
the  meconium  tend  to  obscure  the  larval  remains  so  that  care  must  be  taken  not  to  lose  the 
cephalic  structures  in  any  of  the  study  procedures. 

Description  of  larva.  - The  cephalic  structures  of  the  final  instar  larva  (Fig.  42)  with  a 
complete  epistomal  arch  which  is  very  lightly  sclerotized  and  difficult  to  recognize.  Superior 
mandibular  processes  sclerotized  and  small.  Inferior  mandibular  processes  well  developed. 
Lacinial  sclerite  absent.  Hypostoma  long,  narrow,  straight,  and  heavily  sclerotized.  Hyposto- 
mal  spur  about  Wi  times  as  long  as  wide  at  base  and  meets  straight,  well-developed  stipital 
sclerite  at  about  midpoint.  Stipital  sclerite  meets  top  of  labial  sclerite.  Labial  sclerite  widest 
at  one-third  of  way  from  base  and  narrowed  to  a rounded  point  at  end.  End  of  labial  sclerite 
very  lightly  sclerotized  and  may  appear  lost.  Silk  press  large,  wide,  well  developed  and  ter- 
minated by  a long  narrow  spur.  Mandibles  small  with  a short  blade  which  curves  directly 
from  base.  Labral  sclerite  absent.  Suspensorial  sclerite  short  and  narrow.  Maxillary  palpi 
large  and  well  developed  and  protrude  in  a large  membranous  sack.  Antennae  large  and  well 
developed. 

Biology.  - C.  atkinsoni  is  the  only  recorded  parasitoid  of  E.  ochrogaster  which  at- 
tacks the  early  instar  larvae.  It  usually  kills  the  host  in  the  third  or  fourth  instar.  For 
this  reason,  this  species  is  usually  overlooked  or  not  found  in  studies  based  solely  on 
outbreak  conditions.  The  first  evidence  of  the  presence  of  a C.  atkinsoni  larva  in  a host 
appears  2 to  4 days  before  the  host  is  killed.  At  this  point,  the  host  is  noticeably  more 
sluggish,  eats  less  than  normal,  and  tends  to  bulge  abnormally.  Subsequently,  the  host 
stops  eating,  loses  its  mobility,  and  begins  to  lose  its  normal  shape.  Just  before  the  pa- 
rasitoid emerges,  it  can  be  seen  moving  about  in  the  host,  as  the  host’s  integument  is 
very  limp.  The  C.  atkinsoni  larva  devours  the  entire  contents  of  the  host  so  that  after 
it  rips  its  way  out  of  the  host,  the  remains  lie  very  flat  and  are  nearly  transparent.  The 
parasitoid  larva  begins  to  spin  its  cocoon  immediately  after  emergence  and  usually  needs 
some  object  to  crawl  against  so  that  it  can  complete  its  cocoon.  The  cocoon  is  stuck 
to  the  substrate  with  silk  fibers.  The  adult  C.  atkinsoni  emerge  7 to  10  days  after  the 
completion  of  the  cocoon.  All  the  specimens  studied  in  1968  emerged  from  the  host 
during  the  last  few  days  of  May  and  the  first  week  of  June,  and  the  adults  emerged 
during  the  first  3 weeks  of  June.  Although  all  available  cutworm  species  found  in  the 
test  area  were  reared  in  both  1967  and  1968,  no  alternative  host  was  found  later  in 
the  season. 

Hosts.  — E.  ochrogaster  appears  to  be  the  only  recorded  host  of  C.  atkinsoni.  Graham 
(1965)  lists  several  noctuid  hosts  of  Campoletis  spp.  and  one  yponomeutid,  the  diamond- 
back  moth,  Plutella  maculipennis  Curtis.  As  C.  atkinsoni  attacks  only  the  early  instars  of  its 
cutworm  hosts,  its  host  range  will  likely  only  be  found  in  studies  which  are  based  upon 
endemic  cutworm  populations  rather  than  epidemic  populations. 


Parasitoids  of  Euxoa 


101 


Netelia  Gray 

The  genus  Netelia  formerly  was  named  Paniscus  Schrank.  The  name  change  was  necessi- 
tated because  of  the  misapplication  of  the  name  Paniscus  by  Gravenhorst  (Townes  et  al. , 
1961).  Townes  et  al.  provide  taxonomic  information  on  the  adults  of  this  genus  and  give 
keys  and  characters  to  separate  specimens  to  the  sub-generic  level.  Using  his  characters,  it 
was  found  that  the  specimens  reared  from  E.  ochrogaster  were  in  the  sub-genus  Netelia. 

Description  of  cocoon.  - The  cocoons  of  three  borrowed  specimens  of  Netelia  sp.  were 
examined  but  none  contained  any  larval  remains.  One  cocoon  found  in  the  field  study  at 
Calahoo  closely  resembled  those  of  Netelia , and  it  also  lacked  any  larval  remains.  It  is  possi- 
ble that  the  larval  exuviae  were  removed  from  the  borrowed  specimens  by  a previous  worker. 
It  is  also  possible  that  the  remains  are  left  loose  in  the  cocoon  and  easily  lost.  While  the 
literature  contains  generalized  descriptions  of  Netelia  larvae,  no  mouth  part  drawings  appear 
to  be  available.  Cushman  (1926)  and  Strickland  (1923)  provide  illustrations  of  larvae  and 
eggs  of  various  members  of  the  genus. 

The  cocoons  studied  averaged  4x12  mm  in  size.  The  cocoon  (Fig.  41)  consists  of  a very 
sparse  outer  covering  of  silk  which  appears  as  a fluffy  mass  at  the  ends,  and  a tough, 
tightly  constructed  inner  layer.  The  over-all  color  is  shiny  black,  but  Strickland  (1923) 
noted  that  it  is  originally  a light  color  which  changes  with  maturity.  When  viewed  laterally, 
the  cocoons  had  a slightly  curved  shape.  The  adults  emerged  from  the  end  of  the  cocoon 
leaving  a very  ragged  and  irregular  exit  hole. 

Biology.  — Netelia  sp.  is  the  only  recorded  external  parasitoid  of  E.  ochrogaster . Cushman 
(1926),  Vance  (1927),  and  Strickland  (1923)  provided  information  on  the  biology  of  spe- 
cies in  this  genus.  Only  a generalized  summary  of  the  life  history  of  Netelia  spp.  is  given 
here. 

The  females  deposit  one  to  four  stalked  eggs  on  the  thoracic  region  of  late  instar  lepidop- 
terous  larvae.  Of  these  only  one  survives  to  maturity.  Strickland  (1923)  showed  that  the  egg 
would  remain  attached  even  though  the  host  molted.  Cushman  (1926)  stated  that  the  host 
larvae  were  attacked  when  they  were  very  large  and  when  they  were  about  to  pupate  in 
some  protective  medium.  The  egg  hatches  generally  after  the  host  has  entered  a pupation 
site.  The  parasitoid  larva  feeds  by  puncturing  the  host  integument,  attaching  its  mouthparts, 
and  remaining  in  one  place  until  new  punctures  are  made  necessary.  The  parasitoid  larva 
remains  attached  to  the  egg  for  most  of  its  life,  at  least  till  the  final  instar.  Attachment  to 
the  egg  shell  is  accomplished  by  a special  spined  area  on  the  terminal  abdominal  segment. 
The  cast  larval  skins  all  remain  attached  to  the  egg  shell  and  provide  a convenient  record  of 
the  larval  morphology  of  each  instar.  The  parasitoid  develops  rapidly  and  kills  the  host 
leaving  a dry  skin.  Shortly  after  this  it  spins  a cocoon. 

As  the  species  which  has  been  reared  from  cutworms  has  not  been  identified,  it  is  not 
possible  to  construct  a host  list.  The  genus  is  too  large  and  widespread  to  be  considered  in 
this  paper.  Townes  et  al.  (1961)  state  that  the  hosts  of  Netelia  are  exposed,  medium-sized 
lepidopterous  larvae  that  pupate  in  the  ground.  As  Netelia  spp.  attack  the  larvae  just  before 
they  are  to  pupate,  it  is  likely  that  any  species  is  a potential  host  if  it  crawls  on  the  soil 
looking  for  a pupation  site.  The  examined  specimens  reared  from  E.  ochrogaster  were  from 
Saskatoon  and  Red  Deer,  Saskatchewan.  One  empty  cocoon  which  likely  was  of  Netelia  was 
found  at  Calahoo. 


Gravenhorstia  propinqua  (Cresson) 


Originally,  King  and  Atkinson  (1928)  recorded  Erigorgus  sp.  as  being  reared  from  E. 


102 


Schaaf 


ochrogaster.  Erigorgus  is  now  considered  to  be  a synonym  of  Gravenhorstia  (Townes  et  al. , 
1961).  Specimens  of  Gravenhorstia  propinqua  which  were  reared  from  E.  ochrogaster  were 
found  in  the  C.N.C. 

Description  of  cocoon.  — The  cocoon  of  this  species  is  spun  inside  the  host  pupa.  It 
appears  to  be  lightly  constructed  of  a single  layer  of  silk  which  likely  offers  little  protection 
to  the  parasitoid  pupa.  The  emerging  adult  destroys  the  entire  anterior  end  of  the  host  pupa 
and  leaves  much  of  the  cocoon  visible  (Fig.  65).  The  larval  remains  are  stuck  in  the  posterior 
end  of  the  cocoon  and  are  easily  recovered  and  handled. 

Description  of  larva.  — The  cephalic  structures  of  the  last  larval  instar  (Fig.  43)  are  dis- 
tinct from  the  other  pupal  parasitoids  and  are  described  as  follows:  The  epistoma  is  heavy 
and  complete,  with  superior  mandibular  processes  short,  wide  and  directed  ventrally.  In- 
ferior mandibular  processes  are  reduced  to  one  which  is  broad  and  heavy.  Lacinial  sclerite  is 
absent.  Long,  heavy  hypostomal  arms  are  curved  ventrally  in  a wide  semi-circle  so  that  the 
ends  form  a nearly  straight  line  with  ends  of  the  labial  sclerite.  Hypostomal  spur  absent. 
Stipital  sclerite  long,  narrow,  extending  from  one-third  of  way  along  the  hypostoma  ventro- 
medially  to  a point  three-quarters  along  its  length,  where  it  bends  sharply  dorso-medially 
to  touch  the  labial  sclerite.  Labial  sclerite  incomplete  with  long  narrow  arms  extending 
ventrally.  Silk  press  present,  very  lightly  sclerotized  forming  a wide  U-shape.  Labral  sclerite 
straight,  short  and  irregular  in  outline  with  several  vacuoles.  Suspensorial  sclerite  very  wide 
and  well  developed.  Mandibles  large,  well  developed,  with  distinct  blade  clearly  marked  off 
from  base.  Maxillary  and  labial  palps  clearly  distinct.  Antennae  not  observed. 

Biology.  — Except  that  it  kills  the  pupal  stage  of  E.  ochrogaster,  little  is  known  of  the 
biology  of  Gravenhorstia  propinqua.  King  and  Atkinson  (1928)  noted  that  the  species  they 
recorded  overwintered  in  the  host  pupa.  One  interesting  fact  appears  in  the  host  lists  for 
the  genus.  Two  tortricids  have  been  recorded  as  hosts,  and  both  larvae  were  killed  by  the 
Gravenhorstia  sp.  In  both  noctuid  hosts,  the  pupal  stage  was  killed.  It  is  possible  that  when 
this  genus  is  studied  more  carefully,  more  than  one  genus  will  be  found  within  the  present 
concept. 

Hosts.  — The  following  hosts  have  been  recorded  for  Gravenhorstia  spp.:  Polia  purpurissa- 
ta  Grote  (Wood  et  al.,  1954),  New  Brunswick:  Archips  argyrospilus  Walker  (Paradis,  1960), 
Quebec;  Tortrix  alleniana  Fern  (Martin,  1958),  Ontario;  Agrotis  orthogonia  (King  and 
Atkinson,  1928).  The  specimens  reared  from  E.  ochrogaster  were  from  Saskatchewan. 

BRACONIDAE 


Morphology  of  immature  braconids 

Basically,  the  methods  used  to  separate  the  immature  stages  of  ichneumonids  apply  to 
those  of  braconids.  In  addition,  important  characters  are  found  in  the  color,  size,  shape, 
and  number  of  cocoons  per  host  in  each  species.  The  appearance  of  the  cocoon  mass  is 
also  of  importance. 

Four  species,  Microplitus  kewleyi,  Apanteles  laeviceps,  A.  griffini,  and  A.  acronyctae, 
from  the  subfamily  Microgastrinae  have  been  recorded  as  reared  from  E.  ochrogaster.  Short 
(1952)  stated  that  the  final  instar  larvae  are  characterized  as  follows:  hypostoma,  stipital 
sclerite,  and  labial  sclerite  present;  hypostomal  spur  reduced;  pleurostoma  weakly  sclero- 
tized; epistoma  always  absent;  antennae  not  distinct;  setae  present  on  body  but  spines  not 
present.  Capek  (1970)  gives  the  same  basic  characters  but  makes  no  definite  statement 
about  the  epistoma.  I believe  that  Short  is  incorrect  in  stating  that  the  epistoma  is  always 
missing,  as  it  is  present  in  M.  kewleyi.  Capek  states  that  members  of  the  group  of  genera  to 
which  Apanteles  and  Microplitus  belong  are  endoparasites  of  lepidopterous  larvae,  are  often 


Parasitoids  of  Euxoa 


103 


gregarious,  emerge  as  mature  larvae  to  pupate,  and  that  the  emergence  hole  is  regular  due  to 
a cap. 


Apan  teles  Foerst 
Apart  teles  acronyctae  Riley 

Apanteles  acronyctae  was  recorded  as  being  reared  from  E.  ochrogaster  by  King  and 
Atkinson  (1928).  The  record  was  based  on  only  two  specimens  reared  by  the  authors.  A. 
acronyctae  is  normally  associated  only  with  arctiid  hosts  and  likely  does  not  attack  cut- 
worms (Mason,  pers.  comm.).  This  record  is  likely  based  on  a mistaken  identification. 

Apanteles  laeviceps  Ashmead 

Apanteles  laeviceps  was  recorded  as  reared  from  E.  ochrogaster  by  Strickland  (1923). 

Description  of  cocoon.  — The  cocoon  mass  (Fig.  45)  contains  22  to  28  individual  cocoons. 
The  mass  is  compact  and  only  rarely  was  an  isolated  cocoon  observed.  The  cocoons  are 
tightly  woven  together  and  are  united  in  a single  irregular  deep  mass.  The  individual  cocoon 
(Fig.  46)  is  3.0  to  3.2  mm  long,  cylindrical  with  bluntly  rounded  ends,  and  is  lightly  con- 
structed. When  treated  with  a mild  KOH  solution  the  cocoon  structure  is  completely  des- 
troyed. The  loose  outer  layer  of  silk  is  tightly  interwoven  with  that  of  the  other  cocoons. 
As  with  the  other  braconids,  a distinct  cap  is  formed  and  breaks  off  when  the  adult  emerges. 
The  color  of  the  mass  changes  from  a pale  yellow  to  white  as  the  cocoons  mature.  In  many 
cocoons,  the  contents  can  be  seen  through  their  walls. 

Description  of  larvae.  — The  penultimate  larval  stage  of  A.  laeviceps  (Fig.  47),  which  is 
found  in  the  host,  differs  from  the  same  stage  of  Meteorus  vulgaris  in  the  following  ways. 
The  cephalic  region  of  A.  laeviceps  is  more  clearly  defined  than  that  of  M.  vulgaris.  The 
body  of  A.  laeviceps  is  long,  narrow,  and  terminated  by  a bulbous  caudal  appendage.  Overall 
length  is  approximately  5 mm,  with  the  widest  point  measuring  1.8  mm.  As  in  M.  vulgaris, 
the  larvae  molt  to  the  final  instar  as  they  escape  from  the  host.  The  final  instar  larva  was 
described  by  Strickland  (1923).  It  differs  from  that  of  M.  vulgaris  in  that  it  lacks  a definite 
caudal  appendage  and  has  a series  of  small  black  spines  on  its  body.  It  differs  from  the 
penultimate  stage  by  not  having  the  globular  caudal  appendage.  In  both  of  the  last  two 
larval  instars  of  A.  laeviceps  the  mouthparts  appear  similar.  The  cephalic  structures  of  the 
final  instar  larva  with  epistoma  missing,  pleurostoma  apparently  missing,  lower  mandibular 
process  blunt  and  rounded  (Fig.  44).  Hypostoma  heavily  sclerotized,  long,  narrow  and 
curved  medially.  Hypostomal  spur  small,  distinct,  pointed.  Stipital  sclerite  heavily  sclero- 
tized, wide  with  a distinct  twist  in  the  mid-area.  Labial  sclerite  complete,  well  developed, 
thick  at  top,  narrowed  at  bottom,  sometimes  widened  at  end.  Silk  press  very  obscure  but 
large.  Maxillary  and  labial  palpi  small.  Mandibles  with  long  curved  blade  arising  low  on  man- 
dibular base.  Antennae  and  spiracles  not  apparent. 

Biology.  — The  females  of  A.  laeviceps  oviposit  in  early  instar  host  larvae.  This  was 
demonstrated  in  the  current  study  by  rearing  early  instar  cutworms  which  had  been  captured 
in  the  field,  and  finding  them  to  be  attacked  by  A.  laeviceps.  The  earliest  instar  cutworm 
found  attacked  was  an  early  third  instar.  The  host  is  usually  killed  in  the  fifth  or  early  sixth 
instar.  The  larvae  of  A.  laeviceps  exert  a strong  influence  over  the  behaviour  of  the  host. 
Strickland  (1923)  noted  that  until  the  A.  laeviceps  larvae  began  to  attach  to  the  host’s 
cuticle,  no  ill  effects  were  evident.  At  this  stage,  the  host  leaves  the  soil  and  climbs  some 
convenient  object  such  as  a grain  stem  or  a large  clod  of  earth.  I have  observed  E.  ochro- 


104 


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gaster  larvae  climbing  only  when  attacked  by  A.  laeviceps.  While  the  host  is  on  the  object, 
the  parasitoid  larvae  emerge  from  all  sides  of  it.  The  host  then  crawls  away  leaving  the  para- 
sitoids  behind.  The  parasitoids  begin  to  spin  their  cocoons  almost  immediately.  When  study- 
ing cutworm  outbreaks,  cocoon  masses  of  A.  laeviceps  are  easily  found  by  examining  marker 
stakes  or  emergence  traps,  both  of  which  serve  as  climbing  points  for  the  hosts.  The  host 
larva  returns  to  the  soil  but  always  dies  within  a short  time.  In  the  laboratory,  I have  seen 
E.  ochrogaster  larvae  live  up  to  3 days  following  A.  laeviceps  emergence.  These  host  larvae 
are  recognizable  by  the  presence  of  the  exuvia-plugged  emergence  holes  of  the  parasitoid, 
the  presence  of  one  or  more  A.  laeviceps  larvae  in  the  body  cavity  which  failed  to  emerge, 
and  by  the  general  lack  of  damage  to  the  muscles  and  nerves. 

Hosts.  — The  following  hosts  have  been  recorded  in  the  literature:  Eucirrhoidea  pampina 
Gn.  (Wood  and  Nielson,  1957),  New  Brunswick;  Spaelotis  clandestina  Harrison  (Wood, 
1951),  New  Brunswick;  Syngrapha  epigaea  Grote  (Wood,  1951),  New  Brunswick;  E.  ochro- 
gaster, Chorizagrotis  auxiliaris  (Strickland,  1923),  Alberta;  Meliana  albilinea  Hiibner  (Web- 
ster, 1911);  Loxostege  sticticalis  L.  (Vierick,  1916 ),  Pseudaletia  unipuncta  (Guppy,  1967) 
Ontario.  During  the  current  study  A.  laeviceps  was  reared  from  E.  ochrogaster  in  1967  and 
1968  and  from  Feltia  ducens  in  1967. 

Apart  teles  griffini  Viereck 

Schaffner  and  Griswold  (1934)  recorded  Apart  teles  griffini  as  being  reared  from  E.  ochro- 
gaster in  the  north-eastern  part  of  the  United  States. 

Description  of  cocoon.  — The  cocoon  mass  of  A.  griffini  (Fig.  49)  is  deeply  divided 
longitudinally  so  that  two  distinct  but  connected  portions  are  evident.  The  construction  and 
density  of  the  mass  is  very  similar  to  that  of  A.  laeviceps.  The  groove  is  likely  formed  be- 
cause the  larvae  pupate  almost  immediately  after  emergence  and  do  not  move  together.  The 
groove  corresponds  to  the  location  of  the  host  at  the  time  of  emergence  of  the  parasitoid 
larvae. 

Description  of  larvae.  — No  specimens  of  the  entire  larvae  were  available  for  study. 
Cephalic  structures  of  final  instar  larvae  (Fig.  48)  similar  to  those  of  A.  laeviceps.  Hypo- 
stoma  less  distinctly  curved  than  in  A laeviceps.  The  labial  sclerite  more  narrowly  developed 
than  that  of  A.  laeviceps.  Mandible  with  a long  narrow  blade  which  arises  near  the  center  of 
the  base.  The  base  is  more  triangular  than  that  of  A.  laeviceps.  The  hypostomal  spur  is  short 
and  bluntly  developed. 

Biology.  — A.  griffini  is  generally  found  in  the  more  southern  range  of  E.  ochrogaster 
(Mason,  pers.  comm.).  Little  is  known  of  the  biology  of  this  species. 

Hosts.  — Walkden  (1950)  recorded  the  following  hosts  of  A.  griffini  from  the  central 
great  plains,  U.S.A.:  Agrotis  orthogonia,  A.  gladiaria  Morrison,  Chorizagrotis  auxiliaris, 
Peridroma  margaritosa  Haworth. 

Microplitus  kewleyi  Muesebeck 

Schaffner  and  Griswold  (1934)  recorded  Microplitus  kewleyi  as  being  reared  from  E. 
ochrogaster . Only  one  specimen  was  available  for  study. 

Description  of  cocoon.  — The  cocoon  (Fig.  51)  is  solitary,  small  (3.1  x 1.4  mm),  has 
very  little  outer  silk,  has  a very  tough  inner  layer  of  silk,  and  is  an  opaque  buff  color. 

Description  of  larvae.  — Cephalic  structures  (Fig.  50)  of  the  final  instar  larva  with  a very 
weakly  sclerotized,  incomplete  epistoma.  Superior  mandibular  process  very  indistinct,  as 
well  as  the  rest  of  pleurostoma.  Inferior  mandibular  processes  each  small  and  bluntly  devel- 


Parasitoids  of  Euxoa 


105 


ig.  40,  42.  Campoletis  atkinsoni.  40.  cocoon.  42.  cephalic  structures  of  final  instar  larva.  Fig.  41.  Cocoon  of  Netelia  sp. 
Fig.  43.  Gravenhorstia  propingua  final  instar  larva  cephalic  structures.  Fig.  44-47.  Apanteles  laeviceps.  44.  cephalic 
structures  of  final  mstar  larva.  45.  cocoon  mass.  46.  individual  cocoon.  47.  penultimate  larva.  Scale  in  millimeters. 


106 


Schaaf 


oped.  Hypostoma  heavily  sclerotized,  curved  medially,  with  a reduced,  pointed  hypostomal 
spur.  Stipital  sclerite  long,  narrow,  and  with  irregular  outline.  Labial  sclerite  heavily  sclero- 
tized, widest  at  midpoint  and  narrowed  to  a point  at  distal  end.  Silk  press  lightly  sclerotized, 
long  and  narrow.  Suspensorial  sclerite  lightly  sclerotized  and  irregular  in  form.  Mandibles 
with  short  narrow  blades  arising  from  bottom  of  mandible  bases.  The  mandible  open  poste- 
riorly in  a distinct  groove. 

Biology.  — Extremely  little  is  known  of  the  biology  of  this  species  other  than  that  it 
attacks  larval  cutworms.  The  examined  specimen  was  reared  from  a cutworm  at  College 
Park,  Maryland. 


Meteorus  Haliday 

Two  species  of  Meteorus,  dimidiatus  Cresson  and  vulgaris  Cresson  have  been  recorded  as 
reared  from  E.  ochrogaster.  Strickland  (1923)  recorded  the  former  as  being  a common 
parasitoid  of  prairie  cutworms.  He  noted,  however,  that  difficulty  had  been  encountered 
when  the  series  were  identified  in  distinguishing  them  from  M.  vulgaris , and  Dr.  W.R.M. 
Mason  also  is  sceptical  of  this  record.  The  current  host  lists  (Graham,  1965)  do  not  include 
M.  dimidiatus.  All  of  the  Meteorus  specimens  reared  from  E.  ochrogaster  in  my  study  were 
M.  vulgaris.  Strickland  dealt  at  length  on  the  biology,  hosts,  and  development  of  M.  dimi- 
diatus. This  information  on  the  biology  of  M.  dimidiatus  appears  now  to  be  valid  for  M. 
vulgaris. 

Meteorus  vulgaris  Cresson 

This  species  was  recorded  as  reared  from  E.  ochrogaster  by  King  (1926),  and  has  sub- 
sequently been  recorded  several  times. 

Description  of  cocoon.  — The  cocoons  are  4.9-5 .4  mm  x 1.9-2. 3 mm,  elliptical  in  shape 
with  the  widest  point  between  the  posterior  one-third  and  the  midpoint  (Fig.  55).  The 
anterior  end  of  the  cocoon  terminates  in  a distinct  cap  which  breaks  off  at  adult  emergence. 
Specimens  which  were  observed  spinning  their  cocoons  formed  the  cap  only  after  the  rest  of 
the  cocoon  was  completed.  While  the  rest  of  the  cocoon  turns  brown  with  maturity,  the 
cap  remains  a much  lighter  color.  The  cocoons  are  usually  translucent  so  that  the  contents 
are  easily  seen.  An  outer  layer  of  fine  silk  binds  the  cocoon  to  the  substrate  and  to  sur- 
rounding cocoons.  The  cocoon  mass  (Fig.  54)  is  very  loose  and  irregular,  and  it  is  common 
to  find  cocoons  completely  separated  from  the  rest  of  the  mass.  The  larval  remains  are  easily 
found  and  removed  from  the  cocoons. 

Description  of  larvae.  — Cephalic  structures  of  the  final  instar  (Fig.  52)  with  incomplete, 
lightly  sclerotized  epistoma.  Each  superior  mandibular  process  well  developed,  inferior  man- 
dibular processes  small  and  blunt,  lacinial  sclerite  small  and  pointed.  Hypostomal  arms 
short,  narrow  curved,  and  only  lightly  sclerotized.  Pleurostoma  distinctly  sclerotized  as  op- 
posed to  light  sclerotization  of  the  epistoma  and  hypostoma.  Stipital  sclerite  long,  narrow, 
almost  straight,  reaching  upper  end  of  labial  sclerite.  Labial  sclerite  large,  heavy  and  greatly 
thickened  in  ventral  portion.  Silk  press  well  developed  and  moderately  sclerotized.  Well- 
developed  pharyngeal  region  with  lightly  sclerotized  ridges  behind  silk  press  and  top  of 
labial  sclerite.  Mandibles  small,  with  short,  pointed,  conical  blades  well  set  off  from  base. 
Maxillary  palps  large  and  with  well-developed  membranous  projection.  Antennae  large  and 
well  developed. 

Strickland  provided  Figures  of  the  entire  final  instar  larva  and  of  the  penultimate  larva. 
The  larva  normally  molts  as  it  emerges  from  the  host  leaving  the  exuvia  of  penultimate  larva 
in  the  emergence  hole.  The  penultimate  larva  as  shown  by  Strickland  has  a long  caudal 


Parasitoids  of  Euxoa 


107 


appendage  whereas  the  final  instar  larva  has  a much  reduced  caudal  appendage.  No  larvae 
similar  to  that  described  by  Strickland  as  the  penultimate  instar  were  found  during  my 
study.  The  shape  of  the  final  instar  larva  (Fig.  53)  changes  as  the  cocoon  is  spun.  The  larva 
shortens  and  thickens,  the  segmentation  becomes  less  defined,  and  the  caudal  appendage 
becomes  more  obscure.  When  dissecting  host  specimens  from  which  M.  vulgaris  larvae  had 
emerged,  it  was  found  that  one  to  three  larvae  were  usually  remaining  in  the  host.  These 
appeared  the  same  as  the  final  instar  larvae  and  likely  molted  even  though  they  failed  to 
emerge. 

Biology.  — The  biology  of  the  immature  stages  of  M.  vulgaris  is  well  documented  by 
Strickland  (1923).  Normally,  24  to  28  pupae  were  reared  from  a single  host  but  as  many  as 
36  were  reached  during  my  study.  The  hosts  killed  by  M.  vulgaris  during  1967  and  1968 
were  in  the  late  stages  of  the  sixth  instar.  In  most  cases,  the  host  larva  crawled  away  from 
the  M.  vulgaris  pupae  and  died.  Dissection  of  the  host  cutworm  after  the  emergence  of  the 
M.  vulgaris  larvae  revealed  that  the  muscle  layers  were  partially  destroyed,  as  is  the  case 
with  other  parasitoids.  The  exuviae  of  the  penultimate  instar  larvae  formed  blackened  areas 
on  the  cutworm  integument  indicating  the  emergence  points.  These  usually  occur  on  the 
ventro-lateral  portion  of  the  cutworm’s  body.  The  presence  of  the  cast  exuviae  gives  the 
cutworm  a characteristic  appearance,  aiding  in  recognition  of  the  cutworm  after  the  M. 
vulgaris  larvae  have  emerged,  but  before  the  death  of  the  host.  In  two  cases,  M.  vulgaris 
pupal  masses  were  found  in  the  loose  upper  soil.  The  adults  emerge  from  the  pupae  14  to  17 
days  after  pupation. 

Hosts.  — The  following  hosts  have  been  recorded  for  M.  vulgaris:  Euxoa  ochrogaster  (King 
and  Atkinson,  1928;  King,  1926)  Saskatchewan;  E.  tristicula  (King,  1926)  Saskatchewan; 
Peridroma  saucia  (Fletcher,  1901);  Syngrapha  epigaea  (Wook  and  Nielson,  1960)  New 
Brunswick.  Strickland  (1923)  reared  M.  vulgaris  (or  dimidiatus ) from  the  following  hosts 
captured  in  Alberta:  A.  orthogonia,  E.  ochrogaster,  C.  auxiliaris,  E.  tristicula,  S.  devestator 
Brace,  Actebia  fennica.  Walkden  (1950)  listed  the  following  hosts  of  M.  vulgaris  from  the 
central  great  plains,  U.S.A.:  A.  orthogonia,  A.  gladiaria,  A.  ypsilon,  C.  auxiliaris,  Feltia 
subgothica  Haworth,  Euxoa  messoria,  Peridroma  margaritosa.  I reared  M.  vulgaris  from 
Feltia  ducens  in  the  summer  of  1967. 

ENCYRTIDAE 
Copidosoma  bakeri  (Howard) 

Copidosoma  bakeri  is  probably  the  most  important  single  parasitoid  of  E.  ochrogaster 
over  most  of  its  range.  This  species  was  first  described  in  the  genus  Berecyntus  and  most  of 
the  literature  associated  with  this  species  is  found  under  Berecyntus  bakeri.  Originally  C. 
bakeri  was  described  as  a single  species  with  several  varieties  including  gemma  Girault, 
arizonensis  Girault,  euxoae  Strickland,  and  bakeri  Girault.  Other  workers  gave  these  varieties 
subspecific  rank  (Gibson,  1917;  Peck,  1951).  Peck  (1963)  states  that  C.  bakeri  cannot  be 
subdivided  into  varieties  or  subspecies,  as  the  earlier  groupings  are  merely  a reflection  of 
color  patterns  and  not  of  population  differences. 

Biology.  — Work  on  the  biology  and  development  of  C.  bakeri  has  been  published  by 
Gibson,  1915;  Strickland,  1916;  King  and  Atkinson,  1928;  and  Cook,  1930.  The  most 
extensive  work  done  on  this  species  was  that  of  McMillan  (1930)  in  an  unpublished  masters 
thesis. 

C.  bakeri  oviposits  in  the  eggs  of  its  many  hosts  but  the  host  is  not  killed  until  the 
approximate  time  that  host  pupation  occurs.  The  polyembryonic  development  of  C.  bakeri 


108 


Schaaf 


50 


0.1 


54 


. l.O- 


Fig.  48,  49.  Apanteles  griffmi.  48.  cephalic  structures  of  final  instar  larva.  49.  cocoon  mass.  Fig.  50,  51.  Microplitus 
kewleyi.  50.  cephalic  structures  of  final  instar  larva.  51.  cocoon.  Fig.  52-55 . Meteorus  vulgaris.  52.  cephalic  structures 
of  final  instar  larva.  53.  final  instar  larva.  54.  cocoon  mass.  55.  cocoon.  Scale  in  millimeters. 


Parasitoids  of  Euxoa 


109 


will  be  discussed  later. 

McMillan  (1930)  observed  C.  bakeri  adults  in  the  field  in  central  Saskatchewan  from 
early  May  until  the  end  of  August,  while  I observed  them  during  the  latter  portion  of  July 
and  most  of  August.  The  great  length  of  time  during  which  adults  occur  in  the  field  is  due 
to  the  emergence  from  different  host  species  completing  development  at  different  times 
during  the  summer.  C.  bakeri  adults  emerge  from  E.  ochrogaster  larvae  at  approximately  the 
same  time  as  do  the  E.  ochrogaster  adults.  In  the  laboratory  the  maximum  time  which  C. 
bakeri  adult  females  can  be  kept  alive  is  16  days,  with  a mean  of  1 1 days  (McMillan,  1930). 
It  is  likely  that  eggs  of  E.  ochrogaster  are  exposed  to  only  a small  portion  of  the  C.  bakeri 
adults  which  emerged  from  E.  ochrogaster  larvae  because  of  the  long  preoviposition  period 
of  E.  ochrogaster  females.  A detailed  biology  of  adult  C.  bakeri  may  be  found  in  McMillan 
(1930). 

The  parasitic  egg  develops  for  approximately  45  days  within  the  host  egg  till  it  reaches  an 
overwintering  stage.  During  this  time  it  has  approximately  doubled  in  size  and  is  transformed 
into  a syncytium  of  cleavage  nuclei  surrounded  by  a trophamnion  (McMillan,  1930).  The 
trophamnion  provides  nutriment  for  the  embryonic  mass  from  the  host  tissues  as  the  eggs 
of  polyembryonic  parasitoids  are  relatively  free  of  yolk  (Chapman,  1969).  After  the  host 
emerges  from  its  egg,  the  growth  of  the  polyembryonic  mass  resumes.  The  actual  mechanism 
of  polyembryonic  development  is  described  by  McMillan  (1930)  and  Leiby  (1922). 

The  earliest  that  a polyembryonic  body  (p.e.b.)  was  detected  in  my  dissections  of  red- 
backed  cutworms  was  in  the  third  instar  of  the  host.  The  polyembryonic  body  at  this  time 
is  very  small,  compact  and  is  usually  flattened  and  little  internal  differentiation  is  apparent. 
It  grows  in  size  in  the  next  instars  of  the  host  until  it  fills  a major  portion  of  the  area  be- 
tween the  gut  and  the  muscle  layer  under  the  integument.  At  this  point  it  may  be  present 
as  a simple  flattened  structure  or  it  may  be  lobed  or  divided  into  smaller  bodies.  If  more 
than  one  p.e.b.  is  present  in  the  host  then  one  is  usually  much  more  developed  than  the 
other.  There  were  never  two  distinct  p.e.b. ’s  in  the  same  area  of  the  host’s  body.  The  body 
remains  largely  undifferentiated  until  either  the  sixth  or  seventh  instar  of  the  host,  depen- 
ding upon  whether  an  extra  instar  occurs.  The  p.e.b.  then  begins  to  divide  into  smaller 
embryonic  units  which  are  marked  by  density  changes  in  the  p.e.b.  These  embryonic  units 
change  to  spherical  structures  and  give  the  entire  p.e.b.  the  appearance  of  a bag  of  marbles. 
These  structures  change  into  the  form  of  larvae  while  still  in  the  intact  p.e.b.  The  p.e.b.  in 
the  meantime  becomes  larger  and  more  deformed  till  it  is  very  lobed.  When  the  larvae  are 
fully  developed  within  the  body,  they  begin  to  escape  from  it.  Several  dissections  were  made 
at  the  time  when  the  p.e.b.  had  begun  to  disintegrate  and  the  larvae  spread  throughout  the 
host  body.  Not  all  the  larvae  appeared  to  leave  the  p.e.b.  simultaneously;  in  fact,  some  of 
the  remaining  larvae  were  less  developed  and  likely  did  not  complete  their  development 
before  the  earlier-developing  larvae  destroyed  both  the  host  and  the  remains  of  the  p.e.b. 
Occasionally,  some  larvae  were  found  free  of  the  p.e.b.  before  the  majority  of  the  larvae 
were  mature  enough  to  leave  the  p.e.b.  These  likely  correspond  to  the  pseudolarvae  referred 
to  by  Leiby  (1922,  1926).  The  pseudolarvae  are  actually  larvae  which  failed  to  obtain 
sufficient  nutrition  while  in  the  p.e.b.,  and  will  not  survive  to  the  pupal  stage. 

Once  the  larvae  break  free  of  the  p.e.b.,  they  begin  to  actively  ingest  the  host  body 
contents.  This  is  a very  rapid  process  taking  2 to  4 days. 

The  behaviour  of  the  host  changes  radically  during  the  last  stages  of  intact  p.e.b.  and  the 
beginning  of  the  parasitoid  larval  attack.  The  host  eats  more  during  the  last  period  than  does 
the  normal  cutworm  (McMillan,  1930).  As  the  p.e.b.  breaks  down  the  host  is  very  active 
and  restless.  Strong  turning  and  twisting  activity  is  often  noticed.  Feeding  ceases  during 
this  period.  The  parasitoid  larvae  distribute  themselves  throughout  the  body  and  rapidly 


110 


Schaaf 


destroy  the  internal  organs  leaving  the  external  musculature  and  nerve  network  until  last. 
The  host  is  usually  capable  of  reflexes  i.e.  curling  movements  even  if  the  entire  internal 
structure  is  destroyed.  Finally,  the  muscles,  brain,  and  nervous  system  are  eaten  leaving 
only  a ‘plastic’  bag  of  host  integument  full  of  parasitoid  larvae.  In  one  case,  even  this 
integument  bag  was  destroyed.  The  external  appearance  also  changes  radically  from  the 
attack  of  the  larvae.  Sometimes  the  intact  p.e.b.  can  be  seen  through  the  intersegmental 
membranes  of  the  host  as  a solid  white  mass  which  does  not  move  like  the  surrounding  fat 
body.  The  host  at  this  stage  is  a typical  dorsal  red  and  ventral  clear  light  grey.  The  crawling 
and  curling  movements  appear  normal.  As  the  p.e.b.  begins  to  break  down,  the  color  of  the 
body  changes.  The  reds  become  lighter  and  pass  through  a pale  pink  and  then  become  an 
off-grey.  The  ventral  surface  changes  from  the  clear  grey  to  a mottled  grey  as  the  parasitoid 
larvae  can  be  seen  through  the  integument  (Fig.  59).  As  the  last  muscle  and  nerve  tissue  is 
destroyed,  a fluid  discharge  is  emitted  from  the  body  of  the  host  which  leaves  a brown  stain 
on  the  filter  paper  of  a culture  dish.  The  host  is  now  a uniform  buff-grey  shade  and  the 
larvae  are  packed  into  every  portion  of  the  body  including  the  prolegs,  brain  and  eyes  (Fig. 
58).  The  body  of  the  host  sags  to  the  most  stable  shape,  which  in  culture  dishes  fits  the 
pattern  of  the  objects  on  which  it  lies.  Specimens  found  in  the  field  were  flattened,  screw- 
shaped,  or  almost  normally  curved.  The  C.  bakeri  larvae  pupate  within  2 days  of  the  host 
body  collapse. 

As  the  pupae  mature,  the  color  of  the  host  carcass  darkened  noticeably  due  to  the  color 
change  of  the  individual  pupae.  After  14  to  26  days  the  adults  emerge  in  a period  of  6 to  12 
hours.  The  host  carcass  is  perforated  with  holes  on  all  sides  except  the  bottom.  Each  adult 
does  not  form  a new  hole  but  will  use  an  old  hole  if  one  exists  near  it.  After  emergence  the 
host  resumes  a dull  grey  or  tan  color. 

The  individual  larvae  (Fig.  56)  of  C.  bakeri  have  very  few  distinctive  characters.  McMillan 
(1930)  gives  a figure  of  the  mandibles  of  the  larvae,  but  I found  the  mandibles  so  hard  to 
obtain  and  study  that  I regard  them  as  being  essentially  useless  as  an  identifying  character. 
C.  bakeri  larvae  are  easily  distinguished  from  other  larvae  likely  to  be  found  attacking  E. 
ochrogaster  by  their  great  numbers  and  by  their  lack  of  sclerotized  characters.  Late  larvae  or 
prepupae  (Fig.  57)  are  also  easily  recognized  as  the  host  is  essentially  destroyed  by  the  time 
of  their  appearance. 

Hosts.  - C.  bakeri  has  a wide  range  of  natural  hosts.  The  following  hosts  were  recorded 
by  Peck  (1963)  with  the  authors  he  cited  and  additional  references.  Location  by  province  or 
state  is  included  to  provide  additional  information  on  the  distribution  of  both  the  host  and 
the  parasitoid:  a.  Agrotis  orthogonia  (Cook,  1930),  Montana;  b.  A.  venerabilis  Walker 
(King  and  Atkinson,  1928),  Saskatchewan;  c.  Amathes  smithi  Snellen  (Wood  and  Nielson, 
1957), New  Brunswick;d.  Chorizagrotis  auxiliaris  (Strickland,  19 16;  Girault,  1917),  Alberta; 
(Snow,  1925),  Utah;  (Walkden,  1950),  Kansas;  e.  C.  thanatologia  (King  and  Atkinson,  1928), 
Saskatchewan;  f.  Chorizagrotis  sp.  Sm.  (Girault,  1916),  Arizona;  g.  Cry  modes  devastator 
(Gibson,  1915,  1917;  Girault,  1916;  Treheme,  1915),  Ontario;  h.  Euxoa  deter sa  Walker 
(King  and  Atkinson,  1928),  Saskatchewan;  i.  E.  flavicollis  (King  and  Atkinson,  1928), 
Saskatchewan;  j.  E.  intrita  Morrison  (Cook,  1930),  Montana;  k.  E.  messoria  (Walkden, 
1950),  Kansas;  1.  E.  ochrogaster  (King  and  Atkinson,  1928),  Saskatchewan;  m.  E.  scandens 
(Walkden,  1950),  Kansas;  n.  E.  tristicula  (Strickland,  1921;  WhitehouSe,  1922),  Alberta; 
(King  and  Atkinson,  1928),  Saskatchewan;  o.  Euxoa.  sp.  (Girault,  1916;  Gibson,  1917), 
Ontario;  p.  Feltia  ducens  (King  and  Atkinson,  1928),  Saskatchewan;  q.  Feltia  subgothica 
(V/alkden,  1943,  1950),  Kansas;  (Peck,  1951),  Kansas,  Alberta,  New  Mexico;  r.  Lacinopolia 
renigera  Stephens  (Walkden,  1950),  Kansas,  s . Pissodes  strobi  (Taylor,  1929),  Massachusetts. 

McMillan’s  (1930)  thesis  lists  nine  of  the  above  species  as  hosts  of  C.  bakeri.  In  addition 


Parasitoids  of  Euxoa 


111 


Fig.  56-59.  Copidosoma  bakeri.  56.  feeding  larva.  57.  prepupa.  58.  cross-section  of  host  at  completion  of  C.  bakeri 
feeding.  59.  surface  view  of  above.  Fig.  60-65.  E.  ochrogaster  pupae.  60.  normal  before  moth  emergence.  61.  normal 
after  moth  emergence.  62.  after  Gonia  spp.  emergence.  63.  after  Poecilanthrax  alcyon  emergence.  64.  after  ichneumo- 
nine  emergence.  65.  after  Gravenhorstia  propingua  emergence.  Scale  in  millimeters. 


112 


Schaaf 


six  more  species  are  given  in  which  parasitism  was  induced.  They  are  Euxoa  tesselata,  E. 
verticalis  Grote,  E.  dargo  Stkr.,  E.  divergens  Walker,  E.  campestris  Grote,  L.  renigera.  McMil- 
lan felt  that  natural  parasitism  does  not  occur  in  these  species  because  they  normally  lay 
their  eggs  in  hard  packed  prairie  soil.  He  proved  that  firmly  packed  soil  provides  an  effec- 
tive barrier  to  C.  bakeri  oviposition. 

Of  the  20  species  which  I have  listed  as  recorded  hosts,  I believe  that  one  is  in  error. 
The  species  Pissodes  strobi,  the  white  pine  weevil,  attacks  the  shoots  of  white  pine  trees. 
( Pinus  strobus ).  The  list  from  which  Peck  (1963)  derived  his  information  is  headed  ‘Parasites 
reared  from  P.  strobi  or  weeviled  material’  (Taylor,  1929)  and  is  data  from  several  different 
workers.  All  other  recorded  hosts  are  cutworms  which  are  typically  found  in  open  areas  and 
whose  eggs  are  laid  on  or  near  the  soil,  while  P.  strobi  is  a weevil  and  occurs  in  wooded  areas. 
Also,  P.  strobi  lays  its  eggs  in  the  tips  of  white  pine  shoots  so  that  it  is  very  unlikely  that  one 
parasitoid  would  attack  such  a wide  range  of  hosts  in  such  divergent  habitats. 

During  the  present  study  C.  bakeri  was  reared  from  Feltia  ducens  during  the  summer  of 
1967,  and  from  E.  ochrogaster  in  1967  and  1968  and  was  dissected  from  E.  ochrogaster 
larvae  in  1969. 


DISCUSSION 

Twenty-seven  species  of  parasitoids  have  been  recorded  as  reared  from  Euxoa  ochro- 
gaster. Three  of  these  records,  Gonia  capitata,  Apanteles  acronyctae,  and  Meteorus  dimidi- 
atus  are  incorrect.  Of  the  24  species  of  confirmed  parasitoids,  the  immature  stages  of  15 
are  described.  Of  the  remaining  species,  Brooks  (1952)  has  provided  descriptions  of  four 
of  the  bombyliids.  Pseudambly teles  subfuscus  and  Gonia  sp.  were  not  described  as  I was 
not  able  to  find  what  species  these  records  are  now  valid  for  and  thus  could  not  locate 
specimens.  Gonia  fuscicollis,  Villa  lateralis,  and  Agamermis  sp.  were  not  described  because 
I could  obtain  no  specimens  of  any  of  the  immature  stages.  Because  of  the  possibility  that 
both  Periscepsia  laevigata  and  Gonia  sequax  are  parasitoids  of  the  red-backed  cutworm, 
descriptions  of  the  available  immature  stages  of  these  species  have  been  included. 

The  known  biology  of  each  species  of  parasitoid  is  given  in  the  text  and  some  of  this 
information  is  summarized  in  Table  1.  It  should  be  noted  that  the  stage  of  the  host  in  which 
the  parasitoid  lays  its  egg  was  determined  either  by  literature  records  or  by  using  rearing 
data  from  field-collected  specimens.  The  earliest  instars  selected  by  the  parasitoid  can  be 
calculated  by  determining  both  the  instar  and  the  time  of  the  season  in  which  the  host, 
which  later  produced  a parasitoid,  was  captured.  The  latest  instars  can  be  found  by  dissec- 
tions of  host  material  and  sometimes  by  observing  the  oviposition  behaviour  of  the  para- 
sitoid. For  example,  no  specimens  of  the  red-backed  cutworm  collected  before  the  fourth 
instar  produced  ichneumonine  adults.  When  sixth  instar  cutworms  were  attacked  in  the 
laboratory,  the  ichneumonine  females  were  unable  to  lay  eggs  in  the  host  because  of  its 
defense  reactions. 

Figure  66  summarizes  the  parasitoid  complex  and  the  stages  at  which  the  parasitoid  eggs 
are  laid  and  those  at  which  the  hosts  are  killed. 

Alternative  hosts 

All  the  better  known  species  of  parasitoids  in  this  study  have  several  hosts  from  which 
they  have  been  reared  and  recorded.  Indubitably,  for  each  of  these  species  of  parasitoids, 
there  are  more  hosts  which  have  yet  to  be  recorded.  Most  host  records  for  parasitoids  result 
from  the  rearing  of  economic  species,  and  there  is  little  reason  to  suspect  that  these  para- 
sitoids attack  only  hosts  which  are  of  economic  importance.  With  the  exception  of  Villa 


Parasitoids  of  Euxoa 


1 13 


Table  1 . Summary  of  the  biology  of  E.  ochrogaster  parasitoids. 


Species 

Status 

Stage 

attacked 

Stage 

killed 

No.  of 
known  hosts 

Tachinidae 

Gonia  aldrichi 

confirmed 

IV,  V,  VI 

pupa 

2 

Gonia  capita ta 

rejected 

— 

— 

— 

Gonia  fuscicollis 

confirmed 

IV,  V,  VI 

pupa 

1 

Gonia  sequax 

suspected 

IV,  V,  VI 

pupa 

1 

Bonne tia  comta 

confirmed 

III,  IV 

prepupa 

7 

Periscepsia  helymus 

confirmed 

larva 

VI 

17 

Peris  cep  sia  laevigata 

suspected 

larva 

V 

5 

Bombyliidae 

Villa  alternata 

confirmed 

IV,  V,  VI 

pupa 

6 

Villa  fulviana 

confirmed 

IV,  V,  VI 

pupa 

1 

Villa  lateralis 

confirmed 

IV,  V,  VI 

pupa 

1 

Villa  mo ro ides 

hyper  parasitoid 

— 

— 

— 

Poecilanthrax  alcyon 

confirmed 

IV,  V,  VI 

pupa 

5 

Poecilanthrax  willistonii 

confirmed 

IV,  V,  VI 

pupa 

9 

Ichneumonidae 

Eutanyacra  suturalis 

confirmed 

IV,  V,  ?VI 

pupa 

4 

Diphyus  No.  1 

confirmed 

IV,  V,  ?VI 

pupa 

3 

Spilichneumon  superbus 

confirmed 

IV,  V,  ?VI 

pupa 

6 

Pseudambly  teles  subfuscus 

? 

— 

Comp  ole  tis  atkinsoni 

confirmed 

I,  II 

III,  IV 

1 

Netelia  sp. 

confirmed 

V,  VI 

prepupa 

— 

Gravenhorstia  propingua 

confirmed 

larva 

pupa 

1 

Braconidae 

Apan  teles  acronyctae 

rejected 

— 

Apan  teles  laeviceps 

confirmed 

11.  Ill 

V 

8 

Apan  teles  griffini 

confirmed 

?II,  III 

?V 

4 

Microplitus  kewleyi 

confirmed 

?III,  IV 

?IV,  V 

1 

Meteorus  vulgaris 

confirmed 

III,  IV 

VI 

14 

Meteorus  dimidiatus 

rejected 

- 

- 

- 

Encyrtidae 

Copidosoma  bakeri 

confirmed 

egg 

VI  or  VII 

19 

Nematoda  Merinthidae 

Agamerimis  sp. 

confirmed 

? 

1 

— 

114 


Schaaf 


alternata , all  parasitoids  recorded  as  reared  from  E.  ochrogaster  were  mainly  restricted  to 
noctuid  larvae  as  additional  hosts.  I believe  it  is  necessary  to  consider  the  type  of  habitat  in 
which  all  of  the  species  in  this  host-parasitoid  complex  existed  before  agricultural  practices 
modified  large  areas.  Large  numbers  of  cutworm  species  probably  existed  in  low  densities  in 
the  prairie  and  parkland  areas  of  Canada.  The  densities  of  individual  host  species  were 
probably  low  enough  to  prevent  host-parasitoid  complexes  restricted  to  small  numbers  of 
species  from  occurring.  In  order  to  survive,  each  parasitoid  species  probably  had  to  attack 
any  host  in  a given  taxonomic  range  in  a given  habitat  type.  The  hosts  of  some  of  the  species 
include  ground  cutworms  and  climbing  cutworms  which  are  found  in  grasslands,  fields  or 
low  bush  open  areas. 

Cutworms  such  as  E.  ochrogaster  were  probably  adapted  to  feeding  in  areas  of  recently 
disturbed  soil.  This  is  reflected  by  the  preference  of  the  female  to  lay  her  eggs  in  loose  soil. 
One  of  the  parasitoids,  Copidosoma  bakeri,  is  also  restricted  to  areas  of  loose  soil.  McMillan 
(1930)  showed  that  C.  bakeri  was  capable  of  completing  development  in  species  normally 
found  only  in  packed  soil  but  that  in  the  field  they  are  unable  to  penetrate  the  soil  and  find 
the  eggs.  It  is  likely  that  the  high  degree  of  polyembryony  in  C.  bakeri  is  the  result  of  the 
unstable  nature  of  the  habitat  it  requires  and  the  relative  difficulty  of  finding  such  a habitat. 
As  agriculture  has  increased  the  amount  of  disturbed  soil  and  at  the  same  time  made  the 
cutworm  habitat  more  stable,  the  densities  of  both  the  host  species  and  their  parasitoids 
have  probably  increased. 

Regulation  of  E.  ochrogaster  populations  by  parasitoids 

Nine  species  of  parasitoids  were  reared  from  E.  ochrogaster  during  this  study.  The  esti- 
mated percentages  of  hosts  killed  in  the  1967  and  1968  seasons  are  given  in  Table  2.  These 
estimates  were  calculated  using  the  number  of  hosts  killed  by  a parasitoid  (or  which  con- 
tained immature  stages  of  that  parasitoid)  out  of  the  total  number  of  hosts  entering  the 
stage  which  the  parasitoid  normally  killed.  As  was  found  by  King  and  Atkinson  (1928), 
Gonia  aldrichi,  Meteorus  vulgaris , Campoletis  atkinsoni,  and  Copidosoma  bakeri  were  impor- 
tant parasitoids.  Interestingly,  King  and  Atkinson  found  no  ichneumonine  species  as  para- 
sitoids of  E.  ochrogaster,  while  I found  three.  Of  these,  Spilichneumon  superbus  was  the 
most  important  species  and  Diphyus  No.  1 may  be  important  in  other  years.  The  total 
ichneumonine  complex  is  an  important  regulatory  factor  in  the  population  fluctuations  of 
the  red-backed  cutworm.  A.  laeviceps  was  recorded  by  Strickland  (1923)  as  killing  5%  of 
the  red-backed  cutworms  in  1915  and  less  than  1%  in  1916.  In  1967,  the  number  of  fifth 
instar  larvae  collected  was  too  low  to  permit  estimation  of  the  mortality  caused  by  A.  laevi- 
ceps. In  1968,  only  one  collected  red-backed  cutworm  was  killed  by  A.  laeviceps.  It  does 
not  appear  as  if  this  species  was  important  in  regulating  the  numbers  of  E.  ochrogaster 
during  1967  and  1968. 

It  appears  from  my  study  and  from  the  work  of  King  and  Atkinson  (1928),  that  the 
parasitoids  are  not  the  chief  controlling  factors  of  E.  ochrogaster  populations,  but  are 
important  regulating  factors. 

Figure  67  shows  the  decrease  in  size  of  the  E.  ochrogaster  population  at  Calahoo  in  the 
summer  of  1968.  The  population  estimates  were  made  weekly  and  were  based  on  20  one- 
half  square  meter  samples.  These  samples  were  taken  using  a modified  random  sampling 
plan  within  a 100  quadrat  sampling  area.  The  sampling  area  was  100  meters  square.  The 
soil  was  removed  to  a depth  of  15  cm  and  sieved  using  a mechanical  shaker. 

The  decrease  shown  is  far  more  rapid  than  if  parasitoids  were  the  most  important  regula- 
ting factors.  Frank  (1971a)  does  not  feel  that  carabid  predators  are  the  controlling  factor 
either.  It  is  likely  that  the  reduction  of  population  is  due  to  a complex  interaction  of 


Parasitoids  of  Euxoa 


115 


predators,  diseases,  and  parasitoids.  All  of  these  factors  are  likely  directly  influenced  by  the 
weather  conditions  within  any  season.  Until  more  intensive  research  is  carried  out  on  the 
total  regulatory  complex,  the  most  important  controlling  factor  must  remain  in  doubt. 


Table  2.  The  estimated  mortality  caused  by  the  species  of  parasitoids  reared  from  E.  ochro- 
gaster  at  Calahoo  in  1967  and  1968. 


Species 

1967 

1968 

Gonia  aldrichi 

5 - 10% 

9% 

Eutanyacra  suturalis 

<5% 

3% 

Diphyus  No.  1 

<5% 

5% 

Spilichneumon  superbus 

5-10% 

10% 

Campoletis  atkinsoni 

- 

9% 

Apan  teles  laeviceps 

= 5% 

<1% 

Meteorus  vulgaris 

10% 

5% 

Copidosoma  bakeri 

35% 

22% 

Agamermis  sp. 

<1% 

0 

Economic  benefits  of  parasitoids 

In  addition  to  analyzing  the  effect  of  any  parasitoid  on  the  yearly  population  fluctuations 
of  E.  ochrogaster  the  influence  of  this  parasitoid  on  the  amount  of  damage  caused  in  any 
given  year  must  be  considered.  To  reduce  the  damage  done,  feeding  by  the  cutworms  must 
be  reduced  or  prevented.  Because  it  kills  the  early  instar  larvae  Campoletis  atkinsoni  essen- 
tially prevents  economic  damage,  and  its  rate  of  killing  hosts  is  its  rate  of  economic  return. 
Apart  teles  laeviceps,  which  kills  during  the  host’s  fifth  instar,  reduces  the  damage  done  by 
the  hosts  which  it  attacks.  A.  laeviceps  has  shown  a very  low  rate  of  attack  during  my  study. 
While  C.  atkinsoni  and  A.  laeviceps  reduce  the  loss  caused  by  their  hosts,  Copidosoma 
bakeri  increases  the  loss  by  the  current  generation.  McMillan  (1930)  found  that  hosts 
attacked  by  C.  bakeri  consumed  27.5%  more  food  than  did  normal  hosts.  If  a seventh  instar 
occurs,  this  extends  the  feeding  period  of  each  host.  As  C.  bakeri  attacks  at  a fairly  high 
rate,  the  increased  loss  due  to  its  presence  probably  exceeds  that  prevented  by  C.  atkinsoni 
and  A.  laeviceps.  The  other  parasitoids  kill  at  such  low  rates,  or  kill  after  the  host  damage 
has  occurred  and  appear  to  be  economically  neutral  i.e.  they  neither  increase  nor  decrease 
the  loss  during  that  season. 

Future  research 

My  approach  has  been  to  study  each  of  the  known  parasitoids  of  E.  ochrogaster  and  its 
geographic  range.  The  knowledge  of  the  parasitoid  complex  is  a reflection  of  the  area  where 
this  host  is  most  commonly  a pest,  namely  western  Canada.  It  appears  that  E.  ochrogaster , 
or  in  fact  any  economic  cutworm,  is  attacked  by  the  normal  parasitoid  complex  which 
attack  cutworms.  The  parasitoid  complex  of  any  given  host  could  change  or  be  very  different 
in  different  regions.  I believe  that  it  is  very  difficult  at  this  point  to  study  all  the  parasitoids 
of  any  host  which  covers  a wide  geographic  range.  A more  valuable  approach  may  be  to 


Campoletis  atkinsoni 


Bonne tia  comta 
Meteorns  vulgaris 
Apan  teles  laeviceps 
j ' Campoletis  atkinsoni 

ichneumonines 

Gonia  spp 
bombyliid  species  _ 
e§8  Netelia  sp. 

\ Copidosoma  bakeri 
adult  ^ 


Gonia  aldrichi 
Gonia  fusciocollis 
Villa  alternata 
Villa  fulviana 
Villa  lateralis 


pupa 

f Gonia  spp 

Poecilanthrax  willistoni 
Poecilanthrax  alcyon 
Gravenhorstia  propingua 
Eutanyacra  suturalis 
Spilichneumon  superbus 
Diphyus  No.  1 
Pseudambly  teles  subfuscus 


Microplitus  kewleyi 
Apan  teles  laeviceps 

Apan  teles  griff  ini 
Agamermis  sp. 
Periscepsia  helymus 
Periscepsia  laevigata 


Meteorus  vulgaris 


Copidosoma  bakeri 


Netelia  sp. 
Bonnetia  comta 

x 

Villa  moroides 


Fig.  66.  The  parasitoid  complex  of  Euxoa  ochrogaster.  Arrows  entering  the  circle  indicate  oviposition  by  the  parasitoids. 
Arrows  leaving  the  circle  indicate  the  stage  at  which  the  host  is  killed. 


NO/m2 
• 2.0  O 


o 


□ □ 


- 1 .5 


-1.0 


o 


o 


□ 


□ larvae 
V pupae 

O total 


-0.5 


V 


V 


V 

□ 


O 

V 


JUNE 

i 

1 4 


V 


2 1 


JULY  □ 

i i i i 

2 8 5 12  1 9 


vO  AUGUST 

— 1 □_! Cj i _J 

26  3 10 


Fig.  67.  The  decline  of  the  Euxoa  ochrogaster  population  at  Calahoo,  Alberta  in  the  summer  of  1968. 


Parasitoids  of  Euxoa 


1 17 


study  the  complex  of  hosts  and  parasitoids  in  any  given  region.  In  such  a study,  all  of  the 
parasitoids  of  all  the  cutworm  hosts  which  are  found  in  that  region  would  be  studied  and 
described.  As  a result  of  such  a study,  all  the  potential  parasitoids  of  any  cutworm  species 
found  within  that  region  would  be  known.  The  advantages  of  such  an  approach  are  that 
when  any  one  species  is  studied  some  of  its  actual  parasitoids  may  not  be  present  in  sampled 
populations.  If  several  species  of  hosts  are  studied  than  any  one  parasitoid  is  less  likely  to  be 
overlooked  and  thus  will  be  recognized  in  a future  outbreak  of  cutworms.  Also,  if  a different 
species  of  cutworm  appears  in  a region,  most  of  the  potential  parasitoids  will  be  easily 
recognized,  allowing  a meaningful  analysis  of  the  role  of  the  parasitoids  as  regulation  factors. 

ACKNOWLEDGEMENTS 

I would  like  to  acknowledge  the  help  which  W.  G.  Evans,  my  supervisor,  provided  during 
this  study.  I would  also  like  to  thank  J.  H.  Frank  for  his  valuable  help  and  advice.  Also,  I 
would  like  to  thank  M.  Wook,  F.  J.  McAlpine,  O.  Peck,  and  W.  R.  Mason  of  the  Canadian 
National  Collection  for  their  help  and  advice.  This  study  was  supported  in  part  by  a grant 
(No.  AR  67-31)  from  The  Alberta  Agriculture  Research  Trust. 

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COLLEMBOLA  ON  FLOWERS  ON  BANKS  ISLAND,  N.  W.  T. 


PETER  G.  KEVAN 
Plant  Research  Institute 
Central  Experimental  Farm 
Ottawa,  Canada 


Quaes tiones  entomologicae 
8:  121  1972 


Two  species  of  Collembola,  Entomobrya  comparata  Folsom  and  Corynothrix  borealis 
Tullberg  ( Entomobryiidae)  were  collected  from  the  flowers  of  Lesquerella  arctica  (Worm- 
skjold) S.  Watson  (Cruciferae)  where  they  were  feeding  on  pollen  directly  from  the  anthers. 
The  observations  compare  well  with  those  reported  earlier  from  northern  Ellesmere  Island. 

Deux  especes  de  collembole,  Entomobrya  comparata  Folsom  et  Corynothrix  borealis 
Tullberg  (Entomobryiidea)  ont  ete  recueillis  alors  qu’ils  se  nourrissaient  du  pollen  directe- 
ment  des  antheres  des  fleurs  de  Lesquerella  arctica  (Wormskjold)  S.  Watson  ( Cruciferae).  Ces 
observations  sont  en  accord  avec  celles  rapportees  precedement  du  nord  de  Vile  d' Ellesmere. 

Kevan  and  Kevan  (1970)  have  reviewed  the  literature  on  Collembola  as  visitors  to  flowers 
and  pollen  feeders,  and  reported  on  observations  from  around  Lake  Hazen,  Ellesmere  Island, 
N.  W.  T.  On  5 July  1970  on  Banks  Island,  N.  W.  T.,  I collected  1 1 Collembola  associated 
with  the  flowers  of  Lesquerella  arctica  (Wormskjold)  S.  Watson  (Cruciferae),  the  same  plant 
species  as  for  the  Lake  Hazen  Collembola-flower  association.  The  following  observations 
were  made  on  a stony  well-drained  ridge  of  the  coastal  escarpment  2 km  east  of  the  village 
of  Sachs  Harbour  (71°  59'N.,  125°  1 l'W.).  Lesquerella  arctica  was  flowering  elsewhere,  but 
Collembola  were  not  found  in  association. 

Two  specimens  of  Entomobrya  comparata  Folsom  (Entomobryiidae)  were  collected  and 
observed,  in  the  same  circumstances  as  described  for  this  species  around  Lake  Hazen.  Anoth- 
er was  seen,  but  it  escaped.  Eight  specimens  of  the  darker  Corynothrix  borealis  Tullberg 
(Entomobryiidae)  were  collected  from  the  flowers.  Of  these,  six  were  inside  the  corolla  and 
two  were  crawling  on  the  outside  of  the  petals.  One  other  was  found  on  the  ground  beside 
a flowering  plant.  Of  the  six  within  the  corollas,  three  were  watched  as  they  fed  on  pollen 
directly  from  the  anthers.  Their  postures  were  different  from  those  assumed  by  E.  com- 
parata. They  gripped  the  anther  and  filament  of  the  stamen  being  fed  at  so  that  their  bodies 
were  parallel  to  the  filament,  rather  than  curled  around  the  anther  as  E.  comparata.  One 
other  individual  of  C.  borealis  was  in  a similar  posture  but  its  mouthparts  were  applied  to 
the  stigma  and  its  body  parallel  to  the  style. 

Dr.  K.  Christiansen,  Grinnell  College,  Iowa,  kindly  identified  the  specimens  for  me  and 
examined  their  gut  contents  according  to  my  suggestions.  Both  specimens  of  E.  comparata 
and  seven  of  the  specimens  of  C.  borealis  had  pollen  of  L.  arctica  in  their  guts,  and  most 
were  well  stuffed.  One  individual  of  C.  borealis  had  also  ingested  some  xylem  vessel  ele- 
ments and  fungal  hyphae. 

The  dates  of  these  records  coincide  almost  exactly  with  the  suggested  “sensitive  period” 
for  E.  comparata  around  Lake  Hazen  (Kevan  and  Kevan,  1970).  Data  are  insufficient  to 
make  a statement  about  C.  borealis  in  this  regard.  Corynothrix  borealis  is  the  first  species  in 
this  genus  recorded  visiting  flowers  and  feeding  on  pollen  and  fits  within  the  group  of  light 
coloured  Collembola  (albeit  a little  browner  than  E.  comparata)  in  exposed  flowers  where 
they  would  be  least  conspicuous.  The  day  on  which  the  Banks  Island  collection  was  made 
was  heavily  overcast,  so  that  there  would  have  been  no  raised  intra-floral  temperature  to 
hold  the  Collembola  in  the  flowers. 


Kevan,  P.  G.  and  D.  K.  McE.  Kevan.  1970.  Collembola  as  pollen  feeders  and  flower  visitors 
with  observations  from  the  high  arctic.  Quaest.  ent.  6:31 1-326. 


REFERENCE 


123 


Book  Review 

JACOT-GUILLARMOD,  C.  F.  Catalogue  of  the  Thysanoptera  of  the  World.  Annals  of  the 
Cape  Provincial  Museums  (Natural  History).  Vol.  7,  Part  1 (1970),  p.  i-iii,  1-216;  Part  2 
(1971),  p.  217-515.  Published  jointly  by  the  Cape  Provincial  Museums  at  the  Albany  Mu- 
seum, Grahamstown,  South  Africa.  (No  price  given.) 

Judging  by  the  treatment  given  them  in  most  textbooks  of  general  entomology,  one  might 
think  the  Thysanoptera  to  be  a poorly  known  order  of  insects.  That  they  are  at  least  as 
studied  as  other  groups  of  comparable  size  should  quickly  become  apparent  on  perusal  of 
these,  the  first  two  of  a projected  six-  or  seven-  part  catalogue  of  the  order. 

In  his  introduction  to  Part  1,  Jacot-Guillarmod  outlines  his  methods  and  indicates  that 
his  goal  is  to  list  all  literature  on  the  species  treated  in  each  part  up  to  the  time  of  publica- 
tion. The  rest  of  Part  1 treats  the  sub-order  Terebrantia  except  for  the  family  Thripidae,  and 
Part  2 the  subfamilies  Panchaetothripinae  (=Heliothripinae),  and  Thripinae  (in  part)  in  the 
family  Thripidae.  Systematically-arranged  are  the  names  of  superorder,  order,  suborder, 
superfamilies,  families,  subfamilies,  tribes  and  subtribes.  Generic  names  are  listed  alphabeti- 
cally under  the  next  higher  category;  specific  names  under  the  genus  or  subgenus,  and  sub- 
specific names  under  the  species.  Every  publication  that  cites  a name  is  listed  under  the 
valid  name  of  the  species  in  chronological  order  and  each  reference  is  complete  except  for 
title.  Type-species  for  valid  and  invalid  genera  are  indicated;  the  locations  of  type-specimens 
are  shown,  and  distribution,  type-locality  and  habitat  are  given  for  each  species.  Invalid 
names  are  cross-indexed. 

This  is  a very  difficult  work  to  use  because  there  is  no  index  and  because  the  headings  of 
all  categories  above  the  genus  are  printed  in  similar-sized  type.  For  the  benefit  of  my  readers 
I here  list  the  names  of  the  higher  categories  and  the  pages  on  which  they  are  found:  Part  1 . 
Thysanoptera  = p.l;  Terebrantia  = 9;  Aeolothripoidea  = 15;  Aeolothripidae  =17;  Melan- 
thripinae  = 22;  Mymarothripinae  = 60;  Aeolothripinae  = 62;  Orothripini  = 63;  Franklino- 
thripini  = 86;  Aeolothripini  = 94;  Mesothripidae  (fossil)  = 174;  Palaeothripidae  (fossil)  = 
174;  Permothripidae  (fossil)  = 175;  Merothripoidea  = 176;  Merothripidae  =176;  Erotido- 
thripinae  = 177;  Merothripinae  = 178;  Thripoidea  = 185;  Heterothripidae  = 186;  Hetero- 
thripini=  188;  Opadothripini  = 211;  Uzelothripidae  = 216.  Part  2.  Thripidae  = 217;  Panchae- 
tothripinae = 225;  Thripinae  = 322;  Dendrothripini  = 324;  Sericothripini  = 356;  Sericothri- 
pina  = 358;  Chirothripini  = 436. 


Bruce  S.  Heming 
Department  of  Entomology 
University  of  Alberta 
Edmonton  7,  Alberta 


124 


Book  Review 

GILLETT,  J.  D.  1971.  Mosquitos.  (The  World  Naturalist  Series,  Ed.  Richard  Carrington), 
pp.  xiii  + 274+  22  Figs  + 38  plates.  468  refs.  London,  Weidenfeld  and  Nicolson.  Price  £5.90. 

Professor  Gillett  has  given  us  a readable  and  well-illustrated  account  of  the  life  of  mosqui- 
tos and  their  effects  on  human  affairs.  The  emphasis  is  on  their  ecology  and  behaviour,  and 
on  the  need  to  consider  the  population  as  well  as  the  individual. 

An  introductory  chapter  gives  an  outline  classification  of  mosquitos  and  notes  on  the 
history  and  distribution  of  the  family.  One  chapter  each  is  devoted  to  eggs,  larvae,  pupae, 
adults,  and  flight.  The  next  three  chapters,  on  the  ovarian  cycle,  the  circadian  rhythm,  and 
strains  and  species,  are  especially  interesting  as  Professor  Gillett  has  worked  extensively  in 
these  fields.  After  two  more  chapters,  on  parasites  and  predators  and  mosquitos  as  nuisance, 
the  remainder  of  the  text,  about  one  fifth  of  the  book,  is  devoted  to  mosquitos  and  disease 
and  mosquitos  and  history.  The  author  worked  for  many  years  at  the  East  African  Virus 
Research  Institute,  Entebbe,  Uganda,  and  was  one  of  those  responsible  for  elucidating  the 
roles  of  Aedes  africanus  and  Aedes  simpsoni  in  sylvan  yellow  fever.  The  account  of  this 
work,  and  of  other  mosquito-borne  viruses  is  well  worth  reading.  Malaria,  by  contrast,  is 
rather  briefly  dealt  with. 

The  author  states  that  he  set  out  to  draw  on  personal  experience  wherever  possible,  and 
the  book  is  enhanced  by  some  accounts  of  the  practical  difficulties  of  studying  mosquitos 
in  nature,  such  as  building  tree  platforms  25  meters  above  the  forest  floor  to  study  biting 
cycles  in  the  canopy. 

The  drawings  and  photographs  are  well  chosen  and  the  jacket  design,  a painting  of  the 
Brazilian  Sabethes  belisarioi  Neiva,  is  strikingly  attractive.  There  are  no  tables  of  data  in  the 
text  but  for  those  looking  for  more  detail  there  is  a list  of  some  500  references.  A curious 
feature  of  the  book  is  a series  of  9 appendices  listing  by  geographical  region  the  2,500  or  so 
known  species,  with  symbols  to  indicate  if  they  have  been  found  to  transmit  filariae,  malar- 
ia, or  viruses  over  any  part  of  their  range.  Since  these  appendices  cover  32  pages,  an  extra 
page  or  two  of  analysis  of  their  contents  would  have  been  helpful. 

I noticed  few  typographical  errors.  The  name  of  Dr.  C.  B.  Cuellar  is  misspelled  on  page 
ix  and  that  of  Jack  Colvard  Jones  on  page  104.  Male  Culiseta  inornata  (Will.)  do  not  find 
their  females  only  by  touch,  as  stated  on  page  104.  Kliewer  et  al.  (1966.  Ann.  Ent.  Soc. 
Amer.  59:  530)  have  shown  that  the  female  produces  a pheromone  which  attracts  the  male 
and  releases  his  sexual  response. 

In  his  final  chapter  Professor  Gillett  reminds  us  that  in  spite  of  the  great  expenditure  of 
money  and  effort  on  mosquito  eradication  schemes  no  species  has  actually  been  eradicated 
by  man.  The  only  complete  success  was  the  eradication  of  Anopheles  gambiae  from  Brazil, 
a species  introduced  there  only  ten  years  previously.  Only  by  understanding  the  ways  of 
mosquitos  can  man  learn  to  avoid  their  ravages.  It  is  noteworthy  that  four  of  the  most  severe 
biters  and  transmitters  of  disease,  A.  gambiae,  Aedes  aegypti,  Culex  pipiens  fatigans,  and 
Culex  tarsalis  owe  much  of  their  present  success  to  conditions  that  man  has  created  for 
them.  The  case  of  C.  tarsalis  will  be  particularly  hard  to  solve  since  its  spread  is  associated 
with  irrigation  schemes,  otherwise  worthwhile  ventures.  Insecticides  are  hardly  mentioned  in 
this  book,  a refreshing  change  for  those  of  us  who  are  accustomed  to  seeing  mosquitos  as 
figures  on  mortality  tables. 


J.  E.  Hudson 
Department  of  Entomology 
University  of  Alberta 
Edmonton  7,  Alberta 


125 


Book  Review 

WEISS,  E.  (Editor).  1971.  Arthropod  cell  cultures  and  their  application  to  the  study  of 
viruses.  Current  topics  in  Microbiology  and  Immunology.  Vol.  55.  Springer-Verlag,  Berlin, 
Heidelberg,  New  York,  xx  + 288  pp.,  151  figs.,  author  and  subject  indices.  Cloth.  $22.60 
(U.  S.). 

Although  this  book  presents  the  proceedings  of  a symposium,  it  is  much  more  than  the 
collection  of  papers  on  rather  unrelated  topics  that  usually  emanates  from  such  conferences. 
Its  contents  are  organized  into  1 1 chapters  with  the  following  titles:  1 . The  culture  of  cells 
from  insects  and  ticks;  2.  Analysis  of  cells  from  established  insect  cell  lines;  3.  Physiology  of 
cultivated  arthropod  cells;  4.  Arthropod  tissue  culture  in  the  study  of  arboviruses  and 
Rickettsiae:  A review;  5.  Propagation  of  arboviruses  in  Singh’s  Aedes  cell  lines;  6.  Growth  of 
arboviruses  in  arthropod  cell  cultures:  comparative  studies;  7.  Growth  of  viruses  in  arthro- 
pod cell  cultures:  applications;  8.  Homoptera  cell  culture  and  its  application  to  the  study 
of  plant  pathogens;  9.  Lepidoptera  cell  culture  and  its  application  to  the  study  of  plant 
viruses  and  animal  parasites;  10.  Drosophila  cell  culture  and  its  application  for  the  study  of 
genetics  and  virology;  and  1 1 . New  opportunities  in  biological  research  offered  by  arthropod 
cell  cultures. 

Each  chapter  contains  several  papers  on  topics  related  to  the  chapter  title.  References 
are  gathered  together  at  the  end  of  the  book.  The  approach  of  different  authors  varies:  some 
contributions  are  short  research  reports,  while  others  are  full  accounts  with  summaries  of 
previous  work  and  discussion  of  implications  for  other  fields  and  for  the  future. 

In  Chapter  2,  J.  L.  Vaughn  emphasizes  the  difficulties  inherent  in  research  in  invertebrate 
tissue  culture.  Accidental  contamination  of  cell  lines  with  microorganisms  or  with  cells  from 
other  lines  will  probably  become  a problem  as  more  lines  are  introduced  and  as  the  use  of 
these  proliferates.  Since  cells  in  culture  look  similar  regardless  of  their  source,  contamination 
of  this  kind  cannot  be  recognized  by  differences  in  cell  morphology.  A.  E.  Greene  and  J. 
Charney  received  a cell  culture  supposedly  from  Aedes  aegypti  in  their  lab  in  1967.  Using 
agar  gel  immunodiffusion  and  isoenzyme  analysis  they  showed  that  this  culture  had  been 
contaminated  and  replaced  by  cells  of  a moth,  Antherea  eucalypti.  They  indicate  also  how 
these  techniques  can  be  used  in  identifying  cultures  of  mammalian,  piscine  and  avian  origin 
and  in  separating  these  from  cells  of  arthropod  origin. 

Since  the  organs  and  tissues  of  animals  are  comprised  of  cells  organized  and  specialized  in 
particular  directions  and  because  the  form  of  these  structures  is  determined  by  the  genetic 
makeup  of  the  cells  interacting  with  the  environment  during  and  after  embryonic  develop- 
ment, T.D.C.  Grace  suggests  that  a good  way  to  study  these  phenomena  is  in  tissue  culture 
where  the  investigator  has  some  control  over  the  cells’  surroundings.  R.  L.  Seecof  and  R.  L. 
Teplitz  monitored  the  development  of  individual  cells  from  dissociated  embryos  of  Droso- 
phila. Some  of  these  divided  unequally  and  produced  long  extensions  in  culture.  Events 
similar  to  these  occur  in  the  development  of  the  central  nervous  system  from  neuroblasts 
during  normal  embryogenesis.  When  the  neuro  endocrine  organs  of  the  cockroach,  Leuco- 
phaea  maderae  were  explanted  in  culture,  they  continued  to  function  in  vitro  for  some  time 
(E.  P.  Marks). 

J.  H.  Conover  and  his  colleagues  succeeded  in  producing  bi-nucleate,  somatic  cell  hybrids 
between  mosquito  (Aedes  aegypti ) and  human  (He  La)  cell  lines.  When  control  He  La  cul- 
tures were  innoculated  with  polio  virus,  they  died  within  three  days  whereas  mosquito- 
human  hybrids  persisted  until  day  10.  Mosquito  (Aedes  aegypti)  cell  cultures  were  little 
affected  when  treated  with  organo-phosphate,  carbamate,  chlorinated  hydrocarbon,  arseni- 
cal, nicotine  and  pyrethrin  insecticides,  but  similar  amounts  of  these  chemicals  applied  to 
larvae  killed  them  (T.D.C.  Grace  and  J.  Mitsuhashi). 


126 


There  are  two  reviews  in  the  book:  one  by  C.  E.  Yunker  on  the  culture  of  arboviruses 
and  Rickettsiae  in  cultured  cells  and  the  other  by  H.  Hirumi  on  the  use  of  homopteran  cell 
cultures  in  the  study  of  plant  pathogens.  Yunker  points  out  that  two  thirds  of  all  published 
work  in  the  area  of  his  review  was  done  in  1968  and  1969.  From  this  work  he  concludes, 
among  other  things,  that  primary  cultures  of  arthropod  tissues  will  support  growth  of 
viruses  that  that  particular  donor  arthropod  or  a relative  can  transmit.  Arboviruses  may 
propagate  to  a higher  degree  in  cultures  of  vector  tissues  than  in  the  cells  of  the  intact  ar- 
thropod. Since  primary  tick  cultures  and  established  insect  cell  lines  are  very  sensitive  to 
many  arboviruses  and  Rickettsiae,  they  may  be  used  to  detect  these  pathogens  at  lower  con- 
centrations than  do  techniques  (animal,  egg,  or  vertebrate  culture)  now  used. 

Since  there  is  little  knowledge  of  how  plant  virus  particles  penetrate  and  multiply  in  the 
cells  of  their  vector  species,  Hirumi  suggests  that  the  successful  culture  of  these  microorgan- 
isms in  vector  cell  cultures  is  a promising  avenue  of  research.  Both  virus-vector  and  myco- 
plasma-vector interactions  have  been  studied  in  cultured  embryonic  cells  of  leafhoppers, 
aphids  and  planthoppers. 

Although  most  articles  in  the  book  deal  with  virus-cell  culture  interactions,  one  by  T.  J. 
Kurtti  and  M.  A.  Brooks  summarizes  their  successful  culture  of  Glugea  disstriae,  a micro- 
sporidian  protozoan,  in  cell  cultures  of  Malacosoma  disstria  and  M.  americanum.  Since  this 
microorganism  is  a naturally-occurring  parasite  of  these  insects,  eventually  we  may  be  able 
to  use  tissue-culture  techniques  in  the  mass-production  of  this  and  other  protozoan  parasites 
for  biological  control  of  pest  species. 

C.  Barigozzi  summarizes  the  advantages  of  using  Drosophila  cell  culture  for  genetical 
studies.  Molecular  biologists  are  beginning  to  switch  their  activities  from  procaryote  to 
eucaryote  organisms  because  of  their  increasing  interest  in  the  factors  controlling  develop- 
ment in  higher  organisms.  Drosophila  cell  lines  have  advantages  over  vertebrate  ones  for 
biochemical  studies  of  this  kind  because  the  genetics  of  this  genus  is  so  well  understood. 
Particularly  useful  is  the  occurrence  of  somatic  pairing  and  the  ability  to  induce  crossing- 
over  in  these  cells  by  X-ray  irradiation  as  has  been  shown  by  H.  A.  Schneiderman  and  his 
students  at  Irvine,  California.  Such  techniques  could  be  put  to  good  advantage  in  Drosophila 
cell  lines  since  an  accurate  genetic  analysis  is  aided  by  crossing-over. 

The  future  of  tissue  culture  is  speculated  upon  by  B.  W.  Schlesinger  and  W.  Trager  in  the 
final  chapter.  Schlesinger  suggests  that  culture  work  with  viruses  may  eventually  shed  some 
light  on  the  evolutionary  origin  and  relationships  between  different,  arthropod-transmitted 
plant  and  animal  viruses.  These  viruses,  as  he  emphasizes,  are  the  only  ones  known  to  bridge 
the  evolutionary  gaps  between  kingdoms  (animals  and  plants)  and  phyla  (arthropods  and 
vertebrates).  The  ability  to  switch  such  viruses  back  and  forth  between  vertebrate  and  ar- 
thropod cells  under  controlled  conditions  should  help  us  to  understand  both  of  these  sub- 
jects. He  speculates  on  the  evolutionary  origin  of  viruses  and  asks  questions  which  may  be 
answered  with  culture  techniques. 

Since  Trager  was  the  first  (1935)  to  culture  viruses  (nuclear  polyhedrosis)  in  invertebrate 
( Bombyx  mori)  tissue  culture,  it  is  fitting  that  he  should  have  the  final  say  in  this  book. 
He  predicts  that  major  breakthroughs  in  the  understanding  of  parasitic  protozoan  life  cycles 
(e.g.  malaria,  sleeping  sickness),  insect  mycetomes  and  their  function,  and  insect  devel- 
opment will  follow  from  increased  work  in  insect  tissue  culture.  Why,  he  asks,  do  the  cells 
of  larval  Cyclorrhapha  stop  dividing  in  the  egg  and  subsequently  develop  through  an  increase 
in  cell  size  and  chromosomal  polyteny?  This  does  not  occur  when  embryonic  cells  of  these 
insects  are  cultured;  when  explanted  they  continue  to  increase  in  number  by  mitosis.  He 
completes  his  discussion  with  reference  to  the  work  of  Hadorn  and  his  students  on  determi- 
nation and  transdetermination  in  Drosophila.  The  whole  theory  of  structural  homology,  so 


127 


important  in  comparative  morphological,  palaentological,  evolutionary,  and  systematic  stud- 
ies, can  be  thrown  into  question,  if,  as  has  been  shown  experimentally,  cells  determined  to 
form  one  structure,  can  be  “transdetermined”  in  nature  to  form  almost  any  other  in  the 
body. 

This  book  should  have  wide  appeal.  Plant  pathologists,  microbiologists,  parasitologists, 
medical  entomologists,  and  developmental  biologists  will  profit  from  a careful  reading  of 
pertinent  parts  of  it.  The  book  is  well  produced  and  is  amazingly  free  of  typographical 
errors  (I  found  three)  considering  that  its  English  text  was  printed  in  Germany.  With  a few 
exceptions,  the  photomicrographs  are  excellent.  However  the  price  ($22.60)  indicates  why 
it  is  that  violation  of  copyright  is  an  increasing  problem. 


Bruce  S.  Heming 
Department  of  Entomology 
University  of  Alberta 
Edmonton  7,  Alberta 


ANNOUNCEMENT 

An  English  translation  of  Rohdendorf  s Historical  Development  of  the  Diptera  edited  by 
Harold  Oldroyd  and  Brian  Hocking  will  be  published  by  the  University  of  Alberta  Press  in 
the  Fall  of  1972.  A further  announcement  concerning  price  and  details  will  follow. 


Publication  of  Quaestiones  Entomologicae  was  started  in  1965  as  part 
of  a memorial  project  for  Professor  E.  H.  Strickland,  the  founder  of  the 
Department  of  Entomology  at  the  University  of  Alberta  in  Edmonton 
in  1922. 

It  is  intended  to  provide  prompt  low-cost  publication  for  accounts  of 
entomological  research  of  greater  than  average  length,  with  priority 
given  to  work  in  Professor  Strickland’s  special  fields  of  interest  including 
entomology  in  Alberta,  systematic  work,  and  other  papers  based  on  work 
done  at  the  University  of  Alberta. 

Copy  should  conform  to  the  Style  Manual  for  Biological  Journals 
published  by  the  American  Institute  of  Biological  Sciences,  Second 
Edition,  1964,  except  as  regards  the  abbreviations  of  titles  of  periodicals 
which  should  be  those  given  in  the  World  List  of  Scientific  Periodicals, 
1964  Edition.  The  appropriate  abbreviation  for  this  journal  is  Quaest.  ent. 
An  abstract  of  not  more  than  500  words  is  required.  All  manuscripts  will 
be  reviewed  by  referees. 

Illustrations  and  tables  must  be  suitable  for  reproduction  on  a page 
size  of  93/4x6%  inches,  text  and  tables  not  more  than  7%X43/4  inches, 
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when  proofs  are  returned,  and  will  be  supplied  at  cost.  Subscription  rates 
are  the  same  for  institutions,  libraries,  and  individuals,  $4.00  per 
volume  of  4 issues,  normally  appearing  at  quarterly  intervals;  single 
issues  $1.00.  An  abstract  edition  is  available,  printed  on  one  or  both 
sides  (according  to  length)  of  3X5  inch  index  cards  (at  $1.00  per 
volume)  or  on  5X8  inch  standard  single  row  punched  cards  ($1.50  per 
volume) . 

Communications  regarding  subscriptions  and  exchanges  should  be 
addressed  to  the  Subscription  Manager  and  regarding  manuscripts  to: 

The  Editor,  Quaestiones  Entomologicae, 
Department  of  Entomology, 

University  of  Alberta,  Edmonton,  Canada. 


E.b.-  qtil| 


Quaestiones 

MU3.  COMP.  ZOOL. 


entomologicae 


LIBRARY 

AUG  1 9 


HARVARD 

UNIVERSITY 


A periodical  record  of  entomological  investigations, 
published  at  the  Department  of  Entomology, 
University  of  Alberta,  Edmonton,  Canada. 


VOLUME  VIII 


NUMBER  3 


JULY  1972 


QUAESTIONES  ENTOMOLOGICAE 


A periodical  record  of  entomological  investigation  published  at  the  Department  of 
Entomology,  University  of  Alberta,  Edmonton,  Alberta. 

Volume  8 Number  3 18  July  1972 

CONTENTS 


Book  Review 129 

Whitehead  — Classification,  phylogeny,  and  zoogeography  of 

Schizogenius  Putzeys  (Coleoptera:  Carabidae:  Scaritini)  131 

Announcements  349 


Book  Review 

WILSON,  Edward  O.  1971.  The  Insect  Societies.  The  Belknap  press  of  Harvard  University 
Press,  Cambridge,  Massachusetts,  U.S.A.  pp.  x + 548.  $22.00. 

This  book  is  a very  impressive  achievement  for  a single  author.  E.O.  Wilson  states  at  the 
beginning  of  the  work  that  he  is  attempting  to  provide  a modern  synthesis  of  insect  socio- 
logy, and  to  present  the  knowledge  within  the  framework  of  the  concepts  of  population 
biology.  I think  he  has  succeeded  to  a remarkable  degree.  Almost  all  of  the  information 
reviewed  is  from  the  works  of  other  authors  and  the  range  covered  is  enormous.  The  biblio- 
graphy extends  over  55  pages  and  includes  references  published  from  the  seventeenth 
century  up  to  1971.  Such  coverage  has  often  resulted  in  books  becoming  a dull  catalogue 
of  abstracts,  but  the  author’s  control  of  his  material  has  produced  a very  readable  and 
coherent  account,  which  is  consistently  interesting  and  clear.  In  some  parts  the  book  is 
even  compulsive. 

Michener’s  classification  of  the  degrees  of  social  behavior  (Ann.  Rev.  Ent.,  14:299-342) 
is  adopted  for  use  throughout  the  book.  Four  chapters  are  devoted  to  descriptions  of  the 
social  organizations  found  among  wasps,  ants,  bees  and  termites  respectively.  The  accounts 
also  include  the  taxonomy  of  the  social  species,  what  is  known  of  the  fossil  records,  and 
current  and  previous  hypotheses  about  the  evolution  of  sociality  within  the  groups. 

An  interesting  chapter  on  pre-social  insects  follows,  bringing  together  information  on 
parental,  co-operative,  sub-  and  quasi-social  behavior  among  several  orders  of  insects  and  in 
spiders. 

Three  chapters  are  given  to  caste  in  ants,  bees  and  wasps,  and  termites,  respectively,  in 
which  the  evolution  and  determination  of  castes  and  the  division  of  labor  among  them  are 
discussed.  As  in  other  parts  of  the  book,  the  limits  of  current  knowledge  are  always 
stressed. 

The  sensory  physiology  and  mental  capacities  of  the  social  insects  are  reviewed  as  a 
prelude  to  discussions  of  the  communication  systems  employed  in  alarm  and  assembly, 
recruitment,  recognition,  food  exchange  and  grooming  which  make  up  the  next  three 
chapters.  The  author  emphasizes  the  way  in  which  complex  social  behavior  is  created  out 
of  the  relatively  simple  individual  reactions  of  colony  members  to  stimuli  from  the  rest  of 
the  colony  and  from  the  environment.  These  discussions  lead  into  the  chapters  on  group 
effects  and  the  control  of  nestmates  and  on  social  homeostasis  and  the  superorganism. 

Wilson  gives  considerable  attention  to  the  historical  importance  of  the  idea  of  an  insect 
colony  as  a superorganism  and  concludes  that  the  concept  has  lost  popularity  not  because  it 
is  wrong  but  because  it  has  become  irrelevant.  It  was  valuable  in  stimulating  interest  and 
research,  but  does  not  itself  contribute  towards  understanding  the  phenomena  which  have 


130 


Whitehead 


been  discovered  through  that  research. 

Hamilton’s  idea  of  the  importance  of  haplodiploidy  in  the  development  of  insect  sociality 
(J.  Theoret.  Biol.,  7:1-52,  1964)  is  among  those  dealt  with  in  a chapter  on  the  genetic 
theory  of  social  behavior.  The  intriguing  suggestion  that  because  hymenopteran  males  are 
haploid,  and  a female  thus  shares  more  genes  with  her  sisters  than  with  her  offspring, 
social  behavior  improves  her  chances  of  perpetuating  her  own  genes,  is  subjected  to  close 
scrutiny.  Predictions  which  should  follow  from  it  are  examined  using  available  data,  and 
Wilson  concludes  that  the  idea  can  be  provisionally  accepted.  But  he  stresses  that  multiple 
mating  by  the  queen  can  cancel  the  bias  unless  the  population  shows  low  dispersal  or 
much  interbreeding. 

A chapter  entitled  “Compromise  and  Optimization  in  Social  Evolution”  discusses  how 
social  organization  is  affected  by  the  environmental  circumstances  of  the  colony.  It  includes 
a review  of  Wilson’s  own  earlier  work  on  the  hypothesis  that  the  proportions  of  castes  in  a 
mature  colony  represent  an  ‘optimal  mix’  which  minimizes  the  ‘production  cost’  of  the 
new  virgin  queens.  The  constitution  of  the  optimal  mix  depends  on  the  degree  of  specializa- 
tion of  the  castes  and  varies  with  changes  in  the  environment.  The  small  amount  of  data 
which  supports  the  hypothesis  is  quoted,  but  it  will  be  very  difficult  to  prove  or  disprove. 

Two  chapters  on  symbioses  follow,  treating  relationships  among  the  social  insects  and 
with  other  arthropods  respectively.  Wilson  thinks  that  permanent  parasitism  of  one  ant 
species  on  another  can  be  reached  by  any  of  three  routes:  via  the  slave-making  habit,  via 
temporary  parasitism  in  colony  foundation,  or  via  xenobiosis,  the  habit  of  one  species  of 
living  within  the  nest  of  another.  He  gives  examples  to  support  his  opinion. 

The  penultimate  chapter  is  on  the  population  dynamics  of  colonies,  a study  which  the 
author  considers  to  be  the  next  essential  stage  in  accounting  for  the  observed  social  pheno- 
mena of  insects,  following  work  on  their  physiology.  The  chapter  covers  survivorship  of 
colony  members  and  of  colonies,  regulation  of  colony  growth,  competition  and  territori- 
ality, control  of  colony  destiny  and  species  diversity,  and  dispersal  of  colonies. 

The  concluding  chapter  relates  the  study  of  insect  societies  to  that  of  vertebrate  societies 
and  looks  forward  to  the  founding  of  a general  theory  of  sociobiology. 

Throughout  the  text  entomological  terms  are  explained  where  they  occur  and  no  assump- 
tions are  made  about  the  reader’s  background.  A glossary  is  provided.  It  is  clear  that  the  book 
is  intended  for  wider  readership  than  entomologists  only.  Each  chapter  can  be  read  and 
understood  independently,  although  references  are  made  to  other  chapters  which  discuss  in 
detail  themes  mentioned  in  passing.  This  design  leads  to  some  repetition  of  data,  especially 
in  the  section  on  behavior,  although  on  most  occasions  a new  facet  of  interest  is  revealed. 

All  the  many  illustrations  are  borrowed  from  earlier  works,  either  reproduced  directly 
or  modified,  and  are  mostly  good.  Some  take  the  form  of  original  drawings  by  Sarah 
Landry  composed  from  one  or  more  sources,  and  these  are  both  clear  and  pleasing. 

Wilson  also  quotes  many  passages  of  description  or  argument  directly  from  earlier  authors. 
Such  inserts  provide  a valuable  change  of  pace  for  the  reader  and  enhance  the  author’s  own 
style.  He  also  gives  his  personal  assessment  of  the  value  of  other  people’s  work,  and  in  cases 
where  no  data  are  available  to  enable  a mystery  to  be  explained,  makes  a suggestion  of  his 
own  towards  a solution. 

I estimate  that  it  took  me  about  thirty-one  hours  to  read  through  the  entire  book.  The 
price  of  the  volume  may  seem  high,  but  for  so  much  solid  entertainment  and  enlighten- 
ment it  compares  well  with  other  media,  even  without  counting  re-reading  time.  I would 
recommend  The  Insect  Societies  to  anyone.  You  would  actually  read  it. 

Doreen  Watler 
University  of  Alberta 


CLASSIFICATION,  PHYLOGENY,  AND  ZOOGEOGRAPHY  OF 
SCHIZO GENIUS  PUTZEYS  (COLEOPTERA:  CARABIDAE:  SCARITINI)* 


DONALD  R.  WHITEHEAD 
Department  of  Entomology 
University  of  Alberta 
Edmonton  7,  Alberta 


Quaes tiones  entomologicae 
8 : 131-348  1972 


North  and  Middle  American  species  of  Schizogenius  Putzeys  are  reviewed  in  detail,  and 
South  American  species  are  treated  provisionally.  The  genus  is  redefined,  characterized,  and 
illustrated.  Two  subgenera  are  Genioschizus  new  subgenus,  with  three  species  groups,  and 
Schizogenius  s.  str.,  with  21  species  groups.  In  all,  I recognize  68  species  and  two  subspecies. 
Keys  are  given  to:  subgenera;  described  species  groups,  species,  and  subspecies  of  Genio- 
schizus; same  of  North  and  Middle  American  Schizogenius  s.  str.;  and  species  groups  and 
most  described  species  of  South  American  Schizogenius  s.  str. 

I describe  as  new  the  following  27  taxa:  S.  crenulatus  chiapatecus,  S.  sculptilis,  S.  im- 
puncticollis,  S.  suturalis,  S.  jacarensis,  S.  bicolor,  S.  cearaensis,  S.  negrei,  S.  costipennis,  S. 
reichardti,  S.  ocellatus,  S.  lindrothi,  S.  brevisetosus,  S.  seticollis  vandykei,  S.  plurisetosus, 
S.  kulti,  S.  dilatus,  S.  tibialis,  S.  ozarkensis,  S.  planuloides,  S.  neovalidus,  S.  chiricahuanus, 
S.  pacificus,  S.  emdeni,  S.  scopaeus,  S.  falli,  and  S.  ochthocephalus.  Synonymies  proposed 
for  the  first  time  are:  S.  peninsularis  Van  Dyke  (=  S.  auripennis  Bates),  S.  angusticollis 
Putzeys  (=  S.  arechavaletae  Putzeys),  S.  validus  Fall  (=  S.  longipennis  Putzeys),  and  S. 
championi  Kult  (=  S.  pygmaeus  Van  Dyke). 

Given  for  each  species  are,  as  appropriate:  synonymic  list,  diagnostic  combination,  de- 
scription, discussion  of  variation,  etymological  derivation,  geographic  distribution  list,  col- 
lecting notes,  taxonomic  notes,  and  illustrations  of  important  structural  characteristics. 
Geographic  distributions  are  mapped  for  all  North  and  Middle  American  species.  Descrip- 
tions of  most  North  and  Middle  American  species  are  augmented  with  tables  of  descriptive 
statistics.  Results  of  detailed  statistical  analyses  of  geographic  variation  for  members  of 
some  North  and  Middle  American  species  groups  are  tabulated,  mapped,  and  discussed. 

A phytogeny  is  reconstructed  for  the  genus,  and  carefully  integrated  with  historical  zoo- 
geography. No  geographic  evidence  is  used  to  reconstruct  the  phytogeny  of  major  groups 
and  lineages;  phytogenies  derived  from  contrasting  phyletic  and  phenetic  techniques  are 
favorably  compared.  To  derive  the  phytogeny  of  members  of  the  truquii  lineage,  which 
includes  most  North  and  Middle  American  species  of  Schizogenius,  use  of  zoogeographic 
evidence  is  essential ; simple  cladistic  techniques  are  not  used  because  character  states  of  too 
many  characteristics  are  reversible.  An  average  time  between  dichotomies  in  the  recon- 
structed phytogeny  of  Schizogenius  is  about  3,000,000  years.  This  interval  is  used  to  inte- 
grate Schizogenius  phytogeny  and  zoogeography.  On  the  same  basis,  phytogenies  and  zoo- 
geographies of  Brachinus  and  Evarthrus  are  compatible.  Zoogeographies  o/Brachinus,  Evar- 
thrus,  and  Schizogenius  are  compared  for  North  and  Middle  American  faunas. 

Ancestral  Schizogenius  evolved  in  South  America  about  Middle  Eocene.  Ancestors  of  the 
ferrugineus  group,  the  truquii  lineage,  and  the  crenulatus  group  entered  North  America  in 
Late  Eocene,  Middle  Oligocene,  and  Middle  Miocene.  The  ancestor  of  the  truquii  lineage 
ultimately  evolved  into  eight  species  groups,  and  their  evolutionary  zoogeographies  are  dis- 
cussed in  detail.  Since  Early  Pleistocene,  members  of  the  tenuis,  optimus,  and  lindrothi 
groups  have  crossed  the  Panamanian  land  bridge  to  Middle  America,  and  members  of  the 
depressus  group  have  spread  southward  into  South  America. 

* Revised  version  of  thesis  submitted  to  the  Faculty  of  Graduate  Studies  and  Research  in  partial  fulfillment  of  require- 
ments for  the  degree  of  Doctor  of  Philosophy  at  the  University  of  Alberta.  The  author’s  present  address  is  c/o  Department 
of  Entomology,  United  States  National  Museum,  Washington,  D.  C.  20560. 


132 


Whitehead 


En  este  trabajo  se  revisan,  en  detalle,  las  especies  de  Schizogenius  Putzeys  de  Norte  y 
Centro  America,  en  tanto  que  las  especies  de  Sur  America  son  revisadas  provisionalmente.  El 
genero  es  redefinido,  sehalado,  e ilustrado.  Dos  subgeneros  son  el  Genioschizus,  subgenero 
nuevo,  con  tres  grupos  de  especies;  y el  Schizogenius  s.  str.,  con  21  grupos.  En  total,  yo 
reconozco  68  especies  y dos  subespecies.  Se  han  ofrecido  claves  para  los  subgeneros,  grupos 
de  especies,  especies,  y subespecies  del  Genioschizus;  lo  mismo  que  para  el  Schizogenius 
s.  str.  de  Norte  y Centro  America,  asi  como  para  los  grupos  de  especies,  y la  mayor  parte 
de  las  especies  descritas  en  Sur  America. 

Yo  describo  como  nuevos  los  siguientes:  S.  crenulatus  chiapatecus,  S.  sculptilis,  S.  impunc- 
ticollis,  S.  suturalis,  S.  jacarensis,  S.  bicolor,  S.  cearaensis,  S.  negrei,  S.  costipennis,  S.  reich- 
ardti,  S.  ocellatus,  S.  lindrothi,  S.  brevisetosus,  S.  seticollis  vandykei,  S.  plurisetosus,  S.  kulti, 
S.  dilatus,  S.  tibialis,  S.  ozarkensis,  S.  planuloides,  S.  neovalidus,  S.  chiricahuanus,  S.  pacifi- 
cus,  S.  emdeni,  S.  scopaeus,  S.  falli,  y S.  ochthocephalus.  Son  sinonimias  nuevas:  S.  penin- 
sularis  Van  Dyke  (~  S.  auripennis  Bates),  S.  angusticollis  Putzeys  (=  S.  arechavaletae  Putzeys), 
S.  validus  Fall  (=  S.  longipennis  Putzeys),  y S.  championi  Kult  (=  S.  pygmaeus  Van  Dyke). 

Para  cada  especie  se  ha  ofrecido,  como  conviene:  lista  de  sinonimias,  combinacion  de 
diagnostico,  descripcion,  discurso  de  variacion,  derivacion  etimologica,  lista  de  localidades, 
notas  sobre  coleccion  y taxonomia,  e ilustraciones  de  caract  eristic  as  esenciales.  Igualmente 
se  dan  cartas  de  distribucion  para  las  especies  de  Norte  y Centro  America.  Las  especies  de 
Norte  y Centro  America  han  sido  provistas,  en  su  descripcion,  con  tablas  de  estadisticas 
descriptivas;  y los  resultados  de  analisis  estadisticos  detallados,  son  catalogados,  delineados 
en  cartas,  y discutidos. 

Se  ha  reconstruido  una  filogenia  para  el  genero,  y,  cuidadosamente,  se  ha  integrado  esta 
con  una  zoogeografia  historica.  No  se  ha  usado  ninguna  evidencia  geografica  para  la  recons- 
truccion  de  la  filogenia  de  grupos  y los  linajes  mayores;  las  filogenias  derivadas  de  tecnicas 
fileticas  y feneticas  son  comparadas  favorablemente.  Para  derivar  la  filogenia  de  miembros 
del  linaje  truquii,  que  contiene  la  mayor  parte  de  las  especies  del  Schizogenius  de  Norte  y 
Centro  America,  el  uso  de  evidencia  zoogeografica  es  indispensable.  No  se  han  usado  tecni- 
cas feneticas  sencillas  debido  a que  los  estados  de  caracter  con  demasiadas  caracteristicas, 
son  reversibles.  Un  intervalo  proporcional  entre  dicotomias  en  la  filogenia  reconstruida  del 
Schizogenius  se  aproxima  a los  3,000,000  de  anos.  Este  intervalo  es  usado  para  integrar  la 
filogenia  con  la  zoogeografia  del  Schizogenius.  Igualmente,  las  filogenias  y zoogeografias  del 
Brachinus  y el  Evarthrus  son  compatibles.  Las  zoogeografias  del  Brachinus,  el  Evarthrus,  y 
el  Schizogenius  son  comparadas  para  las  faunas  de  Norte  y Centro  America. 

El  Schizogenius  ancestral  se  desarrollo  en  Sur  America  alrededor  del  Eoceno  Medio.  Los 
predecesores  del  grupo  ferrugineus,  del  linaje  truquii,  y del  grupo  crenulatus,  penetraron  en 
Norte  America  durante  el  Eoceno  Ultimo,  Oligoceno  Medio,  y Mioceno  Medio.  El  predece- 
sor  del  linaje  truquii  finalmente  desarrollado  en  ocho  grupos  de  especies,  y sus  zoogeografias 
evolutivas  son  discutidos  en  detalle.  Desde  el  Pleistoceno  Primario,  miembros  de  los  grupos 
tenuis,  optimus,  y lindrothi  han  atravesado  el  puente  de  tierra  de  Panama  hacia  Centro 
America,  y miembros  del  grupo  depressus  se  han  distribuido  en  el  interior  de  Sur  America. 

CONTENTS 


Materials 133 

Methods  135 

Taxonomy 142 

Phylogeny 302 

Zoogeography 326 

References 345 


Classification,  phylogeny  and  zoogeography  of  Schizogenius 


133 


Introduction.  — Small  American  beetles  of  the  tribe  Scaritini  have  received  little  careful 
attention.  This  is  unfortunate  since  as  a result  no  other  aspects  of  their  biologies  have  been 
studied,  but  fortunate  in  that  comparatively  few  taxonomic  errors  have  been  introduced  into 
the  literature.  Lindroth  (1961 ) wrote  a fine  revision  of  the  Canadian  and  Alaskan  members  of 
this  fascinating  group  of  beetles.  His  treatment  of  Schizogenius,  the  subject  of  my  investiga- 
tions, is  excellent  for  the  limited  Canadian  fauna  but  is  not  adequate  to  identify  specimens 
from  most  regions  in  the  United  States.  The  genus,  after  all,  is  mainly  Neotropical,  and  in  fact 
a good  taxonomy  for  it  must  be  based  on  the  entire  fauna.  Except  for  Lindroth’s  (1961 ) key 
to  described  species  from  America  north  of  Mexico,  the  most  recent  full  revisions  of  Schizo- 
genius were  done  by  Putzeys  (1866)  for  Mexico  and  southward,  and  by  LeConte  (1879) 
for  America  north  of  Mexico. 

How  much  is  known  about  these  beetles?  Since  Thomas  Say  named  the  first  species  in 
1823,  little  more  than  descriptive  morphological  work  has  been  done.  Immature  stages 
remain  completely  unknown,  and  aside  from  collector’s  notes  on  habitat  we  have  no  in- 
formation on  adult  biology.  The  earliest  names  for  species  now  placed  in  Schizogenius  were 
proposed  by  Say  (1823),  Castelnau  (1835),  and  Haldeman  (1843).  Putzeys  (1846)  intro- 
duced the  generic  name  Schizogenius,  and  in  this  and  subsequent  important  papers  (1863, 
1866,  1878)  he  named  numerous  species.  Meanwhile,  LeConte  (1848,  1852,  1857,  1863) 
studied  Nearctic  species,  and  in  1879  summarized  his  knowledge  about  them  in  a key  and 
synonymic  list.  Bates  (1881,  1891)  treated  species  of  the  Mexican  and  Middle  American 
faunas  as  known  to  him.  Darlington  (1934,  1939),  Fall  (1901),  Kirsch  (1873),  Steinheil 
(1869),  and  Van  Dyke  (1925,  1949)  each  introduced  one  or  more  new  names  to  the  liter- 
ature. Kult  (1950)  produced  the  first  major  report  on  the  Neotropical  fauna  since  the 
nineteenth  century,  and  Lindroth  (1961)  did  likewise  for  the  Nearctic  fauna.  Since  I began 
my  studies  on  the  genus  I have  named  two  South  American  Schizogenius  species  as  new 
(1966a),  and  transferred  Darlington’s  S.  arenarius  to  Halocoryza,  a closely  related  genus 
which  I revised  and  characterized  (1966b,  1969). 

In  this  study  I treat  in  detail  the  taxonomy  and  distribution  of  Schizogenius  in  North 
and  Middle  America,  and  provide  the  basis  for  a similar  study  of  the  South  American  fauna. 
I define  “North  America”  as  America  north  of  Guatemala,  “Middle  America”  as  America 
from  Guatemala  to  Costa  Rica  plus  the  West  Indies,  and  “South  America”  as  South  America 
proper  plus  Panama.  I hope  that  my  work  will  not  only  make  possible  the  identification  of 
specimens,  but  that  it  will  stimulate  more  detailed  investigations  into  the  many  problems  in 
evolutionary  biology  that  render  members  of  this  genus  so  interesting.  Thus,  I attempt  to 
unravel  some  of  the  myriad  puzzles  in  speciation  in  North  and  Middle  American  Schizoge- 
nius, and  point  out  others  that  particularly  demand  attention. 

MATERIALS 

This  revision  is  based  on  the  study  of  9326  adult  specimens  of  North  and  Middle  Amer- 
ican Schizogenius,  plus  932  adults  from  South  America.  I have  collected  these  insects  along 
streams  in  much  of  the  United  States  and  Mexico,  and  other  collections  were  made  espe- 
cially for  this  study  by  G.  E.  Ball,  T.  L.  Erwin,  D.  H.  Kavanaugh,  D.  J.  Larson,  and  H.  B. 
Leech.  Other  specimens  were  loaned  to  me  by  various  museums  and  private  collectors;  the 
following  abbreviations  represent  these  collections  and  their  respective  curators. 

AMNH  American  Museum  of  Natural  History,  New  York,  New  York  10024;  P.  Vaurie. 

1 ANSP  Academy  of  Natural  Sciences,  Philadelphia,  Pennsylvania  19103;  H.  R.  Roberts. 

BMNH  British  Museum  (Natural  History),  London,  England;  P.  Hammond,  R.  D.  Pope. 

! CAS  California  Academy  of  Sciences,  San  Francisco,  California  94118;  H.  B.  Leech. 

! CNC  Canadian  National  Collection  of  Insects,  Entomology  Research  Institute,  Ottawa, 


134 


Whitehead 


Ontario;  E.  C.  Becker,  W.  J.  Brown. 

CNHM  Chicago  Natural  History  Museum,  Chicago,  Illinois  60605;  H.  Dybas. 

CPBo  C.  Bolivar  y Pieltain,  Instituto  Politecnico  Nacional,  Mexico,  D.  F. 

CUNY  Cornell  University,  Ithaca,  New  York  14850;  H.  Dietrich,  L.  L.  Pechuman. 

DHKa  D.  H.  Kavanaugh,  University  of  Alberta,  Edmonton  7,  Alberta. 

DJLa  D.  J.  Larson,  University  of  Calgary,  Calgary,  Alberta. 

DRWh  D.  R.  Whitehead,  University  of  Alberta,  Edmonton  7,  Alberta. 

FDAG  Florida  Department  of  Agriculture,  Gainesville,  Florida  32601;  R.  E.  Woodruff. 
GRNo  G.  R.  Noonan,  University  of  Alberta,  Edmonton  7,  Alberta. 

HFHo  H.  F.  Howden,  Carleton  University,  Ottawa,  Ontario. 

HGou  H.  Goulet,  University  of  Alberta,  Edmonton  7,  Alberta 

INHS  Illinois  State  Natural  History  Survey,  Urbana,  Illinois  61803;  L.  K.  Gloyd. 

IRSB  Institut  Royal  des  Sciences  Naturelles  de  Belgique,  31,  Rue  Vautier,  Bruxelles  4, 
Belgium;  R.  Damoiseau. 

JHen  J.  Hendrichs  S.,  Apartado  Postal  1 1-774,  Mexico  1 1,  D.  F. 

JNeg  J.  Negre,  9,  Boulevard  de  Lesseps,  Versailles,  France. 

KHSt  K.  H.  Stephan,  3038  East  Eastland,  Tucson,  Arizona  85716. 

KSUM  Kansas  State  University,  Manhattan,  Kansas  66502;  H.  D.  Blocker. 

LACM  Los  Angeles  County  Museum,  Exposition  Park,  Los  Angeles,  California  90007; 
C.  L.  Hogue. 

LBSC  Long  Beach  State  College,  Long  Beach,  California  90804;  E.  L.  Sleeper. 

MCZ  Museum  of  Comparative  Zoology,  Cambridge,  Massachusetts  02138;  P.  J.  Dar- 
lington. Jr.,  J.  F.  Lawrence. 

MGFT  Museum  G.  Frey,  Entomologisches  Institut,  Tutzing  bei  Munchen,  Germany;  M. 
von  Falkenhayn. 

MHNG  Museum  d’histoire  Naturelle,  Geneva,  Switzerland;  C.  Besuchet. 

MNHP  Museum  National  d’histoire  Naturelle,  Paris,  France;  A.  Bons,  J.  Negre. 

MSUL  Michigan  State  University,  East  Lansing,  Michigan  48823;  T.  F.  Hlavac. 

MZSP  Museu  de  Zoologia  da  Universidade  de  Sao  Paulo,  Caixa  Postal  7172,  Sao  Paulo, 
Brazil;  H.  Reichardt. 

OSUC  Ohio  State  University,  Columbus,  Ohio;  C.  A.  Triplehom. 

PSUU  Pennsylvania  State  University,  University  Park,  Pennsylvania  16802;  W.  W.  Boyle, 
S.  W.  Frost. 

RTBe  R.  T.  Bell,  University  of  Vermont,  Burlington,  Vermont  05401. 

RUNB  Rutgers  University,  New  Brunswick,  New  Jersey  08903;  J.  B.  Schmitt. 

TCBa  T.  C.  Barr,  University  of  Kentucky,  Lexington,  Kentucky  40506. 

TLEr  T.  L.  Erwin,  United  States  National  Museum,  Washington,  D.  C.  20560. 

UAFA  University  of  Arkansas,  Fayetteville,  Arkansas  72701 ; R.  T.  Allen. 

UASM  University  of  Alberta,  Strickland  Museum,  Edmonton  7,  Alberta;  G.  E.  Ball. 
UATA  University  of  Arizona,  Tucson,  Arizona  85721;  F.  G.  Werner. 

UCB  University  of  California,  Berkeley,  California  94720;  J.  A.  Powell. 

UCD  University  of  California,  Davis,  California  95616;  R.  O.  Schuster. 

UKSM  University  of  Kansas,  Snow  Museum,  Lawrence,  Kansas  66045;  G.  W.  Byers. 
UMCG  University  of  Miami,  Coral  Gables,  Florida  33124;  H.  F.  Strohecker. 

USNM  United  States  National  Museum,  Washington,  D.  C.  20560;  P.  J.  Spangler. 

UWLW  University  of  Wyoming,  Laramie,  Wyoming  82070;  N.  L.  Marston. 

VMKi  V.  M.  Kirk,  North  Grain  Insects  Research  Laboratories,  Brookings,  South  Dakota 
57006. 

ZMLS  Zoological  Institut,  University  of  Lund,  Lund,  Sweden;  C.  H.  Lindroth. 


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135 


METHODS 


Dissecting  techniques 

Representative  female  specimens  of  subgenera  and  some  species  groups  were  relaxed  in 
boiling  water,  then  dissected  with  fine  forceps;  mouthparts,  wings,  and  ovipositors  were 
mounted  in  Hoyer’s  medium  on  slides,  studied  with  a compound  microscope,  and  drawn. 
As  I found  no  characteristics  useful  in  species  recognition,  these  dissections  were  not  done 
routinely  at  species  level. 

Male  genitalia  including  parameres  were  dissected  from  the  abdominal  apices  of  relaxed 
specimens,  examined  for  peculiarities  in  form,  and  mounted  in  Hoyer’s  medium  on  slides. 
Proper  positioning  was  obtained  by  spreading  the  drop  of  mountant  to  a thin  layer  before 
placing  the  cover  slip.  Two  or  three  days  after  mounting,  each  preparation  had  cleared  suf- 
ficiently for  study,  and  no  additional  clearing  techniques  were  used.  The  endophallus  was 
everted  in  at  least  one  specimen  of  nearly  every  North  and  middle  American  species,  speci- 
mens permitting,  by  hooking  and  pulling  the  apex  of  the  virga  with  a fine  pin.  I did  not 
attempt  this  for  most  South  American  species,  because  in  older  material,  particularly  speci- 
mens collected  in  other  than  ethyl  acetate,  incidence  of  damage  was  prohibitive.  Nor  was 
this  procedure  used  routinely  for  members  of  the  depressus  group,  because  of  their  general- 
ly small  sizes.  After  genitalia  were  studied  and  drawn,  they  were  removed  from  slides,  glued 
to  cards,  and  pinned  with  the  specimens. 

Measurements  and  statistics 

Mensurable  and  meristic  data.  — Certain  routine  measurements  used  in  species  descrip- 
tions and  in  variation  analyses  were  made  with  a Leitz  stereoscopic  microscope  at  a mag- 
nification of  50  diameters,  using  a micrometer  eyepiece  with  a scale  interval  of  0.025  mm. 
These  measurements  and  their  abbreviations  are:  DP,  maximum  depth  of  thorax  from 
intercoxal  process  of  prostemum  to  basal  carina  of  pronotum;  LE,  length  of  left  elytron 
along  suture  from  basal  tubercle  to  apex;  LH,  length  of  head  from  base  of  eye  to  antero- 
lateral angle  of  clypeus;  LP,  length  of  pronotum  along  midline;  PS,  minimum  distance 
from  apex  of  left  paramedian  longitudinal  sulcus  to  base  of  pronotum.  Ta,  length  of  hind 
tarsus,  claws  excluded;  Ti,  length  of  hind  tibia;  TL,  total  length,  combining  LH,  LP,  and 
LE;  WE,  maximum  width  across  closed  elytra;  WF,  minimum  width  of  head  between  eyes; 
WH,  maximum  width  of  head  across  eyes;  WP,  maximum  width  of  pronotum,  hind  angles 
excluded. 

Most  of  these  measurements  can  be  made  accurately,  but  fixed  reference  points  are  not 
available  for  intercoxal  process  in  DP,  apex  of  paramedian  sulcus  in  PS,  or  base  of  tibia  in 
Ti,  and  thus  these  measurements  are  less  accurate.  Least  accurate  is  the  Ta  measurement, 
made  from  base  of  article  one  to  apex  of  article  five  regardless  of  expansion  or  contraction 
between  articles. 

Some  of  these  measurements  were  used  to  obtain  ratios  which  help  express  body  propor- 
tions. These  ratios  are:  DP/LP,  relative  depth  or  convexity  of  thorax;  LE/WE,  form  of 
elytra;  LP/WE,  relative  size  of  pronotum;  LP/WP,  form  of  pronotum.  PS/LP,  relative  length 
of  paramedian  pronotal  sulcus;  Ta/Ti,  relative  length  of  hind  tarsus;  and  WF/WH,  relative 
eye  size.  The  range  of  variation  in  any  given  sample  depends  in  part  on  the  accuracy  of 
measurements  used  to  form  ratios;  for  example,  Ta/Ti  is  quite  inaccurate  and  thus  too 
variable  for  use  in  infraspecific  comparisons.  Except  for  LE/WE,  a ratio  used  only  for 
species  and  species  groups  for  which  it  had  been  used  before  (Kult,  1950),  these  proportions 
are  intended  to  represent  different,  independent  aspects  of  body  form. 

In  those  species  having  more  than  three  or  four  setae  on  the  elytral  disc,  counts  were 


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made  of  numbers  of  setae  on  intervals  three,  five,  and  seven  of  the  left  elytron.  These 
counts  were  totalled  when  used  for  statistical  analysis.  Similar  counts  were  made  of  num- 
bers of  marginal  pronotal  setae,  in  the  few  forms  having  more  than  the  standard  two  pairs. 

Although  I am  not  sure  that  variation  in  numbers  of  elytral  setae  exactly  follows  a normal 
distribution,  especially  when  numbers  are  small,  I treated  these  data  in  the  same  way  as  I 
did  the  mensurable  data.  Sokal  and  Rohlf  (1969)  suggest,  as  a rule  of  thumb,  that  the 
number  of  unit  steps  in  a range  of  variables  should  range  from  about  30  to  300.  Such  a 
broad  range  in  the  proportion  variables  would  require  greater  accuracy  in  the  original 
measurements  than  I found  practical.  Nor  do  numbers  of  setae  per  elytral  disc,  at  least 
in  most  species,  vary  to  this  amount.  But,  since  I use  statistics  more  to  discern  relation- 
ships than  to  determine  similarities,  I believe  descriptive  statistics  based  on  these  data  are 
adequate. 

Descriptive  statistics.  — Except  for  the  poorly  known  and  unusually  variable  S.  tibialis,  I 
give  a comprehensive  set  of  descriptive  statistics  for  one  sample  of  each  described  North  and 
Middle  American  species;  I do  not  do  so  for  South  American  species  because  of  inadequate 
material.  Mensurable  and  meristic  data  so  treated  include,  where  appropriate,  TL,  LE,  WH, 
WP,  WE,  setae  on  intervals  three,  five,  and  seven  of  the  left  elytron  and  their  total,  WF/WH, 
LP/WP,  DP/LP,  LP/WE,  LE/WE,  Ta/Ti,  and  PS/LP.  When  at  least  eight  specimens  made  up 
a sample,  I give  range  of  variation,  mean,  1 .5  standard  deviations  and  two  standard  errors 
(Hubbs  and  Hubbs,  1953),  and  coefficient  of  variability.  I use  1.5  rather  than  one  standard 
deviation,  since  it  is  more  critical  as  an  analytical  tool  (Mayr,  et  al.,  1953).  When  fewer  than 
eight  specimens  were  used  in  a sample,  I give  only  range  and  mean.  I do  not  use  coefficient 
of  difference,  a statistic  suggested  by  Mayr  (1969)  for  subspeciation  analysis,  because  my 
criteria  for  subspecies  differ  from  his. 

Ideally,  each  sample  included  at  least  20  males  collected  at  one  time  and  place.  These 
conditions  were  often  not  entirely  satisfied,  usually  because  of  inadequate  numbers.  In  those 
species  belonging  to  species  groups  which  do  not  have  reliable  external  characteristics  to 
distinguish  sexes,  sex  was  ignored.  For  some  species,  samples  of  adequate  size  could  be  made 
only  by  combining  specimens  collected  over  a more  or  less  extensive  area.  When  too  few 
specimens  were  available  for  adequate  statistical  treatment,  the  sample  comprised  all  speci- 
mens on  hand  regardless  of  sex  or  provenance. 

These  statistics  complement  species  characterizations,  and  are  used  for  comparisons  be- 
tween closely  related  species  or  between  species  groups.  They  are  not  intended  for  compar- 
isons between  less  closely  related  species.  Thus,  the  statistical  treatment  is  as  uniform  as 
possible  for  species  within  a species  group,  but  sample  compositions  or  characteristics 
studied  may  differ  for  different  species  groups. 

Analysis  of  geographic  variation.  — Where  problems  in  species  recognition  were  evident,  I 
carefully  studied  variation  among  population  samples  before  making  taxonomic  decisions. 
I chose  for  analysis  characteristics  which  could  be  easily  and  accurately  measured,  and  which 
promised  to  tell  the  most  about  geographic  variation.  These  were  LE,  WF/WH,  LP/WP, 
total  number  of  setae  on  disc  of  left  elytron,  and,  for  the  pluripunctatus  group,  PS/LP  and 
LP/WE.  If  two  samples  differ  from  one  another  by  non-overlap  of  two  standard  errors  from 
the  means,  I term  them  “statistically  significantly  different;”  this  is  equivalent  to  the  t-test 
at  0.05  probability.  I use  the  term  “taxonomically  significantly  different”  if  two  samples 
differ  from  one  another  by  non-overlap  of  1 .5  standard  deviations  from  the  means,  to  imply 
that  90%  or  more  specimens  from  one  sample  can  be  distinguished  from  90%  or  more  from 
the  other  sample  (Mayr  et  al.,  1953). 

Samples  were  composed  of  equal  numbers  of  females  and  males,  despite  probably  in- 
creasing sample  variance  and  thereby  decreasing  statistical  sensitivity,  in  order  to  increase 


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137 


the  numbers  of  samples  suitable  in  size  for  useful  comparisons.  These  samples,  designated  by 
numbers,  are  plotted  on  maps;  details  of  locality  and  composition  may  be  derived  by 
consulting  maps  and  distribution  lists.  Ideally,  each  sample  included  at  least  ten  of  each 
sex,  and  samples  having  fewer  than  five  of  each  were  not  analyzed.  To  insure  that  samples 
represented  biological  populations,  I tried  to  form  each  sample  from  specimens  collected  at 
one  time  and  place.  If  this  could  not  be  done,  then  date  of  collection  was  ignored.  When 
still  no  adequate  sample  could  be  made,  I drew  from  a wider  geographic  area.  These  latter 
samples  probably  represent  composite  biological  populations,  but  are  approximations  which 
can  be  modified  after  additional  collections  are  made.  Henceforth,  I use  the  word  “popula- 
tion” to  mean  the  statistical  population  from  which  a sample  was  drawn,  unless  otherwise 
specified.  Because  samples  were  limited  by  numbers  and  distributions  of  available  specimens, 
geographic  coverage  is  less  than  ideal.  Samples  form  a loose  network  over  the  geographic 
area  covered,  and  proximate  samples  may  not  represent  truly  proximate  biological  popula- 
tions. Thus,  statistical  data  among  proximate  samples  only  approximately  suggest  biologi- 
cal relationships,  and  should  be  reinterpreted  if  geographically  intermediate  samples  become 
available. 

Understanding  relationships  between  taxa  requires  knowledge  of  similarities,  but  within  a 
species,  or  between  closely  related  allopatric  phena,  similarity  alone  may  not  be  a useful 
yardstick  of  relationship  (Mayr,  1969).  Statistical  analysis  of  single  characteristics  yields 
limited  direct  information  about  relationships.  If  statistics  are  used  to  obtain  such  direct 
information,  statistical  data  should  represent  the  total  phenotype  and  be  studied  by  various 
procedures  of  multivariate  analysis  or  numerical  taxonomy  (e.g.,  Sokal  and  Sneath,  1963). 
In  an  analysis  of  geographic  variation,  however,  each  characteristic  should  first  be  considered 
independently,  since  characteristics  within  a species  may  vary  independently.  This  is  clinal 
variation,  or  the  geographic  variation  of  single  characteristics  (e.g.,  Mayr,  1969).  Only  after 
characteristics  have  been  studied  separately  may  they  be  profitably  studied  in  combination, 
as  for  example  by  hybrid  index  techniques  (e.g.,  Freitag,  1965).  I limited  my  statistical 
studies  of  geographic  variation  to  clinal  analyses;  more  sophisticated  procedures  were  un- 
necessary. 

Statistically  significant  differences  in  one  or  more  characteristics  between  population 
samples  of  a species,  or  between  closely  related  allopatric  phena,  are  evidence  of  evolution- 
ary divergence  between  them.  Indeed,  if  evolution  is  a dynamic  continuing  process,  some 
evolutionary  divergence  between  any  two  biological  populations  of  a species  should  take 
place,  given  sufficient  time.  Its  extent  would  depend  on  geographic  relationships  of  the 
populations,  duration  and  completeness  of  their  isolation,  and  various  environmental  factors. 
Speciation  occurs  when  two  populations  or  groups  of  populations  have  diverged  sufficiently 
to  acquire  reproductive  isolation,  and  when  they  have  lost  reproductive  links  through  other 
populations. 

Statistically  or  taxonomically  significant  differences  in  one  or  more  characteristics  be- 
tween samples  of  geographically  distant  populations  provide  no  information  about  presence, 
absence,  or  amount  of  gene  flow  between  them.  They  do  give  information  about  amount  of 
divergence  or  similarity,  but  not  about  biological  relationships,  and  without  additional  data 
from  intermediate  population  samples  they  yield  little  useful  evidence  either  for  or  against 
conspecificity.  Furthermore,  since  any  two  proximate  samples  may  not  represent  contiguous 
biological  populations,  the  biological  significance  of  whatever  statistical  difference,  or  indeed 
or  any  difference,  between  them  cannot  readily  be  interpreted.  More  meaningful  compari- 
sons may  be  made  of  two  samples  via  one  or  more  intermediate  samples.  From  them,  one 
may  predict  the  relative  extent  of  gene  flow  between  populations  represented  by  these 
samples.  The  intent  of  this  discussion  is  to  emphasize  that  the  terms  “statistically  significant” 


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Whitehead 


and  “taxonomically  significant”  indicate  amount  of  difference  or  similarity,  not  any  precise 
relationship.  These  points  are  illustrated  in  my  analysis  of  S.  pygmaeus  and  S.  scopaeus. 

I use  statistics  to  study  clinal  relationships  among  populations  within  species  and  be- 
tween closely  related  allopatric  species  or  phena,  in  order  to  determine  probable  biological 
relationships  of  samples  rather  than  similarities  or  differences.  These  clinal  studies  yield 
evidence  about  possible  gene  flow,  and  consequently  about  possible  reproductive  isolation. 
From  them,  one  can  reasonably  predict  whether  or  not  populations  are  conspecific,  given 
sufficient  data.  Mayr  (1969)  states  that  isolated  allopatric  populations  may  be  either  species 
or  subspecies,  and  that  a taxonomic  decision  may  be  arbitrary;  he  suggests  that  such  isolates 
are  best  treated  as  subspecies.  However,  I think  information  on  clinal  variation  is  relevant  in 
such  decisions.  If  the  nearest  populations  in  two  isolated  groups  of  populations  are  the  most 
similar,  or  at  least  show  no  evidence  of  divergence,  then  I would  expect  that  some  gene 
exchange  does  exist  and  that  the  populations  are  conspecific.  If  these  two  populations  are 
the  most  different  in  one  or  more  characteristics,  or  even  if  only  one  of  them  is  divergent, 
then  I think  that  gene  flow  is  unlikely  and  that  the  two  groups  of  populations  are  not 
conspecific.  Within  the  pluripunctatus  group,  an  example  of  the  former  situation  is  the 
taxon  S.  seticollis,  with  two  subspecies;  and  as  an  example  of  the  second,  I regard  the  taxa 
S.  plurisetosus  and  S.  multisetosus  as  separate  species.  If  intermediate  samples  diverge  in 
one  or  more  characteristics,  then  the  more  distant  samples,  regardless  how  similar,  may  not 
be  conspecific;  see  for  example  my  treatment  of  the  taxa  S.  pygmaeus  and  S.  scopaeus. 
And  if  the  intermediate  samples  converge,  then  the  more  distant  samples  probably  are 
conspecific,  regardless  how  dissimilar;  see  my  disposition  of  the  names  S.  pygmaeus  and 
S.  championi. 

In  a statistical  study  of  character  dines,  I think  the  most  useful  statistic  is  the  sample 
mean,  compared  with  means  of  proximate  samples.  Statistics  of  dispersion  from  the  mean 
are  more  useful  to  estimate  the  relative  significance  of  observed  differences  between  means 
than  to  estimate  absolute  differences.  They  suggest  whether  differences  between  means  are 
or  are  not  the  result  of  coincidence.  If  two  samples  differ  significantly  in  a particular 
characteristic,  but  an  intermediate  sample  differs  significantly  from  neither,  I conclude  not 
only  that  the  first  two  populations  have  diverged  but  also  that  gene  flow  still  exists  between 
them.  I think  this  pattern  is  unlikely  to  result  from  coincidence. 

Illustrations 

Line  drawings  were  made  on  paper  squared  to  a one  cm  grid,  with  the  aid  of  an  ocular 
grid  mounted  in  a Leitz  stereoscopic  dissecting  microscope.  I used  much  care  in  preparing 
these  drawings,  since  I used  them  to  make  direct  comparisons  of  details  of  form  and  struc- 
ture, and  to  supplant  verbal  description.  Homologous  structures  are  drawn  consistently  to 
the  same  scale  to  facilitate  comparisons.  Male  genitalia,  endophalli,  and  other  small  organs 
mounted  on  slides  were  drawn  from  an  optical  magnification  of  150  diameters,  and  their 
details  checked  at  still  higher  magnifications  with  a compound  microscope.  Partial  and 
entire  habitus  drawings  were  made  from  a magnification  of  50  diameters,  and  fine  detail  and 
shading  added  as  needed. 

Distribution  maps  are  given  for  all  North  and  Middle  American  species;  I do  not  give 
such  maps  for  South  American  species  since  most  are  known  from  too  few  localities  to  even 
begin  to  picture  their  distributions.  Special  maps  were  prepared  to  illustrate  clinal  relation- 
ships, where  studied,  and  also  to  illustrate  broad  zoogeographic  patterns. 

Criteria  for  species-group  and  genus-group  taxa 

For  deciding  interrelationships  of  specimens  and  populations  at  species  level,  I follow  a 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


139 


slightly  qualified  version  of  the  biological  species  definition  proposed  by  Mayr  (1963,  1969): 
species  are  populations  or  groups  of  populations  through  which  gene  flow  actually  or 
potentially  exists,  but  which  are  reproductively  isolated  from  all  other  populations.  The 
biological  species,  if  bisexual,  is  the  only  taxon  which  has  nonarbitrary  boundaries  at  any 
given  time  and  place;  it  is  nonarbitrary  as  to  what  is  included  and  as  to  what  is  excluded 
(Simpson,  1961).  However,  in  contemporary,  bisexual  organisms,  nonarbitrariness  may  be 
difficult  to  demonstrate  for  geographically  distant  populations,  since  information  about 
gene  flow  or  potential  reproductive  isolation  may  not  be  available.  For  example,  consider 
a series  of  populations  the  most  distant  of  which  are  similar  to  one  another  and  may  not 
be  reproductively  isolated,  while  the  most  proximate,  especially  when  parapatric  or  sym- 
patric,  are  dissimilar  and  reproductively  isolated.  The  entire  aggregate  of  populations  may 
fit  the  biological  species  definition,  but  the  most  potent  available  taxonomic  evidence 
is  the  reproductive  isolation  of  the  near  populations.  The  biological  species  definition 
cannot  here  be  strictly  applied,  for  lack  of  evidence,  and  I think  the  best  taxonomic 
solution  is  to  recognize  two  biological  species  where  there  may  really  be  only  one.  But, 
while  sympatry  is  a test  for  reproductive  isolation  between  two  populations,  it  may  not 
always  indicate  specific  distinctness.  If,  for  example,  the  two  sympatric  populations  were 
shown  to  be  end  points  in  a circle  of  races,  then  I would  regard  them  as  members  of  a 
single  species. 

Beyond  the  question  of  species  definition  is  that  of  practical  species  recognition.  For 
Schizogenius,  the  only  data  presently  available  are  from  adult  morphology  and  distribution; 
there  is  no  direct  information  about  reproductive  isolation,  at  least  between  allopatric 
populations,  and  indeed  the  limits  of  populations  are  unknown.  In  carabid  beetles,  the 
structures  of  greatest  use  in  species  recognition,  because  of  diversity  in  form,  are  often  in 
the  male  genitalia.  In  Schizogenius,  major  differences  in  male  genitalia  among  related  species 
are  exceptional,  and  so  I could  not  rely  upon  finding  them  to  distinguish  species.  In  the 
few  species  groups  where  major  differences  were  found,  however,  and  assuming  no  contrary 
evidence,  I accept  them  as  suitable  criteria  for  species  recognition.  When  species  are  sym- 
patric, there  is  usually  little  difficulty  in  their  recognition.  If  two  forms  differ  constantly  in 
one  or  more  ways,  and  if  there  is  no  geographic  or  biologic  evidence  to  the  contrary,  I 
assume  the  differences  are  maintained  through  reproductive  isolation  and  treat  the  two 
forms  as  separate  species.  Generally,  differences  between  sympatric  species  are  numerous 
and  well  marked.  When  similar  differences  distinguish  allopatric  forms,  and  again  if  there  is 
no  contrary  evidence,  I consider  the  allopatric  forms  to  represent  distinct  species.  When 
allopatric  forms  differ  less  markedly,  I use  analytical  statistics  to  determine  if  reproductive 
isolation  is  likely,  specimens  permitting.  When  two  allopatric  forms  cannot  be  linked  in 
some  reasonable  geographic  way  because  adjacent  or  intermediate  samples  are  divergent 
in  one  or  more  characteristics,  I regard  them  as  separate  species.  If  these  samples  do  not 
diverge,  and  particularly  if  they  converge,  I regard  the  allopatric  forms  they  represent  as 
conspecific. 

I use  the  subspecies  category  only  for  taxa  of  uncertain  status,  and  then  only  for  com- 
pleteness. All  or  nearly  all  individuals  of  a subspecies  are  morphologically  distinguishable 
from  those  of  an  allopatric  form,  yet  no  judgement  may  yet  be  made  as  to  biological  status 
in  terms  of  reproductive  isolation.  That  is,  there  is  a substantial  statistical  or  morphological 
gap  between  the  two  forms,  but  one  which  could  easily  be  bridged  by  collections  made  in 
intermediate  areas  if  the  two  forms  are  truly  conspecific.  If,  on  the  other  hand,  I have 
strong  evidence  of  gene  flow  between  allopatric  forms,  or  if  I have  the  direct  evidence  of 
hybridization  between  them,  I recognize  only  one  geographically  variable  taxon,  the  species. 
Aside  from  this  use  of  the  subspecies  category  as  an  expression  of  uncertainty,  I share 


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Whitehead 


Erwin’s  (1970)  view  that  naming  subspecies  is  undesirable. 

Unlike  species-group  taxa,  genus-group  taxa  are  arbitrarily  limited  as  to  inclusiveness 
(Simpson,  1961).  Most  taxonomists  agree  that  the  subgenus  or  genus  should  contain  an 
aggregate  of  related  species  separated  from  similar  aggregates  by  a gap.  It  is  not  agreed  how 
large  this  gap  should  be,  nor  is  it  agreed  how  inclusive  should  be  the  aggregates  so  separated. 
In  the  study  of  single  genera,  I think  the  existing  concept  of  the  genus  should  be  accepted 
unless  it  is  poorly  defined  or  clearly  unreasonable.  I have  elsewhere  (1966b)  segregated  and 
defined  the  genus  Halocoryza.  I see  no  reason  to  otherwise  modify  Putzeys’  (1846,  1863, 
1866)  concept  of  Schizogenius. 

A supraspecific  taxon  should  be  strictly  monophyletic,  according  to  Hennig  (1966). 
However,  especially  when  no  fossil  evidence  is  available,  I doubt  that  monophyly  can  be 
definitely  demonstrated  for  all  groupings;  and  other  objections  to  this  strict  requirement 
for  monophyly  have  been  expressed  by  Mayr  (1969).  I have  already  (1966b)  suggested  that 
Halocoryza  and  Schizogenius  share  a common  ancestry.  I think  that  Schizogenius  really  is 
monophyletic  but  Halocoryza  may  be  either  paraphyletic  or  monophyletic,  and  if  my 
interpretations  of  its  origin  and  relationships  were  correct  it  is  indeed  paraphyletic.  If  so, 
the  requirement  for  monophyly  indicates:  1,  a single  genus  comprising  Halocoryza  + Schizo- 
genius; 2,  the  Old  World  genus  Halocoryza  and  the  New  World  genus  Schizogenius;  or  3, 
three  genera,  the  Old  World  Halocoryza , New  World  Halocoryza ,”  and  Schizogenius.  I 
chose  none  of  these,  since  my  separation  of  Halocoryza  and  Schizogenius  better  reflects 
known  biological  and  zoogeographic  peculiarities.  Also,  the  relationships  I suggested  are 
based  more  on  conviction  than  on  demonstrable  fact;  quite  possibly  the  two  genera,  as  I 
define  them,  really  are  monophyletic  assemblages. 

Lindroth  (1969)  notes  that  recognition  of  subgenera  may  lead  to  chaos,  and  he  uses  in- 
formal “species  groups.”  I agree  that  one  should  not  formally  recognize  numerous  closely 
related  subgenera,  but  believe  that  use  of  the  category  may  be  justified  for  larger  genera 
if  done  sparingly  and  carefully,  and  in  accord  with  Simpson’s  (1961)  suggestions  about 
ranking  taxa.  I use  the  subgenus  category  for  major  monophyletic  lineages  which  are  ap- 
proximately equally  divergent  from  one  another,  yet  subordinate  in  my  concept  of  the 
genus.  I use  the  informal  species  group  category  for  minor  monophyletic  lineages  within 
subgenera. 

Taxonomic  methods 

Specimens  of  North  and  Middle  American  Schizogenius  were  first  sorted  into  presump- 
tively related  or  conspecific  complexes  based  on  external  morphological  similarity.  These 
complexes  were  further  sorted  by  geographic  locality,  and  the  original  sorting  then  refined. 
A preliminary  survey  of  the  male  genitalia  within  each  complex  was  then  made,  and  the 
complexes  refined  further,  as  necessary.  Relationships  and  status  of  specimens  within  each 
complex  were  then  decided,  based  on  my  criteria  for  species  and  subspecies.  If  sufficient 
specimens  were  available,  and  if  there  was  a problem  in  deciding  whether  forms  were  conspe- 
cific or  not,  I used  statistical  methods  to  analyze  variation  and  to  determine  probable  gene 
flow  patterns.  I examined  at  least  six  male  genitalia  for  each  species,  specimens  permitting, 
and  as  many  more  as  necessary  to  reach  appropriate  conclusions  about  variation.  Naming  of 
specimens  was  based  on  knowledge  of  type  specimens  of  all  previously  described  taxa,  and 
lectotypes  were  designated  as  required.  The  diacritical  mark  “(!)”  is  used  to  indicate  that 
I examined  the  type  specimen. 

Similar  methods  could  not  be  applied  throughout  to  the  South  American  specimens 
studied,  because  of  insufficient  numbers.  More  reliance  was  placed  on  absolute  superficial 
or  genitalic  similarity,  and  none  on  statistical  methods,  but  criteria  for  species  recognition 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


141 


remained  the  same.  No  subspecies  were  recognized.  When  taxonomic  decisions  could  not  be 
made,  specimens  were  left  undescribed  or  unassigned  to  species.  I studied  type  material  of 
some,  but  not  all,  described  South  American  species,  and  may  therefore  have  made  some 
errors  in  association.  I think  this  fault  is  outweighed  by  the  need  to  provide  the  basis  for 
future  study  of  the  South  American  fauna,  and  also  to  aid  in  understanding  the  North  and 
Middle  American  faunas. 

Simpson  (1961)  clearly  defined  zoological  classification:  it  is  the  ordering  of  animals 
into  groups  based  on  their  relationships.  Accordingly,  it  should  reflect  the  evolutionary 
history  of  animals.  Mayr  (1969)  wrote  that  the  major  purpose  of  zoological  classifica- 
tion is  to  provide  a system  for  maximum  information  storage  and  retrieval,  and  that  the 
most  efficient  classification  for  that  purpose  is  based  on  evolutionary  relationships.  I be- 
lieve that  the  classification  should  allow  one  to  make  and  test  predictions  about  zoo- 
geographic, biochemical,  chromosomal,  larval,  or  other  relationships  not  already  directly 
embodied  in  the  classification.  For  this  purpose,  a classificaiton  based  on  hypotheses  about 
evolutionary  relationships  is  the  most  efficient,  since  only  it  can  theoretically  contain  and 
yield  more  information  than  was  used  in  its  creation.  The  difference  between  a classifica- 
tion based  on  suspected  or  real  relationships  and  one  based  only  on  similarities  is  that 
the  former  is  structured  around  evolutionary  theory,  while  the  latter  is  a mechanical  struc- 
ture with  no  intended  theoretical  basis.  This  difference  may  be  not  or  scarcely  evident  in 
the  end  product,  but  I think  it  important.  I have  tried  to  provide  for  Schizogenius  a 
classification  with,  I hope,  some  predictive  value,  and  I hope  it  will  be  tested  in  future 
investigations. 

Mayr  (1969)  summarized  methods  used  in  zoological  classification.  I have  used  various  of 
them  in  various  combinations,  depending  on  the  particular  problem  at  hand.  I obtained 
much  useful  information  about  relationships  of  components  of  some  species  complexes  by 
using  methods  of  analytical  statistics.  These  methods  did  not,  however,  give  useful  informa- 
tion about  less  closely  related  forms.  Perhaps  some  of  the  various  techniques  of  multivariate 
analysis  would  be  useful  to  detect  these  more  distant  relationships,  but  I remain  uncon- 
vinced of  either  their  validity  or  utility  (Mayr,  1969).  Similarly,  I have  not  used  other 
numerical  or  deliberately  phenetic  methods,  except  some  methods  of  numerical  cladistics 
(Camin  and  Sokal,  1965).  I did  use  some  of  the  weighted  character  methods  suggested  by 
Mayr  (1969),  together  with  some  of  the  more  rigidly  cladistic  approaches  preferred  by 
Hennig  (1966).  I give  a more  detailed  discussion  of  phylogenetic  methods,  and  discuss 
relationships  between  classification  and  phylogeny,  in  later  sections. 

Taxonomic  literature  and  synonymic  lists 

Pertinent  taxonomic  works  are  listed  for  all  previously  described  taxa  in  Schizogenius  in 
synonymic  lists  for  those  taxa.  These  synonymic  lists  do  not  pretend  to  completeness.  In 
particular,  most  faunal  and  catalogue  listings  are  excluded. 

I made  no  effort  to  verify  locality  records  cited  in  faunal  works,  and  unless  such  records 
are  particularly  important  and  probably  correct,  references  to  those  works  are  excluded. 
Many  old  records  are  doubtless  erroneous;  for  example,  some  reports  of  S.  lineolatus  from 
eastern  North  America  are  doubtless  based  on  specimens  of  S.  sulcifrons.  Important  faunal 
works  that  I normally  omitted  from  synonymic  lists  include  those  by  Blatchley  (1910), 
Brimley  (1938),  Fattig  (1949),  Leonard  (1926),  and  Smith  (1910). 

Catalogue  listings  are  excluded,  since  they  neither  contribute  new  taxonomic  information 
nor  contain  useful  summaries  of  then  available  taxonomic  information  other  than  literature 
references.  Such  catalogues  are  those  by  Blackwelder  (1944),  Csiki  (1927),  Gemminger  and 
Harold  (1868),  Leng  (1920),  and  Leng  and  Mutchler  (1933). 


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TAXONOMY 

Genus  Schizogenius  Putzeys 

Schizogenius  Putzeys  1846:650.  Type  species.  - Schizogenius  strigicollis  Putzeys  1846: 
650  (subsequent  designation  by  Lindroth,  1961:164).  LeConte  1857:82.  Putzeys  1863: 
24.  Putzeys  1866:222.  LeConte  1879:34.  Kult  1950: 139.  Lindroth  1961:164. 

Diagnostic  combination.  — Clypeus  tridentate  apically;  frons  with  four  or  five  pairs  of 
longitudinally  directed  carinae  between  eyes;  lacinia  plurisetose  on  inner  and  outer  margins; 
gula  not  more  than  0.3  width  of  mentum;  pygidium  with  numerous  longitudinally  directed 
crenulate  carinae  near  middle;  stylus  and  coxite  of  ovipositor  fused,  plurisetose. 

Description.  — Small  to  medium  size,  LE  1.65  to  5.00  mm.  Body  pedunculate,  elongate, 
depressed  to  cylindrical.  Color  various,  maculate  or  not,  not  to  strongly  aeneous  or  metallic. 
Integument  shiny  or  dull;  microsculpture  varied  in  extent,  isodiametric,  often  useful  in 
species  recognition. 

Head.  Large,  prognathous.  Labrum  Fig.  1-3,  biemarginate  to  deeply  emarginate;  dorsal 
surface  with  seven  setae  in  front,  median  and  two  outer  ones  longest;  lateral  margins  each 
with  five  to  about  fifteen  pairs  of  frayed  or  bifid  setae,  anterior  pairs  curved  forward  and 
inward  over  mandibles.  Clypeus  with  two  strong  paramedian  teeth;  median  tooth  prominent 
in  most  species;  paramedian  carinae  varied,  oblique  or  arcuate,  joined  at  apex  with  median 
tooth  or  not;  median  field  triangular  or  hemicircular,  basal  width  varied;  clypeus  with  one 
pair  of  setae  basad  and  laterad  to  carinae.  Clypeal  suture  obsolete  to  sharply  engraved. 
Frontal  lobes  prominent.  Frons  with  four  or  five  pairs  of  longitudinally  directed  carinae 
between  eyes,  neither  perfectly  equidistant  nor  equally  raised,  appearance  of  frons  not 
evenly  convex;  median  sulcus  broader  than  outer  paramedian  sulci,  with  or  without  median 
carina;  carina  five  variable,  obsolete  or  not;  carina  six  obsolete  except  on  frontal  lobes,  not 
raised  above  dorsal  margin  of  eye.  Anterior  supraorbital  seta  set  in  front  of  carina  five; 
posterior  seta  set  between  bases  of  frontal  carinae  four  and  five.  Eyes  varied  in  size,  promi- 
nence, and  size  and  number  of  facets.  Neck  punctate  at  least  on  sides,  orbit  extended 
laterad  along  posterior  margin  of  eye.  Antennal  articles  five  to  ten  slightly  transverse  to 
quite  elongate,  moniliform  to  filiform;  scape  with  one  subapical  dorsal  seta;  pedicel  with 
one  ventral  seta,  or  bisetose  or  plurisetose  in  some  species;  articles  three  to  four  plurisetose 
or  pubescent;  articles  five  to  eleven  pubescent,  without  glabrous  areas.  Mandibles  Fig.  5-7, 
stout  and  broadly  curved  along  lateral  margin;  inner  ventral  margin  of  right  mandible  with 
small  tooth  near  middle;  scrobe  oblique.  Maxilla  Fig.  9-11,  terminal  article  of  palpus  swollen 
basally;  lacinia  with  apex  acute  and  abruptly  bent,  setose  on  outer  and  inner  margins. 
Labium  Fig.  13-15,  penultimate  article  of  palpus  bisetose.  Mentum  Fig.  17-20,  deeply 
emarginate  at  middle,  with  one  anterior  pair  of  paramedian  setae;  median  tooth  obsolete  to 
acute;  lateral  lobes  truncate  or  with  antero-lateral  angles  acutely  produced;  ventral  surface 
with  broad,  abruptly  depressed  concave  area,  limited  behind  by  arcuate  carina;  base  of 
mentum  with  one  pair  of  large,  pouch-like  sensory  pits,  and  one  pair  of  paralateral  setae. 
Submentum  with  one  paramedian  and  one  postero-lateral  pair  of  setae,  plurisetose  in  S. 
strigicollis  Putzeys.  Gula  narrow,  at  narrowest  part  0.05-0.30  width  of  mentum. 

Thorax.  Pronotum  slightly  to  moderately  transverse;  median  longitudinal  sulcus  not  bor- 
dered by  carinae;  paramedian  longitudinal  sulci  well  developed  in  most  species,  hooked 
basally;  paralateral  sulci  present  in  some  species;  lateral  grooves  shallow,  bordered  by  distinct 
carinae  in  some  species;  anterior  and  posterior  pairs  of  marginal  setae  present,  additional 
marginal  setae  present  in  some  species;  basal  carina  moderately  to  strongly  elevated  above 
margin;  disc  flattened  to  moderately  convex.  Prothoracic  pleuron  smooth,  without  longi- 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


143 


tudinal  ridge,  rugose  or  punctate  in  some  species.  Prosternum  strongly  compressed  between 
front  coxae;  posterior  process  broadened  and  convex,  without  setae  or  carinae.  Metepis- 
ternum  slender,  elongate.  Anterior  coxal  cavities  closed-separated-unbridged;  middle  and 
hind  coxal  cavities  disjunct-confluent  (see  Bell,  1967). 

Elytra.  Lateral  channel  varied  in  form,  slightly  narrowed  at  apex,  or  flared  and  with  one 
or  more  deep  subapical  pits  (Fig.  22);  umbilicate  series  of  punctures  unbroken.  Elytron 
with  whip-like  seta  at  base  of  interval  three;  disc  in  most  species  with  two  or  more  setigerous 
punctures  on  interval  three,  in  many  species  with  variable  numbers  on  intervals  five  and 
seven.  Intervals  one  to  six  flat  to  convex,  in  some  species  with  short  apical  carinae,  in  some 
species  entirely  carinate;  interval  seven  carinate  or  not;  interval  eight  carinate  at  apex,  in 
most  species  not  joined  at  apex  by  other  intervals.  Striae  punctate  at  least  basally,  faintly  so 
in  some  species;  striae  evident  to  apex,  in  most  species  deeply  impressed,  outer  and  inner 
striae  equally  engraved. 

Hind  wings.  Macropterous  and  probably  functional  in  all  North  and  Middle  American 
species,  brachypterous  in  at  least  one  South  American  species.  Venation  (Fig.  31-34)  of 
usual  carabid  type  except  wedge  cell  absent  and  oblongum  cell  hooked  in  front. 

Legs.  Front  tibia  anisochaetous,  with  four  evident  external  teeth;  subapical  spurs  varied 
in  size  and  form,  not  grossly  unequal;  ventral-basal  margin  with  three  or  four  setae.  Middle 
and  hind  tibiae  with  apical  spurs  slender,  inconspicuous.  Anterior  and  middle  tarsi  of  males 
of  many  species  slightly  dilated  and  more  densely  pubescent  ventrally  than  in  females,  but 
without  distinctive  adhesive  pads.  Hind  tarsi  narrow  and  nearly  glabrous.  Tarsi  moderately 
to  markedly  elongate,  hind  tarsus  more  than  0.55  length  of  hind  tibia;  article  one  of  hind 
tarsus  2.0  to  4.0  times  as  long  as  article  two.  Paronychia  obsolete  to  nearly  as  long  as  tarsal 
claws. 

Abdomen.  Sternum  three  with  one  pair  of  strong,  oblique  paramedian  carinae,  in  many 
species  strongly  curved  outward  at  apices.  Sterna  four  to  six  each  with  one  pair  of  para- 
median ambulatory  setae.  Sternum  seven  with  or  without  one  pair  of  paramedian  ambula- 
tory setae,  sexually  dimorphic  or  not,  and  two  pairs  of  approximately  equidistant  apical 
marginal  setae.  Sterna  four  to  seven  without  basal  transverse  impressions.  Pygidium  (Fig.  23- 
25)  with  two  or  more  pairs  of  paramedian  setae,  one  pair  of  large  setae  near  pygidial  glands, 
variable  numbers  of  microsetae,  and  conspicuous  crenulate  carinae  along  midline;  margin  of 
pygidium  entire  in  males,  but  serrate  or  crenulate  in  some  or  all  females  of  many  species. 

Male  genitalia.  Parameres  subequal,  slightly  asymmetric,  each  with  one  to  three  large 
apical  setae.  Median  lobe  arcuate  to  angulate,  in  most  species  nearly  symmetric,  not  con- 
stricted near  base,  base  not  lobate;  apical  third  compressed,  varied  in  form,  in  many  species 
strongly  deflected.  Endophallus  doubly  invaginated;  basal  stylets  of  varied  form;  no  flagel- 
lum; apical  brush  or  virga  without  large  spines;  dorsal  cap  sclerite  present;  basal  collar  spines 
present  in  some  species  (see  Whitehead,  1966a).  Abdominal  segment  nine  of  normal  carabid 
type,  ring  sclerite  complete. 

Female  genitalia  and  ovipositor.  Internal  genitalia  without  sclerotized  structures,  not 
studied  in  detail.  Ovipositor  (Fig.  27-29)  with  stylus  and  coxite  fused,  plurisetose  at  middle; 
setae  of  posterior  margin  of  valvifer  normally  in  two  groups;  paraprocts  absent;  proctiger 
articulated  with  upper  margin  of  valvifers. 

Immature  stages.  Unknown;  see  Vinson  (1956)  for  description  of  the  presumably  similar 
larva  of  Halocoryza. 

Etymology.  — Greek,  schizo  = split,  plus  genio  = chin  or  mentum;  in  reference  to  form  of 
mentum. 

Distribution.  — Members  of  this  genus  are  known  from  continental  North,  Middle,  and 
South  America,  from  southern  Canada  to  central  Argentina;  one  species  is  endemic  to  Cuba. 


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Separation  of  Schizogenius  and  Halocoryza 

In  my  review  of  Halocoryza  (Whitehead,  1966b),  I stated  that  members  of  the  genus 
Schizogenius  were  best  distinguished  by  uni-  or  bisetose  antennal  pedicels  and  striate  pygi- 
dia.  However,  at  least  one  South  American  species  has  plurisetose  antennal  pedicels,  proba- 
bly secondarily  so.  Also,  American  Halocoryza  species  have  striate  pygidia,  but  the  striations 
are  inconspicuous  and  weakly  developed  (Fig.  26).  These  striae  are  actually  rows  of  tuber- 
cles, file-like  ridges  possibly  used  for  stridulation;  in  this  work  I call  them  “crenulate 
carinae.” 

Several  other  differences  listed  in  that  paper  are  apparently  without  exception.  The 
lacinia  of  Schizogenius  is  setose  only  on  the  outer  margin.  The  gula  of  Schizogenius  is  much 
narrower  than  that  of  Halocoryza  (Fig.  21).  Another  important  difference,  not  recognized 
previously,  is  that  in  Halocoryza  the  fused  stylus  and  coxite  of  the  ovipositor  has  one  large 
seta  (Fig.  30).  I found  no  characters  in  labrum  (Fig.  4),  mandibles  (Fig.  8),  labium  (Fig.  16), 
or  wing  useful  to  distinguish  Schizogenius  from  Halocoryza,  but  in  some  forms  of  H.  arena- 
ria  the  apical  part  of  the  wing  is  reduced. 

Key  to  subgenera  of  Schizogenius 

I here  recognize  two  subgenera,  Genioschizus  new  subgenus  and  Schizogenius  s.  str.,  both 
represented  in  North,  Middle,  and  South  America.  Genioschizus  is  a small,  mainly  South 
American  subgenus,  composed  of  three  closely  related  species  groups.  Schizogenius  s.  str. 
is  much  larger  and  more  varied,  and  particularly  in  North  America  is  the  dominant  subgenus. 

1.  Lateral  channel  of  elytron  flared  near  apex,  with  one  or  more  deep  subapical  pits 

Genioschizus,  new  subgenus,  p.  144 

1’  Lateral  channel  of  elytron  narrowed  near  apex,  without  deep  subapical  pits 

Schizogenius  s.  str.,  p.  165 

Subgenus  Genioschizus  new  subgenus 

Type  species.  — Schizogenius  crenulatus  LeConte  1849:197,  here  designated. 

Diagnostic  combination.  — Lateral  channel  of  elytron  flared  near  apex  and  with  one  or 
more  deep  subapical  pits.  Also:  paramedian  clypeal  carinae  tuberculate,  not  joined  to  medi- 
an clypeal  tooth;  clypeal  suture  not  sharply  engraved;  antennae  moniliform;  paralateral 
longitudinal  sulci  present  on  pronotum  or  not;  front  tarsi  slightly  dilated  in  males  only; 
paramedian  ambulatory  setae  on  sternum  seven  in  both  sexes;  and  pygidium  not  serrate  or 
crenulate  in  either  sex. 

Description.  - Small  beetles,  LE  1.65-2.75  mm.  Body  cylindrical.  Color  testaceous  to 
piceous,  aeneous  or  not,  elytra  sellate  or  not.  Integument  shiny;  microsculpture  reduced  on 
or  absent  from  median  frontal  sulcus,  prothoracic  pleura,  and  middle  of  abdominal  sterna, 
present  in  small  patches  in  coxal  depressions  of  sternum  three. 

Head.  Labrum  (Fig.  1)  slightly  emarginate  or  biemarginate  apically,  margined  laterally 
with  five  or  six  pairs  of  setae.  Clypeus  with  median  tooth  reduced;  paramedian  carinae 
short,  ended  before  median  tooth,  either  oblique  and  tuberculate  or  arcuate  and  with  apices 
nearly  joined  in  an  arc;  median  field  triangular  or  hemicircular,  at  base  more  than  1.5  » 

greatest  width  of  median  frontal  sulcus.  Clypeal  suture  obsolete.  Frontal  carinae  one  to  four 
irregular,  nearly  parallel,  confused  basally  or  not,  carinae  one  and  four  more  strongly  raised, 
carina  five  reduced  or  obsolete;  median  sulcus  wider  than  at  least  outer  paramedian  sulci, 
not  limited  in  front  by  transverse  carina,  with  no  trace  of  median  carina,  sides  parallel  or 
slightly  divergent  behind.  Eyes  prominent  (WF/WH,  0.49-0.63),  multifaceted,  facets  uniform 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


145 


Fig.  1-4.  Labrum,  dorsal  aspect.  1.  Schizogenius  crenulatus  LeConte.  2.  S.  optimus  Bates.  3.  S.  sallei  Putzeys.  4.  Halo- 
coryza  acapulcana  Whitehead.  Fig.  5-8.  Mandibles,  dorsal  aspect.  5.  S.  crenulatus  LeConte.  6.  S.  optimus  Bates.  7.  S. 
sallei  Putzeys.  8.  H.  acapulcana  Whitehead.  Fig.  9-12.  Left  maxilla,  ventral  aspect.  9.  S.  crenulatus  LeConte.  10.  S. 
optimus  Bates.  11.  S.  sallei  Putzeys.  12.  H.  acapulcana  Whitehead.  Fig.  13-16.  Labium,  ventral  aspect.  13.  S.  crenulatus 
LeConte.  14.  S.  optimus  Bates.  15.  S.  sallei  Putzeys.  16.  H.  acapulcana  Whitehead. 


146 


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Fig.  17-21.  Mentum,  ventral  aspect.  17.  S.  crenulatus  LeConte.  18.  S.  tenuis  Bates.  19.  S.  optimus  Bates.  20.  S.  sallei 
Putzeys.  21.  H.  acapulcana  Whitehead.  Fig.  22.  Elytral  apex,  postero-lateral  aspect,  S.  crenulatus  LeConte.  Fig.  23-26. 
Female  pygidium,  dorsal  aspect.  23.  S.  crenulatus  LeConte.  24.  S.  optimus  Bates.  25.  S.  sallei  Putzeys.  26.  H.  arenaria 
Darlington.  Fig.  27-30.  Ovipositor,  lateral  aspect.  27.  S.  crenulatus  LeConte.  28.  S.  optimus  Bates.  29.  S.  sallei  Putzeys. 
30.  H.  arenaria  Darlington. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


147 


Fig.  31-34.  Hind  wing.  31.  S.  crenulatus  LeConte.  32.  S.  tenuis  Bates.  33.  S.  optimus  Bates.  34.  S.  sallei  Putzeys. 


148 


Whitehead 


or  inner  facets  enlarged.  Antennal  articles  five  to  ten  square  to  slightly  elongate,  moniliform; 
pedicel  unisetose;  articles  three  to  eleven  pubescent.  Mandibles  (Fig.  5)  stout.  Mentum  (Fig. 
17-18)  deeply  emarginate  at  middle;  median  tooth  either  small  and  sharp  or  obsolete; 
antero-lateral  angles  of  lateral  lobes  either  broadly  rounded  or  acutely  produced;  anterior 
paramedian  setae  close  together.  Submentum  without  accessory  setae.  Gula  narrow,  its 
narrowest  part  0.15-0.20  width  of  mentum. 

Thorax.  Pronotum  in  most  species  slightly  transverse  (LP/WP,  0.91-1.01);  paramedian 
sulci  well  developed;  paralateral  sulci  present  or  absent;  lateral  grooves  not  bordered  by 
distinct  carinae;  accessory  marginal  setae  absent;  hind  angles  reduced  or  nearly  obsolete. 
Prothoracic  pleuron  impunctate. 

Elytra.  Lateral  channel  broad  and  deep  at  apex,  with  one  to  three  or  four  large  pits  near 
apex  above  umbilicate  series  (Fig.  22).  Disc  with  two  to  four  setae  on  interval  three,  zero  to 
two  basally  on  interval  five,  and  none  on  interval  seven.  Interval  eight  strongly  carinate, 
especially  near  apex;  interval  seven  carinate  at  least  in  basal  half;  intervals  two  to  seven 
subequal  in  width,  convex,  sharply  carinate  at  extreme  apices  or  not;  interval  eight  joined  by 
combined  apices  of  intervals  five  and  seven  or  not;  interval  seven  normally  joined  by  apex 
of  interval  five  and  either  ended  before  apex  of  interval  three  or  joined  to  it.  Striae  deeply 
engraved  and,  particularly  near  base  of  elytron,  coarsely  punctate. 

Hind  wings.  Fully  developed  and  probably  functional  in  all  species.  Venation,  Fig.  31-32. 

Legs.  Front  and  middle  tarsi  at  most  slightly  more  dilated  and  densely  pubescent  ventral- 
ly  in  male  than  in  female;  hind  tarsus  narrow,  moderately  elongate  (Ta/Ti,  0.59-0.75); 
article  one  of  hind  tarsus  2. 0-2. 5 times  as  long  as  article  two.  Paronychia  conspicuous,  about 
as  long  as  tarsal  claws.  Front  tibia  narrowed  evenly  to  base  where  much  narrower  than  at 
level  of  subapical  spur;  distal  tooth  nearly  straight,  stout,  and  blunt;  apical  and  subapical 
spurs  subequal,  slender  and  acute;  posterior  ventral  margin  with  three  setae  proximad  to 
spur. 

Abdomen.  Sternum  seven  with  paired  paramedian  ambulatory  setae  in  both  sexes.  Para- 
median carinae  of  sternum  three  not  rounded  at  apices.  Margin  of  pygidium  entire  in  both 
sexes. 

Male  genitalia.  Median  lobe  arcuate,  symmetric.  Endophallus  without  distinct  basal  collar 
spines;  membrane  around  virga  reduced;  basal  stylets  various,  useful  in  species  recognition. 

Female  ovipositor.  Number  and  position  of  setae  variable,  of  no  value  in  species  recog- 
nition (Fig.  27). 

Etymology.  — Genioschizus  is  an  anagram  of  Schizogenius,  and  bears  the  same  meaning: 
Greek,  genio  = mentum;  schizo  = split. 

Distribution.  — Members  of  this  subgenus  range  from  southern  Arizona  and  New  Mexico 
in  the  north,  southward  at  least  to  northern  Argentina  and  Brazil.  None  are  known  from 
the  West  Indies.  I examined  326  specimens  of  this  subgenus. 

Taxonomic  notes.  - Members  of  this  subgenus  are  here  placed  in  three  quite  easily 
distinguished  and  apparently  natural  groups,  but  some  species  are  difficult  to  distinguish. 
The  crenulatus  group  contains  one  polytypic  species  found  from  southwestern  United  States 
to  Honduras.  The  quinquesulcatus  group,  known  only  from  South  America,  includes  three 
recognized  species.  The  tenuis  group  includes  one  species  distributed  from  Mexico  to 
Colombia,  one  species  endemic  to  southern  Mexico,  and  three  additional  species  described 
from  Colombia.  Six  additional  specimens  of  the  tenuis  group  from  Argentina,  Bolivia, 
Brazil,  and  Peru  probably  represent  at  least  two  undescribed  species,  but  I defer  formal 
treatment  until  more  material  is  available  for  study.  I tentatively  assign  to  the  tenuis  group 
a species  described  from  Brazil,  S.  maculatus  Kult,  but  I have  seen  neither  the  type  nor 
any  other  specimens  which  match  its  description. 


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149 


Key  to  described  species  groups,  species,  and  subspecies  of  the  subgenus  Genioschizus 

1.  Pronotum  with  two  pairs  of  paramedian  longitudinal  sulci 2 

1’  Pronotum  with  one  pair  of  paramedian  longitudinal  sulci  ( tenuis  group) 6 

2. (1).  Lateral  lobes  of  mentum  truncate,  broadly  rounded  in  front;  median  tooth  of 

mentum  obsolete.  North  and  Middle  America  ( crenulatus  group) 3 

2’  Lateral  lobes  of  mentum  angulate,  acutely  produced;  median  tooth  of  mentum 

sharp,  prominent.  South  America  ( quinquesulcatus  group) 4 

3. (2).  Abdominal  sterna  without  coarse  lateral  microsculpture;  base  of  interval  five 

normally  asetose.  Arizona  and  California  to  Sinaloa  and  Nayarit 

S.  crenulatus  crenulatus  LeConte,  p.  150 

3’  Abdominal  sterna  with  coarse  lateral  microsculpture;  base  of  interval  five  normal- 
ly with  one  or  two  setae.  Jalisco  to  Honduras 

S.  crenulatus  chiapatecus  new  subspecies,  p.  152 

4. (2’).  Elytra  uniformly  dark.  Brazil,  Ecuador 

S.  quinquesulcatus  Putzeys,  p.  153 

4’  Elytra  pale,  with  dark  sutural  macula 5 

5. (4’).  Sides  of  pronotum  broadly  rounded  (Fig.  37).  Colombia,  Argentina 

S.  szekessyi  Kult,  p.  155 

5’  Sides  of  pronotum  not  strongly  rounded  (Fig.  38).  Brazil 

S.  janae  Kult,  p.  155 

6. (1’).  Base  of  interval  five  asetose;  elytra  with  sutural  macula.  Brazil 

S.  maculatus  Kult,  p.  165 

6’  Base  of  interval  five  with  one  seta.  Colombia  and  northward  (specimens  from 

south  of  Colombia  not  keyed) 7 

7. (6’).  Interval  six  of  elytron  conspicuously  narrowed  and  carinate  in  apical  third;  ab- 

dominal sterna  normally  with  coarse  lateral  microsculpture.  Chiapas  and  Tabasco. 

S.  sculptilis  new  species,  p.  156 

7’  Interval  six  of  elytron  not  narrowed  in  apical  third,  finely  carinate  only  at  extreme 

apex  if  at  all;  abdominal  sterna  four  to  six  usually  without  continuous,  coarse 
lateral  microsculpture 8 

8. (7’).  Median  longitudinal  sulcus  of  frons  not  narrowed  toward  apex;  median  lobe  of 

male  genitalia  with  apex  sharply  deflexed  (Fig.  53).  Colombia 

S.  impressicollis  Putzeys,  p.  161 

8’  Median  longitudinal  sulcus  of  frons  narrowed  toward  apex;  median  lobe  of  male 

genitalia  with  apex  not  sharply  deflexed 9 

9. (8’).  Head  with  neck  nearly  or  quite  impunctate  along  midline.  Colombia 

S.  impuncticollis  new  species,  p.  161 

9’  Head  with  neck  distinctly  punctate  along  midline 10 

10. (9’).  Disc  of  elytron  with  distinct  sutural  macula.  Colombia v 

S.  suturalis  new  species,  p.  162 

10’  Disc  of  elytron  without  sutural  macula;  eyes  with  inner  facets  larger  than  outer 

facets.  Mexico  to  Colombia S.  tenuis  Bates,  p.  158 

The  crenulatus  group 

Diagnostic  combination.  — Clypeal  carinae  strongly  arcuate;  mentum  with  median  tooth 
obsolete,  lateral  lobes  truncate;  pronotum  with  short  but  evident  paralateral  longitudinal 
sulci;  elytron  with  three  or  four  conspicuous  pits  near  apex  of  lateral  channel;  interval  five 
with  or  without  basal  setae;  intervals  two  to  seven  not  carinate  apically;  interval  eight  free 


150 


Whitehead 


to  apex;  and  parameres  normally  uni-  or  bisetose. 

The  mentum,  with  lateral  lobes  truncate,  is  diagnostic  within  the  genus.  Members  of  this 
group  differ  from  other  North  and  Middle  American  members  of  the  subgenus  by  ecarinate 
apices  of  elytral  intervals  two  to  seven. 

Distribution.  — This  group  includes  one  species,  S.  crenulatus  LeConte,  with  two  sub- 
species distributed  from  extreme  southwestern  United  States  to  Honduras.  Other  groups  of 
the  subgenus  have  more  tropical  distributions,  though  one  species  of  the  tenuis  group  ranges 
nearly  as  far  northward.  I studied  64  specimens  of  the  crenulatus  group. 

Schizogenius  crenulatus  LeConte 

Diagnostic  combination.  - Specimens  of  S.  crenulatus  differ  from  other  Schizogenius  by 
truncate  mentum  lateral  lobes.  I recognize  two  subspecies,  distinguished  by  extent  of  micro- 
sculpture  on  abdominal  sterna. 

Schizogenius  crenulatus  crenulatus  new  combination 

Schizogenius  crenulatus  LeConte  1852:197.  Type  locality  “California,”  here  restricted  to 

the  Colorado  River  opposite  Yuma,  Yuma  County,  Arizona;  type  in  MCZ,  specimen 

labelled  MCZ  5480  here  designated  lectotype  (!).  LeConte  1857:82.  Putzeys  1863:24. 

Putzeys  1866:223.  LeConte  1879:34.  Lindroth  1961:165. 

Diagnostic  combination.  - From  specimens  of  the  other  subspecies,  specimens  of  this 
form  are  distinguished  by  reduced  microsculpture  on  sides  of  abdominal  sterna.  Also,  most 
are  paler,  and  most  lack  basal  setae  on  elytral  interval  five. 

Description.  - Color  rufopiceous,  legs  paler,  palpi,  antennae,  and  tarsi  testaceous;  without 
strong  aeneous  luster,  elytra  not  sellate  or  maculate. 

Integument.  Fine  but  conspicuous  micro  sculpture  on  genae,  gula,  mouthparts,  front 
tibiae  and  apical  half  of  anterior  surfaces  of  front  femora,  middle  legs  except  trochanters, 
hind  tibiae  and  posterior  surfaces  of  hind  femora,  elytral  epipleura  at  base  and  on  apical 
two  thirds,  and  in  coxal  depressions  of  sternum  three;  paramedian  frontal  sulci  obscurely 
microsculptured. 

Head.  Fig.  35.  Eyes  prominent,  subglobose,  coarsely  and  uniformly  faceted.  Neck  densely 
and  coarsely  punctate.  Genae  strongly  punctate,  finely  rugose  in  front.  Mentum  (Fig.  17) 
lateral  lobes  truncate,  median  tooth  obsolete.  Antennae  short,  moniliform,  article  five  1.0- 
1 . 1 times  as  long  as  wide. 

Pronotum.  Fig.  35.  Paramedian  sulci  faintly  punctate,  shallow  apically,  deep  and  broadly 
hooked  basally,  basal  tips  abruptly  limited;  paralateral  sulci  usually  deep,  coarsely  punctate, 
no  more  than  half  length  of  paramedian  sulci;  anterior  transverse  impression  finely  to 
coarsely  punctate;  base  transversely  rugose. 

Elytra.  Three  discal  setigerous  punctures  on  interval  three  adjacent  to  second  stria;  inter- 
val five  asetose  or  in  some  specimens  with  one  basal  seta.  Intervals  one  to  six  convex; 
interval  seven  carinate  in  basal  two-thirds;  interval  eight  carinate  throughout,  sharply  so  in 
apical  third.  Interval  eight  fused  with  interval  one  at  apex,  otherwise  free;  intervals  three  and 
five  joined  to  apex  of  interval  seven  in  most  specimens.  Lateral  channel  with  three  or  four 
large  pits  near  apex. 

Male  genitalia.  Median  lobe  (Fig.  44)  variable,  some  specimens  as  in  chiapatecus  (Fig.  45), 
apical  portion  suddenly  deflexed;  basal  stylets,  Fig.  58-59;  virga  (Fig.  57)  without  distin- 
guishing spines  or  scales.  Seven  specimens  studied. 

Measurements  and  proportions.  See  Table  1. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


151 


Table  1.  Descriptive  statistics  for  S.  crenulatus,  based  on  28  specimens  of  undetermined 
sex  from  Arizona  and  California. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

nts,  in  mm. 

TL 

3.52-4.25 

3.962 

0.272 

0.068 

4.57 

LE 

2.16-2.60 

2.426 

0.172 

0.043 

4.47 

WH 

0.69-0.83 

0.776 

0.048 

0.012 

4.08 

WP 

0.94-1.16 

1.075 

0.081 

0.021 

5.05 

WE 

1.10-1.35 

1.246 

0.089 

0.022 

4.74 

B.  Proportions 

WF/WH 

0.54-0.59 

0.569 

0.023 

0.006 

2.78 

LP/WP 

0.94-1.01 

0.975 

0.025 

0.006 

1.70 

DP/LP 

0.80-0.86 

0.831 

0.025 

0.006 

2.00 

LP/WE 

0.81-0.87 

0.842 

0.022 

0.006 

1.73 

Ta/Ti 

0.63-0.73 

0.691 

0.038 

0.010 

3.69 

PS/LP 

0.62-0.69 

0.651 

0.025 

0.006 

2.55 

Variation.  — One  specimen  from  Blythe,  California,  has  a basal  seta  on  interval  five  of 
left  elytron,  but  in  all  other  specimens  interval  five  is  asetose  on  both  elytra.  Paralateral 
pronotal  sulci  of  some  specimens  are  quite  weakly  impressed.  Paramedian  clypeal  carinae  of 
some  specimens  are  short  and  widely  separated  at  apices.  There  is  no  pronounced  external 
sexual  dimorphism,  though  some  males  are  distinguished  by  slightly  broader  front  tarsi. 
Among  specimens  studied,  extremes  in  size  were  found  in  two  females  from  Rio  San 
Lorenzo  in  Sinaloa  (LE,  1.89-2.61  mm).  Variation  in  selected  characteristics  in  specimens 
from  the  Colorado  River  drainage  basin  of  Arizona  and  California  is  given  in  Table  1. 

Etymology.  — Latin,  crenulatus  = minutely  toothed,  in  reference  to  the  minutely  toothed 
elytral  apices. 

Distribution.  — Specimens  of  S.  crenulatus  crenulatus  have  been  collected  at  low  eleva- 
tions in  the  lower  reaches  of  the  Colorado  River  and  environs  in  southern  California  and 
Arizona,  south  to  southern  Sinaloa  and  northern  Nayarit  (Fig.  74).  I studied  57  specimens 
from  the  following  localities. 


UNITED  STATES 

No  locality  (1;  ANSP).  ARIZONA  (7;  CAS,  INHS,  USNM):  Gila  Co.,  Salt  River  (2;  ANSP,  MCZ);  Maricopa  Co.,  Phoenix 
(1;  CUNY);  Pima  Co.  (1;  USNM);  Yuma  Co.  (10;  CAS,  USNM),  Yuma  (9;  CAS,  MCZ,  USNM).  CALIFORNIA:  Riverside 
Co.,  Blythe  (2;  CAS). 


MEXICO 

NAYARIT:  Jesus  Maria  (2;  UCB).  SINALOA:  30.6  mi.  s.  Culiacan  (18;  DRWh,  UASM),  21  mi.  e.  Villa  Union  (2;  CNC),  26 
mi.  ne.  Villa  Union  1000’  (1;  LBSC).  SONORA:  7.2  mi.  se.  Alamos  (1;  GRNo). 


Collecting  notes.  - The  most  recently  collected  specimens  seen  from  the  United  States 
are  from  Blythe  and  Phoenix,  taken  in  1917.  Perhaps,  because  of  environmental  changes 
resulting  from  manipulation  of  the  Colorado  River  system,  & crenulatus  may  now  be  extinct 
or  nearing  extinction  in  the  southwestern  United  States.  All  Mexican  specimens,  however, 
were  collected  more  recently. 


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According  to  label  data,  adults  of  S.  crenulatus  are  probably  active  throughout  the  year. 
Several  specimens  were  taken  at  lights,  so  no  doubt  wings  are  functional.  I have  no  field 
experience  with  adults  of  this  subspecies,  but  suspect  they  normally  live  in  sandy  river  banks 
rather  than  gravel  bars.  G.  E.  Ball,  T.  L.  Erwin,  and  R.  E.  Leech  collected  19  specimens 
near  Culiacan,  Sinaloa,  under  litter  on  moist  sand  along  the  Rio  San  Lorenzo. 

Taxonomic  notes.  — I treat  the  old  name  S.  crenulatus  as  a new  combination,  S.  crenula- 
tus crenulatus,  because  I recognize,  as  a new  subspecies,  S.  crenulatus  chiapatecus . Available 
specimens  of  the  two  forms  are  well  distinguished  morphologically,  and  hence  I think  that 
recognition  of  separate  subspecies  is  well  justified.  Proximate  localities  of  these  allopatric 
forms  are  not  greatly  distant,  but  I can  neither  reject  nor  defend  a proposition  of  reproduc- 
tive isolation. 


Schizogenius  crenulatus  chiapatecus  new  subspecies 

Type  material.  — Holotype,  female,  labelled  “MEXICO.  Chiapas.  3.2  mi.  n.  Arriaga  400’ 
Rte.  195  III.2.1966”  and  “George  E.  Ball,  D.  R.  Whitehead  collectors”  (MCZ).  Four  females 
and  one  male  from  various  localities  in  Chiapas,  Guerrero,  Jalisco,  and  Oaxaca  are  paratypes 
(FDAG,  MCZ,  UASM). 

Diagnostic  combination.  — Sterna  four  to  six  with  continuous  microsculpture  on  each 
side.  All  specimens  seen  have  one  or  two  basal  setigerous  punctures  on  interval  five  on  at 
least  one  elytron. 

Description.  — As  in  S.  crenulatus  crenulatus  except  as  follows.  Color  darker,  most 
specimens  with  faint  aeneous  luster,  strongest  in  Chiapas  specimens;  antennae,  maxillae, 
labial  palpi,  and  tarsi  dark  testaceous. 

Integument.  Conspicuous  microsculpture  on  sternum  two,  sternum  three  except  median 
field,  sterna  four  to  six  at  sides,  and  sternum  seven  along  margin. 

Pronotum.  Anterior  transverse  impression  more  finely  punctate;  base  of  pronotum  often 
more  strongly  rugose,  basal  tips  of  paramedian  sulci  not  sharply  limited. 

Elytra.  Interval  five  normally  with  one  or  two  basal  setae. 

Male  genitalia.  Median  lobe,  Fig.  45;  basal  stylets,  Fig.  60;  virga  as  in  S.  crenulatus 
crenulatus.  One  specimen  studied. 

Measurements  and  proportions.  Holotype:  TL,  4.36  mm;  LE,  2.72  mm;  WH,  0.85  mm; 
WP,  1.14  mm;  WE,  1.38  mm;  WF/WH,  0.55;  LP/WP,  0.98;  DP/LP,  0.84;  LP/WE,  0.81; 
Ta/Ti,  0.72;  PS/LP,  0.67.  Holotype  plus  paratypes:  TL,  3.80-4.09-4.31  mm;  LE,  2.35-2.54- 
2.67  mm;  WH,  0.75-0.80-0.85  mm;  WP,  1.02-1.11-1.18  mm;  WE,  1.17-1.29-1.36  mm;WF/ 
WH,  0.55-0.57-0.58;  LP/WP,  0.94-0.96-0.98;  DP/LP,  0.82-0.85-0.86;  LP/WE,  0.8 1-0.82-0.84; 
Ta/Ti,  0.65-0.70-0.75;  PS/LP,  0.63-0.68-0.70. 

Variation.  — Variations  in  numbers  of  discal  setae  on  elytra  include:  absence  of  middle 
seta  of  interval  three  of  left  elytron  in  Chiapas  paratype;  absence  of  seta  of  interval  five  of 
left  elytron  in  Oaxaca  male;  and  presence  of  two  setae  near  base  of  fifth  interval  on  both 
elytra  in  Acapulco  specimen.  Otherwise  I found  no  noteworthy  variation. 

Etymology.  — The  name  chiapatecus  is  given  in  reference  to  the  type  locality,  in  Chiapas. 
Distribution.  — Seven  specimens  of  S.  crenulatus  chiapatecus  have  been  collected  at  low 
elevations  from  Jalisco  south  to  Honduras  (Fig.  74). 

MEXICO 

CHIAPAS:  3.2  mi.  n.  Arriaga  (2;  MCZ,  UASM).  GUERRERO:  Acapulco  (1;  MCZ).  JALISCO:  Pitillal  (1;  UASM). 
OAXACA:  Rio  Jaltepec  (2;  FDAG). 

HONDURAS 

CORTES:  La  Lima  (1;  FDAG). 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


153 


Collecting  notes.  — Specimens  of  this  subspecies  have  been  collected  in  January,  March, 
May,  June,  and  August.  The  Chiapas  specimens  were  taken  in  coarse  sand  along  a small 
stream.  Other  specimens  were  collected  at  lights,  and  probably  arrived  by  flight. 

Taxonomic  notes.  — Since  the  Chiapas  locality  is  distant  from  the  known  range  of  S. 
crenulatus  crenulatus,  I chose  one  of  the  two  females  from  there  as  holotype.  I found  no 
diagnostic  characteristics  in  the  male  genitalia.  S.  crenulatus  chiapatecus  may  be  a biologi- 
cally distinct  species,  but  evidence  from  morphological  characteristics  reported  here  is  in- 
conclusive. That  the  possibility  exists,  however,  is  sufficient  reason  to  distinguish  this  form 
taxonomically.  As  all  known  specimens  of  this  subspecies  are  distinguishable  from  all  known 
specimens  of  the  other,  the  names  refer  to  distinctive  phena  which  may  eventually  prove  to 
be  reproductive ly  isolated. 


The  quinquesulcatus  group 

Diagnostic  combination.  — Head  with  clypeal  carinae  straight  to  moderately  arcuate; 
mentum  with  median  tooth  small  and  sharp,  lateral  lobes  acutely  produced  at  antero-lateral 
angles;  pronotum  with  short  but  evident  paralateral  longitudinal  sulci;  elytron  with  at  least 
two  conspicuous  pits  near  apex  of  lateral  channel;  interval  five  with  basal  seta;  intervals  two 
to  seven  finely  carinate  at  extreme  apices;  interval  eight  free  to  apex;  and  parameres  normal- 
ly unisetose. 

The  acutely  produced  lateral  lobes  of  the  mentum  and  the  deep  paralateral  sulci  of  the 
pronotum  are,  in  combination,  diagnostic  of  the  group  within  the  genus. 

Distribution.  — I here  recognize  three  species,  all  closely  related,  from  nearly  throughout 
continental  South  America.  I studied  20  specimens  of  the  group  from  northern  Colombia 
south  to  northern  Argentina. 

Schizogenius  quinquesulcatus  Putzeys 

Schizogenius  quinquesulcatus  Putzeys  1863:26.  Type  locality  “Amazone”,  Brazil;  type  fe- 
male in  IRSB  (!).  Putzeys  1866:232. 

Schizogenius  exaratus  Putzeys  1863:27.  Type  locality  Nova  Friburgo,  Brazil;  type  female 

in  IRSB  (!).  Putzeys  1866:232,  established  synonymy. 

Diagnostic  combination.  — Within  the  quinquesulcatus  group,  specimens  of  this  species 
are  distinguished  by  dark,  non-sellate  elytra. 

Description.  — Color  dark  piceous;  femora  rufous  or  infuscated,  legs  and  antennae  other- 
wise ferrugineous;  palpi  testaceous;  elytra  with  slight  aeneous  luster,  not  sellate  or  maculate, 
apex  pale  or  not. 

Integument.  Fine  microsculpture  on  genae,  gula,  mouthparts,  front  tibiae  and  anterior 
surfaces  of  front  femora,  middle  legs  except  trochanters,  hind  tibiae  and  posterior  surfaces 
of  hind  femora,  elytral  epipleura  on  base  and  apical  two-thirds,  sides  of  sterna  two  to  six, 
and  apex  of  sternum  seven;  paramedian  frontal  sulci  obscurely  microsculptured. 

Head.  Fig.  36;  paramedian  clypeal  carinae  straight  to  arcuate,  apices  widely  separated, 
median  field  triangular  or  hemicircular;  inner  paramedian  frontal  carinae  nearly  parallel. 
Eyes  prominent,  subglobose,  coarsely  and  uniformly  faceted.  Neck  densely,  finely  to  coarse- 
ly punctate.  Genae  strongly  punctate,  finely  rugose  in  front.  Mentum  with  lateral  lobes 
acutely  produced,  median  tooth  conspicuous  but  short  and  sharp.  Antennae  short,  monili- 
form,  article  eight  1 .0-1.2  times  as  long  as  wide. 

Pronotum.  Fig.  36;  sides  broadly  rounded;  paralateral  longitudinal  sulci  and  anterior 
transverse  impression  punctate;  base  transversely  rugose. 


154 


Whitehead 


Elytra.  Three  discal  setigerous  punctures  on  interval  three,  near  or  adjacent  to  stria  two; 
interval  five  with  one  seta  near  base.  Striae  deep  and  sharply  engraved  throughout,  evidently 
punctate  except  near  apex  of  elytron,  coarsely  punctate  in  basal  half.  Intervals  one  to  six 
broad,  convex,  narrowed  and  finely  carinate  at  apices;  interval  six  not  carinate  basally; 
interval  seven  carinate  throughout;  interval  eight  carinate  in  apical  half.  Interval  eight  free  to 
apex;  intervals  seven,  five,  and  three  joined  at  apices  in  most  specimens.  Lateral  channel 
with  two  large  conspicuous  pits  near  apex. 

Male  genitalia.  Median  lobe,  Fig.  46-47;  basal  stylets.  Fig.  61-63;  virga  without  distin- 
guishing spines  or  scales.  Three  specimens  studied. 

Measurements  and  proportions.  Ranges  of  variation  in  measurements  and  proportions 
among  eight  specimens  studied  are:  TL,  3.24-3.95  mm;  LE,  1.99-2.50  mm;  WH,  0.65-0.76 
mm;  WP,  0.92-1.07  mm;  WE,  1.00-1.28  mm;  WF/WH,  0.57-0.62;  LP/WP,  0.88-0.9 1 ; DP/LP, 
0.84-0.9 1;LP/WE,  0.74-0.82;  LE/WE,  1 .94-1 .99;  Ta/Ti,  0.67-0.79;  PS/LP,  0.64-0.73. 

Variation.  — The  specimen  from  Ecuador  differs  from  Brazilian  specimens  by  more 
coarsely  punctate  neck,  longer  antennae,  and  shorter  paramedian  pronotal  sulci,  but  is 
otherwise  not  distinctive.  Paramedian  clypeal  carinae  vary  in  form,  but  this  variation  is 
probably  not  taxonomically  important  since  it  forms  no  evident  geographic  pattern. 

Etymology.  - For  quinquesulcatus,  Latin,  quinque  = five,  plus  sulcatus  = grooved,  in 
reference  to  the  five  longitudinal  grooves  on  the  pronotum.  For  exaratus,  Latin,  ex  = out, 
plus  aratus  = plow,  in  reference  to  the  fossorial  habitus. 

Distribution.  — I studied  14  specimens  of  this  species  from  Atlantic  and  Pacific  drainage 
systems  from  the  following  continental  South  American  localities. 

BRAZIL 

“Amazones”  (1;  IRSB,  Type  of  quinquesulcatus).  CEARA:  Fortaleza  (1;  IRSB).  RIO  DE  JANEIRO:  Nova  Friburgo  (1; 
IRSB,  type  of  exaratus).  SANTA  CATARINA  (2;  IRSB):  Nova  Teutonia  (6;  DRWh,  MCZ).  SAO  PAULO:  Piracicaba  (2; 
OSUC,  UASM). 


EL  ORO:  9 mi.  s.  Santa  Rosa  ( 1 ; CAS). 


ECUADOR 


Collecting  notes.  — Specimens  of  this  species  have  been  collected  in  January,  February, 
April,  August,  September,  October,  and  November,  and  individuals  may  therefore  be  active 
throughout  the  year. 

Taxonomic  notes.  - The  names  quinquesulcatus  and  exaratus  were  both  originally  pro- 
posed for  unique  specimens.  The  specimen  listed  here  as  type  of  S.  quinquesulcatus  is 
certainly  so,  but  label  data  on  the  S.  exaratus  specimen  (“S.  exaratus.  P,”  handwritten  on 
green  paper,  and  “Dup.  8/8  47  Bres.,”  handwritten  on  white  paper)  are  equivocal.  Neither 
specimen  matches  its  original  description  in  detail,  but  since  Putzeys  often  erred  in  his 
descriptions  I think  this  may  be  ignored.  My  reasons  for  suggesting  that  the  S.  exaratus 
specimen  is  indeed  the  type  are  the  following.  Putzeys  gave  no  indication  that  the  type  was 
deposited  elsewhere  than  in  his  collection,  and  no  other  specimen  in  IRSB  can  be  the  type. 
The  specimen  was  collected  well  before  the  date  of  the  original  description,  and  thus  was 
likely  available  to  Putzeys  since  he  corresponded  with  Dupont.  Most  important,  the  speci- 
men is  the  only  one  labelled  as  S.  exaratus  in  Putzeys’  own  hand,  something  he  probably 
would  not  have  done  with  subsequent  material  as  his  discovery  of  a third  specimen  of  the 
species  prompted  him  to  synonymize  the  two  names.  Quite  possibly  the  third  specimen 
mentioned  by  Putzeys  (1866)  is  the  Ceara  specimen. 

I believe  all  of  the  Brazilian  specimens  are  conspecific,  despite  variation  in  form  of  clypeal 
carinae.  The  Ecuadorian  specimen  may  well  not  be  conspecific,  but  I am  unable  to  decide 
without  additional  material. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


155 


Schizogenius  szekessyi  Kult 

Schizogenius  szekessyi  Kult  1950:144.  Type  locality  Aracataca,  Colombia;  type  in  Budapest 

Museum,  not  studied. 

Diagnostic  combination.  - From  specimens  of  S.  janae,  the  only  other  described  member 
of  the  quinquesulcatus  group  having  pale  elytra  with  a dark  sutural  macula,  specimens  of 
this  form  are  distinguished  by  broadly  rounded  pronotal  sides. 

Description.  - As  in  S.  quinquesulcatus  except:  body  paler,  rufopiceous;  elytron  rufo- 
testaceous,  with  dark  sutural  macula  extended  outward  to  interval  three  or  four;  head  and 
pronotum,  Fig.  37,  clypeal  carinae  straight  and  neck  coarsely  punctate  in  all  specimens 
studied;  male  median  lobe  (Fig.  48)  and  basal  stylets  (Fig.  64-65)  as  illustrated;  three 
specimens  studied. 

Measurements  and  proportions.  Ranges  of  variation  in  measurements  and  proportions 
among  five  specimens  studied  are:  TL,  3.52-3.78  mm;  LE,  2.20-2.35  mm;  WH,  0.65-0.79 
mm;  WP,  0.96-1.08  mm;  WE,  1.14-1.24  mm;  WF/WH,  0.55-0.58;  LP/WP,  0.89-0.92;  DP/ 
LP,  0.85-0.87;  LP/WE,  0.76-0.78;  LE/WE,  1.90-2.00;  Ta/Ti,  0.65-0.73;  PS/LP,  0.63-0.73. 

Variation  — The  specimen  from  Argentina  differs  from  Colombian  specimens  by  smaller 
sutural  macula  and  relatively  narrower  elytra.  This  specimen  has  only  two  setae  on  interval 
three  of  the  left  elytron. 

Etymology.  — This  species  was  named  in  honor  of  the  Hungarian  entomologist,  Dr.  W. 
Szekessy. 

Distribution.  — I assign  the  name  S.  szekessyi  to  four  topotypic  specimens  from  Colombia, 
and  one  specimen  from  Argentina. 


ARGENTINA 

SALTA:  Oran(l;MCZ). 

COLOMBIA 

MAGDALENA:  Aracataca  (4;  DRWh,  MCZ). 


Collecting  notes.  — Colombian  specimens  were  collected  from  January  to  May,  and  the 
Argentinian  specimen  was  taken  in  September;  thus,  individuals  are  probably  active  as  adults 
throughout  the  year. 

Taxonomic  notes.  — I have  not  studied  type  material  of  this  species,  but  think  there  is  no 
question  of  identity  since  topotypic  material  agrees  with  the  original  description.  I see  no 
reason  to  treat  the  specimen  from  Argentina  as  other  than  conspecific. 

Schizogenius  janae  Kult 

Schizogenius  janae  Kult  1950: 145.  Type  locality  Brazil;  type  in  Kult  Collection,  not  studied. 

Diagnostic  combination.  - The  sides  of  the  pronotum  are  less  broadly  rounded  in  speci- 
mens of  S.  janae  than  in  those  of  S.  szekessyi,  the  only  other  member  of  the  quinquesulcatus 
group  having  a sutural  macula  on  the  elytra;  differences  in  form  of  median  lobe  of  male 
genitalia  may  also  be  useful  in  species  recognition. 

Description.  — As  in  S.  quinquesulcatus  except  as  follows.  Color  of  body  paler,  rufopi- 
ceous; elytra  rufotestaceous,  each  with  a dark  sutural  macula  extended  outward  to  interval 
three  or  four.  Head  and  pronotum,  Fig.  38;  pronotal  sides  weakly  rounded,  clypeal  carinae 
straight,  and  neck  coarsely  punctate.  Male  median  lobe  (Fig.  49)  and  basal  stylets  (Fig.  66) 
as  illustrated;  one  specimen  studied. 

Measurements  and  proportions.  TL,  3.25  mm;  LE,  2.00  mm;  WH,  0.68  mm;  WP,  0.92 


156 


Whitehead 


mm;  WE,  1.02  mm;  WF/WH,  0.59;  LP/WP,  0.91,  DP/LP,  0.80;  LP/WE,  0.82;  LE/WE,  1.96; 
Ta/Ti,  0.68;  PS/LP,  0.72. 

Etymology.  — This  species  was  dedicated  by  Kult  to  his  wife. 

Distribution.  — I studied  one  specimen  of  this  species  from  Brazil. 

BRAZIL 

MATO  GROSSO:  Jacare  (1;  MGFT). 


Collecting  notes.  — The  Brazilian  specimen  reported  here  was  collected  in  September. 

Taxonomic  notes.  — I have  not  studied  the  type,  but  think  this  specimen  represents  S. 
janae  because  it  agrees  well  with  the  original  description  and  comes  from  a suitable  locality. 
In  particular,  the  specimen  is  smaller  and  paler  than  specimens  of  S.  szekessyi,  and  the 
form  of  the  pronotum  is  correct.  Other  characters  given  in  the  original  description  are 
unreliable. 

I think  it  highly  probable  that  the  names  S.  janae  and  S.  szekessyi  are  synonyms,  despite 
differences  illustrated  for  form  of  pronotum  and  male  median  lobe.  More  specimens  are 
needed  from  throughout  the  range  to  determine  extent  of  individual  and  geographic  varia- 
tion. 


The  tenuis  group 

Diagnostic  combination.  — Head  with  clypeal  carinae  normally  arcuate;  mentum  with 
median  tooth  prominent,  lateral  lobes  acutely  produced;  pronotum  without  paralateral  sulci; 
elytron  with  one  to  three  conspicuous  subapical  pits  in  lateral  channel;  interval  five  with  or 
without  basal  seta;  intervals  two  to  seven  carinate  apically;  interval  eight  free  to  apex  or 
not;  and  parameres  in  most  species  bi-  or  trisetose. 

The  lack  of  paralateral  longitudinal  sulci  from  the  pronotum  is  unique  within  the  sub- 
genus. 

Distribution.  — Specimens  of  this  group  have  been  collected  from  northern  Mexico  to 
northern  Argentina.  I here  recognize  one  species  known  only  from  Mexico,  another  from 
Mexico  to  Colombia,  and  three  others  from  Colombia.  Six  unplaced  specimens  from  widely 
separated  localities  elsewhere  in  South  America  probably  represent  at  least  two  additional, 
undescribed,  species.  I tentatively  place  as  a member  of  this  group  a species  described  by 
Kult  from  Brazil,  S.  maculatus.  I examined  244  specimens  of  the  tenuis  group. 

Schizogenius  sculptilis  new  species 

Type  material.  — Holotype,  male,  and  allotype,  female,  labelled  “MEXICO.  Chiapas  San 
Quintin  700'  91°20'-16°24'  stream  margins  II.  5-20.  1966”  and  “George  E.  Ball  D.  R. 
Whitehead  collectors”  (MCZ).  Twenty  additional  specimens  from  two  localities  in  Chiapas 
and  Tabasco  are  paratypes  (BMNH,  CAS,  CNC,  DRWh,  IRSB,  UASM,  USNM). 

Diagnostic  combination.  — Most  reliably  distinguished  from  S.  tenuis  by  narrowed,  cari- 
nate apices  of  elytral  interval  six,  and  by  characteristic  form  of  basal  stylets  of  median  lobe. 
On  most  specimens,  the  abdomen  is  microsculptured  laterally,  while  in  most  Middle  Ameri- 
can specimens  of  S.  tenuis  the  abdomen  lacks  extensive  microsculpture. 

Description.  — Color  dark  piceous,  legs,  palpi,  and  antennae  paler;  no  definite  aeneous 
luster;  elytra  not  sellate  or  maculate,  apices  normally  not  pale. 

Integument.  As  described  for  S.  tenuis  except  on  abdomen;  sides  of  sterna  four  to  six 
and  margin  of  sternum  seven  strongly  microsculptured  in  most  specimens. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


157 


Head.  Fig.  39.  Generally  as  described  for  S.  tenuis.  Frontal  carina  one  broad,  carinae  two 
to  four  narrow,  carina  five  short  but  well  developed.  Inner  facets  of  eyes  much  larger  than 
marginal  facets.  Mentum  with  median  tooth  minute. 

Pronotum.  Fig.  39.  Generally  as  described  for  S.  tenuis  except  disc  flatter,  paramedian 
sulci  longer,  and  anterior  transverse  impression  coarsely  punctate. 

Elytra.  Discal  setae  and  striae  as  in  S.  tenuis.  Intervals  one  to  five  broad,  convex,  carinate 
at  extreme  apices;  interval  six  narrowed,  carinate  in  apical  third;  interval  seven  carinate 
throughout;  interval  eight  carinate  in  apical  half.  Apex  of  interval  eight  fused  with  interval 
one,  nearly  fused  with  interval  two,  otherwise  free;  intervals  three,  five,  and  seven  in  many 
specimens  more  or  less  fused  at  apex.  Lateral  channel  with  one  small  but  conspicuous  sub- 
apical  pit. 

Male  genitalia.  Median  lobe  (Fig.  50)  arcuate,  apical  part  not  suddenly  deflexed;  basal 
stylets  (Fig.  67)  characteristic;  virga  without  distinguishing  spines  or  scales.  Four  specimens 
studied. 

Measurements  and  proportions.  See  Table  2.  Holotype:  TL,  3.83  mm;  LE,  2.30  mm;  WH, 
0.78  mm;  WP,  1.08  mm;  WE,  1.21  mm;  WF/WH,  0.59;  LP/WP,  0.96;  DP/LP,  0.82;  LP/WE, 
0.86;  Ta/Ti,  0.73;  PS/LP,  0.71.  Allotype:  TL,  3.82  mm;  LE,  2.30  mm;  WH,  0.78  mm;  WP, 
1.11  mm;  WE,  1.24  mm;  WF/WH,  0.60;  LP/WP,  0.93;  DP/LP,  0.81;  LP/WE,  0.83;  Ta/Ti, 
0.63;  PS/LP,  0.74. 


Table  2.  Descriptive  statistics  for  S.  sculptilis,  based  on  22  unsexed  specimens  from  Chiapas 
and  Tabasco. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measurem 
TL 

ents,  in  mm. 
3.33-3.98 

3.658 

0.271 

0.077 

4.93 

LE 

1.94-2.44 

2.210 

0.184 

0.052 

5.56 

WH 

0.70-0.81 

0.749 

0.048 

0.014 

4.27 

WP 

0.93-1.12 

1.036 

0.078 

0.022 

5.02 

WE 

1.09-1.26 

1.176 

0.075 

0.021 

4.28 

B.  Proportions. 


WF/WH 

0.57-0.63 

0.592 

0.028 

0.008 

3.15 

LP/WP 

0.91-0.99 

0.945 

0.027 

0.008 

1.91 

DP/LP 

0.80-0.89 

0.830 

0.037 

0.011 

3.00 

LP/WE 

0.81-0.86 

0.834 

0.021 

0.006 

1.67 

Ta/Ti 

0.59-0.73 

0.666 

0.053 

0.015 

5.32 

PS/LP 

0.68-0.74 

0.715 

0.029 

0.008 

2.67 

Variation.  — Two  specimens  from  Chiapas  and  one  from  Tabasco  have  reduced  abdominal 
microsculpture.  Another  Chiapas  specimen  has  an  extra  seta  on  interval  three  of  the  left 
elytron,  and  one  Tabasco  specimen  has  pale  elytral  apices.  Aside  from  slightly  broader  front 
tarsi  in  males,  there  is  no  evident  secondary  sexual  differentiation. 

Etymology.  — Latin,  sculptilis  = carved,  in  reference  to  form  of  apices  of  elytral  intervals. 


158 


Whitehead 


Distribution.  - Specimens  of  this  species  have  been  collected  at  two  localities  (Fig.  75) 
in  lowland  rain  forests  of  Chiapas  and  Tabasco.  I studied  22  specimens  from  the  following 
localities. 


MEXICO 

CHIAPAS:  San  Quintin  (20;  BMNH,  CAS,  CNC,  IRSB,  MCZ,  UASM,  USNM). 

TABASCO:  59.4  mi.  se.  Villahermosa  (2;  DRWh). 

Collecting  notes.  — Adults  of  S.  sculptilis  are  probably  active  throughout  the  year,  since 
specimens  were  collected  in  February  and  June.  No  specimens  were  seen  to  fly,  and  none 
were  taken  at  lights,  but  like  other  North  and  Middle  American  Schizogenius,  adults  of  S. 
sculptilis  probably  are  capable  of  flight.  Among  North  and  Middle  American  members  of 
Schizogenius , habits  of  this  species  are  unusual  in  that  the  preferred  habitat  is  in  sand,  along 
shaded  forest  streams.  Three  specimens  were  taken  under  leaf  litter  on  sand  along  the  Rio 
Jatate,  but  all  other  San  Quintin  specimens  were  collected  in  deep  forest  in  a sand-gravel 
substrate  along  stream  margins.  Specimens  of  S.  tenuis  also  found  at  San  Quintin  were 
collected  on  sandy,  sunlit  shores  of  the  Rios  Jatate  and  Perlas. 

Taxonomic  notes.  — In  addition  to  characters  mentioned  in  the  diagnostic  combination, 
specimens  of  S.  sculptilis  generally  differ  from  specimens  of  S.  tenuis  by:  uniformly  dark 
elytra;  second  and  eighth  elytral  intervals  united  at  apex;  and  subapical  pit  in  lateral  channel 
smaller.  Ecological  and  morphological  differences,  and  sympatry  at  least  in  Chiapas,  indicate 
reproductive  isolation.  S.  sculptilis  agrees  with  S.  tenuis,  and  differs  from  other  South 
American  species,  by  having  enlarged  inner  eye  facets,  a peculiarity  suggesting  possible  close 
relationship  with  S.  tenuis.  But  S.  tenuis  is  otherwise  more  similar  to  other  South  American 
species,  and  I suspect  the  relationship  between  S.  sculptilis  and  S.  tenuis  is  remote. 

Schizogenius  tenuis  Bates 

Schizogenius  tenuis  Bates  1881 :38.  Type  locality  Paso  Antonio,  Guatemala;  type  in  BMNH, 

specimen  labelled  as  holotype  here  designated  lectotype  (!). 

Diagnostic  combination.  — Specimens  of  this  species  differ  from  those  of  S.  sculptilis, 
the  only  other  member  of  the  tenuis  group  known  from  Middle  America,  most  conspicu- 
ously by  differences  in  elytral  structure:  interval  six  not  carinate  except  finely  at  extreme 
apex,  and  subapical  pits  of  lateral  channel  larger.  Also,  the  basal  stylets  of  the  median  lobe 
are  strikingly  different.  Specimens  of  S.  tenuis  differ  from  other  South  American  specimens 
of  the  group  seen  by  me  by  the  combination  of:  neck  punctate  medially;  elytra  not  sellate 
or  maculate;  and  eyes  in  most  specimens  with  inner  facets  enlarged. 

Description.  — Color  dark  piceous,  legs,  palpi,  antennae,  and  in  most  specimens  elytral 
apices  paler;  elytra  with  slight  aeneous  luster,  not  sellate. 

Integument.  Fine  but  conspicuous  microsculpture  on  paramedian  sulci  of  frons,  genae, 
gula,  mouthparts,  sides  and  base  of  pronotum,  front  tibiae  and  anterior  surfaces  of  front 
femora,  middle  legs  except  trochanters,  hind  tibiae  and  posterior  surfaces  of  hind  femora, 
elytral  epipleura  on  base  and  apical  two-thirds,  and  coxal  depressions  of  sternum  three. 
Abdomen  without  extensive  microsculpture  on  sides  of  sterna  four  to  six  or  margin  of 
sternum  seven. 

Head.  Fig.  40.  Clypeus  with  apices  of  paramedian  carinae  fused  to  form  an  arc,  or  nearly 
so;  median  field  hemicircular,  width  at  base  more  than  2.0  apical  width  of  median  field  of 
frons.  Frons  with  median  longitudinal  sulcus  narrowed  in  front.  Eyes  prominent,  subglobose, 
coarsely  faceted,  inner  facets  usually  larger  than  marginal  facets.  Neck  densely  and  coarsely 
punctate.  Genae  strongly  punctate,  finely  rugose  in  front.  Mentum  (Fig.  18)  with  lateral 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


159 


lobes  acutely  produced,  median  tooth  conspicuous  but  short  and  sharp.  Antennae  short, 
moniliform,  article  five  1.0- 1.1  times  as  long  as  wide. 

Pronotum.  Fig.  40.  Disc  convex,  slightly  transverse,  greatest  width  near  middle;  parame- 
dian sulci  short,  faintly  punctate,  deep  and  abrupt  apically,  deep  and  broadly  hooked 
basally,  basal  tips  normally  confused  laterad  with  basal  rugosity;  paralateral  sulci  absent; 
anterior  transverse  impression  impunctate  to  finely  punctate. 

Elytra.  Three  discal  setigerous  punctures  on  interval  three,  first  adjacent  to  second  stria, 
others  in  middle  of  interval;  interval  five  with  one  seta  at  base.  Intervals  one  to  six  broad, 
convex,  carinate  at  most  at  extreme  apices,  interval  six  not  carinate  basad  oi  posterior  discal 
seta;  interval  seven  carinate  throughout;  interval  eight  carinate  in  apical  half.  Apex  of  interval 
eight  often  fused  with  apices  of  intervals  three,  five,  and  seven,  as  well  as  with  interval  one. 
Lateral  channel  with  at  least  one  large  subapical  pit,  some  specimens  with  one  or  more 
additional  smaller  pits,  or  with  two  or  three  large  pits. 

Male  genitalia.  Median  lobe  (Fig.  51)  arcuate,  apical  part  not  suddenly  deflexed;  basal 
stylets,  Fig.  68;  endophallus  without  distinguishing  spines  or  scales  on  virga.  Thirteen  speci- 
mens studied. 

Measurements  and  proportions.  See  Table  3. 


Table  3.  Descriptive  statistics  for  S.  tenuis , based  on  22  unsexed  specimens  from  San  Quin- 
tin,  Chiapas. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measuren 

lents,  in  mm. 

TL 

3.38-3.98 

3.645 

0.263 

0.075 

4.82 

LE 

2.09-2.49 

2.255 

0.169 

0.048 

5.00 

WH 

0.68-0.92 

0.747 

0.055 

0.016 

4.89 

WP 

0.88-1.05 

0.957 

0.072 

0.021 

5.05 

WE 

1.04-1.25 

1.135 

0.084 

0.024 

4.93 

B.  Proportio 

ns. 

WF/WH 

0.49-0.57 

0.544 

0.031 

0.009 

3.78 

LP/WP 

0.95-1.00 

0.968 

0.015 

0.004 

1.06 

DP/LP 

0.80-0.87 

0.828 

0.032 

0.009 

2.58 

LP/WE 

0.79-0.85 

0.818 

0.027 

0.008 

2.16 

Ta/Ti 

0.60-0.75 

0.675 

0.070 

0.020 

6.87 

PS/LP 

0.62-0.68 

0.655 

0.027 

0.008 

2.73 

Variation.  — I found  no  important  geographic  variation  in  size  or  body  proportions,  and 
because  of  paucity  of  material  did  no  statistical  analysis  of  variation.  The  smallest  specimen 
studied  (LE,  1.86  mm)  is  from  Paso  Antonio,  Guatemala,  and  the  largest  (LE,  2.50  mm)  is 
from  Villahermosa,  Tabasco.  Variation  in  discal  setae  on  elytra  includes  reduction,  with  the 
middle  seta  of  interval  three  lacking  from  both  elytra  in  one  specimen  from  San  Luis  Potosi, 
and  addition,  with  four  setae  on  interval  three  of  one  or  both  elytra  in  several  specimens. 
All  specimens  have  at  least  one  large  subapical  pit  in  the  lateral  channel  of  the  elytron. 


160 


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Many  have  one  or  more  additional,  but  much  smaller,  pits  as  well.  Small  specimens,  espe- 
cially from  the  Paso  Antonio  and  El  Coyul  samples,  tend  to  have  unusually  large  secondary 
pits,  and  a strongly  marked  tendency  for  the  apices  of  elytral  intervals  three,  five,  and  seven 
to  converge  on  interval  eight.  I think  these  differences  lack  geographic  significance.  In  some 
Colombian  specimens,  and  less  noticeably  in  some  Costa  Rican  specimens,  the  inner  eye 
facets  are  not  as  distinctly  enlarged  as  in  specimens  from  more  northern  areas,  but  the 
transition  is  gradual. 

Etymology.  — Latin,  tenuis  = thin,  in  reference  to  body  form. 

Distribution.  — The  known  distribution  of  S.  tenuis  (Fig.  76)  extends  from  northern 
Mexico  to  Colombia,  from  low  elevations  to  as  high  as  4000'  in  Puebla.  I studied  203 
specimens  from  the  following  localities. 


MEXICO 

CHIAPAS:  San  Quin  tin  (23;  UASM).  DURANGO:  Ventanas  (1;  BMNH).  JALISCO:  18  km.  n.  Puerto  Vallarta  (25;  UASM). 
NAYARIT:  Acaponeta  (1;  CAS),  Jesus  Maria  (5;  UCB),  Rio  Santiago  Ferry  (2;  CAS),  5 mi.  s.  Rio  Santiago  Ferry  (4;  CAS). 
OAXACA:  17.7  mi.  w.  El  Camaron  (2;  DRWh),  29.4  mi.  e.  El  Coyul  (3;  AMNH,  CNC),  11.1  mi.  n.  Matias  Romero  (34; 
MGFT,  UASM),  Valle  Nacional  (5;  UASM).  PUEBLA:  Tepexco  (7;  UASM).  SAN  LUIS  POTOSI:  Tamazunchale  (3;  CNHM, 
DRWh).  SINALOA:  30.6  ml  s.  Culiacan  (2;  IRSB).  SONORA:  10  mi.  w.  Alamos  (1;  AMNH).  TABASCO:  Teapa  (3; 
BMNH),  Villahermosa  (2;  MCZ).  VERACRUZ:  20  mL  nw.  Huatusco  (1;  FDAG). 

GUATEMALA 

ESCU1NTLA:  Paso  Antonio  (2;  BMNH).  IZABAL:  Los  Amates  (2;  MCZ). 

HONDURAS 

CORTES:  La  Lima  (24;  FDAG). 

COSTA  RICA 

GUANACASTE:  Las  Canas  (2;  UASM).  LIMON:  Los  Diamantes  (18;  FDAG),  Reventazon  (9;  USNM).  PUNTARENAS: 
Palmar  Sur  (2;  UAFA). 


PANAMA 

CANAL  ZONE:  San  Pablo  (1;  USNM).  CHIRIQUI:  Tole  (6;  IRSB,  MCZ). 

COLOMBIA 

MAGDALENA:  Aracataca  (2;  MCZ),  Rio  Frio  (10;  MCZ).  TOLIMA:  Coyaima  (1;  CAS). 

Collecting  notes.  — I have  seen  specimens  of  S.  tenuis  collected  nearly  throughout  the 
year,  except  during  the  latter  part  of  the  dry  season  from  early  April  to  mid-June.  Numerous 
specimens  were  taken  at  black  lights,  which  probably  indicates  flight. 

I collected  numbers  of  this  species  only  at  Matias  Romero  and  San  Quintin.  Specimens  of 
several  species  were  collected  along  the  Rio  Malatengo  in  gravel  bars,  but  the  S.  tenuis  were 
found  alone,  in  sand.  All  but  three  of  the  San  Quintin  specimens  were  found  along  the  Rio 
Jatate  in  sandy  banks;  the  others  were  collected  on  the  Rio  Perlas  a short  distance  away, 
perhaps  also  in  sand.  One  specimen  from  Tamazunchale,  two  from  Culiacan,  and  three  from 
El  Coyul  were  also  taken  in  silt  or  fine  sand;  other  specimens  were  hand  collected  by  G.  E. 
Ball  or  me  at  El  Camaron  and  Valle  Nacional,  but  whether  in  sand  or  not  is  unrecorded.  S. 
tenuis  thus  probably  agrees  with  other  species  of  the  subgenus  by  living  in  a sandy  habitat, 
but,  unlike  those  of  S.  sculptilis,  specimens  are  more  likely  to  be  found  along  open  rivers. 

Taxonomic  notes.  — Mexico,  this  species  is  sympatric  with  both  S.  crenulatus,  at  Jesus 
Maria  and  Culiacan,  and  S.  sculptilis , at  San  Quintin.  In  Colombia,  it  is  sympatric  with  at 
least  two  other  members  of  the  tenuis  group;  specimens  of  S.  tenuis,  S.  impuncticollis,  and 
S.  suturalis  were  taken  at  Aracataca  by  P.  J.  Darlington. 

Possibly  related  but  probably  distinct  are  a female  from  Tucuman,  Argentina  (JNeg),  a 
male  (Fig.  52,  69)  from  Santa  Isabel  do  Araguaia,  Brazil  (CAS),  and  a specimen  of  unde- 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


161 


termined  sex  from  Tabilas,  Salta,  Argentina  (CAS).  In  addition  to  differences  in  male  genita- 
lia as  illustrated,  eyes  are  uniformly  faceted,  and  elytral  apices  are  pale  in  sharp  contrast  to 
the  rest  of  the  elytron. 


Schizogenius  impressicollis  Putzeys 

Schizogenius  impressicollis  Putzeys  1846:653.  Type  locality  Colombia;  lectotype  here  desig- 
nated, male  in  the  Institut  Royal  des  Sciences  Naturelles  de  Belgique,  so  labelled  (!). 

Putzeys  1863:24.  Putzeys  1866:223. 

Diagnostic  combination.  — Within  the  tenuis  group,  basal  stylets  (Fig.  70),  form  of  median 
lobe  (Fig.  53),  and  form  of  median  frontal  sulcus  (Fig.  41),  of  the  lectotype  are  distinctive. 
Also,  lateral  lobes  of  male  genitalia  are  broad  and  trisetose  apically,  rather  than  bisetose  as 
in  other  members  of  the  group.  The  elytra  probably  are  not  sellate,  the  eyes  are  uniformly 
faceted,  and  the  neck  is  sparsely  punctate  along  the  midline.  Since  I examined  only  one  old, 
discolored  specimen,  I prefer  not  to  give  a detailed  description. 

Distribution.  — The  specific  locality  at  which  the.  type  specimens  were  collected  in 
Colombia  is  unknown,  and  I have  seen  no  additional  specimens. 

Etymology.  — Latin,  impressus  = impressed,  plus  collum  = neck  in  reference  to  the  sulcate 
pronotum. 

Taxonomic  notes.  — In  the  original  description  of  this  species,  Putzeys  wrongly  stated  it 
to  have  three  or  four  setae  on  intervals  three,  five,  and  seven;  this  was  an  error  he  repeated 
in  several  other  species  descriptions. 

Schizogenius  impuncticollis  new  species 

Type  material.  — Holotype,  male,  labelled  “Aracataca,  Mgd.  Colombia  III  2 29  Darling- 
ton” (MCZ).  Allotype  female,  label  data  same  as  in  holotype  except  III  3 29  (MCZ).  Two 
additional  specimens  from  the  same  locality  are  paratypes  (MCZ). 

Diagnostic  combination.  — Specimens  of  this  species  are  readily  distinguished  from  other 
members  of  the  tenuis  group  by  combination  of:  neck  nearly  or  quite  impunctate  along 
midline;  basal  transverse  impression  of  pronotum  greatly  deepened;  body  strongly  aeneous 
or  metallic;  and  elytral  apices  contrastingly  pale. 

Description.  — Color  rufopiceous,  strong  aeneous  or  metallic  green  luster;  front  femora 
rufous;  antennae,  palpi,  elytral  apices,  and  legs  except  front  femora  testaceous. 

Integument  as  described  for  S.  tenuis. 

Head  and  pronotum.  Fig.  42.  Generally  as  described  for  S.  tenuis,  except  eyes  uniformly 
faceted,  neck  not  or  hardly  punctate  along  midline,  anterior  transverse  impression  strongly 
punctate,  and  basal  transverse  impression  unusually  deep. 

Elytra.  As  in  S.  tenuis  except  more  strongly  bicolored. 

Male  genitalia.  Median  lobe,  Fig.  54;  basal  stylets,  Fig.  71;  virga  without  distinguishing 
spines  or  scales.  Three  specimens  studied. 

Measurements  and  proportions.  Holotype:  TL,  3.35  mm;  LE,  2.05  mm;  WH,  0.69  mm; 
WP,  0.90  mm;  WE,  1 .09  mm;  WF/WH,  0.62;  LP/WP,  0.98;  DP/LP,  0.77;  LP/WE,  0.81 ; Ta/Ti, 
0.70;  PS/LP,  0.69.  Holotype  plus  paratypes:  TL,  3.23-3.36-3.49  mm;  LE.  1.99-2.06-2.15 
mm;  WH,  0.65-0.68-0.70  mm;  WP,  0.88-0.90-0.92  mm;  WE,  1.05-1.10-1.16  mm;  WF/WH, 
0.59-0.60-0.62;  LP/WP,  0.93-0.96-0.98;  DP/LP,  0.77-0.79-0.82;  LP/WE,  0.76-0.78-0.81 ; Ta / 
Ti,  0.66-0.70-0.72;  PS/LP,  0.67-0.68-0.70. 

Etymology.  — Latin,  im  = not,  plus  punctus  = punctate,  plus  collum  = neck,  in  allusion  to 
one  diagnostic  characteristic  of  specimens  of  this  species,  lack  of  punctures  along  midline 


162 


Whitehead 


of  neck. 

Distribution.  — Only  the  type  specimens  from  Aracataca,  in  northern  Colombia,  are 
known. 

Collecting  notes.  — Specimens  of  this  species  were  collected  in  March.  As  wings  are  fully 
developed,  adults  probably  can  fly. 

Taxonomic  notes.  - The  range  of  this  species  overlaps  that  of  at  least  S.  tenuis  and  S. 
suturalis  among  other  members  of  the  tenuis  group.  Also,  specimens  of  this  species  differ 
sufficiently  from  those  of  other  described  members  of  the  group  that  they  undoubtedly 
represent  a distinct  species. 

A female  from  Yoay,  Bolivia  (CAS),  a male  from  Yurac,  Peru  (CAS),  and  an  unsexed 
specimen  from  El  Cidral,  Bolivia  (MCZ)  agree  with  the  Colombian  specimens  in  having  the 
neck  impunctate.  These  specimens  are  much  less  strongly  metallic  and  have  less  strongly 
bicolored  elytra,  and  the  basal  transverse  impression  of  the  pronotum  is  shallower.  The 
female  (LE,  2.43  mm)  is  markedly  larger  than  specimens  from  Colombia,  and  the  male 
differs  in  form  of  median  lobe  (Fig.  55)  and  basal  stylets  (Fig.  72).  For  these  reasons  I 
doubt  that  these  specimens  represent  S.  impuncticollis.  Final  judgment  as  to  their  relation- 
ships must  await  study  of  additional  material. 

Schizogenius  suturalis  new  species 

Type  material.  - Holotype,  male,  and  allotype,  female,  labelled  “Aracataca,  Mgd.  Co- 
lombia III  2 29  Darlington”  (MCZ).  Five  additional  specimens  from  the  same  locality  are 
paratypes  (DRWh,  MCZ,  UASM). 

Diagnostic  combination.  — Specimens  of  this  species  differ  from  others  in  the  tenuis  group 
by  having,  in  combination,  sellate  elytra  and  one  seta  near  base  of  interval  five. 

Description.  — Color  rufopiceous,  with  strong  aeneous  or  metallic  green  luster;  front 
femora  rufous;  elytra  rufotestaceous,  central  macula  aeneopiceous;  antennae,  palpi,  and  legs 
except  front  femora  testaceous. 

Integument.  As  described  for  S',  tenuis. 

Head  and  pronotum.  Fig.  43.  Generally  as  described  for  S.  tenuis  except  eyes  uniformly 
faceted. 

Elytra.  Except  for  coloration,  as  described  for  S.  tenuis. 

Male  genitalia.  Median  lobe,  Fig.  56;  basal  stylets,  Fig.  73;  virga  without  distinguishing 
spines  or  scales.  Two  specimens  studied. 

Measurements  and  proportions.  Holotype:  TL,  3.56  mm;  LE,  2.18  mm;  WH,  0.70  mm; 
WP,  0.94  mm;  WE,  1.15  mm;  WF/WH,  0.57;  LP/WP,  0.97;  DP/LP,  0.81;  LP/WE,  0.79;  Ta/ 
Ti,  0.62;  PS/LP,  0.60.  Holotype  plus  paratypes:  TL,  3.19-3.58-3.77  mm;  LE,  1.96-2.21- 
2.35  mm;  WH,  0.65-0.70-0.75  mm;  WP,  0.83-0.95-1.02  mm;  WE,  1.01-1.15-1.23  mm;  WF/ 
WH,  0.55-0.57-0.61;  LP/WP,  0.94-0.95-0.97;  DP/LP,  0.81-0.82-0.84;  LP/WE,  0.77-0.79-0.81; 
Ta/Ti,  0.62-0.67-0.7 1 ; PS/LP,  0.57-0.62-0.66. 

Variation.  — One  paratype  has  an  extra  seta  on  left  margin  of  pronotum,  and  another  has 
four  setae  on  interval  three  of  right  elytron. 

Etymology.  — Latin,  sutura  = seam,  in  reference  to  sutural  macula. 

Distribution.  — The  only  known  specimens  are  from  Aracataca,  in  northern  Colombia. 

Collecting  notes.  — Specimens  of  this  species  were  collected  in  March  and  May.  Wings 
are  fully  developed,  so  individuals  probably  fly. 

Taxonomic  notes.  — S.  suturalis,  S.  tenuis,  and  S.  impuncticollis  are  sympatric  in  Colom- 
bia, so  are  undoubtedly  specifically  distinct.  The  only  other  described  species  included  in 
the  tenuis  group  which  has  the  sutural  macula  is  S.  maculatus  Kult.  If  that  species  really 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


163 


Fig.  35-43.  Head  and  pronotum,  dorsal  aspect.  35.  S.  crenulatus  crenulatus  LeConte,  Culiacan,  Sinaloa.  36.  S.  quin- 
quesulcatus  Putzeys,  Nova  Teutonia,  Brazil.  37.  S.  sze/cessyi  Kult,  Aracataca,  Columbia.  38.  S.  janae  Kult,  Jacare,  Brazil. 
39.  S.  sculptilis  new  species,  San  Quintin,  Chiapas.  40.  S.  tenuis  Bates,  Matias  Romero,  Oaxaca.  41.  5.  impressicollis 
Putzeys,  Columbia.  42.  S.  impuncticollis  new  species,  Aracataca,  Columbia.  43.  S.  suturalis  new  species,  Aracataca, 
Columbia.  Fig.  44-55.  Male  median  lobe,  lateral  aspect.  44.  S.  crenulatus  crenulatus  LeConte,  Yuma,  Arizona.  45.  S. 
crenulatus  chiapatecus  new  subspecies,  Rio  Jaltepec,  Oaxaca.  46.  S.  quinquesulcatus  Putzeys,  Nova  Teutonia,  Brazil. 
47.  Same,  Santa  Rosa,  Ecuador.  48.  S.  szekessyi  Kult,  Aracataca,  Columbia.  49.  S.  janae  Kult,  Jacare,  Brazil.  50.  S. 
sculptilis  new  species,  San  Quintin,  Chiapas.  51.  S.  tenuis  Bates,  Rio  Santiago,  Nayarit.  52.  S.  nr.  tenuis  Bates,  Santa 
Isabel,  Brazil.  53.  S.  impressicollis  Putzeys,  Colombia.  54.  S.  impuncticollis  new  species,  Aracataca,  Colombia.  55.  S. 
nr.  impuncticollis  new  species,  Yurac,  Peru. 


Whitehead 


Fig.  56.  Male  median  lobe,  lateral  aspect,  S.  suturalis  new  species,  Aracataca,  Colombia.  Fig.  57.  Male  endophallus, 
S.  crenulatus  crenulatus  LeConte,  Culiacan,  Sinaloa.  Fig.  58-73.  Basal  stylets  of  male  endophallus.  58,  59.  S.  crenulatus 
crenulatus  LeConte,  Culiacan,  Sinaloa.  60.  S.  crenulatus  chiapatecus  new  subspecies,  Rio  Jaltepec,  Oaxaca.  61,  62. 
S.  quinquesulcatus  Putzeys,  Nova  Teutonia,  Brazil.  63.  Same,  Santa  Rosa,  Ecuador.  64,  65.  S.  szekessyi  Kult,  Araca- 
taca, Colombia.  66.  S.  janae  Kult,  Jacare,  Brazil.  67.  S.  sculptilis  new  species,  San  Quintin,  Chiapas.  68.  S.  tenuis  Bates, 
Rio  Santiago,  Nayarit.  69.  S.  nr.  tenuis  Bates,  Santa  Isabel,  Brazil.  70.  S.  impressicollis  Putzeys,  Colombia.  71.  S.  im- 
puncticollis  new  species,  Aracataca,  Columbia.  72.  S.  nr.  impuncticollis  new  species,  Yurac,  Peru.  73.  S.  suturalis  new 
species,  Aracataca,  Colombia.  Fig.  74-76.  Known  distributions.  74.  S.  crenulatus  crenulatus  LeConte,  circles,  and  S. 
crenulatus  chiapatecus  new  subspecies,  triangles.  75.  S.  sculptilis  new  species.  76.  S.  tenuis  Bates. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


165 


belongs  in  the  group,  and  if  the  original  description  is  correct,  then  it  differs  from  S.  sutu- 
ralis  by  not  having  a basal  seta  on  interval  five. 

Schizogenius  maculatus  Kult 

Schizogenius  maculatus  Kult  1950:  143.  Type  locality  Para,  Amazonas,  Brazil;  type  in  Kult 

collection,  present  location  not  known,  not  seen  by  me. 

Diagnostic  combination.  — Assuming  the  original  description  is  correct,  and  if  this  species 
really  is  a member  of  the  tenuis  group,  specimens  of  it  should  be  readily  recognized  by  not 
having  a basal  seta  on  elytral  interval  five.  The  only  other  described  species  in  the  group 
having  a dark  sutural  macula  is  S.  suturalis,  which  may  differ  further  by  having  the  striae 
coarsely  punctate  basally. 

Taxonomic  notes.  — Since  I have  not  studied  specimens  which  I could  identify  with  this 
name,  I am  unable  to  give  further  data  on  S.  maculatus.  My  allocation  to  the  tenuis  group, 
and  indeed  to  the  subgenus  Genioschizus,  is  tentative,  since  critical  subgeneric  characteristics 
were  not  mentioned  in  Kult’s  description.  However,  the  only  small  species  of  Schizogenius 
with  a sutural  macula,  reduced  elytral  setation,  and  moniliform  antennae  known  to  me  are 
Genioschizus,  and  absence  of  paralateral  pronotal  sulci  indicates  placement  in  the  tenuis 
group.  Absence  of  basal  setae  from  interval  five  of  the  elytron  is  not  otherwise  known  in  this 
group,  but  is  characteristic  of  S.  crenulatus  crenulatus  in  the  crenulatus  group.  I do  not 
suspect  Kult  misinterpreted  this  characteristic,  since  he  correctly  described  it  in  S.  szekessyi 
and  S.  janae.  The  names  S.  maculatus  and  S.  suturalis  may  be  synonymous,  but  the  informa- 
tion available  to  me  indicates  otherwise. 

Subgenus  Schizogenius  sensu  stricto 

Type  species.  — Schizogenius  strigicollis  Putzeys  1846:650,  subsequent  designation  by 
Lindroth,  1961:  164. 

Diagnostic  combination.  — Lateral  channel  of  elytron  not  flared  near  apex,  without  deep 
subapical  pits;  and  females,  except  of  S.  pluripunctatus,  and  species  of  basalis  group,  nor- 
mally without  paramedian  ambulatory  setae  on  sternum  seven.  Also:  paramedian  clypeal 
carinae  tuberculate  or  not;  antennae  moniliform  to  filiform;  front  tarsi  markedly  dilated 
in  many  species,  more  strongly  so  in  males  than  females;  male  endophallus  of  many  species 
with  well  developed  basal  collar  spines;  paralateral  longitudinal  sulci  not  present  on  prono- 
tum;  clypeal  suture  obsolete  to  sharply  engraved;  and  pygidium  serrate  or  crenulate  in  fe- 
males of  many  species. 

Description.  — Size  variable,  LE  1.65-3.70  mm  in  North  and  Middle  America,  up  to  5.00 
mm  in  South  America.  Body  cylindrical  to  strongly  depressed.  Color  testaceous  or  ferrugi- 
neous  to  darkpiceous,  aeneous  or  metallic  in  some  species,  elytra  maculate  or  sellate  in  some 
South  American  species.  Integument  shiny  in  most  species,  dorsum  without  extensive  micro- 
sculpture except  in  some  South  American  species;  coxal  depressions  of  sternum  three  of 
most  species  with  small  patches  of  particularly  coarse  microsculpture;  microsculpture  iso- 
diametric  or  nearly  so. 

Head.  Labrum  (Fig.  2-3)  weakly  emarginate  or  biemarginate  apically,  or  in  some  South 
American  species  deeply  emarginate;  margined  laterally  with  six  or  seven  pairs  of  setae,  to 
about  fifteen  pairs  in  S.  optimus.  Clypeus  with  median  tooth  prominent,  nearly  or  quite  as 
large  as  paramedian  teeth,  except  in  some  South  American  species;  paramedian  carinae 
straight  or  arcuate,  joined  to  median  tooth  or  not;  median  field  triangular  or  hemicircular, 
base  of  varied  width.  Clypeal  suture  obsolete  to  deeply  engraved  at  middle.  Frons  with 


Whitehead 


1 66 


carinae  one  to  four  straight  and  nearly  parallel,  or  arcuate  in  some  South  American  species; 
carinae  one  and  four  more  strongly  elevated  or  not;  carina  five  distinct  or  not;  median  sulcus 
wider  than  at  least  outer  paramedian  sulci,  limited  in  front  by  transverse  carina  or  not,  with 
median  longitudinal  carina  in  some  species;  sides  of  median  sulcus  parallel,  divergent  behind, 
or  bowed  outward  at  middle.  Eyes  flat  to  prominent,  multifaceted,  inner  facets  not  enlarged 
in  most  species;  eyes  in  S.  ocellatus  each  reduced  to  an  apparently  single,  bubble-like  facet. 
Antennal  articles  five  to  ten  moniliform  to  elongate;  pedicel  unisetose,  or  bisetose  or  pluri- 
setose  in  some  South  American  species;  articles  three  and  four  pubescent,  or  plurisetose  in 
some  South  American  species;  articles  five  to  eleven  pubescent.  Mandibles  (Fig.  6-7)  elon- 
gate, prominent.  Mentum  (Fig.  19-20)  deeply  emarginate  at  middle,  or  shallowly  in  some 
South  American  species,  median  tooth  large  and  sharp,  or  blunt  in  some  South  American 
species;  antero-lateral  angles  of  lateral  lobes  acutely  produced;  anterior  paramedian  setae 
widely  spaced.  Submentum  without  accessory  setae,  except  in  S.  strigicollis.  Gula  narrow,  its 
narrowest  part  0.15-0.30  width  of  mentum,  or  less  than  0.10  in  some  South  American 
species. 

Thorax.  Pronotum  convex  to  flat,  transverse  to  elongate;  paramedian  longitudinal  sulci 
usually  present,  paralateral  sulci  absent;  lateral  grooves  narrow  and  deep,  or  broad  and 
shallow  in  some  South  American  species;  lateral  grooves  not  bordered  internally  by  carinae, 
except  in  some  South  American  species;  accessory  marginal  setae  present  or  absent;  hind 
angles  obsolete  to  prominent.  Prothoracic  pleura  impunctate  except  ins',  amphibius. 

Elytra.  Lateral  channel  narrow  to  broad  at  apex,  shallow,  not  flared,  without  foveae  or 
pits  above  umbilicate  series.  Disc  without  setae,  with  setae  on  interval  three,  intervals  three 
and  five,  or  on  intervals  three,  five,  and  seven.  Elytral  intervals  broad,  flat  to  moderately 
convex,  carinate  in  some  South  American  species;  interval  eight  and  in  some  species  intervals 
five  and  seven  carinate  at  apex;  interval  eight  free,  interval  seven  in  most  species  joined  by 
apices  of  intervals  three  and  five.  Striae  in  most  species  deeply  engraved,  in  most  species 
distinctly  punctate  at  least  in  basal  half. 

Hind  wings.  Fully  developed  except  in  S.  ocellatus.  Venation  as  in  Fig.  33-34;  oblongum 
cell  broad,  proximal  transverse  vein  not  broken. 

Legs.  Front  and  middle  tarsi  expanded  and  with  relatively  dense  ventral  pubescence  in 
many  species,  particularly  in  males,  or  narrow  in  both  sexes.  Hind  tarsi  slender,  short  to 
elongate;  article  one  of  hind  tarsus  2. 5-4.0  times  as  long  as  article  two.  Paronychia  varied 
from  about  half  to  quite  as  long  as  tarsal  claws,  or  obsolete  in  some  South  American  species. 
Front  tibia  narrowed  evenly  to  base  or  not;  distal  tooth  varied  in  form;  spurs  subequal, 
slender  and  acute;  posterior  ventral  margin  with  three  setae  proximad  to  spur,  or  four  in 
some  South  American  species. 

Abdomen.  Sternum  seven  with  paramedian  ambulatory  setae  present  in  males  of  most 
species,  absent  from  females  except  of  S.  pluripunctatus  and  of  members  of  basalis  group. 
Paramedian  carinae  of  sternum  three  curved  outward  at  apices  or  not.  Margin  of  pygidium 
(Fig.  24-25)  crenulate  or  not  in  females,  not  in  males. 

Male  genitalia.  Median  lobe  arcuate,  nearly  symmetric  in  most  species,  form  of  apex 
various.  Endophallus  with  distinctive  basal  collar  spines  in  many  species;  membrane  around 
virga  reduced  or  not;  basal  stylets  various,  distinctive  in  some  species. 

Female  ovipositor.  Number  and  position  of  setae  variable,  of  no  value  in  species  recogni- 
tion (Fig.  28-29). 

Distribution.  — Members  of  this  subgenus  range  from  southern  Canada  southward  to 
central  Argentina.  One  species  is  known  from  Cuba  in  the  West  Indies.  I examined  9930 
specimens  of  the  subgenus  Schizogenius. 

Taxonomic  notes.  — The  subgenus  is  most  diverse  in  South  America.  Since  I treat  only 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


167 


the  North  and  Middle  American  fauna  in  detail,  I deliberately  mention  characteristics  pecu- 
liar to  South  American  forms  in  the  description.  These  characteristics  are  not  repeated  in 
species  descriptions,  except  in  species  groups  where  appropriate. 


Key  to  described  North  and  Middle  American  species  groups,  species,  and  subspecies  of 
subgenus  Schizogenius 

1.  Pronotum  with  paramedian  longitudinal  sulci  obsolete;  clypeal  suture  obsolete; 

neither  sex  with  paramedian  ambulatory  setae  on  sternum  seven  ( optimus  group) 

S.  optimus  Bates,  p.  173 

1’  Pronotum  with  deep  paramedian  longitudinal  sulci;  clypeal  suture  sharply  en- 

graved; at  least  males  with  paramedian  ambulatory  setae  on  sternum  seven  . . . 
2 


2.(1’). 

2’ 

3.  (2). 
3’ 

4. (2’). 
4’ 

5. (4’). 
5’ 

6. (5’). 
6’ 

7. (6). 
7’ 

8. (7). 
8’ 

9. (8). 
9’ 

10.(7’). 


Paramedian  clypeal  carinae  tuberculate,  not  or  hardly  extended  to  median  tooth; 

antennal  articles  four  to  ten  moniliform  {ferrugineus  group)  3 

Paramedian  clypeal  carinae  extended  to  median  tooth;  antennal  articles  four  to 

ten  moderately  to  strongly  elongated,  filiform 4 

Abdomen  strongly  microsculptured;  body  without  metallic  luster 

S.  ferrugineus  Putzeys,  p.  179 

Abdomen  not  extensively  microsculptured;  body  with  distinct  aeneous  luster  . . . . 

S.  auripennis  Bates,  p.  182 

Paramedian  pronotal  sulci  extended  forward  nearly  to  anterior  transverse  impres- 
sion; front  tarsi  slender  in  both  sexes  ( truquii  group) 

S.  truquii  Putzeys,  p.  204 

Paramedian  pronotal  sulci  shorter;  front  tarsi  slightly  to  strongly  dilated,  at  least 

in  males 5 

Interval  seven  with  at  most  one  seta,  usually  with  none  ( lindrothi  group)  

S.  lindrothi  new  species,  p.  199 

Interval  seven  of  elytron  with  at  least  three  discal  setae 6 

Discal  setae  of  elytra  average  1 .2  or  more  times  longer  than  maximum  width  of 
interval  two;  striae  usually  indistinctly  punctate;  body  form  cylindrical  to  sub- 
cylindrical;  pronotum  with  or  without  accessory  marginal  setae  {pluripunctatus 

group) 7 

Discal  setae  of  elytra  average  no  more  than  1 .0  times  longer  than  maximum  width 
of  interval  two;  striae  usually  distinctly  punctate;  body  form  various;  pronotum 

without  accessory  marginal  setae 12 

Pronotum  with  accessory  marginal  setae  8 

Pronotum  without  accessory  marginal  setae 10 

Front  and  middle  tarsi  strongly  expanded  in  both  sexes.  California  and  Baja 

California 9 

Front  and  middle  tarsi  not  strongly  expanded.  Northeastern  Mexico 

S.  plurisetosus  new  species,  p.  214 

Disc  of  elytron  with  fewer  than  60  setae.  California 

S.  seticollis  seticollis  Fall,  p.  209 

Disc  of  elytron  with  more  than  60  setae.  Southern  Baja  California 

S.  seticollis  vandykei  new  subspecies,  p.  212 

Females  with  paramedian  ambulatory  setae;  apex  of  median  lobe  weakly  de- 
flected; disc  of  elytron  with  35  or  more  setae.  Arizona  and  New  Mexico  south  to 
Durango  and  northern  Nayarit,  north  of  Rio  Grande  de  Santiago  drainage  basin 
S.  pluripunctatus  LeConte,  p.  221 


168 


Whitehead 


10’ 

11.(10’). 

11’ 


12.(6’). 

12’ 

13. (12’). 
13’ 

14. (13). 
14’ 

15. (14’). 
15’ 

16. (15’). 
16’ 

17. (16). 
17’ 

18. (17’). 
18’ 

19. (16’). 
19’ 

20. (13’). 
20’ 

21. (20’). 
21’ 


Females  without  paramedian  ambulatory  setae;  apex  of  median  lobe  arcuate  or 
sharply  deflected.  Not  known  from  north  of  Rio  Grande  de  Santiago  drainage 

basin 11 

Apex  of  median  lobe  sharply  deflected;  elytron  normally  with  more  than  40 
discal  setae,  none  seen  with  fewer  than  37.  San  Luis  Potosi  south  to  Puebla  and 

Oaxaca S.  multisetosus  Bates,  p.  216 

Apex  of  median  lobe  not  angularly  deflected;  elytron  normally  with  fewer  than 
35  setae,  none  seen  with  more  than  41.  Central  Nayarit  in  west  and  Veracruz  in 

east,  south  at  least  to  Guatemala,  near  coasts  or  not 

S.  kulti  new  species,  p.  223 

Elytron  with  35  or  more  short  discal  setae;  abdomen  not  extensively  microsculp- 
tured;  body  strongly  depressed;  color  castaneous;  eyes  prominent  ( brevisetosus 

group) S.  brevisetosus  new  species,  p.  206 

Elytron  with  fewer  than  35  discal  setae,  or  abdomen  extensively  microsculp- 

tured 13 

Abdomen  with  entire  ventral  surface  microsculptured,  or  nearly  so;  paramedian 
frontal  sulci  not  or  hardly  closed  behind;  body  moderately  to  strongly  depressed; 

not  in  West  Indies  {tristriatus  group)  14 

Abdomen  not  with  entire  ventral  surface  microsculptured,  or  paramedian  frontal 

sulci  strongly  closed  behind,  or  in  West  Indies 20 

Prothoracic  pleura  distinctly  punctate;  small,  LE  under  2.50  mm  

S.  amphibius  Haldeman,  p.  236 

Prothoracic  pleura  not  distinctly  punctate;  larger,  LE  over  2.50  mm 15 

Prothoracic  pleura  strongly  microsculptured  

S.  tibialis  new  species,  p.  234 

Prothoracic  pleura  not  or  weakly  microsculptured 16 

Range,  United  States;  eyes  flattened,  WF/WH  0.65  or  more;  each  elytron  normal- 
ly with  25  or  more  discal  setae 17 

Range,  Mexico;  eyes  prominent,  WF/WH  0.65  or  less;  each  elytron  normally  with 

fewer  than  25  discal  setae 19 

Range,  east  of  Mississippi  River;  median  lobe.  Fig.  175  

S.  planulatus  LeConte,  p.  238 

Range,  west  of  Mississippi  River 18 

Range,  north  of  Red  River;  median  lobe,  Fig.  176 

S.  ozarkensis  new  species,  p.  240 

Range,  south  of  Red  River;  median  lobe,  Fig.  177 

S.  planuloides  new  species,  p.  241 

Elytra  rufopiceous,  apices  reddish;  front  tibia  not  dilated  proximally 

S.  tristriatus  Putzeys,  p.  23 1 

Elytra  castaneous,  concolorous;  front  tibia  distinctly  expanded  proximally 

S.  dilatus  new  species,  p.  232 

Abdomen  distinctly  microsculptured  along  midline  {sallei  group)  

S.  sallei  Putzeys,  p.  229 

Abdomen  not  distinctly  microsculptured  along  midline,  or  if  so  then  entire  abdo- 
men microsculptured  21 

Sternum  three  without  small  paralateral  patch  of  distinct  microsculpture  on  each 

side  {longipennis  group) 22 

Sternum  three  with  small  paralateral  patch  of  distinct  microsculpture  on  each  side 
25 


22.(21). 

22’ 

23. (22’). 
23’ 

24. (23’). 
24’ 

25. (21’). 
25’ 

26. (25’). 
26’ 

27. (26). 
27’ 

28. (26’). 
28’ 

29. (28’). 
29’ 

30. (29). 
30’ 

31. (29’). 
31’ 

32. (31’). 
32’ 

33. (32’). 
33’ 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius  169 


Range,  Arizona  and  New  Mexico;  front  femur  piceous;  frontal  carina  three  in 
most  specimens  abbreviated  behind;  apex  of  median  lobe  (Fig.  197)  deflected  at 

nearly  right  angle S.  neovalidus  new  species,  p.  252 

If  in  Arizona  and  New  Mexico,  then  front  femur  reddish;  frontal  carina  three 
in  most  specimens  not  abbreviated  behind;  median  lobe  with  apex  less  sharply 

deflected 23 

Elytra  rufous  to  rufopiceous;  endophallus  (Fig.  204)  with  basal  collar  spines 

broad.  Southern  Arizona  S.  chiricahuanus  new  species,  p.  257 

Elytra  normally  piceous 24 

Endophallus  (Fig.  203)  with  basal  collar  spines  broad;  virga  small;  most  specimens 

smaller,  stockier,  and  with  tibiae  paler.  Western  Mexico 

S.  pacificus  new  species,  p.  258 

Endophallus  (Fig.  205)  with  basal  collar  spines  narrow;  virga  large.  Arizona  and 

Tamaulipas  south  to  Costa  Rica S.  longipennis  Putzeys,  p.  254 

Pronotal  hind  angles  sharply  developed;  elytra,  except  in  specimens  from  Rio 
Grande  drainage,  piceous  with  aeneous  tinge  (in  Rio  Grande  area,  elytra  casta- 
neous  with  clearly  paler  apices).  Rio  Grande  drainage  and  northward,  east  of 

Rocky  Mountains  ( lineolatus  group) S.  lineolatus  Say,  p.  246 

Pronotal  hind  angles  not  sharply  developed,  or  if  so  then  elytra  pale,  unicolorous 


( depressus  group) 26 

Paramedian  frontal  sulci  strongly  closed  behind  by  fused  paramedian  carinae; 

elytra  testaceous  to  brunneous 27 

Paramedian  frontal  sulci  not  or  hardly  closed  behind,  or  elytra  piceous 28 

Abdomen  extensively  microsculptured.  Northern  and  central  California 


S.  ochthocephalus  new  species,  p.  285 

Abdomen  not  extensively  microsculptured.  Not  in  northern  and  central  California 

S.  falli  new  species,  p.  281 

Abdomen  extensively  microsculptured.  Color  piceous.  West  Indies 

S.  arimao  Darlington,  p.  264 


Abdomen  not  extensively  microsculptured.  Not  in  West  Indies 29 

Elytra  piceous,  and  range  United  States  and  Canada 30 

Elytra  testaceous  to  brunneous,  or  range  Mexico  and  southward 31 


Range,  east  of  Mississippi  River  S.  sulcifrons  Putzeys,  p.  265 

Range,  Pacific  coastal  drainage  systems  S.  litigiosus  Fall,  p.  268 

Pronotal  hind  angles  normally  prominent;  color  brunneous;  larger,  LE  normally 

over  2.15  mm S.  depressus  LeConte,  p.  287 

Pronotal  hind  angles  normally  rounded;  color  various;  size  various,  LE  under  or 

over  2.15  mm .'32 

Piceous;  sternum  three  with  paralateral  patches  of  microsculpture  reduced,  indis- 
tinct; elytron  with  12  to  14  discal  setae.  Guatemala  to  Costa  Rica;  median  lobe, 

Fig.  223 S.  emdeni  new  species,  p.  265 

Color  various;  sternum  three  with  paralateral  patches  of  microsculpture  distinct; 

elytron  with  14  or  more  discal  setae 33 

Color  testaceous  to  brunneous,  not  piceous;  range,  northeastern  Mexico,  and 
United  States  from  Rio  Grande  River  northward,  east  of  Rocky  Mountains;  me- 
dian lobe,  Fig.  226,  227  S.  scopaeus  new  species,  p.  278 

Color  various;  range,  United  States  west  of  Rocky  Mountains,  and  Mexico  south 
to  Colombia;  in  northeastern  Mexico,  color  piceous;  median  lobe,  Fig.  228,  229 
S.  pygmaeus  Van  Dyke,  p.  270 


170 


Whitehead 


Partial  key  to  South  American  species  groups  and  species  of  the  subgenus  Schizogenius 


1. 


2.(1). 


3. (2’). 
3’ 

4. (3). 
4’ 


5. (3’). 
5’ 

6. (1’). 


6’ 

7.(6). 


8.(7’). 


8’ 


9.(8’). 


10.(6’). 

10’ 

11.(10). 

ir 

12.(10’). 

12’ 

13. (12). 
13’ 

14. (12’). 
14’ 

15. (14). 


Hind  tarsi  elongate,  Ta/Ti  0.75  or  more;  paramedian  frontal  sulci  not  micro- 
sculptured;  clypeal  suture  not  strongly  engraved;  elytral  disc  asetose  or  with  setae 

on  interval  three  only 2 

Hind  tarsi  shorter,  Ta/Ti  under  0.75;  paramedian  frontal  sulci  microsculptured  in 

most  species;  clypeal  suture  weakly  to  strongly  engraved 6 

Paramedian  pronotal  sulci  well  developed;  interval  three  of  elytron  bi-  or  trisetose 

( jacarensis  group) S.  jacarensis  new  species,  p.  172 

Combination  of  characters  not  as  above  ( optimus  group) 3 

Paramedian  pronotal  sulci  obsolete  or  nearly  so 4 

Paramedian  pronotal  sulci  well  developed;  elytral  disc  asetose 5 

Elytra  piceous,  metallic;  clypeal  carinae  straight,  apices  abbreviated;  elytra  ovate 

S.  dyschirioides  Putzeys,  p.  175 

Elytra  brunneous,  unmetallic;  clypeal  carinae  arcuate,  apices  joined  and  extended 

by  stem  to  median  tooth;  elytra  elongate,  not  ovate 

S.  clivinoides  Putzeys,  p.  176 

Elytron  with  large  sutural  macula S.  grossus  Whitehead,  p.  176 

Elytron  not  maculate S.  bicolor  new  species,  p.  177 

Elytral  disc  with  setae  on  intervals  three,  five,  and  seven;  antennal  articles  five 
to  ten  nearly  moniliform;  paramedian  clypeal  carinae  tuberculate,  apices  not 
extended  to  median  tooth;  clypeal  field  broad,  at  base  more  than  1.5  apical  width 

of  median  frontal  sulcus  ( basalis  group) 7 

Combination  of  characters  not  as  above 10 

Pygidium  not  strongly  crenulate  in  either  sex;  color  ferrugineous,  not  or  slightly 

aeneous S.  negrei  new  species,  p.  186 

Pygidium  strongly  crenulate  in  females;  at  least  pronotum  aeneopiceous  to  metal- 
lic   8 

Elytral  striae  not  distinctly  punctate  in  basal  half;  occiput  densely  punctate 

S.  multipunctatus  Kult,  p.  185 

Elytral  striae  distinctly  punctate  in  basal  half;  occiput  not  densely  punctate  . . . 

9 

Elytral  disc  with  more  than  40  setae;  elytral  length  over  2.25  mm;  pronotum 

piceous,  metallic  at  sides  and  base  only  S.  basalis  Putzeys,  p.  184 

Elytral  disc  with  fewer  than  40  setae;  elytral  length  under  2.25  mm;  pronotum 
entirely  metallic  green  S.  cearaensis  new  species,  p.  185 

Paramedian  pronotal  sulci  evident  only  at  base;  elytral  intervals  three,  five,  and 

seven  asetose  ( elongatus  group)  11 

Paramedian  pronotal  sulci  more  elongated,  developed  well  in  front  of  base 

12 

Metallic  green;  total  length  over  5 mm S.  costiceps  Steinheil,  p.  1 88 

Less  strongly  metallic;  total  length  about  4 mm S.  elongatus  Kult,  p.  188 

Elytral  disc  asetose,  intervals  carinate  ( carinatus  group) 13 

Elytral  disc  with  setae  at  least  on  interval  three 14 

Body  color  piceous S.  carinatus  Whitehead,  p.  189 

Body  color  rufotestaceous S.  costipennis  new  species,  p.  190 

Elytral  disc  with  setae  on  interval  three  only 15 

Elytral  disc  with  setae  on  interval  five  16 

Eye  normal,  multifaceted  ( quadripunctatus  group) 

S.  quadripunctatus  Putzeys,  p.  202 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


171 


15’ 

16. (14’). 
16’ 

17. (16). 
17’ 

18. (17). 
18’ 

19. (17’). 
19’ 

20. (19’). 
20’ 

21. (16’). 


21’ 


Eye  ocellate,  reduced  to  one  bubble-like  facet  ( ocellatus  group)  

S.  ocellatus  new  species,  p.  196 


Elytral  disc  with  setae  on  intervals  three  and  five  only 17 

Ely tral  disc  with  setae  on  intervals  three,  five,  and  seven 21 

Elytral  interval  five  with  setae  in  basal  half  only  {are  chav  ale  tae  group) 18 

Elytral  interval  five  with  setae  evenly  distributed 19 


Humeral  angles  of  elytra  prominent S.  reichardti  new  species,  p.  193 

Humeral  angles  of  elytra  rounded S.  arechavaletae  Putzeys,  p.  1 92 

Inner  paramedian  frontal  carinae  grossly  enlarged;  median  frontal  sulcus  micro- 

sculptured,  with  or  without  median  carina {darlingtoni  group),  p.  197 

Inner  paramedian  frontal  carinae  normal;  median  frontal  sulcus  not  microsculp- 

tured,  without  median  carina 20 

Antennal  articles  five  to  ten  short,  nearly  moniliform  {lindrothi  group) 

S.  banningeri  Kult,  p.  201 

Antennal  articles  five  to  ten  elongate,  filiform {capitalis  group),  p.  1 98 

Submen  turn  with  numerous  accessory  setae;  pronotum  with  lateral  grooves  bor- 
dered internally  by  distinct  carinae  ( strigicollis  group) 

S.  strigicollis  Putzeys,  p.  191 

Submen  turn  with  two  standard  pairs  of  setae  only;  pronotum  without  lateral 
carinae {depressus  group),  p.  263 


As  I am  unfamiliar  with  some  named  South  American  species  of  subgenus  Schizogenius 
and  cannot  evaluate  their  characteristics  from  original  descriptions,  this  key  is  provisional 
and  incomplete.  In  particular,  I have  no  useful  concepts  of  forms  named  S.  canaliculatus 
Putzeys,  S.  gracilis  Putzeys,  S.  leprieuri  Castelnau,  S.  sellatus  Putzeys,  and  S.  sulcatulus 
Putzeys,  and  cannot  place  them  to  species  group.  From  original  descriptions,  I associate  the 
names  S.  clivinoides  Putzeys  and  S.  interstriatus  Putzeys  with  species  groups,  but  not  with 
specimens.  I tentatively  associate  the  names  S.  costiceps  Steinheil  and  S.  putzeysi  Kirsch 
with  actual  specimens.  All  other  associations  are  based  either  on  type  material  or  on  topo- 
typic  or  nearly  topotypic  specimens  which  closely  match  good  original  descriptions.  I did 
not  key  South  American  species  belonging  to  the  darlingtoni,  capitalis,  or  depressus  groups, 
since  I am  insufficiently  familiar  with  them. 

The  jacarensis  group 

Diagnostic  combination.  — Members  of  this,  group  may  be  distinguished  by  the  following 
combination  of  characters:  hind  tarsi  elongate,  Ta/Ti  over  0.75;  clypeal  suture  weakly 
engraved;  paramedian  frontal  sulci  not  microsculptured;  elytral  disc  with  two  or  three  setae 
on  interval  three;  paramedian  pronotal  sulci  well  developed;  basal  sensory  bristles  of  termi- 
nal article  of  maxillary  palpus  arranged  transversely;  and  apex  of  median  lobe  of  male 
genitalia  neither  elongate  nor  strongly  deflexed.  Also:  size  large  (LE  over  2.75  mm);  integu- 
ment not  extensively  microsculptured;  labrum  strongly  biemarginate,  fringed  laterally  with 
six  or  seven  pairs  of  setae;  clypeal  carinae  straight  and  abbreviated;  clypeal  field  at  base 
more  than  1.5  apical  width  of  median  frontal  sulcus;  clypeus  with  median  tooth  reduced; 
frontal  carinae  not  arcuate;  antennal  articles  five  to  ten  moniliform,  pedicel  unisetose, 
articles  three  and  four  pubescent;  mentum  deeply  emarginate,  median  tooth  blunt;  front 
and  middle  tarsi  slender  in  both  sexes;  paronychia  nearly  obsolete;  posterior  ventral  margin 
of  front  tibia  with  three  setae  near  base;  paramedian  carinae  of  sternum  three  straight; 
paramedian  ambulatory  setae  present  on  sternum  seven  of  males  only;  pygidium  not  crenu- 


172 


Whitehead 


late  in  either  sex;  and  endophallus  without  enlarged  basal  collar  spines. 

Distribution.  — This  monobasic  group  is  known  only  from  three  specimens  from  central 
Brazil. 


Schizogenius  jacarensis  new  species 

Type  material.  — Holotype  male  and  allotype  female  labelled  “Jacare  P.  N.  Xingu  M. 
Grosso  — Bras.  XI.  1961  leg.  M.  Alvarenga”  (MGFT).  A specimen  from  Goias  state  is  a 
paratype  (DRWh). 

Diagnostic  combination.  — S.  jacarensis  is  the  only  known  member  of  the  jacarensis 
group. 

Description.  — Body  stout,  subcylindrical,  head  proportionately  large.  Color  rufocasta- 
neous,  elytra  strongly  aeneous  or  weakly  metallic,  not  maculate;  legs  and  mouthparts  rufous; 
tarsi  and  antennae  testaceous. 

Integument.  Fine  microsculpture  on  genae,  mouthparts,  pronotal  base,  front  tibiae  and 
anterior  surfaces  of  front  femora,  middle  legs  except  trochanters,  hind  tibiae  and  posterior 
surfaces  of  hind  femora,  sterna  two  and  three,  sides  of  sterna  four  to  six,  and  margin  of 
sternum  seven.  Sternum  three  without  patches  of  particularly  coarse  microsculpture  in  coxal 
depressions. 

Head.  Fig.  77.  Labrum  strongly  biemarginate,  fringed  laterally  with  six  or  seven  pairs  of 
setae.  Paramedian  clypeal  carinae  straight,  attenuate;  median  clypeal  field  triangular,  width 
at  base  much  more  than  1.5  apical  width  of  median  field  of  frons;  median  clypeal  tooth 
much  shorter  than  paramedian  teeth.  Clypeal  suture  obsolete  or  nearly  so.  Frontal  carinae 
weakly  arcuate,  median  frontal  sulcus  narrowed  from  base  to  apex.  Antennal  articles  five  to 
ten  moniliform,  pedicel  unisetose,  articles  three  and  four  pubescent.  Eye  subglobose,  uni- 
formly faceted.  Terminal  article  of  maxillary  palpus  with  basal  sensory  bristles  arranged 
transversely.  Mentum  deeply  emarginate,  median  tooth  blunt. 

Pronotum.  Fig.  77.  Sides  bisetose,  hind  angles  reduced;  base  not  rugose;  paramedian  sulci 
sharply  engraved;  anterior  transverse  impression  impunctate.  Pronotum  widest  well  in  front 
of  middle. 

Legs.  Front  and  middle  tarsi  slender  in  both  sexes,  without  dense  ventral  pubescence; 
hind  tarsus  slender,  elongate.  Front  tibia  narrowed  evenly  to  base;  postero-ventral  margin 
with  three  setae  near  base.  Front  femur  not  strongly  constricted  near  apex.  Paronychia  much 
less  than  half  length  of  tarsal  claws. 

Elytra.  Disc  with  two  or  three  setigerous  punctures  on  interval  three.  Striae  deep  and 
sharply  engraved,  finely  punctate  in  basal  two-thirds.  Intervals  one  to  eight  broad,  convex, 
apices  not  carinate;  intervals  three,  five,  and  seven  joined  at  apices. 

Abdomen.  Paramedian  carinae  of  sternum  three  straight.  Sternum  seven  with  paramedian 
ambulatory  setae  in  males  only.  Pygidium  with  apical  margin  entire  in  both  sexes. 

Male  genitalia.  Median  lobe,  Fig.  81,  apex  neither  elongate  nor  sharply  deflexed;  endo- 
phallus without  distinctive  spines.  One  specimen  examined. 

Measurements  and  proportions.  Of  holotype:  TL,  5.36  mm;  LE,  3.25  mm;  WH,  1.42  mm; 
WP,  1.60  mm;  WE,  1.68  mm;  WF/WH,  0.75;  LP/WP,  0.80;  DP/LP,  1.02;  LP/WE,  0.76;  Ta/ 
Ti,  0.81;  PS/LP,  0.60.  Of  allotype:  TL,  4.94  mm;  LE,  3.07  mm;  WH,  1.24  mm;  WP,  1.43 
mm;  WE,  1.56  mm;  WF/WH,  0.73;  LP/WP,  0.82;  DP/LP,  1.06;  LP/WE,  0.75;  Ta/Ti,  0.83; 
PS/LP,  0.58. 

Variation.  — Two  specimens  have  three  setae  on  interval  three  of  the  left  elytron. 

Etymology.  - S.  jacarensis  is  named  after  the  type  locality,  Jacare  National  Park. 

Distribution.  — I studied  three  specimens  of  this  species,  from  the  following  localities. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


173 


BRAZIL 

GOIAS:  Santa  Isabel  do  Morro  (1;  DRWh).  MATO  GROSSO:  Jacare  P.  N.  Xingu  (2;  MGFT). 


Collecting  notes.  — Specimens  of  this  species  were  collected  in  June  and  November. 

Taxonomic  notes.  — This  species  probably  represents  an  early  side  branch  of  the  optimus 
group,  since  the  weakly  engraved  clypeal  suture,  elongate  hind  tarsi,  and  unmicrosculptured 
paramedian  frontal  sulci  are  all  characteristics  shared  with  members  of  that  group.  Males  do 
not  have  the  elongated  apex  characteristic  in  genitalia  of  members  of  the  optimus  group, 
and  the  head  is  proportionately  much  larger,  the  eyes  smaller,  and  the  pronotum  more 
transverse.  As  in  all  other  members  of  Schizogenius  except  those  of  the  optimus  group, 
basal  sensory  bristles  of  terminal  articles  of  maxillary  palpi  in  S.  jacarensis  are  transversely 
arranged. 


The  optimus  group 

Diagnostic  combination.  — Members  of  this  group  are  distinguished  by  the  following 
combination  of  characters:  hind  tarsi  elongate,  Ta/Ti  over  0.75;  clypeal  suture  weakly  en- 
graved; paramedian  frontal  sulci  not  microsculptured;  elytral  disc  asetose  or  with  setae  on 
interval  three  only;  if  interval  three  setose,  then  paramedian  pronotal  sulci  obsolete  or  nearly 
so;  basal  sensory  bristles  of  terminal  article  of  maxillary  palpus  arranged  longitudinally;  and 
apex  of  median  lobe  of  male  genitalia  elongate  and  strongly  deflexed.  Also:  size  medium  to 
large  (LE  2.75-5.00  mm);  integument  extensively  microsculptured  or  not,  testaceous  to 
dark  piceous,  metallic  or  not,  elytra  maculate  in  one  species;  labrum  weakly  to  strongly 
emarginate  or  biemarginate,  fringed  laterally  with  six  or  seven  to  about  15  pairs  of  setae; 
clypeal  carinae  straight  and  abbreviated,  or  fused  in  an  arc  and  joined  to  median  tooth  by 
common  stem;  clypeal  field  at  base  more  than  1.5  apical  width  of  median  frontal  sulcus; 
clypeus  with  median  tooth  reduced  or  not;  antennal  articles  five  to  ten  moniliform,  pedicel 
uni-  or  bisetose,  articles  three  and  four  plurisetose  to  pubescent;  mentum  shallowly  to 
deeply  emarginate,  median  tooth  blunt  to  sharp;  gula  0.05-0.20  width  of  mentum;  anterior 
and  middle  tarsi  slender  in  both  sexes;  paronychia  obsolete  to  about  half  length  of  tarsal 
claws;  posterior  ventral  margin  of  front  tibia  with  three  to  four  setae  near  base;  paramedian 
carinae  of  sternum  three  straight;  paramedian  ambulatory  setae  present  on  sternum  seven  of 
males  only,  or  absent;  pygidium  not  crenulate  in  either  sex;  and  endophallus  without  en- 
larged basal  collar  spines. 

Distribution.  — One  Middle  American  species  extends  from  Chiapas  south  to  Costa  Rica. 
Four  other  species  occur  in  Panama  and  much  of  continental  South  America.  I studied  1 17 
specimens  of  the  optimus  group. 

Schizogenius  optimus  Bates 

Schizogenius  optimus  Bates  1881 : 37.  Type  locality  Rio  Naranjo,  Guatemala;  type  in  BMNH, 

specimen  labelled  lectotype  by  G.  E.  Ball  here  so  designated  (!). 

Diagnostic  combination.  — This  is  the  only  Middle  American  species  of  Schizogenius 
having  obsolete  paramedian  pronotal  sulci.  Specimens  of  S.  optimus  are  distinguished  from 
others  in  the  optimus  group  by  the  following  character  combination:  elytra  piceous,  metal- 
lic, not  extensively  microsculptured,  not  maculate;  antennal  pedicel  unisetose;  clypeal  cari- 
nae arcuate,  fused  apically  and  joined  by  common  stem  to  median  tooth;  elytral  interval 
three  asetose;  and  labrum  not  deeply  emarginate,  margined  laterally  with  no  more  than 
seven  pairs  of  setae. 


174 


Whitehead 


Description.  — Body  stout,  subcylindrical.  Color  piceous,  elytra  and  pronotum  strongly 
metallic,  legs,  mouthparts,  and  antennae  testaceous. 

Integument.  Fine  microsculpture  on  gena,  mouthparts,  pronotal  base,  front  tibiae  and 
anterior  surfaces  of  front  femora,  middle  legs  except  trochanters,  hind  tibiae  and  posterior 
surfaces  of  hind  femora,  sternum  two,  coxal  depressions  of  sternum  three,  sides  of  sterna 
four  to  six,  and  margin  of  sternum  seven.  Sternum  three  without  patches  of  particularly 
coarse  microsculpture  in  coxal  depressions. 

Head.  Fig.  78.  Labrum  strongly  emarginate,  fringed  laterally  with  six  or  seven  pairs  of 
setae.  Paramedian  clypeal  carinae  arcuate,  fused  apically  and  extended  by  common  stem  to 
median  tooth;  median  clypeal  field  hemicircular,  width  at  base  over  1.5  apical  width  of 
median  field  of  frons;  median  clypeal  tooth  shorter  than  paramedian  teeth.  Clypeal  suture 
obsolete  to  weakly  engraved.  Frontal  carinae  not  arcuate,  median  frontal  sulcus  narrowed 
from  base  to  apex.  Antennal  articles  five  to  ten  moniliform,  pedicel  unisetose,  articles  three 
and  four  pubescent.  Eyes  subglobose,  uniformly  faceted.  Terminal  article  of  maxillary 
palpus  with  basal  sensory  bristles  longitudinally  arranged.  Mentum  deeply  emarginate,  me- 
dian tooth  sharp. 

Pronotum.  Fig.  78.  Sides  bisetose,  hind  angles  reduced;  base  not  rugose;  paramedian  sulci 
absent  or  barely  evident  at  base;  anterior  transverse  impression  finely  punctate. 

Legs.  Front  and  middle  tarsi  slender  in  both  sexes,  without  dense  ventral  pubescence; 
hind  tarsus  slender,  elongate.  Front  tibia  narrowed  evenly  to  base  where  much  narrower 
than  at  base  of  antennal  cleaner;  posterior  ventral  margin  of  front  tibia  with  three  setae  near 
base.  Front  femur  not  strongly  constricted  near  apex.  Paronychia  short,  about  half  length 
of  tarsal  claws. 

Elytra.  Disc  without  setigerous  punctures,  one  seta  at  base  of  interval  three.  Striae  deep 
and  sharply  engraved,  finely  punctate  in  basal  two-thirds.  Intervals  one  to  eight  broad,  con- 
vex; intervals  five,  seven,  and  eight  carinate  apically;  interval  seven  joined  by  apex  of  inter- 
val five,  otherwise  free. 

Abdomen.  Paramedian  carinae  of  sternum  three  straight.  Sternum  seven  without  para- 
median ambulatory  setae  in  either  sex.  Pygidium  with  apical  margin  entire  in  both  sexes. 

Male  genitalia.  Median  lobe  (Fig.  82)  with  form  of  apex  characteristic;  endophallus  (Fig. 
85)  with  enlarged  scales  on  dorsal  side  of  virga.  Four  specimens  examined. 

Measurements  and  proportions.  See  Table  4. 

Variation.  — I found  two  males  with  one  paramedian  ambulatory  seta  on  sternum  seven. 

Etymology.  — Latin,  optimus  = best,  a reference  to  the  large  size  of  individuals  of  this 
species. 

Distribution.  — The  known  range  of  S.  optimus  extends  from  southern  Chiapas  to  Costa 
Rica  (Fig.  86).  I studied  46  specimens  from  the  following  localities. 

MEXICO 

CHIAPAS:  Pijijiapan  (31;  CAS,  CNC,  DRWh,  IRSB,  UASM). 

GUATEMALA 

GUATEMALA:  Agua  Caliente  (2;  MCZ).  SAN  MARCOS:  Rio  Naranjo  (2;  BMNH). 


HONDURAS 

COMAYAGUA:  Rancho  Chiquito  (2;  FDAG).  CORTES:  La  Lima  (1;  FDAG).  EL  PARAISO:  El  Paraiso  (1;  UCD). 

COSTA  RICA 

No  locality  (1;USNM).  LIMON:  Rio  Reventazon  (2;  BMNH,  USNM). 

PUNTARENAS:  3-7  mi.  ne.  Puerto  Viejo  (4;  UAFA). 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


175 


Table  4.  Descriptive  statistics  for  S.  optimus,  based  on  31  unsexed  specimens  from  Piji- 
jiapan,  Chiapas. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measurem 

ents, 

in  mm. 

TL 

4.98-6.17 

5.634 

0.390 

0.093 

4.61 

LE 

3.00-3.70 

3.387 

0.238 

0.057 

4.69 

WH 

1.12-1.42 

1.258 

0.095 

0.023 

5.03 

WP 

1.43-1.88 

1.668 

0.128 

0.031 

5.11 

WE 

1.58-2.06 

1.803 

0.146 

0.035 

5.41 

B.  Proportion 

s 

WF/WH 

0.66-0.69 

0.675 

0.012 

0.003 

1.20 

LP/WP 

0.86-0.92 

0.892 

0.022 

0.005 

1.66 

DP/LP 

0.85-0.91 

0.876 

0.021 

0.005 

1.62 

LP/WE 

0.78-0.87 

0.826 

0.030 

0.007 

2.41 

Ta/Ti 

0.80-0.92 

0.849 

0.053 

0.013 

4.16 

Collecting  notes.  - I found  specimens  of  this  species  at  Pijijiapan,  Chiapas,  in  gravel 
along  k river.  They  were  concentrated  near  the  edge  of  the  water  in  a strip  only  a few  feet 
in  length,  for  no  reason  apparent  to  me;  specimens  of  several  other  Schizogenius  species 
were  found  along  the  same  stream  but  were  not  so  concentrated.  Adult  individuals  of  S. 
optimus  are  probably  active  throughout  the  year,  and  I have  seen  specimens  collected  in 
March,  May,  June,  August,  and  October. 

Schizogenius  dyschirioides  Putzeys 

Schizogenius  dyschirioides  Putzeys  1863:28.  Type  locality  “Amazone,”  Brazil;  type  speci- 
mens not  seen,  probably  in  the  Chevrolat  Collection  of  the  Hope  Museum  at  Oxford,  or 

in  the  Bates  Collection  in  MNHP.  Putzeys  1866:222. 

Diagnostic  combination.  — Specimens  of  the  optimus  group  with  combination  of  metallic 
elytra,  bisetose  second  antennal  articles,  bisetose  elytral  interval  three,  and  obsolete  parame- 
dian pronotal  sulci  probably  all  represent  this  species. 

Description.  - Body  form,  color,  and  integument  generally  as  described  for  S.  optimus, 
except  sides  of  sterna  four  to  six  more  strongly  and  extensively  microsculptured. 

Head.  Fig.  79.  As  described  for  S.  optimus  except  clypeal  carinae  straight,  neither  fused 
apically  nor  extended  to  median  tooth;  and  antennal  pedicel  bisetose. 

Pronotum.  Fig.  79;  generally  as  described  for  S',  optimus. 

Legs.  As  in  S.  optimus  except  paronychia  shorter,  much  less  than  half  length  of  tarsal 
claws. 

Elytra.  As  described  for  S.  optimus  except  interval  three  bisetose,  and  intervals  three, 
five,  and  seven  joined  at  apices. 

Abdomen.  As  in  S.  optimus  except  males  with  paramedian  ambulatory  setae  on  sternum 
seven. 

Male  genitalia.  Median  lobe,  Fig.  83;  endophallus  not  studied  in  detail,  without  conspic- 
uously enlarged  spines.  Two  specimens  studied. 


176 


Whitehead 


Measurements  and  proportions.  Based  on  five  specimens  from  Brazil,  Colombia,  and 
Panama.  TL,  4.66-5.10-5.33  mm;  LE,  2.85-3.10-3.23  mm;  WH,  0.98-1.09-1.15  mm;  WP, 
1.32-1.48-1.56  mm;  WE,  1.54-1.66-1.72  mm;  WF/WH,  0.63-0.65-0.67;  LP/WP,  0.87-0.90- 
0.92;  DP/LP,  0.85-0.86-0.86;  LP/WE,  0.79-0.80-0.82;  Ta/Ti,  0.82-0.88-0.91 . 

Variation.  — The  Para  specimen,  a female,  is  much  smaller  than  specimens  from  Colombia 
and  Panama,  and  has  an  extra  seta  on  interval  three  of  left  elytron. 

Etymology.  — Latin,  dyschirioides  = Dyschirius- like,  a reference  to  the  quite  oval  elytra. 

Distribution.  — I refer  to  this  species  16  specimens  from  the  following  South  American 
localities. 


BRAZIL 

PARA:  Belem  (1;USNM). 

COLOMBIA 

MAGDALENA:  Rio  Frio  (13;  MCZ),  Sevilla  (1;  MCZ). 

PANAMA 

CANAL  ZONE:  San  Pablo  (1;  USNM). 

Collecting  notes.  — Specimens  of  S.  dyschirioides  were  collected  in  March  and  May. 
Taxonomic  notes.  — Though  I have  not  seen  type  specimens  of  S.  dyschirioides,  there  is 
little  doubt  that  this  association  is  correct.  The  specimens  well  match  the  original  descrip- 
tion, and  the  Para  specimen  may  even  by  topotypic. 

I suspect  that  presence  of  paramedian  ambulatory  setae  on  sternum  seven  in  males  is 
secondary;  these  setae  are  present  in  some  specimens  of  S.  optimus.  The  extra  seta  on  the 
antennal  pedicel  clearly  is  a novelty.  Other  than  these  features,  form  of  clypeal  carinae,  and 
presence  of  setae  on  elytral  interval  three,  S.  dyschirioides  is  not  greatly  different  from  S. 
optimus. 


Schizogenius  clivinoides  Putzeys 

Schizogenius  clivinoides  Putzeys  1866:229.  Type  locality  “Pampas,”  Argentina;  type  not 
seen,  possibly  in  the  Chaudoir  collection  in  MNHP. 

Diagnostic  combination.  - From  the  description,  specimens  of  this  species  should  have 
the  following  combination  of  characteristics:  paramedian  pronotal  sulci  obsolete;  color 
yellowish  or  brownish,  unmetallic;  elytral  interval  three  bisetose;  elytral  striae  punctate  to 
apex;  and  paramedian  frontal  carinae  arcuate,  apices  fused  and  extended  to  median  tooth. 

Taxonomic  notes.  — I have  seen  no  specimens  of  this  species.  Specimens  of  it  should  be 
readily  identified  from  my  diagnosis,  if  the  original  description  is  accurate. 

Schizogenius  grossus  Whitehead 

Schizogenius  grossus  Whitehead  1966:3.  Type  locality  Rio  Madeira,  Brazil;  holotypemale  in 
USNM  (!). 

Diagnostic  combination.  — Specimens  of  this  species  are  the  only  members  of  the  optimus 
group  with  maculate  elytra.  They  are  the  only  members  of  the  genus  known  to  have  four 
rather  than  three  postero-ventral  setae  near  the  base  of  the  front  tibia,  and  the  only  ones  to 
have  more  than  ten  pairs  of  marginal  setae  on  the  labrum.  Other  characteristics  are:  color 
castaneous,  unmetallic;  integument  extensively  microsculptured;  labrum  weakly  biemargin- 
ate;  paramedian  clypeal  carinae  straight,  attenuate;  antennal  articles  three  and  four  pluri- 
setose;  paramedian  pronotal  sulci  present;  paronychia  much  less  than  half  length  of  tarsal 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


177 


claws;  elytral  disc  asetose,  elytral  striae  shallow;  and  sternum  seven  with  paramedian  ambu- 
latory setae  in  neither  sex. 

Description.  — The  original  description  and  illustrations  (Whitehead,  1966a)  are  adequate. 
Etymology.  — Latin,  grossus  = gross,  a reference  to  the  large  size  of  these  insects. 
Distribution.  — In  addition  to  the  type  series  from  Bolivia  and  Brazil,  I studied  the 
following  two  specimens. 


ARGENTINA 

SALTA:  Aguas  Blancas  (1;  MZSP),  San  Martin  (1;  MZSP). 


Schizogenius  bicolor  new  species 

Type  material.  - Holotype  male  and  allotype  female  labelled  “Jacare  P.  N.  Xingu  M. 
Grosso  — Bras.  XI.  1961  leg.  M.  Alvarenga”  (MGFT).  An  additional  40  specimens  from  two 
localities  in  the  Brazilian  state  of  Mato  Grosso  are  paratypes  (CAS,  DRWh,  IRSB,  MCZ, 
MGFT,  MZSP,  UASM). 

Diagnostic  combination.  - Specimens  of  this  species  have  in  combination:  characters  of 
optimus  group;  no  setae  on  elytral  disc;  plurisetose  antennal  articles  three  and  four;  elytra 
paler  than  pronotum,  not  metallic,  not  maculate;  and  pronotum  with  strong  paramedian 
sulci. 

Description.  - Body  stout,  subcylindrical.  Color  castaneous,  elytra  and  appendages  rufo- 
testaceous,  unmetallic,  elytra  not  maculate. 

Integument.  As  described  for  S.  optimus , except  elytra  faintly  microsculptured. 

Head.  Fig.  80.  As  in  S.  optimus  except:  paramedian  carinae  straight,  attenuate;  details  of 
paramedian  frontal  sulci  and  carinae  differ;  antennal  articles  three  and  four  plurisetose. 

Pronotum.  Fig.  80.  Side  margins  indented  at  anterior  marginal  setae  as  in  S.  grossus.  Para- 
median longitudinal  sulci  well  developed.  Sides  bisetose,  hind  angles  reduced;  base  not 
rugose;  anterior  transverse  impression  with  longitudinal  rugae. 

Legs.  As  in  S.  optimus  except  paronychia  much  less  than  half  length  of  tarsal  claws. 

Elytra.  As  in  S.  optimus  except  more  parallel  sided;  intervals  three,  five,  and  seven  not 
carinate,  joined  at  apices;  and  striae  finely  punctate  in  basal  half. 

Abdomen.  As  described  for  S.  optimus. 

Male  genitalia.  Median  lobe,  Fig.  84;  endophallus  not  studied  in  detail.  Two  specimens 
studied. 

Measurements  and  proportions.  Of  holotype:  TL,  5.19  mm;  LE,  3.19  mm;  WH,  1.10  mm; 
WP,  1.50  mm;  WE,  1.64  mm;  WF/WH,  0.66;  LP/WP,  0.93;  DP/LP,  0.83;  LP/WE,  0.85;  Ta / 
Ti,  0.88;  PS/LP,  0.60.  Of  allotype:  TL,  5.48  mm;  LE,  3.40  mm;  WH,  1.17  mm;  WP,  1.61 
mm;  WE,  1.72  mm;  WF/WH,  0.66;  LP/WP,  0.91;  DP/LP,  0.87;  LP/WE,  0.85;  Ta/Ti,  0.91; 
PS/LP,  0.65. 

Variation.  — Six  specimens  from  Piaui  state  are  darker  and  less  distinctly  bicolored  than 
specimens  in  the  type  series. 

Etymology.  — Latin,  bi  = two,  plus  color  = color,  in  reference  to  the  distinctive  coloration 
of  individuals  of  this  species. 

Distribution.  — Specimens  of  this  species  have  been  taken  in  the  Brazilian  states  of  Mato 
Grosso  and  Piaui.  I studied  48  specimens  from  the  following  localities. 

BRAZIL 

MATO  GROSSO:  Barra  de  Tapirape  (2;  MZSP),  Jacare  P.  N.  Xingu  (40;  CAS,  DRWh,  IRSB,  MCZ,  MGFT,  UASM).  PIAUI: 
Terezina  (6;  MGFT). 


178 


Whitehead 


Fig.  77-80.  Head  and  pronotum,  dorsal  aspect.  77.  S.  jacarensis  new  species,  Jacaxe,  Brazil.  78.  S.  optimus  Bates, 
Pijijiapan,  Chiapas.  79.  S.  dyschirioides  Putzeys,  Sevilla,  Colombia.  80.  S.  bicolor  new  species,  Jacare,  Brazil.  Fig.  81-84. 
Male  median  lobe,  lateral  aspect.  81.  S.  jacarensis  new  species,  Jacare,  Brazil.  82.  S.  optimus  Bates,  Pijijiapan,  Chiapas. 
83.  S.  dyschirioides  Putzeys,  Rio  Frio,  Colombia.  84.  S.  bicolor  new  species,  Jacare,  Brazil.  Fig.  85.  Male  endophallus, 
S.  optimus  Bates,  Pijijiapan,  Chiapas.  Fig.  86.  Known  distribution  of  S.  optimus  Bates. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


179 


Collecting  notes.  - Specimens  of  this  species  were  collected  in  November  and  January. 

Taxonomic  notes.  — This  species  is  most  closely  related  to  S.  grossus  Whitehead,  but  is 
less  specialized.  Specimens  are  less  extensively  microsculptured,  have  more  sharply  engraved 
elytral  striae,  have  a deeply  emarginate  mentum,  have  just  three  postero-ventral  setae  on  the 
front  tibia,  and  have  only  six  or  seven  pairs  of  marginal  setae  on  the  labrum.  But  they  are 
similar  in  habitus,  distribution  of  discal  setae,  and  non-pubescent  third  and  fourth  antennal 
articles. 


The  ferrugineus  group 

Diagnostic  combination.  — Within  the  subgenus,  members  of  this  group  are  characterized 
by  the  following  combination  of  characters:  paramedian  clypeal  carinae  tuberculate,  apices 
nearly  or  quite  obsolete;  clypeal  field  triangular,  its  base  1.0  to  more  than  1.5  times  apical 
width  of  median  frontal  sulcus;  clypeal  suture  sharply  impressed;  antennal  articles  five  to  ten 
moniliform;  anterior  tarsi  slender  in  both  sexes;  elytral  disc  with  setae  on  intervals  three, 
five,  and  seven;  paramedian  carinae  of  sternum  three  straight;  paramedian  ambulatory  setae 
present  in  males  only;  pygidium  not  crenulate  in  either  sex;  and  endophallus  without  en- 
larged basal  collar  spines. 

This  group  contains  the  only  North  and  Middle  American  Schizogenius  species  whose 
members  each  have  tuberculate  clypeal  carinae  in  combination  with  discal  setae  present  on 
intervals  three,  five,  and  seven. 

Distribution.  — One  species  ranges  throughout  much  of  eastern  North  America,  south- 
westward  to  Arizona.  A second  species  overlaps  the  first  in  Arizona,  and  ranges  southward  in 
Pacific  drainage  systems  to  at  least  Costa  Rica.  I examined  631  specimens  of  the  ferrugineus 
group. 

Taxonomic  notes.  — The  two  species  included  in  this  group  are  similar  in  habitus  but  are 
not  closely  related.  They  may  even  not  form  a monophyletic  group,  as  I have  found  no 
assuredly  synapomorphic  characteristics  except  perhaps  loss  of  paramedian  anal  setae  from 
sternum  three  of  females,  but  I have  no  reason  to  treat  them  otherwise. 

Schizogenius  ferrugineus  Putzeys 

Schizogenius  ferrugineus  Putzeys  1846:653.  Type  locality  Galveston,  Texas;  location  of 

type  unknown.  LeConte  1857:82.  Putzeys  1863:24.  Putzeys  1866:223.  LeConte  1879: 

34.  Lindroth  1961:168. 

Clivina  sulcata  LeConte  1848:214.  Type  area  New  York  State;  type  in  LeConte  Collection, 

MCZ. 

Schizogenius  sulcatus,  LeConte  1857:83,  suggested  synonymy. 

Diagnostic  combination.  — Specimens  of  this  species  are  readily  distinguished  from  those 
of  S.  auripennis,  the  only  other  known  species  of  the  group,  by  ferrugineous  coloration  and 
extensively  microsculptured  abdomen. 

Head.  Fig.  87.  Clypeal  carinae  tuberculate,  not  or  weakly  extended  to  median  clypeal 
tooth;  median  clypeal  field  triangular,  broad,  width  at  base  about  1 .5  apical  width  of  me- 
dian field  of  frons.  Clypeal  suture  sharply  engraved.  Eye  prominent,  subglobose,  uniformly 
faceted.  Neck  densely  but  finely  punctate.  Gena  rugose  in  front.  Antennal  articles  four  to 
ten  moniliform. 

Pronotum.  Fig.  87.  Sides  bisetose,  hind  angles  reduced,  base  not  rugose.  Paramedian  sulci 
long,  impunctate,  shallow  and  indistinct  apically,  deep  and  strongly  hooked  basally.  Anterior 
transverse  impression  impunctate  or  finely  punctate. 


180 


Whitehead 


Legs.  Front  and  middle  tarsi  slender,  without  dense  ventral  pubescence;  hind  tarsus  slen- 
der, short.  Paronychia  inconspicuous,  about  half  length  of  tarsal  claws.  Front  tibia  evenly 
narrowed  to  base  where  much  narrower  than  at  base  of  antennal  cleaner.  Front  femur  not 
strongly  constricted  near  apex. 

Elytra.  Five  to  seven  setigerous  punctures  each  on  intervals  three  and  Five,  and  three  to 
five  on  interval  seven.  Striae  deep  and  sharply  engraved,  finely  punctate  in  basal  two  thirds. 
Intervals  one  to  seven  broad  and  convex,  interval  eight  carinate  at  apex;  intervals  three  and 
five  broadly  joined  apically  with  interval  seven. 

Abdomen.  Sternum  three  with  paramedian  carinae  straight  at  apices.  Sternum  seven  with 
paramedian  ambulatory  setae  in  male  only.  Pygidium  with  apical  margin  entire  in  both 
sexes. 

Male  genitalia.  Median  lobe,  Fig.  93;  endophallus,  Fig.  98,  without  enlarged  basal  collar 
spines.  Six  specimens  examined. 

Measurements  and  proportions.  See  Table  5. 


Table  5.  Descriptive  statistics  for  S.  ferrugineus,  based  on  20  males  from  Round  Mountain, 
Texas. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

nts,  in  mm. 

TL 

3.13-3.76 

3.434 

0.297 

0.089 

5.77 

LE 

1.89-2.27 

2.072 

0.180 

0.054 

5.80 

WH 

0.66-0.82 

0.738 

0.067 

0.020 

6.07 

WP 

0.83-1.05 

0.944 

0.099 

0.030 

7.03 

WE 

0.91-1.14 

1.006 

0.102 

0.031 

6.79 

B.  Setae  on  lei 

't  elytron. 

Interval  3 

5-  7 

6.2 

Interval  5 

5-  7 

6.3 

Interval  7 

3-  4 

3.5 

Total 

14-18 

16.0 

1.6 

0.5 

6.72 

C.  Proportions 

WF/WH 

0.66-0.72 

0.688 

0.023 

0.007 

2.27 

LP/WP 

0.95-1.01 

0.982 

0.033 

0.010 

2.22 

DP/LP 

0.80-0.86 

0.824 

0.023 

0.007 

1.86 

LP/WE 

0.88-0.96 

0.918 

0.036 

0.011 

2.61 

Ta/Ti 

0.64-0.75 

0.708 

0.052 

0.015 

4.86 

PS/LP 

0.63-0.76 

0.701 

0.050 

0.015 

4.75 

Variation.  - Among  specimens  studied,  the  largest  female  (LE,  2.48  mm)  and  male  (LE, 
2.41  mm)  are  from  Long  Beach,  New  York,  the  smallest  female  (LE,  1.66  mm)  is  from 
Columbus,  Texas,  and  the  smallest  male  (LE,  1.69  mm)  is  from  Clark  Co.,  Kansas.  There  is 
an  apparent  slight  tendency  for  larger  size  toward  the  northeast  than  toward  the  southwest, 


Classification,  phytogeny,  and  zoogeography  of  Schizogenius 


181 


but  population  samples  generally  are  too  small  for  worthwhile  comparisons. 

Etymology.  - Latin,  ferrugineus  = rust  colored,  in  reference  to  the  body  color;  Latin, 
sulcatus  = sulcate,  in  reference  to  the  sulci  on  the  frons  and  pronotum. 

Distribution.  - S.  ferrugineus  ranges  at  tow  elevations  in  sandy  places  from  southern 
Ontario  south  to  northern  Florida  and  west  to  southeastern  Arizona  (Fig.  100).  I examined 
507  specimens  from  the  following  localities. 

CANADA 

No  locality  (1;  UKSM).  ONTARIO  (1;CAS):  Tilbury  (1;  KHSt);  Toronto  (5;  CAS,  CUNY,  KSUM,  MCZ). 

UNITED  STATES 

No  locality  (3;  ANSP,  IRSB).  ALABAMA:  Washington  Co.,  Leroy  (2;  CUNY).  ARIZONA:  Cochise  Co.,  Douglas  (1;UCB); 
Graham  Co.,  Aravaipa  (1;  CAS);  Pima  Co.,  Robles  Ranch  (1;  UCD).  ARKANSAS:  Carroll  Co.,  Eureka  Springs  (1;  UKSM); 
Hempstead  Co.,  Hope  (3;  CUNY,  USNM);  Washington  Co.,  Mount  Sequoyah  (1;  INHS).  COLORADO:  Logan  Co.  (1; 
USNM).  DELAWARE  (1;  MCZ).  FLORIDA:  Nassau  Co.,  Fernandina  Beach  (2;  USNM).  GEORGIA  (2;  ANSP).  ILLINOIS: 
Mason  Co.,  Havana  (1;  USNM);  Pike  Co.,  Pittsfield  (2;  UCD).  INDIANA  (1;  CNHM):  Lake  Co.,  Pine  (6;  CNHM,  MGFT): 
Steuben  Co.,  Fremont  (1;  CNHM);  Tippecanoe  Co.  (1;  UCD).  IOWA:  Johnson  Co.,  Iowa  City  (4;  USNM);  Des  Moines  Co., 
Burlington  (2;  MCZ).  KANSAS  (14;  CNHM,  CUNY,  KSUM,  MCZ,  USNM):  Clark  Co.  (1;  UKSM);  Douglas  Co.  (1;  UKSM), 
Lawrence  (4;  INHS);  Kiowa  Co.  (1;  UKSM);  Leavenworth  Co.  (3;  USNM);  Pottawatomie  Co.,  Onaga  (1;  UASM);  Reno  Co. 
(8;  CAS,  MCZ,  USNM),  Medora  (2;  KSUM);  Riley  Co.  (7;  KSUM,  USNM),  Manhattan  (1;  KSUM);  Sedgwick  Co.  (6;  USNM); 
Mount  Hope  (9;  CAS,  USNM).  KENTUCKY  (1;  AMNH).  LOUISIANA:  Winn  Co.,  Winnfield  (1;  MCZ).  MAINE:  Oxford 
Co.,  Paris  (1;  MCZ).  MARYLAND:  Difficult  (4;  CUNY,  MCZ);  Calvert  Co.,  Chesapeake  Beach  (19;  USNM),  Kenwood 
Beach  (14;  DRWh,  JNeg,  UCD,  USNM);  Montgomery  Co.,  Great  Falls  (1;  CAS),  Plummers  Island  (2;  USNM).  MASSA- 
CHUSETTS: Essex  Co.,  Ipswich  (4;  MCZ);  Hampden  Co.,  Chicopee  (3;  MCZ),  Springfield  (1;  MCZ);  Middlesex  Co., 
Tewksbury  (1;  MCZ),  Tyngsboro  (13;  MCZ).  MICHIGAN:  Huron  Co.  (1;  MSUL);  Monroe  Co.,  Monroe  (10;  MCZ,  MSUL, 
USNM).  MINNESOTA:  Houston  Co.,  Mississippi  Bluff  (1;  MSUL).  MISSISSIPPI:  George  Co.,  Lucedale  (2;  CUNY). 
MISSOURI  (1;  ANSP).  NEBRASKA:  Cuming  Co.,  West  Point  (2;  USNM);  Thomas  Co.,  2.5  mi.  w.  Halsey  (1;  CAS). 
NEW  HAMPSHIRE:  Grafton  Co.,  Rumney  (10;  CNC,  DRWh,  MCZ).  NEW  JERSEY  (2;  ANSP,  MCZ):  Atlantic  Co., 
Atlantic  City  (1;  MCZ),  Brigantine  (3;  CAS,  RUNB);  Burlington  Co.,  Atsion  (8;  CAS);  Camden  Co.,  Cramer  Hill  (1; 
USNM),  Westville  (10;  ANSP,  MCZ,  RUNB,  USNM);  Cape  May  Co.,  Anglesea  (18;  ANSP,  LACM,  MCZ,  RUNB),  Five 
Mile  Beach  (4;  USNM),  Ocean  City  (1;  CAS);  Gloucester  Co.,  Woodbury  (1;  ANSP);  Warren  Co.,  Phillipsburg  (1;  CAS). 
NEW  MEXICO:  Curry  Co.,  Clovis  (2;  UKL).  NEW  YORK  (8;  CAS,  USNM):  New  York  City  and  vicinity  (13;  AMNH, 
CAS,  RUNB);  Long  Island  (4;  CAS,  UNSM);  Brooklyn  Co.,  Coney  Island  (7;  CAS,  MCZ);  Nassau  Co.,  Long  Beach 
(17;  AMNH,  CAS,  USNM);  Queens  Co.,  Rockaway  Beach  (10;  CAS,  CUNY,  USNM);  Suffolk  Co.,  Plum  Island  (1;  MCZ), 
Riverhead  (2;  CUNY),  Sound  Beach  (1;  AMNH);  Tompkins  Co.,  Groton  (2;  CAS).  NORTH  CAROLINA  (1;  MSUL). 
OHIO:  Athens  Co.,  Athens  (2;  UWLW),  Hamilton  Co.,  Cincinatti  (1 ; MCZ).  OKLAHOMA:  Caddo  Co.,  Hinton  (6;  MSUL); 
Love  Co.,  Oswalt  (2;  MSUL);  McCurtain  Co.,  Sherwood  (1;  MSUL);  Woodward  Co.,  Woodward  (3;  USNM).  PENNSYL- 
VANIA (1;  USNM).  SOUTH  CAROLINA  (2;  CAS,  MCZ):  Charleston  Co.,  Charleston  (1;  CAS),  Folly  Beach  (1;  USNM), 
Isle  of  Palms  (5;  ANSP,  RUNB,  USNM).  TEXAS  (65;  AMNH,  ANSP,  CAS,  CNHM,  INHS,  IRSB,  KSUM,  MCZ,  MSUL, 
UKSM,  USNM):  Bexar  Co.,  San  Antonio  (2;  CAS);  Blanco  Co.,  Cypress  Mills  (6;  CAS,  USNM),  Rome  Mountain  (16; 
CAS),  Round  Mountain  (25;  CAS,  MCZ,  RUNB);  Brown  Co.,  Brownwood  (1;  AMNH);  Burnet  Co.  (2;  USNM);  Comal  Co. 
(1;  CAS);  Colorado  Co.,  Columbus  (21;  USNM);  Harris  Co.,  Hockley  (2;  AMNH,  CAS);  Lee  Co.  (2;  MCZ),  Fedor 
(6;  CAS),  Lexington  (1;  MCZ);  Llano  Co.,  Enchanted  Rock  (1;  CNC);  McCulloch  Co.,  16  mi  s.  Brady  (1;  CAS);  Travis 
Co.,  Austin  (1;  USNM);  Wharton  Co.,  Wharton  (2;  CUNY).  VIRGINIA:  Elisabeth  City,  Fort  Monroe  (1;  USNM);  Fairfax 
Co.,  Arlington  (1;  USNM);  Nelson  Co.  (1;  USNM);  Princess  Anne  Co.,  Cape  Henry  (5;  AMNH),  Virginia  Beach  (2; 
USNM).  WISCONSIN:  Grant  Co.,  Boscobel  (1;  FDAG). 


Collecting  notes.  — S.  ferrugineus  differs  from  most  other  species  of  the  subgenus  Schizo- 
genius in  that  individuals  live  in  sand  rather  than  in  gravel  bars.  Specimens  from  Iowa 
City,  Iowa,  are  labelled  “Butler’s  Landing  sand  area.”  Specimens  from  Kenwood  Beach, 
Maryland,  were  found  “under  wash-up.”  P.  J.  Darlington  collected  specimens  “under  tog 
on  beach”  at  Ipswich,  Massachusetts,  and  others  in  “galleries  in  dry  sand  under  togs,  Baker 
River”  at  Rumney,  New  Hampshire.  Additional  specimens  were  collected  in  “drift”  along 
the  Merrimack  River  at  Tyngsboro,  Massachusetts. 

Specimens  of  this  species  have  been  collected  from  13  March  at  Charleston,  South  Caro- 
lina, to  16  October  at  Lawrence,  Kansas.  Many  flew  in  to  lights  at  various  localities.  I can 
add  no  additional  observations,  having  collected  no  specimens  of  this  species. 


182 


Whitehead 


Schizogenius  auripennis  Bates 

Schizogenius  auripennis  Bates  1881:38.  Type  locality  Teleman,  Guatemala;  type  in  BMNH, 

specimen  labelled  “holotype”  here  designated  as  lectotype  (!). 

Schizogenius  peninsularis  Van  Dyke  1949:50.  Type  locality  5 mi.  s.  Miraflores,  Baja  Cali- 
fornia; type  in  CAS,  not  studied.  NEW  SYNONYMY. 

Diagnostic  combination.  — The  shiny,  unmicrosculptured  abdomen  separates  specimens 
of  S.  auripennis  from  those  of  S.  ferrugineus,  the  only  other  known  member  of  the  group. 
The  apex  of  the  male  median  lobe  is  strikingly  different  in  form  from  that  of  any  other 
species  in  the  genus. 

Description.  — Body  cylindrical.  Color  dark  rufopiceous  above  and  dark  rufous  below, 
elytral  apices,  legs,  palpi,  and  antennae  testaceous  or  rufotestaceous;  dorsal  surface  with 
slight  to  strong  aeneous  luster. 

Integument.  Smooth,  shiny.  Fine  microsculpture  on  paramedian  frontal  sulci,  genae, 
mouthparts,  pronotal  base,  front  tibiae  and  anterior  surfaces  of  front  femora,  middle  legs 
except  trochanters,  hind  tibiae  and  posterior  surfaces  of  hind  femora,  elytral  epipleura  at 
extreme  base,  and  at  least  small  patch  in  coxal  depression  of  sternum  three;  some  specimens 
with  microsculpture  on  sternum  two,  sides  of  sterna  three  and  four,  and  margin  of  sternum 
seven. 

Head.  Fig.  88.  As  described  for  S.  ferrugineus  except:  median  field  of  clypeus  narrower 
at  base,  between  1.0  and  1.5  apical  width  of  median  field  of  frons;  second  frontal  carina 
broadened  at  base,  fifth  nearly  obsolete;  eye  larger,  globose;  neck  densely  and  coarsely 
punctate;  and  gena  strongly  rugose  in  front. 

Pronotum.  Fig.  88.  Smaller,  more  transverse  than  in  S.  ferrugineus , paramedian  sulci 
shorter  and  slightly  hooked  basally,  anterior  transverse  impression  finely  punctate. 

Legs.  Paronychia  more  conspicuous,  slightly  shorter  than  tarsal  claws. 

Elytra.  Five  to  eight  setigerous  punctures  on  interval  three,  five  to  seven  on  interval  five, 
and  four  to  seven  on  interval  seven.  Otherwise  as  in  S.  ferrugineus. 

Abdomen.  As  in  S.  ferrugineus. 

Male  genitalia.  Median  lobe  (Fig.  94)  with  form  of  apex  diagnostic;  endophallus,  Fig.  99. 
Eleven  specimens  examined. 

Measurements  and  proportions.  See  Table  6. 

Variation.  — Since  S.  auripennis  has  no  known  close  relatives,  and  as  available  material 
was  limited,  I did  not  study  geographic  variation  in  detail.  Specimens  from  the  south  tend  to 
have  more  abdominal  microsculpture,  more  elytral  setae,  more  elongate  paramedian  prono- 
tal sulci,  and  more  cylindrical  body  form,  but  this  variation  seems  gradual.  The  smallest 
specimens  seen  are  males  and  females  from  Arizona  and  Baja  California  (LE,  2.07  mm)  and 
the  largest  is  a female  from  Tucson,  Arizona  (LE,  2.88  mm). 

Etymology.  — Of  auripennis,  Latin,  aurum  = gold  plus  penna  = wing,  in  reference  to 
elytral  coloration  of  the  slightly  teneral  type;  of  peninsularis,  Latin,  peninsula  = peninsula, 
a reference  to  the  type  locality  in  peninsular  Baja  California. 

Distribution.  — The  known  distribution  of  this  species  (Fig.  101)  extends  from  southern 
Arizona  south  to  Costa  Rica  in  Pacific  drainage  areas,  from  elevations  near  sea-level  to  as 
high  as  4500'  in  Guatemala.  I studied  124  specimens  from  the  following  localities. 

UNITED  STATES 

ARIZONA:  Cochise  Co.,  Douglas  (1;  CAS);  Graham  Co.,  Aravaipa  (1;  CAS);  Maricopa  Co.,  Phoenix  (2;  CUNY,  DRWh), 
Wickenburg  (5;  MCZ,  MSUL);  Pima  Co.,  Arivaca  (1;  KHSt),  Organ  Pipe  National  Monument  (1;  LBSC),  Quitobaquito 
(1;  UATA),  Tucson  (3;  AMNH,  CAS,  USNM);  Santa  Cruz  Co.,  Nogales  (2;  CAS,  UCD),  Pena  Blanca  (4;  UASM,  UATA). 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


183 


MEXICO 

BAJA  CALIFORNIA:  5 mi.  s.  Miraflores  (17;  CAS),  5 mi.  w.  San  Bartolo  (3;  CAS),  6 mi.  sw.  Santiago  (4;  UATA),  Triunfo 
(3;  CAS).  CHIAPAS:  20.9  mi.  n.  Arriaga  (1;  UASM),  Puente  Macuilapa  (2;  FDAG).  JALISCO:  Pitillal  (1;  UASM). 
NAYARIT:  Jesus  Maria  (1;  UCB),  San  Bias  (1;  CAS).  PUEBLA:  Tepexco  (2;  UASM).  SINALOA:  19  mi.  s.  Culiacan  (2; 
UCB),  Real  de  Piaxtla  (2;  AMNH),  4 mi.  s.  Villa  Union  (1;  UCB),  26  mi.  ne.  Villa  Union  (2;  LBSC).  SONORA:  Alamos  (6; 
CAS,  LACM),  5 mi.  w.  Alamos  (2;  UATA),  10  mi.  w.  Alamos  (4;  AMNH),  7 mi.  se.  Alamos  (5;GRNo,  UCB),  Hermosillo 
(34;  CAS),  Minas  Nuevas  (1;  AMNH),  10  mi.  e.  Navajoa  (1;  UATA). 

GUATEMALA 

ALTA  VERAPAZ:  Teleman  (1;  BMNH).  EL  QUICHE:  Sacapulas  (1;  AMNH). 

HONDURAS 

CORTES:  La  Lima  (3;  FDAG,  INHS). 

COSTA  RICA 

PUNTARENAS:  Palmar  Sur  (1;  UAFA).  SAN  JOSE:  Barranca  (1;  JNeg),  3-7  mi.  n.  Puerto  Viejo  (1;  UAFA). 


Collecting  notes.  - Nearly  all  specimens  were  taken  at  lights,  so  little  is  known  of  the 
biology  of  S.  auripennis.  I collected  one  specimen  from  Arriaga  along  a small  stream,  and 
G.  E.  Ball  collected  one  in  gravel  along  a stream  at  Pitillal.  Probably,  individuals  of  this 
species  normally  live  in  sand  rather  than  gravel  bars,  as  do  those  of  S.  ferrugineus ; this 
supposition  is  supported  by  the  cylindrical  body  form,  which  is  not  suited  for  life  in  gravel 
bars.  If  the  normal  habitat  is  sand,  and  particularly  if  it  is  dry  sand,  then  the  distribution  of 
this  species  is  likely  to  be  unaffected  by  location  and  substrate  of  drainage  systems.  Speci- 
mens have  been  collected  throughout  the  year. 


Table  6.  Descriptive  statistics  for  S',  auripennis,  based  on  20  males  from  Hermosillo,  Sonora. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measuremi 

ents,  in  mm. 

TL 

3.46-4.23 

3.824 

0.329 

0.098 

5.73 

LE 

2.14-2.60 

2.362 

0.194 

0.058 

5.47 

WH 

0.73-0.88 

0.789 

0.071 

0.021 

5.98 

WP 

0.97-1.21 

1.080 

0.108 

0.032 

6.64 

WE 

1.10-1.34 

1.201 

0.105 

0.031 

5.85 

B.  Setae  on  le 

:ft  elytron. 

Interval  3 

6-  8 

6.7 

Interval  5 

5-  7 

6.3 

Interval  7 

4-  7 

5.4 

Total 

16-22 

18.4 

1.6 

0.7 

8.64 

C.  Proportion 

is. 

WF/WH 

0.60-0.66 

0.623 

0.029 

0.009 

3.13 

LP/WP 

0.88-0.95 

0.914 

0.027 

0.008 

0.95 

DP/LP 

0.84-0.88 

0.866 

0.018 

0.005 

1.42 

LP/WE 

0.79-0.86 

0.822 

0.032 

0.010 

2.59 

Ta/Ti 

0.63-0.73 

0.680 

0.053 

0.016 

5.18 

PS/LP 

0.56-0.63 

0.599 

0.035 

0.01 1 

3.94 

184 


Whitehead 


Taxonomic  notes.  — I have  seen  topoparatypes  of  both  S.  auripennis  and  S.  peninsularis. 
These  names  are  clearly  synonyms.  Southern  specimens  differ  slightly  as  noted,  but  male 
genitalia  are  constant  throughout  the  known  range.  Exact  relationships  of  this  species  with 
S.  ferrugineus  are  unclear,  but  are  quite  apparently  not  close. 

The  basalis  group 

Diagnostic  combination.  - Specimens  referred  to  this  group  have  the  following  com- 
bination of  characters:  body  convex;  paramedian  clypeal  carinae  not  extended  to  median 
tooth;  clypeal  field  triangular,  at  base  more  than  1 .5  apical  width  of  median  frontal  sulcus; 
clypeal  suture  sharply  impressed;  antennal  articles  five  to  ten  short  but  filiform,  distinctly 
longer  than  wide;  submentum  without  accessory  setae;  pronotum  without  distinct  para- 
lateral  carinae,  with  well  developed  paramedian  sulci;  front  and  middle  tarsi  quite  narrow 
in  both  sexes;  elytra  with  discal  setae  on  intervals  three,  five,  and  seven,  intervals  not  cari- 
nate; sternum  seven  with  paramedian  ambulatory  setae  in  both  sexes;  paramedian  carinae 
of  sternum  three  not  or  slightly  curved  at  apices;  and  pygidium  apex  weakly  to  strongly 
crenulate  in  females. 

Distribution.  — Members  of  this  group  are  known  only  from  Brazil,  Uruguay,  and  Argen- 
tina. I examined  18  specimens  of  the  basalis  group. 

Taxonomic  notes.  — I assign  to  this  group  four  species,  two  described  here  as  new.  I 
examined  the  type  of  S.  basalis  Putzeys,  and  saw  some  Brazilian  specimens  which  agree 
well  with  the  original  description  of  S.  multipunctatus  Kult,  but  I doubt  that  any  additional 
described  species  belong  to  this  group.  I have  not  provided  full  species  descriptions,  as  none 
of  the  species  are  well  represented  in  collections. 

This  group  seems  an  important  phylogenetic  link,  as  it  seems  to  share  a common  ancestry 
with  the  ferrugineus  group  (which  has  no  clear  derivatives),  and  probably  also  with  all  re- 
maining groups  of  Schizogenius. 

Schizogenius  basalis  Putzeys 

Schizogenius  basalis  Putzeys  1866:230.  Type  locality  Santa  Lucia  River,  12  leagues  north 

of  Montevideo,  Uruguay;  holotype  female  in  IRSB  (!).  Kult  1950:148. 

Diagnostic  combination.  - The  only  specimen  of  this  species  seen  by  me  differs  from 
specimens  of  S.  multipunctatus  by  larger  size  (LE  about  2.5  mm),  less  metallic  color- 
ation, and  sparsely  punctate  occiput.  It  differs  from  specimens  of  S.  cearaensis  by  lar- 
ger size,  more  numerous  discal  setae,  and  less  metallic  pronotal  coloration,  and  from 
specimens  of  S.  negrei  by  larger  size,  more  numerous  discal  setae,  and  darker  colora- 
tion. 

Description.  — With  characters  of  the  basalis  group.  Body  coloration  faded  from  aging, 
but  according  to  original  description  aeneopiceous,  with  elytral  base  fuscous;  legs  and 
antennae  ferrugineous;  palpi  testaceous.  Head  and  pronotum.  Fig.  89;  antennal  article  five 
about  1.3  times  longer  than  wide;  occiput  sparsely  punctate.  Elytral  striae  distinctly  punc- 
tate in  basal  half;  left  elytron  with  16,  17,  and  14  setae  on  intervals  three,  five,  and  seven, 
respectively.  Male  genitalia  not  known. 

Measurements  and  proportions.  TL,  4.01  mm;  LE,  2.50  mm;  WH,  0.79  mm;  WP,  1.16 
mm;  WE,  1.33  mm;  WF/WH,  0.67;  LP/WP,  0.87;  DP/LP,  0.86;  LP/WE,  0.76;  PS/LP,  0.61. 

Etymology.  — Latin,  basilaris  - at  the  base,  in  reference  to  the  pale  elytral  base. 

Distribution.  — I examined  only  the  type  specimen  from  near  Montevideo,  Uruguay. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


185 


Schizogenius  cearaensis  new  species 

Type  material.  - Holotype  male  and  allotype  female  labelled  “Fortaleza  Ceara,  BRAZIL 
III  -29/IV-2-63  F.  G.  Werner”  (MCZ).  Nine  additional  specimens  with  the  same  label  data 
are  paratypes  (DRWh,  IRSB,  MCZ,  UASM). 

Diagnostic  combination.  - Within  the  basalis  group,  specimens  of  this  species  are  distin- 
guished by  the  combination  of:  occiput  sparsely  punctate;  elytra  uniformly  pale;  pronotum 
strongly  metallic;  and  elytral  disc  with  fewer  than  40  setae. 

Description.  — With  general  characters  of  basalis  group.  As  in  S.  basalis  except  as  follows. 
Color  ferrugineous;  pronotum  strongly  aeneopiceous  to  metallic  green;  elytra,  antennae, 
palpi,  front  tibiae  and  tarsi,  and  middle  and  hind  legs  testaceous.  Head  and  pronotum,  Fig. 
90.  Left  elytron  with  10  to  12  setae  on  interval  three,  12  to  14  on  interval  five,  and  9 to  13 
on  interval  seven,  total  33  to  38  in  specimens  examined.  Male  genitalia,  Fig.  95;  one  speci- 
men examined. 

Measurements  and  proportions.  Of  holotype,  the  smallest  specimen:  TL,  3.07  mm;  LE, 
1.88  mm;  WH,  0.65  mm;  WP,  0.87  mm;  WE,  1.07  mm;  WF/WH,  0.65;  LP/WP,  0.90;  DP/LP, 
0.88;  LP/WE,  0.73;  PS/LP,  0.54;  Ta/Ti,  0.65.  Of  allotype,  the  largest  specimen:  TL,  3.37 
mm;  LE,  2.09  mm;  WH,  0.67  mm;  WP,  0.85  mm;  WE,  1.12  mm;  WF/WH,  0.67;  LP/WP, 
0.89;  DP/LP,  0.88;  LP/WE,  0.76;  Ta/Ti,  0.73. 

Etymology.  — I name  this  species  for  its  type  locality,  in  the  Brazilian  state  of  Ceara. 

Distribution.  — S.  cearaensis  is  known  only  from  the  type  series  from  Fortaleza,  Ceara, 
Brazil. 

Taxonomic  notes.  — I distinguish  S.  cearaensis  as  a species  distinct  from  the  related  S. 
basalis  because  all  known  specimens  are  smaller,  have  fewer  elytral  setae,  and  differ  in  color- 
ation. It  remains  possible,  however,  that  the  two  forms  are  merely  geographic  variants.  More 
material  of  the  group  is  required  to  study  geographic  variation,  and  to  obtain  more  informa- 
tion about  S.  basalis  including  information  about  male  genitalia. 

Schizogenius  multipunctatus  Kult 

Schizogenius  multipunctatus  Kult  1950:147.  Type  locality  Corumba,  Mato  Grosso,  Brazil; 

holotype  and  paratype  in  Kult  collection,  present  location  not  known. 

Diagnostic  combination.  - Within  the  basalis  group,  specimens  with  occiput  densely 
punctate  belong  to  this  species.  All  known  specimens  of  this  species  differ  further  from 
the  type  of  S.  basalis  by  smaller  body  size  (LE  under  2.3  mm),  and  from  specimens  of 
S.  cearaensis  and  S.  negrei  by  the  dark  elytra. 

Description.  — With  characters  of  basalis  group.  Body  bright  aeneopiceous  to  metallic 
green,  elytra  unicolorous;  legs  and  antennae  ferrugineous;  palpi  testaceous.  Head  and  prono- 
tum, Fig.  91;  antennal  article  five  about  1.3  times  longer  than  wide;  occiput  densely  punc- 
tate. Elytral  striae  finely  punctate  in  basal  half;  left  elytron  with  about  14-18  setae  on  inter- 
val three,  15-17  on  interval  five,  1 1-13  on  interval  seven;  total  41-46  in  specimens  examined. 
Male  genitalia,  Fig.  96;  one  specimen  examined. 

Measurements  and  proportions.  Largest  specimen,  female:  TL,  3.55  mm;  LE,  2.24  mm; 
WH,  0.73  mm;  WP,  0.98  mm;  WE,  1.15  mm;  WF/WH,  0.62;  LP/WP,  0.87;  DP/LP,  0.91 ; LP/ 
WE,  0.74;  Ta/Ti,  0.66;  PS/LP,  0.59.  Smallest  specimen,  male:  TL,  3.17  mm;  LE,  2.00  mm; 
WH,  0.66  mm;  WP,  0.85  mm;  WE,  1.02  mm;  WF/WH,  0.60;  LP/WP,  0.88;  DP/LP,  0.92;  LP/ 
WE,  0.74;  Ta/Ti,  0.58;  PS/LP,  0.57. 

Etymology.  — Latin,  multus  = much,  plus  punctum  = small  hole,  in  reference  to  the 
numerous  discal  setae  on  the  elytra. 


186 


Whitehead 


Distribution.  - Specimens  of  this  species  have  been  collected  in  various  localities  in  cen- 
tral Brazil.  I studied  three  specimens  from  the  following  localities. 

BRAZIL 

GOIAS:  Santa  Isabel  do  Morro  (1;  DRWh).  PIAUI:  Terezina  (2;  MGFT). 


Collecting  notes.  — As  these  specimens  were  collected  in  January  and  June,  adults  of  S. 
multipunctatus  are  probably  active  throughout  the  year. 

Taxonomic  notes.  - Specimens  here  identified  as  S.  multipunctatus  differ  from  specimens 
of  S.  basalis  and  S.  cearaensis  in  coloration  and  in  punctation  of  occiput,  and  doubtless  are 
reproductively  isolated  from  them.  Although  I did  not  see  type  material  of  S.  multipuncta- 
tus, the  specimens  reported  here  fit  the  original  description  quite  well  and  seem  correctly 
assigned. 


Schizogenius  negrei  new  species 

Type  material.  - Holotype  male  (MNHP)  and  allotype  female  (JNeg)  labelled  “Tucuman 
Concepcion  3 1 -xii-46  Coll  A.  Golbach”  ex  collection  J.  Negre.  One  female  specimen  with 
the  same  label  data  is  a paratype  (DRWh). 

Diagnostic  combination.  — Specimens  of  this  species  are  distinguished  from  all  others  in 
the  group  by  entirely  ferrugineous  body  coloration.  Further,  known  females  have  the  pygi- 
dium  apex  only  indistinctly  crenulate. 

Description.  — With  characters  of  the  basalis  group.  As  in  S.  basalis  except  as  follows. 
Color  ferrugineous,  pronotum  slightly  aeneous;  antennae,  palpi,  front  tibiae  and  tarsi,  and 
middle  and  hind  legs  testaceous.  Head  and  pronotum,  Fig.  92;  antennal  article  five  about 
1.2  times  longer  than  wide.  Left  elytron  with  12-13  setae  on  interval  three,  13-17  on  inter- 
val five,  and  10-13  on  interval  seven,  total  35-41  in  specimens  examined.  Male  genitalia,  Fig. 
97;  one  specimen  examined. 

Measurements  and  proportions.  Of  holotype:  TL,  3.38  mm;  LE,  2.08  mm;  WH,  0.71  mm; 
WP,  0.93  mm;  WE,  1.07  mm;  WF/WH,  0.66;  LP/WP,  0.91;  DP/LP,  0.88;  LP/WE,  0.79;  Ta/ 
Ti,  0.73;  PS/LP,  0.62.  Of  allotype:  TL,  3.48  mm;  LE,  2.15  mm;  WH,  0.72  mm;  WP,  0.99 
mm;  WE,  1.13  mm;  WF/WH,  0.65;  LP/WP,  0.89;  DP/LP,  0.89;  LP/WE,  0.78;  Ta/Ti,  0.71; 
PS/LP,  0.63. 

Etymology.  — It  is  with  pleasure  that  I name  this  species  for  my  friend,  J.  Negre. 

Distribution.  — This  species  is  known  only  from  the  type  series  of  three  specimens,  col- 
lected at  Concepcion,  Tucuman,  Argentina.  It  is  the  only  member  of  the  group  known  from 
Argentina. 

Taxonomic  notes.  — S.  negrei  is  morphologically  well  differentiated  from  other  members 
of  the  basalis  group,  and  is  no  doubt  reproductively  isolated  from  them. 

The  elongatus  group 

Diagnostic  combination.  — Specimens  of  this  group  have  the  following  combination  of 
characters:  body  convex;  paramedian  clypeal  carinae  extended  to  median  tooth  or  not; 
clypeal  field  triangular,  at  base  more  than  1 .5  apical  width  of  median  frontal  sulcus;  clypeal 
suture  sharply  impressed;  antennal  articles  five  to  ten  moniliform,  slightly  longer  than  wide; 
submentum  without  accessory  setae;  pronotum  with  paramedian  sulci  distinct  at  base  only, 
and  with  distinct  paralateral  carinae;  front  and  middle  tarsi  slightly  broadened  in  both  sexes; 
elytra  without  discal  setae,  intervals  not  carinate;  sternum  seven  with  paramedian  ambulato- 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


187 


Fig.  87-92.  Head  and  pronotum,  dorsal  aspect.  87.  S.  ferrugineus  Putzeys,  Kenwood  Beach,  Maryland.  88.  S.  auri- 
pennis  Bates,  Arriaga,  Chiapas.  89.  S.  basalis  Putzeys,  Rio  Santa  Lucia,  Uruguay.  90.  S.  cearaensis  new  species,  Forta- 
leza, Brazil.  91.  S.  multipunctatus  Kult,  Terezina,  Brazil.  92.  S.  negrei  new  species,  Concepcion,  Argentina.  Fig.  93-97. 
Male  median  lobe,  lateral  aspect.  93.  S.  ferrugineus  Putzeys,  Logan  County,  Colorado.  94.  S.  auripennis  Bates,  Mira- 
flores,  Baja  California.  95.  S.  cearaensis  new  species,  Fortaleza,  Brazil.  96.  S.  multipunctatus  Kult,  Terezina,  Brazil. 
97.  S.  negrei  new  species,  Concepcion,  Argentina.  Fig.  98-99.  Male  endophallus.  98.  S.  ferrugineus  Putzeys,  Kenwood 
Beach,  Maryland.  99.  S.  auripennis  Bates,  Hermosillo,  Sonora.  Fig.  100-101.  Known  distributions.  100.  S.  ferrugineus 
Putzeys;  hollow  symbols  represent  state  records  only.  101.  S.  auripennis  Bates. 


188 


Whitehead 


ry  setae  in  males,  not  in  females;  paramedian  carinae  of  sternum  three  not  or  hardly  curved 
at  apices;  and  pygidium  apex  strongly  crenulate  in  females. 

Distribution.  — Members  of  this  group  are  known  only  from  Argentina  and  Brazil.  I 
studied  13  specimens  of  the  elongatus  group. 

Taxonomic  notes.  — I assign  here  two  described  species,  S.  costiceps  Steinheil  and  S. 
elongatus  Kult,  though  I have  seen  type  material  of  neither.  Specimens  from  near  the  type 
locality  of  S.  elongatus  agree  well  with  its  original  description.  My  association  of  a specimen 
from  Argentina  with  the  name  S.  costiceps  is  less  definite,  but  I have  no  reason  to  doubt  the 
association.  As  I have  seen  no  type  material  of  the  group,  I give  no  detailed  redescriptions  of 
the  two  described  species  here  assigned  to  it. 

Schizogenius  costiceps  Steinheil 

Schizogenius  costiceps  Steinheil  1869:242.  Type  locality  San  Luis,  Argentina;  location  of 

type  unknown. 

Diagnostic  combination.  — One  specimen  seen  by  me  which  may  belong  to  this  species 
differs  from  specimens  of  S.  elongatus  by  larger  size  (LE  over  2.9  mm),  brighter  green  color, 
more  elongated  paramedian  pronotal  sulci,  and  less  distinctly  punctate  elytral  striae. 

Description.  - With  characters  of  elongatus  group.  Body  piceous,  strongly  metallic  green; 
legs  and  antennae  ferrugineous;  palpi  testaceous.  Head  and  pronotum,  Fig.  102;  clypeal 
carinae  extended  to  median  tooth,  but  weakly  developed  in  apical  third;  antennal  article 
five  about  1.1  times  longer  than  wide.  Elytral  striae  indistinctly  punctate.  Male  genitalia  not 
known. 

Measurements  and  proportions.  TL,  4.69  mm;  LE,  2.96  mm;  WH,  0.91  mm;  WP,  1.27 
mm;  WE,  1.50  mm;  WF/WH,  0.62;  LP/WP,  0.93;  DP/LP,  0.91;  LP/WE,  0.79;  Ta/Ti,  0.66; 
PS/LP,  0.49. 

Etymology.  — Latin,  costa  = rib,  plus  caput  = head,  in  reference  to  the  frontal  carinae. 

Distribution.  - Two  specimens  are  known,  both  from  central  Argentina.  The  type  is  from 
San  Luis,  San  Luis  province.  The  specimen  reported  here  is  labelled  “Rep.  Arg.  Pronuncia- 
mento  Pro-Entre  Rios  XI-63.”  (JNeg). 

Taxonomic  notes.  — The  original  description  of  S.  costiceps  is  inadequate,  as  Steinheil 
thought  that  the  eight  longitudinal  carinae  on  the  frons  were  distinctive.  But  the  Entre  Rios 
specimen  fits  the  description  in  other  characteristics  recorded,  such  as  the  size  and  bright 
green  color,  and  is  from  a sufficiently  close  locality  that  it  may  well  be  conspecific.  Stein- 
heil’s  description  indicates  a more  pale  body  color,  but  his  specimen  was  probably  teneral. 
Further  evidence  that  the  Entre  Rios  specimen  is  conspecific  with  or  closely  related  to  the 
type  of  S.  costiceps  is  in  what  Steinheil  did  not  say:  he  particularly  omitted  any  mention  of 
discal  setae  on  the  elytra. 


Schizogenius  elongatus  Kult 

Schizogenius  elongatus  Kult  1950:146.  Type  locality  Corumba,  Mato  Grosso,  Brazil;  type 
and  paratype  in  Kult  collection,  present  location  not  known. 

Diagnostic  combination.  — Within  the  elongatus  group,  specimens  of  this  species  are  dis- 
tinguished by  small  body  size  (LE  under  2.5  mm),  narrow  body,  aeneopiceous  coloration, 
nearly  obsolete  paramedian  pronotal  sulci,  and  finely  but  distinctly  punctate  elytral  striae. 

Description.  — With  characters  of  elongatus  group.  Body  piceous,  strongly  aeneous;  legs 
and  antennae  ferrugineous;  palpi  testaceous.  Head  and  pronotum,  Fig.  103;  clypeal  carinae 
broken  in  apical  third;  antennal  article  five  about  1.1  times  longer  than  wide.  Elytral  striae 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


189 


finely  but  distinctly  punctate  in  basal  half.  Male  genitalia,  Fig.  108;  1 specimen  examined. 

Measurements  and  proportions.  Largest  specimen,  male:  TL,  3.88  mm;  LE,  2.40  mm; 
WH,  0.79  mm;  WP,  1.07  mm;  WE,  1.20  mm;  WF/WH,  0.59;  LP/WP,  0.94;  DP/LP,  0.86;  LP/ 
WE,  0.84;  Ta/Ti,  0.61;  PS/LP,  0.22.  Smallest  specimen,  female:  TL,  3.39  mm;  LE,  2.13 
mm;  WH,  0.71  mm;WP,  0.94  mm;  WE,  1.04  mm;  WF/WH,  0.58;  LP/WP,  0.91;  DP/LP,  0.92; 
LP/WE,  0.83;  Ta/Ti,  0.64;  PS/LP,  0.26. 

Etymology.  — Latin,  elongatus  = prolonged,  in  reference  to  the  elongate  body  form. 
Distribution.  - Specimens  of  S.  elongatus  have  been  collected  in  two  states  in  central 
Brazil.  I studied  1 2 specimens  from  the  following  localities. 

BRAZIL 

GOIAS:  Santa  Isabel  do  Morro  (8;  DRWh,  MGFT,  UASM).  MATO  GROSSO:  Barra  do  Tapirape  (1;  MZSP),  Caceres  (2; 
MGFT),  Jacare  (1;  MGFT). 


Collecting  notes.  - Adults  of  this  species  have  been  collected  in  June,  November,  and 
December,  and  thus  are  probably  active  throughout  the  year. 

Taxonomic  notes.  - Although  I did  not  study  type  material  of  S.  elongatus,  the  speci- 
mens here  reported  agree  well  with  the  original  description  and  were  collected  near  the  type 
locality.  Kult  erred  in  suggesting  a relationship  with  members  of  the  optimus  group,  which 
lack  distinct  clypeal  suture  and  differ  in  numerous  additional  ways. 

The  carinatus  group 

Diagnostic  combination.  — Members  of  this  group  are  readily  recognized  by  the  following 
combination  of  characters;  antennal  articles  five  to  ten  filiform;  elytral  intervals  strongly 
carinate;  and  elytral  disc  without  setae.  Also:  body  convex;  paramedian  clypeal  carinae  ex- 
tended to  median  tooth  or  not;  clypeal  field  triangular,  at  base  less  than  1 .5  apical  width  of 
median  frontal  sulcus;  clypeal  suture  sharply  impressed;  median  frontal  sulcus  strongly 
microsculptured,  with  median  carina  distinct;  submentum  without  accessory  setae;  prono- 
tum  with  paramedian  sulci  well  developed  and  with  distinct  paralateral  carinae;  front  and 
middle  tarsi  distinctly  expanded,  especially  in  males;  sternum  seven  with  paramedian  ambu- 
latory setae  in  males,  not  in  females;  paramedian  carinae  of  sternum  three  not  curved  at 
apices;  and  pygidium  apex  crenulate  in  females. 

Distribution.  — Seven  specimens  of  this  group  are  known  from  Brazil. 

Taxonomic  notes.  — I assign  here  two  quite  different  species,  S.  carinatus  Whitehead  and 
a new  species  described  below.  Characteristics  of  members  of  this  group  are  so  distinctive 
that  all  described  species  not  seen  by  me  may  safely  be  excluded  from  the  group. 

Schizogenius  carinatus  Whitehead 

Schizogenius  carinatus  Whitehead  1966:2.  Type  locality  Santa  Isabel,  Mato  Grosso,  Brazil; 

holotype  male  in  CAS  (!). 

Diagnostic  combination.  — Within  the  carinatus  group,  specimens  of  this  species  are  dis- 
tinguished from  the  only  known  specimen  of  S.  costipennis  by  the  following:  body  color 
piceous;  clypeal  carinae  clearly  extended  to  median  tooth;  dorsum  not  extensively  micro- 
sculptured;  pronotum  not  rugose;  and  elytral  intervals  more  distinctly  carinate. 

Description.  — I have  nothing  to  add  to  the  original  description  (Whitehead,  1966a), 
except  pygidium  apex  crenulate  in  female. 

Etymology.  - Latin,  carina  = keel,  in  reference  ot  the  carinate  elytra. 


190 


Whitehead 


Distribution.  — I studied  six  specimens  of  S.  carinatus  from  the  following  localities  in 
central  Brazil. 


BRAZIL 

GOIAS:  Santa  Isabel  do  Mono  (2;  DRWh).  MATO  GROSSO:  Jacare  (3;  MGFT),  Santa  Isabel  (1;  CAS). 


Collecting  notes.  — Adults  of  this  species  probably  are  active  throughout  the  year,  as 
specimens  have  been  collected  in  June,  August,  and  November. 

Schizogenius  costipennis  new  species 

Type  material.  — Holotype  female  labelled  “S.  Isabel  do  Morro  Ilha  do  Bananal  Bras. 
Goias  VI.  1961  leg.  M.  Alvarenga”  (MGFT). 

Diagnostic  combination.  — The  only  known  specimen  of  this  species  differs  from  speci- 
mens of  S.  carinatus,  the  only  other  known  member  of  the  carinatus  group,  by  the  follow- 
ing: body  color  rufotestaceous;  integument  extensively  microsculptured;  pronotum  strongly 
rugose;  paramedian  clypeal  carinae  interrupted  before  median  tooth;  and  elytral  intervals 
distinctly  but  weakly  carinate. 

Description.  — Body  broad,  moderately  convex.  Color  rufotestaceous,  without  metallic 
luster. 

Integument.  Entire  body  apparently  microsculptured,  but  under  surface  not  examined  in 
detail;  median  frontal  sulcus  as  well  as  paramedian  sulci  coarsely  microsculptured.  Pronotum 
strongly  rugose. 

Head.  Fig.  104.  Paramedian  clypeal  carinae  straight,  parallel,  abbreviated  at  apices;  me- 
dian field  triangular,  no  wider  at  base  than  apex  of  median  field  of  frons.  Clypeal  suture 
sharply  defined.  Median  frontal  sulcus  divided  by  longitudinal  carina.  Eye  uniformly  fac- 
eted. Neck  rugose-punctate.  Antennal  articles  four  to  ten  elongate,  filiform,  article  five 
about  1 .8  times  longer  than  wide. 

Pronotum.  Fig.  104.  Sides  bisetose,  hind  angles  strongly  reduced,  entire  surface  rugose. 
Paramedian  longitudinal  sulci  long,  impunctate,  sinuate,  shallow  toward  apices,  strongly 
hooked  basally.  Anterior  transverse  impression  finely  punctate.  Paralateral  longitudinal  cari- 
nae strongly  developed. 

Legs.  Front  and  middle  tarsi  strongly  dilated;  hind  tarsi  slender,  short.  Paronychia  about 
half  as  long  as  tarsal  claws.  Front  tibia  narrowed  evenly  to  base.  Front  femur  strongly 
constricted  near  apex. 

Elytra.  Discal  setae  absent.  Striae  deep,  sharply  engraved,  finely  punctate  in  basal  two- 
thirds.  Interval  one  moderately  convex;  intervals  two  to  seven  strongly  raised,  finely  cari- 
nate; interval  eight  sharply  carinate  at  apex;  apices  of  intervals  three,  five,  and  seven  broadly 
joined.  Humeral  denticles  moderately  prominent. 

Abdomen.  Sternum  three  with  paramedian  carinae  straight.  Sternum  seven  without  para- 
median ambulatory  setae.  Pygidium  with  apical  margin  crenulate. 

Male  genitalia.  Not  known. 

Measurements  and  proportions.  Holotype:  TL,  4.15  mm;  LE,  2.68  mm;  WH,  0.92  mm; 
WP,  1.21  mm;  WE,  1.56  mm;  WF/WH,  0.63;  LP/WP,  0.76;  DP/LP,  0.96;  LP/WE,  0.59;  PS / 
LP,  0.70. 

Etymology.  — Latin,  costa  = rib,  plus  penna  = wing,  in  reference  to  the  strongly  sculp- 
tured elytra. 

Distribution.  — This  species  is  known  only  from  the  holotype,  from  central  Brazil. 

Collecting  notes.  — The  holotype  was  collected  in  June.  Specimens  of  the  following 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


191 


additional  species  were  taken  at  the  same  time  and  place:  S.  carinatus,  S.  elongatus,  S.  jacar- 
ensis,  and  S.  multipunctatus.  I presume  that  adults  of  S.  costipennis  are  active  throughout 
the  year,  as  adults  of  the  other  species  probably  are,  and  that  they  probably  live  in  riparian 
gravel  bars. 

Taxonomic  notes.  — S.  costipennis  is  peculiar  in  numerous  ways,  and  perhaps  is  not  close- 
ly related  to  S.  carinatus.  In  addition  to  characteristics  mentioned  in  the  diagnostic  combina- 
tion, the  form  of  the  thorax  is  quite  different,  and  the  elytra  are  much  more  strongly  ovate. 
However,  I doubt  that  the  peculiar  combination  of  elytral  intervals  carinate,  elytral  disc 
without  setae,  and  pronotum  with  strong  paralateral  carinae  is  the  result  of  convergence. 
And  the  two  species  are  sympatric,  so  at  least  there  is  no  evidence  that  this  combination  of 
characteristics  had  independent  origins  in  separate  areas.  If  these  two  species  indeed  are 
related,  then  additional  species  of  the  group  should  exist  and,  if  found,  should  verify  the 
relationship. 


The  strigicollis  group 

Diagnostic  combination.  — Specimens  of  this  group  are  distinguished  from  all  others  of 
the  genus  by  numerous  accessory  setae  on  submentum.  They  have  the  following  additional 
combination  of  characters:  body  convex;  paramedian  clypeal  carinae  extended  to  median 
tooth,  moderately  elevated  basally;  clypeal  field  triangular,  at  base  less  than  1 .5  apical  width 
of  median  frontal  sulcus;  clypeal  suture  sharply  impressed;  antennal  articles  five  to  ten  fili- 
form, distinctly  longer  than  wide;  pronotum  with  well  developed  paralateral  carinae  and 
paramedian  sulci;  front  and  middle  tarsi  moderately  dilated  in  males  and  females;  elytra 
with  numerous  discal  setae  on  intervals  three,  five,  and  seven,  intervals  not  carinate;  sternum 
seven  with  paramedian  ambulatory  setae  in  male  only;  paramedian  carinae  of  sternum  three 
not  curved  at  apices;  and  pygidium  apex  not  crenulate  in  females. 

Distribution.  — This  group  is  known  only  from  three  specimens  from  Colombia. 

Taxonomic  notes.  — Lindroth  (1961)  designated  S.  strigicollis  as  the  type  species  of  the 
genus,  as  it  was  the  first  species  of  the  genus  listed  by  Putzeys  (1846).  This  species  is  the 
only  known  member  of  the  group. 

Schizogenius  strigicollis  Putzeys 

Schizogenius  strigicollis  Putzeys  1846:650.  Type  locality  Colombia,  here  restricted  to  Ara- 

cataca,  Magdalena;  male  specimen  in  IRSB  (!)  labelled  lectotype  here  so  designated. 

Putzeys  1863:24.  Putzeys  1866:222.  Bates  1881 :38.  Lindroth  1961:164. 

Diagnostic  combination.  — S.  strigicollis  is  the  only  known  species  of  Schizogenius  charac- 
terized by  numerous  accessory  setae  on  submentum. 

Description.  - With  characters  of  strigicollis  group.  Body  convex,  bright  aeneopiceous, 
elytra  margined  in  metallic  green;  legs,  antennae,  and  mouthparts  rufotestaceous.  Head  and 
pronotum,  Fig.  105;  antennal  article  five  about  1.8  times  longer  than  wide;  pronotum  dis- 
tinctly rugose  at  sides  and  base.  Elytral  striae  indistinctly  punctate  basally;  left  elytron  with 
about  13-15  setae  on  interval  three,  13  on  interval  five,  9-10  on  interval  seven;  total  35-38 
in  specimens  examined.  Male  median  lobe,  Fig.  109;  one  specimen  examined. 

Measurements  and  proportions.  Largest  specimen,  a female:  TL,  4.90  mm;  LE,  3.12  mm; 
WH,  1.06  mm;  WP,  1.35  mm;  WE,  1 .67  mm;  WF/WH,  0.59;  LP/WP,  0.84;  DP/LP,  0.91 ; LP/ 
WE,  0.68;  Ta/Ti,  0.63;  PS/LP,  0.53.  Smallest  specimen,  a male:  TL,  4.69  mm;  LE,  2.88 
mm ; WH,  1 . 1 0 mm ; WP,  1 .35  mm ; WE,  1 .62  mm ; WF/WH,  0.6 1 ; LP/WP,  0.86 ; DP/LP,  0.89; 
LP/WE,  0.72;  Ta/Ti,  0.62;  PS/LP,  0.59. 


192 


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Etymology.  — Latin,  strigosus  = strigose,  plus  collum  = neck,  in  reference  to  the  rugose 
pronotum. 

Distribution.  — I studied  three  specimens  of  this  species,  all  from  Colombia. 

COLOMBIA 

No  locality  (1;  IRSB).  MAGDALENA:  Aracataca  (2;  MCZ). 


Collecting  notes.  — Specimens  from  Aracataca  were  collected  by  P.  J.  Darlington  in  March 
and  May. 

Taxonomic  notes.  - Records  of  this  species  from  Mexico  (Putzeys,  1846;  Bates,  1881) 
are  no  doubt  erroneous. 


The  arechavaletae  group 

Diagnostic  combination.  — Specimens  of  this  group  differ  from  other  members  of  the 
subgenus  by  having  three  or  four  setae  on  interval  three  of  elytron,  and  two  setae  only  on 
basal  half  of  interval  five.  They  have  the  following  additional  characters:  body  moderately 
convex;  paramedian  clypeal  carinae  abbreviated  before  median  tooth;  clypeal  field  triangu- 
lar, at  base  more  than  1.5  apical  width  of  median  frontal  sulcus;  clypeal  suture  sharply  im- 
pressed; antennal  articles  five  to  ten  filiform,  distinctly  longer  than  wide;  pronotum  without 
paralateral  carinae,  with  short  paramedian  sulci;  front  and  middle  tarsi  moderately  dilated 
in  both  sexes;  elytral  intervals  not  carinate;  sternum  seven  with  paramedian  ambulatory  setae 
in  males  only;  paramedian  carinae  of  sternum  three  curved  outward  at  apices;  and  pygidium 
apex  crenulate  in  females. 

Distribution.  - Two  species  are  known,  one  from  Uruguay  and  the  other  from  north- 
eastern Brazil.  I studied  ten  specimens  of  the  arechavaletae  group. 

Taxonomic  notes.  — The  two. species  included  in  this  group  are  quite  different  in  body 
form,  but  as  they  agree  in  all  important  ways  except  length  of  paronychia  they  probably  are 
quite  closely  related. 


Schizogenius  arechavaletae  Putzeys 

Schizogenius  arechavaletae  Putzeys  1866:227.  Type  locality  Santa  Lucia  River,  north  of 
Montevideo,  Uruguay;  male  specimen  in  IRSB  labelled  lectotype  (!),  here  so  designated. 
Schizogenius  angusticollis  Putzeys  1866:231.  Type  locality  Santa  Lucia  River,  north  of 
Montevideo,  Uruguay;  holotype  male  in  IRSB  (!).  NEW  SYNONYMY. 

Diagnostic  combination.  — Specimens  of  S.  arechavaletae  are  distinguished  from  speci- 
mens of  S.  reichardti,  the  only  other  known  member  of  the  arechavaletae  group,  by  longer 
paramedian  pronotal  sulci,  more  convex  body  form,  and  rounded  humeral  angles  of  elytra. 

Description.  - Body  broad,  convex.  Color  of  type  specimens  faded;  originally  described 
by  Putzeys  as  aeneocupreous,  appendages  fuscous. 

Integument.  Microsculpture  generally  as  in  S.  reichardti  except  on  abdomen:  less  devel- 
oped on  sternum  two,  in  coxal  depressions  and  on  small  paralateral  patches  on  sternum 
three,  otherwise  largely  unmicrosculptured. 

Head.  Fig.  106.  Clypeal  carinae  tuberculate,  in  some  specimens  evidently  but  weakly  ex- 
tended to  median  tooth;  median  clypeal  field  triangular,  broad,  width  at  base  about  1.5 
apical  width  of  median  field  of  frons.  Clypeal  suture  sharply  engraved.  Eye  prominent, 
subglobose,  uniformly  faceted.  Neck  densely,  coarsely  punctate.  Gena  rugose  in  front.  An- 
tenna filiform,  short,  article  five  about  1.5  times  wider  than  long. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


193 


Pronotum.  Fig.  106.  Sides  bisetose,  hind  angles  weakly  developed,  base  and  sides  mo- 
derately rugose.  Paramedian  sulci  short,  im punctate,  distinctly  engraved  apically,  deep  and 
broadly  hooked  basally.  Anterior  transverse  impression  distinctly  punctate. 

Legs.  Front  and  middle  tarsi  distinctly  dilated,  particularly  in  males,  in  males  with  dense 
ventral  pubescence;  hind  tarsus  slender,  short.  Paronychia  nearly  as  long  as  tarsal  claws. 
Front  tibia  evenly  narrowed  to  base  where  much  narrower  than  at  base  of  antennal  cleaner. 
Front  femur  not  strongly  constricted  near  apex. 

Elytra.  Three  or  four  setigerous  punctures  on  interval  three,  two  on  basal  half  of  interval 
five.  Striae  deep  and  sharply  engraved,  finely  punctate  in  basal  half.  Intervals  one  to  seven 
broad  and  convex,  interval  eight  carinate  at  apex;  intervals  three  and  five  broadly  joined 
apically  with  interval  seven.  Humeral  angles  rounded. 

Abdomen.  Sternum  three  with  paramedian  carinae  curved  at  apices.  Sternum  seven  with 
paired  paramedian  ambulatory  setae  in  males  only.  Pygidium  with  apical  margin  crenulate 
in  females. 

Male  genitalia.  Median  lobe,  Fig.  110,  111;  three  specimens  studied. 

Measurements  and  proportions.  Largest  specimen,  a female:  TL,  4.66  mm;  LE,  2.95  mm; 
WH,  0.91  mm;  WP,  1.34  mm;  WE,  1 .60  mm;  WF/WH,  0.65;LP/WP,  0.82;  DP/LP,  0.93;  LP/ 
WE,  0.69;  Ta/Ti,  0.67;  PS/LP,  0.58.  Smallest  specimen,  a male:  TL,  3.61  mm;  LE,  2.27 
mm;  WH,  0.75  mm;  WP,  0.99  mm;  WE,  1.27  mm;  WF/WH,  0.65;  LP/WP,  0.86;  DP/LP,  0.93; 
LP/WE,  0.67;  Ta/Ti,  0.69;  PS/LP,  0.59. 

Etymology.  — The  name  arechavaletae  was  given  in  honor  of  the  collector,  M.  Arechava- 
leta.  The  name  angusticollis  is  derived  from  Latin,  angustus  = narrow,  plus  collum  = neck, 
in  reference  to  the  narrowly  prominent  pronotal  front  angles. 

Distribution.  - I studied  seven  specimens  of  this  species  from  Uruguay,  all  from  type 
series  (IRSB)  of  S.  arechavaletae  and  S.  angusticollis. 

Taxonomic  notes.  — If  the  holotype  specimen  of  S.  angusticollis  is  correctly  labelled,  it 
unquestionably  is  conspecific  with  specimens  named  S.  arechavaletae , and  was  from  the 
same  series.  According  to  original  descriptions,  however,  there  is  some  possibility  of  error; 
the  description  of  S.  angusticollis  suggests  an  animal  more  like  S.  reichardti  than  S.  arecha- 
valetae. This  question  cannot  be  resolved  until  other  specimens  of  the  type  series  are  found, 
and,  for  the  present,  I prefer  to  place  the  names  S.  angusticollis  and  S.  arechavaletae  in 
synonymy. 

Paronychia  apparently  are  primitively  short  in  the  subgenus  Schizogenius , and  secondarily 
elongate  in  S.  arechavaletae  and  all  following  species  groups.  I suspect  that  the  ancestor  of 
the  arechavaletae  group  has  secondarily  elongated  paronychia,  and  that  the  shortened  pa- 
ronychia of  S.  reichardti  represents  a reversion  to  the  ancestral  condition. 

Schizogenius  reichardti  new  species 

Type  material.  - Holotype  female  labelled  “PARAIBA  Corema  - VI-1957  Exp.  Dep. 
Zoologia”  (MZSP).  Two  additional  females  with  the  same  label  data,  from  Brazil,  are  para- 
types  (MZSP,  DRWh). 

Diagnostic  combination.  — Specimens  of  this  species  are  readily  distinguished  from  others 
in  the  arechavaletae  group  by  numerous  characteristics,  including  prominent  humeral  angles 
of  elytra. 

Description.  — Body  broad,  dorsum  flattened.  Color  dark  castaneous,  elytra  with  slight 
aeneous  luster;  legs,  antennae,  and  palpi  ferrugineous. 

Integument.  Distinct  microsculpture  on  paramedian  frontal  sulci,  genae,  mouthparts,  base 
of  pronotum,  anterior  surfaces  of  front  legs,  middle  legs  except  trochanters,  hind  tibiae  and 


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posterior  surfaces  of  hind  femora,  elytral  epipleura  on  base  and  apical  two-thirds,  sternum 
two,  and  portions  of  sterna  three  to  seven. 

Head.  Fig.  107.  Clypeal  carinae  tuberculate,  convergent,  nor  or  weakly  extended  to  me- 
dian tooth;  median  clypeal  field  triangular,  broad,  width  at  base  about  1.5  apical  width  of 
median  field  of  front.  Clypeal  suture  sharply  engraved.  Eye  prominent,  subglobose,  uniform- 
ly faceted.  Neck  densely,  coarsely  punctate.  Gena  rugose  in  front.  Antenna  filiform,  short, 
article  five  about  1.3  to  1.4  times  longer  than  wide. 

Pronotum.  Fig.  107.  Sides  bisetose,  hind  angles  weakly  developed,  base  and  sides  mod- 
erately rugose.  Paramedian  sulci  short,  impunctate,  shallow  and  indistinctly  engraved  apical- 
ly,  deep  and  broadly  hooked  basally.  Anterior  transverse  impression  distinctly  punctate. 

Legs.  Front  and  middle  tarsi  distinctly  dilated,  without  dense  ventral  pubescence;  hind 
tarsus  slender,  short.  Paronychia  distinct,  about  half  length  of  tarsal  claws.  Front  tibia 
evenly  narrowed  to  base  where  much  narrower  than  at  base  of  antennal  cleaner.  Front  femur 
not  strongly  constricted  near  apex. 

Elytra.  Three  or  four  setigerous  punctures  on  interval  three,  two  on  basal  half  of  interval 
five.  Striae  deep  and  sharply  engraved,  distinctly  punctate  in  basal  three-fourths.  Intervals 
one  to  seven  broad  and  moderately  convex,  interval  eight  carinate  at  apex;  intervals  three 
and  five  broadly  joined  apically  with  interval  seven.  Humeral  angles  prominent. 

Abdomen.  Sternum  three  with  paramedian  carinae  curved  at  apices.  Sternum  seven  with- 
out paramedian  ambulatory  setae  in  female.  Pygidium  with  apical  margin  crenulate  in  fe- 
male. 

Male  genitalia.  Unknown. 

Measurements  and  proportions.  Largest  specimen:  TL,  4.99  mm;  LE,  3.09  mm;  WH,  1.15 
mm;  WP,  1.42  mm;  WE,  1.65  mm;  WF/WH,  0.65;LP/WP,  0.88;  DP/LP,  0.90;  LP/WE,  0.73; 
Ta/Ti,  0.73;  PS/LP,  0.44.  Smallest  specimen:  TL,  4.47  mm;  LE,  2.75  mm;  WH,  1.02  mm; 
WP,  1.28  mm;  WE,  1.58  mm;  WF/WH,  0.66;  LP/WP,  0.86;  DP/LP,  0.93;  LP/WE,  0.70;  Ta/ 
Ti,  0.68;  PS/LP,  0.49. 

Variation.  — On  one  paratype  basal  halves  of  all  femora  are  strongly  rugose. 

Etymology.  — I take  pleasure  in  naming  these  distinctive  beetles  for  my  friend,  Hans 
Reichardt,  who  made  specimens  of  them  available  for  study. 

Distribution.  — Only  the  type  specimens  from  northeastern  Brazil  are  known. 

The  ocellatus  group 

Diagnostic  combination.  — Specimens  of  this  group  differ  from  all  others  of  the  genus  by 
remarkably  reduced,  bubble-like  eyes,  and  plurisetose  antennal  pedicels.  They  have  the 
following  additional  combination  of  characters:  body  flattened;  pro  thoracic  pleura  and  ab- 
dominal sterna  microsculptured;  paramedian  clypeal  carinae  extended  to  median  tooth; 
clypeal  field  triangular,  at  base  less  than  1.5  apical  width  of  median  frontal  sulcus,  clypeal 
suture  sharply  impressed;  inner  paramedian  frontal  carinae  not  grossly  thickened;  median 
frontal  sulcus  not  or  weakly  microsculptured,  without  median  longitudinal  carina;  antennal 
articles  five  to  ten  filiform,  elongate;  submentum  without  accessory  setae;  pronotum  with 
elongate  paramedian  sulci,  without  paralateral  carinae;  front  and  middle  tarsi  broadened  in 
both  sexes,  particularly  in  males;  elytra  with  discal  setae  on  interval  three  only;  hind  wings 
brachypterous;  sternum  seven  with  paramedian  ambulatory  setae  in  males,  not  in  females; 
paramedian  carinae  of  sternum  three  strongly  curved  at  apices;  and  pygidium  apex  crenulate 
in  females. 

Distribution.  — Eight  specimens  of  one  species  were  collected  in  a cave  in  southern 
Brazil. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


195 


i Fig.  102-107.  Head  and  pronotum,  dorsal  aspect.  102.  S.  costiceps  Steinheil,  Entre  Rios,  Argentina.  103.  S.  elongatus 

Kult,  Caceres,  Brazil.  104.  S.  costipennis  new  species,  Santa  Isabel  do  Morro,  Brazil.  105.  S.  strigicollis  Putzeys,  Araca- 

| taca,  Colombia.  106.  S.  arechavaletae  Putzeys,  Rio  Santa  Lucia,  Uruguay.  107.  S.  reichardti  new  species,  Corema,  Brazil. 
I Fig.  108-111.  Male  median  lobe,  lateral  aspect.  108.  S.  elongatus  Kult,  Santa  Isabel  do  Morro,  Brazil.  109.  S.  strigi- 
collis Putzeys,  Aracataca,  Colombia.  110,  111.  S.  arechavaletae  Putzeys,  Rio  Santa  Lucia,  Uruguay. 


196 


Whitehead 


Taxonomic  notes.  — The  plurisetose  pedicel  is  reminiscent  of  antennae  in  Halocoryza 
species,  but  is  clearly  a secondary  adaptation.  Relationships  are  uncertain;  castaneous  color, 
ventral  microsculpture,  prominent  hind  angles,  curved  sternal  carinae,  reduced  elytral  seta- 
tion,  and  other  characteristics  suggest  relationship  with  members  of  the  darlingtoni  group. 
The  reduced  eyes  are  quite  unlike  any  others  seen  by  me  in  Carabidae,  and  lend  the  head  a 
peculiar  appearance. 


Schizogenius  ocellatus  new  species 

Type  material  - Holotype  male  and  allotype  female  labelled  “Grutas  das  Areias  Sao 
Paulo  Bresil  30.VII.68  P.  Strinati”  (MHNG).  Six  additional  specimens  with  the  same  label 
data  are  paratypes  (DRWh,  MHNG,  MZSP,  UASM). 

Diagnostic  combination.  — The  peculiarly  reduced  eyes  readily  distinguish  members  of 
this  species  from  all  others  of  the  genus  so  far  known. 

Description.  - Body  flattened,  elytra  ovate.  Color  castaneous,  no  aeneous  luster,  append- 
ages paler. 

Integument.  Conspicuous  microsculpture  on  paramedian  frontal  sulci,  genae,  mouthparts, 
prothoracic  pleura,  front  tibiae  and  anterior  surfaces  of  front  femora,  middle  legs  except 
trochanters,  hind  tibiae  and  posterior  surfaces  of  hind  femora,  elytral  epipleura,  and  most 
of  abdomen  except  on  midline  of  sternum  seven. 

Head.  Fig.  1 12.  Paramedian  clypeal  carinae  straight,  extended  to  median  tooth;  median 
field  narrow,  not  or  hardly  wider  at  base  than  apex  of  median  frontal  sulcus.  Clypeal  suture 
sharply  defined.  Eye  reduced,  bubble-like,  not  apparently  faceted.  Neck  coarsely  punctate. 
Gena  rugose-punctate.  Antennal  articles  five  to  ten  elongate,  article  five  about  2.0  times 
longer  than  wide. 

Pronotum.  Fig.  1 12.  Sictes  bisetose,  hind  angles  prominent,  base  not  rugose.  Paramedian 
longitudinal  sulci  elongate,  impunctate,  deep  throughout,  slightly  hooked  basally.  Anterior 
transverse  impression  punctate.  Front  angles  sharply  produced. 

Legs.  Front  and  middle  tarsi  slightly  dilated  and  pubescent  ventrally  in  females,  strongly 
so  in  males;  hind  tarsus  slender,  short.  Paronychia  conspicuous,  nearly  as  long  as  tarsal  claws. 
Front  tibia  narrowed  evenly  to  base.  Front  femur  not  strongly  constricted  near  apex. 

Elytra.  Three  or  four  setae  on  interval  three,  none  on  intervals  five  or  seven.  Striae  deep 
and  sharply  engraved,  distinctly  punctate  in  basal  two-thirds.  Intervals  one  to  seven  broad 
and  convex,  interval  eight  carinate  at  apex;  apices  of  intervals  three,  five,  and  seven  broadly 
joined.  Hind  wings  brachypterous. 

Abdomen.  Sternum  three  with  paramedian  carinae  curved  at  apices.  Sternum  seven  with 
paired  ambulatory  setae  in  males,  not  in  females.  Apex  of  pygidium  entire  in  males,  crenu- 
late  in  females. 

Male  genitalia.  Median  lobe,  Fig.  116;  one  specimen  examined. 

Measurements  and  proportions.  Holotype:  TL,  3.65  mm;  LE,  2.10  mm;  WH,  0.50  mm; 
WP,  1.07  mm;  WE,  1.17  mm;  WF/WH,  0.79;  LP/WP,  0.98;  DP/LP,  0.78;  LP/WE,  0.90;Ta/Ti, 
0.64;  PS/LP,  0.67.  Allotype:  TL,  3.82  mm;  LE,  2.24  mm;  WH,  0.80  mm;  WP,  1.08  mm; 
WE,  1.21  mm;  WF/WH,  0.81;  LP/WP,  1 .00;  DP/LP,  0.80;  LP/WE,  0.89;  Ta/Ti,  0.63;  PS/LP, 
0.67. 

Etymology.  — Latin,  ocellatus  = having  little  eyes,  in  reference  to  the  ocellus-like  eye. 

Distribution.  — Only  the  type  series  of  eight  specimens  is  known,  from  a cave  in  south- 
eastern Brazil. 

Taxonomic  notes.  - The  only  true  troglobitic  scaritines  heretofore  described  are  Spe- 
laeodytes  mirabilis  Miller  and  Italodytes  stammeri  Muller  from  Europe  and  Antroforceps 


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197 


bolivari  Barr  from  Mexico  (see  Barr,  1967).  Specimens  of  S.  ocellatus  differ  from  all  other 
known  specimens  of  Schizogenius  by  brachyptery,  plurisetose  antennal  pedicel,  and  greatly 
modified  eye  structure,  and  in  these  ways  are  reminiscent  of  A bolivari.  Specimens  of  other 
Schizogenius  species  have  been  taken  in  caves,  and  in  particular  specimens  of  S.  tibialis  from 
Indian  Creek  Cave  in  Texas  have  markedly  reduced  eyes.  But  though  measures  of  relative 
eye  size  (WF/WH)  for  S.  ocellatus  and  cavernicolous  S.  tibialis  are  similar,  they  are  not 
comparable  because  of  grossly  different  eye  structures. 

The  darlingtoni  group 

Diagnostic  combination.  - Specimens  of  this  group  have  the  following  combination  of 
characters:  body  flattened;  paramedian  clypeal  carinae  extended  to  median  tooth;  clypeal 
field  triangular,  at  base  less  than  1.5  apical  width  of  median  frontal  sulcus;  clypeal  suture 
sharply  impressed  or  not;  inner  paramedian  frontal  carinae  grossly  thickened;  median  frontal 
sulcus  microsculptured,  with  or  without  median  longitudinal  carina;  antennal  articles  five 
to  ten  filiform;  submentum  without  accessory  setae;  pronotum  with  elongate  paramedian 
sulci,  without  paralateral  carinae;  front  and  middle  tarsi  broadened  in  both  sexes,  particular- 
ly in  males;  elytra  with  discal  setae  on  intervals  three  and  five  only;  sternum  seven  with 
paramedian  ambulatory  setae  in  males  only;  paramedian  carinae  of  sternum  three  strongly 
curved  at  apices;  and  pygidium  apex  crenulate  in  females  or  not. 

Distribution.  — S.  darlingtoni  Kult  was  described  from  Panama.  I studied  ten  specimens  of 
other  species  from  Colombia,  Venezuela,  and  Peru. 

Taxonomic  notes.  — I studied  type  material  of  no  members  of  this  group,  and  am  there- 
fore unable  to  review  them  at  this  time.  As  here  defined,  the  group  contains  specimens  with 
enlarged  inner  paramedian  frontal  carinae,  microsculptured  frontal  sulcus,  normally  devel- 
oped eyes,  and  setae  on  elytral  intervals  three  and  five.  The  type  specimens  of  S.  darlingtoni 
Kult  (1950:140),  described  from  Volcan  de  Chiriqui,  Panama  (BMNH),  have  strong  micro- 
sculpture on  the  apical  two-thirds  of  the  elytra.  I judge  from  original  descriptions  that  S. 
interstriatus  Putzeys  (1878:54,  from  Medellin,  Colombia)  and  S.  riparius  Putzeys  (1878:54, 
from  Ibague,  Colombia)  are  members  of  this  group.  These  descriptions  are  not  adequate, 
and  I do  not  know  the  present  location  of  type  specimens.  However,  as  an  aid  to  future 
students,  I made  tentative  identifications  of  the  following  specimens. 

Seven  specimens  from  Rio  Frio,  Magdalena,  Colombia  (DRWh,  MCZ)  agree  well  with 
Putzeys’  description  of  S.  interstriatus  in  all  characteristics  reported.  Additional  character- 
istics are:  paramedian  pronotal  sulci  elongate  (PS/LP,  0.76-0.81);  front  femur  strongly  angu- 
late  on  midventral  margin;  elytral  interval  three  with  four  setae,  interval  five  with  five  or 
six  setae;  abdomen  without  extensive  microsculpture;  and  female  pygidium  not  crenulate  at 
apex.  No  other  known  species  in  the  genus  have  the  angulate  front  femur  characteristic  of 
this  species. 

Three  additional  specimens  are  in  general  agreement  with  Putzeys’  description  of  S.  ripa- 
rius, but  may  represent  at  least  two  different  species,  and  as  none  are  from  Colombia 
perhaps  none  are  conspecific  with  S.  riparius.  These  specimens  are  from  the  following  lo- 
calities: Chanchamayo,  Peru  (MCZ);  Cueva  Alfredo  Jahn  Miranda,  Venezuela  (MNHG);  and 
El  Valle,  Venezuela  (CNHM).  The  Venezuelan  specimens  (LE,  2.53-2.66  mm)  agree  in  the 
body  size  reported  for  S.  riparius , but  the  Peruvian  specimen  is  much  larger  (LE,  3.04  mm). 
Additional  characteristics  of  these  specimens  are:  paramedian  pronotal  sulci  less  elongate 
(PS/LP,  0.68-0.71);  front  femur  not  angulate  midventrally;  elytral  interval  three  with  five 
or  six  setae,  interval  five  with  six  setae;  abdomen  extensively  microsculptured;  and  female 
pygidium  crenulate  at  apex. 


198 


Whitehead 


The  capitalis  group 

Diagnostic  combination.  - Specimens  referred  to  this  group  have  the  following  combina- 
tion of  characters:  body  flattened;  paramedian  clypeal  carinae  extended  to  median  tooth; 
clypeal  field  triangular,  at  base  less  than  1.5  apical  width  of  median  frontal  sulcus;  clypeal 
suture  sharply  impressed;  inner  paramedian  frontal  carinae  not  grossly  thickened;  median 
frontal  sulcus  not  microsculptured,  without  median  longitudinal  carina;  antennal  articles  five 
to  ten  elongate,  filiform;  submentum  without  accessory  setae;  pronotum  with  elongate  para- 
median sulci,  without  paralateral  carinae;  front  and  middle  tarsi  broadened  in  both  sexes, 
particularly  in  males;  elytra  with  discal  setae  on  intervals  three  and  five  only;  sternum  seven 
with  paramedian  ambulatory  setae  in  males,  not  in  females;  paramedian  carinae  of  sternum 
three  strongly  curved  at  apices;  and  pygidium  apex  not  crenulate  in  either  sex.  Members  of 
this  group  differ  most  notably  from  members  of  the  related  lindrothi  group  by  less  convex 
body  and  much  more  strongly  filiform  antennae. 

Distribution.  — I have  seen  12  specimens  of  this  group  from  the  following  countries  in 
western  South  America:  Argentina,  Bolivia,  Colombia,  Ecuador,  and  Peru. 

Taxonomic  notes.  - A female  specimen  in  IRSB  (!)  is  labelled  as  the  type  of  S.  capitalis 
Putzeys  (1863:25,  from  Carracas,  Venezuela),  and  is  labelled  “capitalis  P.”  in  Putzeys’ 
script.  However,  the  specimen  is  labelled  “Medellin”  (Colombia),  and  differs  from  the  origi- 
nal description  by  having  four  setae  each  on  intervals  three  and  five  rather  than  five  setae 
each  on  intervals  three,  five,  and  seven.  Despite  these  discrepancies,  the  specimen  otherwise 
fits  the  description,  and  until  shown  otherwise  I think  it  best  to  regard  this  specimen  as 
holotype  of  S.  capitalis. 

I have  not  seen  type  material  of  S.  putzeysi  Kirsch  (1873: 129,  from  Peru),  but  this  seems 
quite  clearly  from  the.  original  description  to  be  a member  of  the  capitalis  group,  and  I sus- 
pect the  type  specimen  is  teneral.  I have  tentatively  identified  as  S.  putzeysi  six  specimens, 
all  smaller  (LE,  2.50-2.79  mm)  than  the  specimen  of  S.  capitalis  (LE,  3.05  mm)  reported 
above,  and  all  with  distinct  microsculpture  on  the  sides  of  the  pronotum.  These  specimens 
are  from  the  following  localities. 


BOLIVIA 

BENI:  Huachi  (2;  USNM). 

ECUADOR 

EL  ORO:  9 mi.  s.  Santa  Rosa  (3;  CAS). 

PERU 

HUANUCO:  24  mi.  e.  Yurac  (1;  CAS). 


I labelled  five  additional  specimens  as  an  undescribed  species  near  S.  putzeysi.  These  are 
also  smaller  (LE,  2.54-2.84  mm)  than  the  specimen  of  S.  capitalis , but  lack  distinct  micro- 
sculpture on  the  sides  of  the  pronotum.  These  specimens  are  from  the  following  localities. 

ARGENTINA 

TUCUMAN  (1;  MCZ):  Tacanas  (3;  MGFT),  Villa  Monti  (1;  CAS). 

As  I have  seen  no  definite  type  material  of  the  capitalis  group,  and  few  specimens  from  a 
broad  geographic  area,  I attempt  no  detailed  revision.  Specimens  reported  above  may  re- 
present as  few  as  one  to  as  many  as  three  species;  tentative  identifications  are  intended  as  an 
aid  to  future  students,  but  the  group  requires  additional  study  before  definite  identifica- 
tions are  possible. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


199 


The  lindrothi  group 

Diagnostic  combination.  - This  group  includes  those  species  with  three  to  six  evenly  dis- 
tributed setae  in  elytral  intervals  three  and  five,  but  none  in  interval  seven;  antennal  articles 
five  to  ten  submoniliform,  not  strongly  elongated;  median  field  of  frons  not  microsculp- 
tured,  not  bounded  by  unusually  thickened  carinae;  and  pygidium  not  crenulate  in  either 
sex.  Also:  body  nearly  cylindrical;  paramedian  clypeal  carinae  extended  to  median  tooth, 
not  strongly  raised  in  basal  half;  clypeal  field  not  or  hardly  wider  at  base  than  apex  of  me- 
dian frontal  sulcus;  clypeal  suture  sharply  impressed;  anterior  tarsi  of  males  broadened  and 
with  dense  ventral  pubescence;  abdomen  not  extensively  microsculptured;  paramedian  ca- 
rinae of  sternum  three  curved  at  apices;  sternum  seven  with  paramedian  ambulatory  setae 
normally  in  males  only;  and  endophallus  with  short  basal  collar  spines. 

Distribution.  — I have  seen  38  specimens  of  this  group,  representing  at  least  two  species, 
from  Florida,  Guatemala,  Costa  Rica,  Venezuela,  and  Brazil. 

Schizogenius  lindrothi  new  species 

Type  material.  — Holotype  male  and  allotype  female  labelled  “FLORIDA.  Bay  Co.  7 mi. 
n.  Southport  10  July  1967  D.  R.  Whitehead”  (MCZ).  An  additional  18  specimens  from  vari- 
ous localities  in  Florida  are  paratypes  (BMNH,  CAS,  CUNY,  DRWh,  IRSB,  UASM,  USNM). 

Diagnostic  combination.  — Specimens  of  this  species  are  the  only  known  specimens  of 
North  and  Middle  American  Schizogenius  with  three  or  four  setae  each  on  elytral  intervals 
three  and  five  but  none  or  only  one  on  interval  seven. 

Description.  — Body  nearly  cylindrical,  elytra  convex.  Color  dark  rufopiceous,  without 
strong  aeneous  luster;  legs  ferrugineous;  palpi  and  antennae  testaceous. 

Integument.  Conspicuous  microsculpture  on  paramedian  frontal  sulci,  genae,  mouthparts, 
pronotal  base,  front  tibiae  and  anterior  surfaces  of  front  femora,  middle  legs  except  tro- 
chanters, hind  tibiae  and  posterior  surfaces  of  hind  femora,  elytral  epipleura  in  apical  two- 
thirds  and  at  extreme  base,  part  of  sternum  two,  and  coxal  depressions  of  sternum  three. 

Head.  Fig.  113.  Paramedian  clypeal  carinae  straight,  evenly  elevated,  extended  to  median 
tooth;  median  field  narrow,  not  or  hardly  wider  at  base  than  apex  of  median  frontal  sulcus. 
Clypeal  suture  sharply  defined.  Eye  globose,  finely  and  uniformly  faceted.  Neck  densely 
punctate.  Gena  rugose  in  front,  strongly  punctate.  Antennal  articles  five  to  ten  slightly  but 
distinctly  elongate,  submoniliform,  article  five  1.3  times  as  long  as  wide. 

Pronotum.  Fig.  1 13.  Sides  bisetose,  hind  angles  not  prominent,  base  not  rugose.  Parame- 
dian longitudinal  sulci  moderately  elongate,  impunctate,  nearly  straight,  deep  throughout, 
slightly  hooked  basally.  Anterior  transverse  impression  finely  punctate. 

Legs.  Front  and  middle  tarsi  slightly  dilated  and  pubescent  ventrally  in  male,  less  so  in 
female;  hind  tarsus  slender,  rather  short.  Paronychia  conspicuous,  more  than  half  length  of 
tarsal  claws.  Front  tibia  narrowed  evenly  to  base.  Front  femur  not  strongly  constricted  near 
apex. 

Elytra.  Three  to  four  setae  each  on  intervals  three  and  five,  none  or  rarely  one  on  interval 
seven.  Striae  deep  and  sharply  engraved,  distinctly  punctate  in  basal  two-thirds.  Intervals 
one  to  seven  broad  and  convex,  interval  eight  carinate  at  apex;  apices  of  intervals  three, 
five,  and  seven  broadly  joined. 

Abdomen.  Sternum  three  with  paramedian  carinae  curved  at  apices.  Sternum  seven  with 
ambulatory  setae  in  males,  normally  not  in  females.  Apex  of  pygidium  entire  in  both  sexes. 

Male  genitalia.  Median  lobe,  Fig.  117;  endophallus,  Fig.  120,  with  short  basal  collar  spines; 
nine  specimens  examined. 


200 


Whitehead 


Measurements  and  proportions.  See  Table  7.  Of  holotype:  TL,  4.19  mm;  LE,  2.52  mm; 
WH,  0.85  mm;  WP,  1.15  mm;  WE,  1 .30  mm;  WF/WH,  0.57;  LP/WP,  0.95;  DP/LP,  0.83;  LP/ 
WE,  0.84;  Ta/Ti,  0.69;  PS/LP,  0.76.  Of  allotype:  TL,  4.15  mm;  LE,  2.52  mm;  WH,  0.68 
mm;  WP,  1.15  mm;  WE,  1.32  mm;  WF/WH,  0.59;  LP/WP,  0.92;  DP/LP,  0.85;  LP/WE,  0.80; 
Ta/Ti,  0.62;  PS/LP,  0.71. 


Table  7.  Descriptive  statistics  for  S.  lindrothi,  based  on  nine  males  from  northern  and 
central  Florida. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

nts,  in  mm. 

TL 

3.76-4.32 

4.006 

0.275 

0.122 

4.58 

LE 

2.28-265 

2.419 

0.180 

0.080 

4.97 

WH 

0.78-0.90 

0.840 

0.053 

0.024 

4.20 

WP 

1.01-1.20 

1.084 

0.096 

0.043 

5.90 

WE 

1.18-1.38 

1.258 

0.095 

0.042 

5.01 

B.  Setae  on  le: 

ft  elytron. 

Interval  3 

3-4 

3.3 

Interval  5 

3-4 

3.3 

Interval  7 

nil 

Total 

6-8 

6.7 

1.3 

0.6 

1.30 

C.  Proportion: 

s. 

WF/WH 

0.55-0.58 

0.564 

0.013 

0.006 

1.54 

LP/WP 

0.94-0.99 

0.958 

0.025 

0.011 

1.73 

DP/LP 

0.83-0.86 

0.841 

0.016 

0.007 

1.26 

LP/WE 

0.80-0.85 

0.830 

0.022 

0.010 

1.81 

Ta/Ti 

0.63-0.69 

0.663 

0.029 

0.013 

2.91 

PS/LP 

0.72-0.79 

0.748 

0.031 

0.014 

2.75 

Variation.  - There  is  not  enough  material  in  collections  to  study  geographic  variation, 
but  if  S.  lindrothi  is  restricted  to  Florida  such  variation  is  probably  negligible.  In  addition  to 
variation  in  the  Florida  specimens  listed  in  Table  7,  I noted  one  female  from  Southport 
with  a single  seta  on  interval  seven  of  the  left  elytron,  and  one  female  from  Enterprise  with 
a pair  of  paramedian  ambulatory  setae  on  sternum  seven. 

I tentatively  associate  with  this  species  single  females  from  Guatemala  and  Costa  Rica. 
The  Guatemala  specimen  differs  by  having  a broad  (LP/WP,  0.88),  deep  (DP/LP,  0.89),  and 
relatively  smaller  (LP/WE,  0.70)  thorax,  and  by  having  more  prominent  pronotal  hind  angles 
and  stronger  humeral  angles  on  the  elytra.  The  Costa  Rican  specimen  falls  outside  the  ob- 
served range  of  variation  in  Florida  specimens  by  having  a smaller  thorax  (LP/WE,  0.71), 
sharper  hind  angles  and  prominent  humeral  denticles,  legs  darker  with  femora  rufopiceous, 
and  more  elytral  setae:  four  or  five  setae  each  on  intervals  three  and  five. 

Etymology.  — I take  pleasure  in  naming  this  new  species  after  C.  H.  Lindroth,  who  first 
mentioned  its  existence  in  his  1961  review  of  the  genus. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


201 


Distribution.  — If  all  specimens  listed  here  are  conspecific,  then  the  distribution  of  S. 
lindrothi  is  disjunct,  in  Florida  and  southern  Middle  America  but  not  in  Mexico  (Fig.  121). 
Inclusion  of  the  Middle  American  specimens  is,  however,  tentative.  I studied  31  specimens 
which  I include  under  the  name  S.  lindrothi , from  the  following  localities. 

UNITED  STATES 

FLORIDA  (2;  CAS,  USNM):  Bay  Co.,  7 mi.  n.  Southport  (9;  BMNH,  DRWh,  IRSB,  MCZ,  UASM);  De  Soto  Co.,  Arcadia 
(1;  CUNY);  Hernando  Co.,  Weeke  Wachee  Springs  (2;  CNC);  Highlands  Co.,  Lake  Placid  (1;  CAS);  Marion  Co.  (1;  CAS); 
Orange  Co.,  Winter  Park  (3;  MCZ);  Sumpter  Co.  (1;  USNM);  Volusia  Co.,  Enterprise  (1;  MCZ,  USNM). 

GUATEMALA 

ALTA  VERAPAZ:  Trece  Aguas  (1;  USNM). 

COSTA  RICA 

LIMON:  Los  Diamantes  (1;  FDAG). 


Collecting  notes.  — I collected  the  Southport  specimens  in  the  sandy  margins  of  a small, 
spring-fed  pond,  and  most  probably  S.  lindrothi  is  exclusively  an  inhabitant  of  sand.  Speci- 
mens of  this  or  related  species  are  not  likely  to  be  found  in  Mexico  or  far  west  of  Florida 
along  the  Gulf  of  Mexico,  if,  as  I suspect,  their  distribution  agrees  with  that  of  the  so-called 
Carribean  pines  (Mirov,  1967).  For  the  same  reason,  it  would  be  no  surprise  to  find  speci- 
mens of  the  lindrothi  group  in  the  West  Indies. 

Taxonomic  notes.  — Specimens  of  this  species  differ  sufficiently  from  those  of  S.  bannin- 
geri  in  details  of  male  genitalia  that  there  is  no  reason  to  suspect  them  conspecific.  Whether 
Middle  American  specimens  are  conspecific  with  S.  lindrothi,  or  even  with  one  another,  is  a 
question  that  cannot  now  be  satisfactorily  answered. 

Schizogenius  banningeri  Kult 

Schizogenius  banningeri  Kult  1950:148.  Type  locality  Corumba,  Mato  Grosso,  Brazil;  type 
in  Kult  Collection,  present  location  not  known;  type  not  seen. 

Diagnostic  combination.  - I assign  to  this  species  all  specimens  of  the  lindrothi  group  seen 
from  South  America.  They  differ  from  specimens  of  S.  lindrothi  by  details  of  male  genitalia 
(median  lobe,  Fig.  1 18;  one  specimen  examined),  and  form  and  sculpture  of  head  and  pro- 
notum  (Fig.  1 14).  In  particular,  the  median  frontal  sulcus  is  less  sharply  defined  and  is  more 
strongly  narrowed  in  front.  As  I have  seen  only  seven  specimens  of  the  lindrothi  group 
from  South  America,  I am  not  certain  they  are  all  conspecific  and  I therefore  do  not  give  a 
detailed  description  here. 

Variation.  — I did  not  study  these  specimens  closely  for  variation,  but  only  the  Mato 
Grosso  specimen  is  as  small  as  specimens  reported  by  Kult. 

Etymology.  — Kult  named  this  species  in  honor  of  M.  Banninger. 

Distribution.  — Just  as  limits  of  the  species  are  unknown,  so  is  the  extent  of  its  distribu- 
tion. I studied  the  following  seven  specimens. 

BRAZIL 

MATO  GROSSO:  Cerceres  (1;  MGFT).  PIAUI:  Terezina  (5;  DRWh,  MGFT). 

VENEZUELA 

CARACAS:  Caracas  (1;  IRSB). 

Collecting  notes.  — Specimens  of  this  species  were  collected  in  January  and  December. 
Taxonomic  notes.  — Though  I have  not  seen  type  material,  the  specimen  from  Cerceres 


202 


Whitehead 


matches  the  original  description  and  is  from  an  appropriate  locality.  Whether  specimens 
from  the  other  two  localities  are  conspecific  is  not  certain. 

The  Caracas  specimen  is  labelled  as  the  type  of  S.  sellatus  Putzeys  (1866:228,  from 
Caracas,  Venezuela),  but  perhaps  incorrectly.  It  does  not  have  maculate  elytra,  though  this 
may  be  the  result  of  aging.  More  important,  it  does  have  a prominent  median  clypeal  tooth, 
in  contradiction  to  Putzeys’  description.  This  is  the  specimen  that  Kult  (1950)  believed  was 
the  type  of  S.  sellatus , and  the  one  with  which  he  compared  his  S.  banningeri.  The  differ- 
ences he  noted  in  body  size  and  convexity  of  elytral  intervals  are  probably  not  important. 
If  this  Caracas  specimen  really  is  the  type  of  S.  sellatus,  then  the  name  S.  banningeri  may 
be  a synonym.  A re-examination  of  all  material  studied  by  Putzeys  will  be  required  to 
resolve  this  question.  For  now,  I regard  the  Caracas  specimen  as  one  of  S.  banningeri,  not  as 
the  type  of  S.  sellatus. 

The  quadripunctatus  group 

Diagnostic  combination.  — Specimens  of  the  only  known  species  of  this  group  have  the 
following  combination  of  characters:  body  moderately  flattened;  paramedian  clypeal  carinae 
extended  to  median  tooth;  antennal  articles  five  to  ten  filiform;  eyes  normal;  discal  setae 
present  on  interval  three  only,  intervals  not  carinate;  and  abdomen  extensively  microsculp- 
tured.  Also:  clypeal  suture  sharply  impressed;  clypeal  field  triangular,  less  than  1.5  apical 
width  of  median  frontal  sulcus;  submentum  without  accessory  setae;  pronotum  with  distinct 
paramedian  sulci,  without  paralateral  carinae;  front  and  middle  tarsi  broadened  and  with 
dense  ventral  pubescence,  especially  in  males;  sternum  seven  with  paramedian  ambulatory 
setae  in  males,  not  in  females;  paramedian  carinae  of  sternum  three  curved  at  apices;  pygi- 
dium  not  crenulate  at  apex  in  either  sex;  and  endophallus  with  basal  collar  spines  distinct. 

Distribution.  — One  species  of  this  group  is  known  from  southern  Brazil  and  northern 
Argentina.  I examined  200  specimens. 

Schizogenius  quadripunctatus  Putzeys 

Schizogenius  quadripunctatus  Putzeys  1866:225.  Type  locality  Parana,  Brazil;  location  of 

type  male  unknown.  Whitehead  1966a: 5. 

Diagnostic  combination.  — Specimens  of  S.  quadripunctatus  are  readily  distinguished  by 
large  size,  dark  color,  filiform  antennae,  microsculptured  abdomen,  and  normally  developed 
eyes. 

Description.  — Since  S.  quadripunctatus  is  the  only  known  member  of  the  quadripuncta- 
tus group,  and  since  specimens  may  readily  be  identified  from  the  original  description,  a 
detailed  redescription  is  not  required.  Head  and  pronotum,  Fig.  1 15.  Left  elytron  with  four 
or  five  discal  setae  on  interval  three.  Male  genitalia,  Fig.  119;  three  specimens  examined. 

Measurements  and  proportions.  Largest  specimen,  female:  TL,  6.20  mm;  LE,  3.97  mm; 
WH,  1.20  mm;  WP,  1.65  mm;  WE,  2.06  mm;  WF/WH,  0.59;  LP/WP,  0.88;  DP/LP,  0.87; 
LP/WE,  0.70;  Ta/Ti,  0.69;  PS/LP,  0.61.  Smallest  specimen,  female:  TL,  5.35  mm;  LE,  3.37 
mm;  WH,  1.05  mm;  WP,  1.39  mm;  WE,  1.81  mm;  WF/WH,  0.60;  LP/WP,  0.91 ; DP/LP,  0.86; 
LP/WE,  0.70;  Ta/Ti,  0.65;  PS/LP,  0.59. 

Etymology.  — Latin,  quadri  = four,  plus  punctum  = small  hole,  in  reference  to  the  normal 
number  of  setae  on  each  elytron. 

Distribution.  — S.  quadripunctatus  is  known  from  two  localities  in  southern  Brazil,  in- 
cluding the  type  locality  in  Parana  state,  and  from  one  locality  in  Argentina.  I examined 
199  specimens  from  Nova  Teutonia,  Santa  Catarina,  Brazil  (DRWh,  MCZ,  MGFT),  and  one 
specimen  from  Pico,  La  Pampa,  Argentina  (MCZ). 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


203 


Fig.  112-115.  Head  and  pronotum,  dorsal  aspect.  112.  S.  ocellatus  new  species,  Grutas  das  Areias,  Brazil.  113.  S. 
lindrothi  new  species,  Southport,  Florida,  114.  S.  banningeri  Kult,  Terezina,  Brazil.  115.  S.  quadripunctatus  Putzeys, 
Nova  Teutonia,  Brazil.  Fig.  116-119.  Male  median  lobe,  lateral  aspect.  116.  S.  ocellatus  new  species,  Grutas  das  Areias, 
Brazil.  117.  S.  lindrothi  new  species,  Southport,  Florida.  118.  S.  banningeri  Kult,  Terezina,  Brazil.  119.  S.  quadri- 
punctatus Putzeys,  Nova  Teutonia,  Brazil.  Fig.  120.  Male  endophallus,  S.  lindrothi  new  species,  Southport,  Florida. 
Fig.  121.  Known  distribution  of  S.  lindrothi  new  species. 


204 


Whitehead 


Collecting  notes.  — Fritz  Plaumann  collected  specimens  of  this  species  in  January,  Feb- 
ruary, March,  April,  August,  September,  and  November,  so  adults  probably  are  active 
throughout  the  year. 

Taxonomic  notes.  — Although  I saw  no  type  material,  specimens  seen  well  match  the 
original  description  of  this  distinctive  species,  and  as  the  locality  in  Santa  Catarina  is  near 
the  type  locality  I have  no  doubt  that  I have  associated  the  name  correctly. 

The  truquii  group 

Diagnostic  combination.  — Members  of  this  group  are  distinguished  from  all  other  in  the 
genus  by  unusually  elongated  paramedian  pronotal  sulci,  which  extend  nearly  to  the  ante- 
rior transverse  impression.  They  are  further  characterized  by  the  following  additional  charac- 
ters: body  subcylindrical;  paramedian  clypeal  carinae  extended  to  median  tooth,  but  more 
strongly  raised  in  basal  half;  clypeal  field  triangular,  its  base  over  1 .5  apical  width  of  median 
frontal  sulcus;  clypeal  suture  sharply  impressed;  antennal  articles  five  to  ten  submoniliform, 
slightly  elongate;  anterior  tarsi  slender  in  both  sexes;  elytral  disc  with  setae  on  intervals 
three,  five,  and  seven;  sternum  seven  with  paramedian  ambulatory  setae  in  males  only; 
paramedian  carinae  of  sternum  three  usually  curved  at  apices;  pygidium  not  crenulate  in 
either  sex;  and  endophallus  without  enlarged  spines. 

Distribution.  - The  only  known  species  in  the  group,  S.  truquii  Putzeys,  is  probably 
restricted  to  the  southern  edge  of  the  Trans-Volcanic  Sierra  of  central  Mexico,  and  north- 
ward along  western  slopes  of  the  Sierra  Madre  Occidental  to  Sinaloa.  I examined  42  speci- 
mens of  the  truquii  group. 


Schizogenius  truquii  Putzeys 

Schizogenius  truquii  Putzeys  1866:224.  Type  locality  Mexico,  here  restricted  to  Cuernavaca, 

Morelos;  type  in  IRSB,  female  specimen  labelled  lectotype  here  so  designated  (!).  Bates 

1881:37. 

Diagnostic  combination.  - S.  truquii  is  the  only  species  of  the  genus  known  to  have  para- 
median pronotal  sulci  extended  forward  nearly  to  anterior  transverse  impression. 

Description.  — Body  subcylindrical.  Color  dark  rufopiceous,  without  metallic  luster, 
appendages  rufous. 

Integument.  Fine  microsculpture  on  paramedian  frontal  sulci,  genae,  mouthparts,  ante- 
rior surfaces  of  front  tibiae  and  femora,  middle  legs  except  trochanters,  hind  tibiae  and 
posterior  surfaces  of  hind  femora,  and  small  patch  near  coxal  depressions  of  sternum  three. 

Head.  Fig.  122.  Clypeal  carinae  extended  to  median  tooth,  more  strongly  raised  in  basal 
half;  clypeal  field  triangular,  narrow,  wider  at  base  than  apex  of  median  frontal  sulcus. 
Clypeal  suture  sharply  engraved.  Frons  with  median  field  smooth,  without  median  carina. 
Eye  globose,  finely  and  uniformly  faceted.  Neck  sparsely,  rather  coarsely  punctate.  Gena 
rugose  in  front.  Antennal  articles  five  to  ten  distinctly  elongate,  submoniliform ; article  five 
1.2-1. 3 times  as  wide  as  long. 

Pronotum.  Fig.  122.  Sides  bisetose,  hind  angles  prominent,  base  not  rugose.  Paramedian 
sulci  elongated  nearly  to  anterior  transverse  impression,  impunctate,  nearly  straight,  deep 
throughout,  slightly  hooked  basally.  Anterior  transverse  impression  punctate.  No  evident 
paralateral  carinae. 

Legs.  Front  and  middle  tarsi  slender,  without  dense  ventral  pubescence;  hind  tarsus  slen- 
der, short.  Paronychia  conspicuous,  nearly  as  long  as  tarsal  claws.  Front  tibia  evenly  nar- 
rowed to  base  where  much  narrower  than  at  base  of  antennal  cleaner.  Front  femur  not 
strongly  constricted  near  apex. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


205 


Elytra.  Six  to  eight  setigerous  punctures  on  interval  three,  five  to  eight  on  interval  five, 
and  three  to  five  on  interval  seven.  Striae  deep  and  sharply  engraved,  distinctly  punctate 
except  at  apex.  Intervals  one  to  eight  broad,  convex;  interval  eight  carinate  at  apex;  intervals 
three,  five,  and  seven  broadly  joined  at  apices. 

Abdomen.  Sternum  three  with  paramedian  carinae  usually  suddenly  curved  outward  at 
apices.  Sternum  seven  with  paramedian  ambulatory  setae  in  males  only.  Pygidium  with 
apical  margin  entire  in  both  sexes. 

Male  genitalia.  Median  lobe,  Fig.  127;  endophallus,  Fig.  137,  with  poorly  developed  basal 
collar  spines;  five  specimens  examined. 

Measurements  and  proportions.  See  Table  8. 

Variation.  — The  smallest  (LE,  2.15  mm)  and  largest  (LE,  2.56  mm)  specimens  are  fe- 
males from  La  Garita,  Jalisco.  A female  from  Sinaloa  has  a flatter  body  (DP/LP,  0.77)  and 
much  smaller  eyes  (WF/WH,  0.67)  than  do  other  specimens. 

Etymology.  — S.  truquii  was  named  to  honor  E.  Truqui,  who  collected  the  specimens 
which  Putzeys  first  reported  as  this  species. 

Distribution.  — Specimens  of  S.  truquii  have  been  found  in  central  and  western  Mexico 
at  moderate  elevations,  from  about  1500'  to  4000'  (Fig.  145).  I studied  42  specimens  from 
the  following  localities. 

MEXICO 

JALISCO:  San  Diego  Cocula  (1;  USNM),  0.4  mi.  w.  Cocula  (2;  DRWh),  8.5  mi.  n.  Juchitlan  (2;  MCZ),  La  Garita  (32; 
BMNH,  CAS,  CNC,  IRSB,  UASM).  MICHOACAN:  8.5  mi.n.  Nueva  Italia  (1;  USNM).  MORELOS:  Cuernavaca  (2;  BMNH). 
SINALOA:  21  mi.  e.  Villa  Union  (1;  CNC). 


Table  8.  Descriptive  statistics  for  S.  truquii , based  on  20  males  from  La  Garita,  Jalisco. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measuremi 

ents,  in  mm. 

TL 

3.67-4.06 

3.854 

0.188 

0.056 

3.26 

LE 

2.25-2.48 

2.347 

0.125 

0.037 

3.54 

WH 

0.75-0.85 

0.805 

0.039 

0.012 

3.26 

WP 

0.93-1.08 

1.000 

0.063 

0.019 

4.19 

WE 

1.15-1.27 

1.204 

0.060 

0.018 

3.40 

B.  Setae  on  le 

;ft  elytron. 

Interval  3 

ON 

00 

6.4 

Interval  5 

00 

6.6 

Interval  7 

3-  5 

4.2 

Total 

14-20 

17.3 

2.0 

0.6 

7.74 

C.  Proportion 

s. 

WF/WH 

0.60-0.64 

0.621 

0.017 

0.006 

2.22 

LP/WP 

0.96-1.02 

0.988 

0.026 

0.008 

1.75 

DP/LP 

0.78-0.85 

0.807 

0.024 

0.007 

1.97 

LP/WE 

0.79-0.86 

0.819 

0.028 

0.008 

2.28 

Ta/Ti 

0.62-0.73 

0.672 

0.052 

0.015 

5.12 

PS/LP 

0.85-0.90 

0.882 

0.022 

0.007 

1.66 

206 


Whitehead 


Collecting  notes.  — At  La  Garita,  the  only  locality  at  which  a long  series  of  this  species 
was  taken,  I found  no  other  species  of  Schizogenius.  1 noted  nothing  peculiar  about  the 
locality,  and  other  carabid  beetles  taken  there  are  found  abundantly  elsewhere  along  similar 
gravel  streams.  At  other  localities  in  Jalisco  and  Michoacan  where  I found  specimens  of  S. 
tniquii,  the  Schizogenius  fauna  included  representatives  of  two  or  three  additional  species. 
Specimens  from  these  states  include  nearly  twice  as  many  males  as  females,  possibly  because 
of  the  season;  all  were  collected  in  January  or  March,  and  none  of  the  collecting  localities 
were  revisited  at  a later  season.  The  Sinaloa  specimen  was  taken  in  July,  at  light. 

Taxonomic  notes.  — Putzeys  (1866)  described  the  body  as  strongly  convex,  and  the 
elytron  as  having  12  setae  each  on  intervals  three,  five,  and  seven.  These  observations  are 
clearly  erroneous.  The  lectotype  specimen  agrees  with  my  description,  and  is  undoubtedly 
one  of  the  specimens  studied  by  Putzeys;  it  has  a printed  label,  “Truqui  Mexique,”  as  well 
as  Putzeys’  handwritten  green  determination  label. 

The  brevisetosus  group 

Diagnostic  combination.  - Specimens  of  this  group  are  distinguished  by  the  following 
combination  of  characters:  elytron  with  35  or  more  short  discal  setae;  abdomen  without 
extensive  microsculpture;  body  depressed;  color  castaneous;  size  large,  LE  over  2.70  mm  in 
all  specimens  seen.  Also:  paramedian  clypeal  carinae  extended  to  median  tooth,  more 
strongly  raised  in  basal  half;  clypeal  field  triangular,  its  base  under  1.5  apical  width  of  me- 
dian frontal  sulcus;  clypeal  suture  sharply  impressed;  antennal  articles  five  to  ten  filiform; 
front  and  middle  tarsi  broadened  and  with  dense  ventral  pubescence,  especially  in  males; 
discal  setae  present  on  intervals  three,  five,  and  seven;  sternum  seven  with  paramedian  am- 
bulatory setae  in  males,  not  in  females;  paramedian  carinae  curved  at  apices;  pygidium  not 
crenulate  in  either  sex;  and  endophallus  with  basal  collar  spines  distinct. 

Distribution.  — S.  brevisetosus  is  known  from  Coahuila,  New  Mexico,  and  Texas.  The 
group  is  probably  most  closely  allied  to  the  pluripunctatus  group,  but  differs  in  various 
ways,  notably  by  short  elytral  setae.  The  two  groups  are  allopatric  but  proximate  in  geog- 
raphic distribution.  I examined  42  specimens  of  the  brevisetosus  group. 

Schizogenius  brevisetosus  new  species 

Type  material.  — Holotype  male  and  allotype  female  labelled  “Sanderson,  TEX.  April 
27,  1959  Becker  & Howden”  (CNC).  An  additional  32  specimens  collected  at  various  times 
and  places  in  Texas  are  paratypes  (CAS,  CNC,  CUNY,  DRWh,  LACM,  MCZ,  UASM,  USNM). 

Diagnostic  combination.  — Specimens  of  this  distinctive  species  are  readily  distinguished 
from  all  others  in  the  genus  by  the  combination  of  large  size,  pale  coloration,  deplanate 
body,  numerous  short  discal  setae  on  elytra,  and  lack  of  extensive  microsculpture  on  abdo- 
men. 

Description.  — Body  deplanate.  Color  dark  castaneous,  without  definite  aeneous  luster, 
appendages  paler. 

Integument.  Conspicuous  microsculpture  on  paramedian  sulci  of  frons,  genae,  mouth- 
parts,  front  tibiae  and  anterior  surfaces  of  front  femora,  middle  legs  except  trochanters, 
hind  tibiae  and  posterior  surfaces  of  hind  femora,  elytral  epipleura  in  apical  two-thirds  and 
at  extreme  base,  and  coxal  depressions  of  sternum  three. 

Head.  Fig.  123.  Labrum  weakly  biemarginate.  Paramedian  carinae  of  clypeus  straight, 
extended  to  median  tooth,  strongly  raised  in  basal  half;  median  field  triangular,  narrow, 
slightly  wider  at  base  than  apex  of  median  frontal  sulcus.  Clypeal  suture  sharply  defined. 
Eye  globose,  finely  and  uniformly  faceted.  Neck  densely,  coarsely  punctate.  Gena  coarsely 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


207 


punctate,  rugose  in  front.  Antennal  articles  four  to  ten  elongate,  filiform;  article  five  1.7 
times  as  long  as  wide. 

Pronotum.  Fig.  123.  Sides  bisetose,  hind  angles  sharp  and  prominent,  base  not  rugose. 
Paramedian  sulci  elongate,  impunctate,  nearly  straight,  deep  throughout,  slightly  hooked 
basally.  Anterior  transverse  impression  finely  punctate. 

Legs.  Front  and  middle  tarsi  slightly  but  distinctly  dilated  and  pubescent  ventrally;  hind 
tarsus  slender,  quite  short.  Paronychia  conspicuous,  more  than  half  length  of  tarsal  claws. 
Front  tibia  narrowed  evenly  to  base.  Front  femur  not  strongly  constricted  near  apex. 

Elytra.  Left  elytron  with  about  13-20  setae  on  interval  three,  12-18  on  interval  five,  and 
10-14  on  interval  seven.  Striae  deep,  sharply  engraved,  finely  but  distinctly  punctate  nearly 
to  apex.  Intervals  one  to  seven  broad  and  flat,  interval  eight  carinate  at  apex;  apices  of 
intervals  three,  five,  and  seven  broadly  joined.  Humeral  denticles  sharp  and  prominent. 

Abdomen.  Sternum  three  with  paramedian  carinae  curved  outward  at  apices.  Sternum 
seven  with  paramedian  ambulatory  setae  in  male  only.  Pygidium  with  apical  margin  entire 
in  both  sexes. 

Male  genitalia.  Median  lobe,  Fig.  128;  endophallus  (Fig.  138)  with  well  developed  basal 
collar  spines.  Three  specimens  examined. 

Measurements  and  proportions.  See  Table  9.  Of  holotype:  TL,  4.87  mm;  LE,  3.05  mm; 
WH,  0.95  mm;  WP,  1.26  mm;  WE,  1.62  mm;  WF/WH,  0.64;  LP/WP,  0.95;  DP/LP,  0.79; 
LP/WE,  0.74;  Ta/Ti,  0.59;  PS/LP,  0.72.  Of  allotype:  TL,  4.63  mm;  LE,  2.98  mm;  WH,  0.85 
mm;  WP,  1.17  mm;  WE,  1.17  mm;  WF/WH,  0.66;  LP/WP,  0.92;  DP/LP,  0.79;  LP/WE,  0.72; 
Ta/Ti,  0.62;  PS/LP,  0.68. 


Table  9.  Descriptive  statistics  forS.  brevisetosus,  based  on  16  males  from  Texas. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

mts,  in  mm. 

TL 

4.40-4.97 

4.699 

0.187 

0.062 

2.65 

LE 

2.79-3.12 

2.949 

0.188 

0.039 

2.66 

WH 

0.82-0.97 

0.913 

0.051 

0.017 

3.75 

WP 

1.11-1.32 

1.220 

0.064 

0.021 

3.48 

WE 

1.46-1.63 

1.549 

0.068 

0.023 

2.92 

B.  Setae  on  le 

ft  elytron. 

Interval  3 

13-20 

15.6 

Interval  5 

12-18 

14.9 

Interval  7 

10-14 

11.9 

Total 

35-50 

42.4 

5.3 

1.8 

8.35 

C.  Proportion 

IS. 

WF/WH 

0.63-0.67 

0.650 

0.017 

0.006 

1.77 

LP/WP 

0.90-0.95 

0.929 

0.022 

0.007 

1.54 

DP/LP 

0.76-0.80 

0.791 

0.017 

0.006 

1.42 

LP/WE 

0.69-0.76 

0.729 

0.028 

0.009 

2.52 

Ta/Ti 

0.57-0.61 

0.591 

0.021 

0.007 

2.38 

PS/LP 

0.67-0.74 

0.694 

0.029 

0.010 

2.83 

208 


Whitehead 


Variation.  — Available  material  is  too  limited  to  permit  study  of  geographic  variation  in 
this  species.  Males  average  about  0.2  mm  smaller  than  females.  The  largest  (LE,  3.33  mm) 
and  smallest  (LE,  2.83  mm)  females  are  from  Hope,  New  Mexico,  and  Limpia  Canyon, 
Texas,  respectively;  the  largest  (LE,  3.05  mm)  and  smallest  (LE,  2.74  mm)  males  are  both 
from  Sanderson,  Texas. 

Etymology.  — Latin,  brevis  = short,  plusse/osws  = bristly,  in  reference  to  the  short  discal 
elytral  setae. 

Distribution.  — Specimens  of  S.  brevisetosus  have  been  collected  from  eastern  New  Mexi- 
co, east  to  central  Texas,  and  south  to  Coahuila,  from  the  Colorado  River  in  the  north  to  the 
Rio  Grande  drainage  system  in  the  south  (Fig.  146).  I examined  42  specimens  from  the 
following  localities. 

UNITED  STATES 

NEW  MEXICO:  Eddy  Co.,  Hope  (2;  UKSM).  TEXAS  (1;  USNM):  Blanco  Co.,  Cypress  Mills  (1;  USNM);  Brewster  Co.  (1; 
MCZ),  Alpine  (3;  CUNY,  MCZ);  Jeff  Davis  Co.,  Davis  Mountains  (3;  CAS),  Limpia  Canyon  (6;  DRWh,  UASM),  Barrel 
Springs  Creek  (1;  DRWh),  Fort  Davis  (4;  CNC);  Kerr  Co.,  Kerrville  (4;  CNC);  Terrell  Co.,  Lozier  Canyon  (3;  MCZ), 
Sanderson  (6;  CNC);  Uvalde  Co.,  Gamer  State  Park  (2;  LACM). 

MEXICO 

COAHUILA:  La  Gloria,  s.  of  Monclova  (3;  AMNH). 

Collecting  notes.  — Specimens  of  this  species  have  been  collected  from  April  to  August, 
frequently  at  light,  apparently  not  always  near  streams  (e.g.,  Lozier  Canyon  and  La  Gloria). 
I collected  two  specimens  in  typical  gravel  streams  near  Fort  Davis,  Texas,  along  with  speci- 
mens of  S.  scopaeus. 

The  pluripunctatus  group 

Diagnostic  combination.  — The  most  obvious  diagnostic  feature  of  members  of  this  North 
and  Middle  American  group  is  unusual  length  of  discal  setae  on  elytron;  these  setae  general- 
ly are  about  1.2  times  as  long  as  the  maximum  width  of  interval  two,  or  longer.  Specimens 
of  some  South  American  species  also  have  long  setae;  from  these,  members  of  the  pluri- 
punctatus group  are  distinguished  by  the  following  additional  characters  in  combination: 
setae  present  on  intervals  three,  five,  and  seven  of  elytra;  pronotum  without  marginal  cari- 
nae;  mentum  without  accessory  setae;  and  pygidium  of  female  not  crenulate.  Most  included 
forms  are  densely  setose,  with  more  than  ten  setae  on  elytral  interval  three.  Additional 
characters  of  members  of  the  group  are:  form  cylindrical  to  subcylindrical;  paramedian 
clypeal  carinae  extended  to  median  tooth,  sometimes  strongly  raised  in  basal  half;  clypeal 
field  triangular,  its  base  under  1.5  apical  width  of  median  frontal  sulcus;  clypeal  suture 
sharply  impressed;  antennal  articles  five  to  ten  filiform;  anterior  and  middle  tarsi  slightly 
to  strongly  broadened,  especially  in  males;  sternum  seven  with  paramedian  ambulatory  setae 
in  males,  sometimes  in  females;  paramedian  carinae  of  sternum  three  curved  at  apices; 
abdomen  without  extensive  microsculpture;  endophallus  with  distinct  basal  collar  spines. 
The  group  contains  the  only  known  species  in  the  genus  with  more  than  two  pairs  of  lateral 
setae  on  the  pronotum. 

Distribution.  — This  group  includes  several  similar,  allopatric  forms,  in  two  geographic 
subgroups.  One  subgroup  ranges  from  northern  California  south  to  southern  Baja  California. 
The  second  subgroup  ranges  from  southern  Arizona  and  New  Mexico,  south  to  Guatemala, 
and  north  in  the  east  to  Nuevo  Leon  and  Tamaulipas.  I examined  585  specimens  of  the 
pluripunctatus  group. 

Taxonomic  notes.  — The  two  geographic  subgroups  could  be  termed  “superspecies,”  and 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


209 


I relied  in  part  on  statistical  analyses  to  sort  out  taxonomic  relationships  within  them.  The 
Californian  subgroup,  with  two  allopatric  forms  here  treated  as  subspecies,  is  distinguished 
by  more  flattened  body  form  and  short  deflexed  apex  of  male  median  lobe.  The  second 
subgroup,  with  four  allopatric  forms  here  recognized  as  species,  is  distinguished  by  more 
cylindrical  body  form  and  relatively  longer  deflexed  apex  of  male  median  lobe. 

Schizogenius  seticollis  Fall 

Diagnostic  combination.  — Specimens  of  this  species,  one  of  two  species  in  the  genus 
characterized  by  accessory  marginal  setae  on  the  pronotum,  differ  from  those  of  S.  plurise- 
tosus  by  form  of  male  median  lobe.  S.  plurisetosus  is  intermediate  between  the  two  sub- 
species of  S.  seticollis  in  numbers  of  elytral  setae,  but  has  fewer  pronotal  setae  and  normally 
only  one  seta  in  recessed  pits  at  pronotal  hind  angles  rather  than  two  or  three.  The  form  of 
the  median  lobe,  with  deflexed  apical  portion  proportionately  short,  is  diagnostic  of  speci- 
mens of  S.  seticollis  within  the  pluripunctatus  group. 

Schizogenius  seticollis  seticollis  new  combination 

Schizogenius  seticollis  Fall  1901 : 209.  Type  locality  Pomona,  California;  type  in  MCZ;  spec- 
imen labelled  MCZ  type  23859  here  designated  lectotype  (!).  Lindroth  1961:165. 

Diagnostic  combination.  - Specimens  of  this  subspecies  are  distinguished  by  fewer  than 
60  setae  per  elytron,  and  most  specimens  have  fewer  marginal  pronotal  setae. 

Description.  — Body  subcylindrical.  Color  piceous;  mandibles,  mentum,  front  coxae, 
trochanters,  tibiae,  femoral  apices,  and  in  some  specimens  margin  and  apex  of  elytron 
rufopiceous;  antennae,  at  least  at  base,  rufous;  and  labial  palpi,  maxillae,  and  tarsi  testa- 
ceous; no  pronounced  metallic  or  aeneous  luster. 

Integument.  Fine  but  conspicuous  microsculpture  on  mouthparts,  genae,  front  legs  except 
posterior  surfaces  of  femora  and  trochanters,  middle  legs  except  anterior  surfaces  of  tro- 
chanters, hind  tibiae  and  posterior  surfaces  of  hind  femora,  extreme  bases  of  epipleura,  and 
small  areas  in  coxal  depressions  of  sternum  three.  Microsculpture  irregular  and  indistinct  in 
paramedian  frontal  sulci. 

Head.  Fig.  124.  Clypeal  carinae  straight,  extended  to  median  tooth,  strongly  elevated  in 
basal  half;  median  field  triangular,  narrow,  no  wider  at  base  than  apex  of  median  frontal 
sulcus.  Clypeal  suture  sharply  defined.  Eye  prominent,  globose,  finely  and  uniformly  fac- 
eted. Neck  densely,  coarsely  punctate.  Gena  coarsely  punctate,  rugose  in  front.  Antennal 
articles  four  to  ten  elongate,  filiform,  article  five  about  1.6  times  as  long  as  wide. 

Pronotum.  Fig.  124.  Sides  plurisetose,  generally  with  seven  to  1 1 setae  on  each  side,  two 
or  three  setae  in  pits  recessed  from  hind  angles;  hind  angles  obsolete;  base  not  rugose.  Para- 
median sulci  rather  short,  impunctate,  nearly  straight,  deep  throughout,  slightly  hooked 
basally.  Anterior  transverse  impression  finely  punctate. 

Legs.  Front  and  middle  tarsi  dilated  and  pubescent  ventrally  in  both  sexes,  slightly  more 
so  in  males;  hind  tarsus  slender,  rather  short.  Paronychia  conspicuous,  more  than  half  length 
of  tarsal  claws.  Front  tibia  narrowed  evenly  to  base.  Front  femur  not  strongly  constricted 
near  apex. 

Elytra.  Discal  setae  about  1.2  times  as  long  as  maximum  width  of  interval  two.  Among  20 
specimens  examined,  10-17  setae  on  interval  three,  12-19  on  interval  five,  and  1 1-23  on  inter- 
val seven;  most  setae  adjacent  to  corresponding  inner  striae,  but  up  to  25%  irregular,  slightly 
biseriate.  Striae  deep  and  sharply  engraved,  finely  to  indistinctly  punctate  in  basal  half  or 
two-thirds.  Intervals  one  to  seven  broad  and  slightly  convex,  interval  eight  carinate  at  apex; 


210 


Whitehead 


apices  of  intervals  three,  five,  and  seven  broadly  joined.  Humeral  denticles  small,  sharp. 

Abdomen.  Sternum  three  with  paramedian  carinae  straight  or  curved  outward  at  apices. 
Sternum  seven  with  paramedian  ambulatory  setae  in  male  only.  Pygidium  with  apical  margin 
entire  in  both  sexes. 

Male  genitalia.  Median  lobe  (Fig.  129)  elongate,  narrow;  apex  deflected  at  weak  angle, 
relatively  short;  endophallus  with  well  developed  basal  collar  spines  (Fig.  139);  four  speci- 
mens examined. 

Measurements  and  proportions.  See  Table  10. 


Table  10.  Descriptive  statistics  for  .S',  seticollis  seticollis,  based  on  1 1 males  from  Los  Angeles 
County,  California. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measurer 

nents,  in  mm. 

TL 

3.73-4.68 

4.381 

0.404 

0.162 

6.14 

LE 

2.29-2.86 

2.686 

0.238 

0.096 

5.91 

WH 

0.78-0.95 

0.896 

0.080 

0.032 

5.97 

WP 

1.00-1.26 

1.187 

0.111 

0.045 

6.23 

WE 

1.22-1.57 

1.451 

0.136 

0.055 

6.24 

B.  Setae  on  1< 

tft  elytron. 

Interval  3 

12-17 

13.8 

Interval  5 

13-19 

16.2 

Interval  7 

12-23 

17.3 

Total 

40-58 

47.3 

8.6 

3.5 

12.11 

C.  Proportioi 

is. 

WF/WH 

0.60-0.65 

0.625 

0.020 

0.008 

2.18 

LP/WP 

0.87-0.94 

0.919 

0.029 

0.012 

2.09 

DP/LP 

0.81-0.87 

0.838 

0.025 

0.010 

1.98 

LP/WE 

0.72-0.78 

0.752 

0.029 

0.012 

2.58 

Ta/Ti 

0.61-0.74 

0.659 

0.059 

0.024 

5.96 

PS/LP 

0.62-0.68 

0.643 

0.029 

0.012 

3.04 

Variation.  - I found  little  noteworthy  variation  in  this  subspecies.  In  five  selected  samples 
(samples  one  to  five),  I found  little  significant  clinal  variation  in  size  (Fig.  148,  Table  1 1); 
slightly  increased  numbers  of  discal  elytral  setae  to  the  south  (Fig.  149,  Table  12);  slightly 
increased  eye  size  to  the  south  (Fig.  150,  Table  13);  and  slightly  narrowed  pronotum  to  the 
south  (Fig.  151,  Table  14). 

Etymology.  — Latin,  seta  ~ bristly,  plus  collis  = hill,  in  reference  to  the  marginal  setae  of 
the  pronotum. 

Distribution.  — Specimens  of  S.  seticollis  seticollis  have  been  collected  in  central  and 
western  California,  from  Shasta  County  in  the  north  to  San  Diego  County  in  the  south 
(Fig.  147).  I examined  232  specimens  from  the  following  localities. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


211 


Table  1 1.  Variation  in  body  size  (LE,  in  mm)  in  selected  samples  of  the  seticollis  subgroup; 
see  Fig.  148. 


Sample 

N 

Males 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

S',  s.  seticollis. 

1 

16 

8 

2.72-3.19 

2.935 

0.204 

0.068 

4.64 

2 

16 

8 

2.65-3.26 

2.942 

0.272 

0.091 

6.17 

3 

10 

5 

2.63-3.35 

3.095 

0.315 

0.133 

6.97 

4 

18 

9 

2.50-3.18 

2.827 

0.314 

0.099 

7.41 

5 

22 

11 

2.29-2.96 

2.747 

0.231 

0.066 

5.61 

S.  s.  vandykei. 

6 

10 

5 

2.54-3.34 

2.886 

0.406 

0.171 

9.39 

Table  12.  Variatioi 
seticollis  subgroup; 

Sample  N 

a in  numbers  of  discal  setae  on  left  elytron  in  selected  samples  of  the 
see  Fig.  149. 

Males  Range  Mean  1.5SD  2SE  CV(%) 

S.  s.  seticollis. 

1 

16 

8 

35-  49 

39.9 

5.4 

1.8 

9.05 

2 

16 

8 

33-  46 

37.4 

5.9 

2.0 

10.47 

3 

10 

5 

42-  51 

46.3 

4.8 

2.0 

6.98 

4 

18 

9 

33-  53 

41.2 

7.7 

2.4 

12.39 

5 

22 

11 

40-  53 

46.7 

7.6 

2.2 

10.91 

S.  s.  vandykei. 

6 

10 

5 

71-104 

83.5 

16.4 

6.9 

12.85 

Table  13.  Variation  in  eye  size  (WF/WH)  in  selected  samples  of  the  seticollis  subgroup;  see 
Fig.  150. 


Sample 

N 

Males 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

S.  s.  seticollis. 

1 

16 

8 

0.62-0.66 

0.631 

0.017 

0.006 

1.81 

2 

16 

8 

0.62-0.66 

0.639 

0.018 

0.006 

1.88 

3 

10 

5 

0.62-0.65 

0.633 

0.017 

0.007 

1.83 

4 

18 

9 

0.59-0.65 

0.629 

0.023 

0.007 

2.41 

5 

22 

11 

0.60-0.65 

0.626 

0.018 

0.005 

1.89 

S.  s.  vandykei. 

6 

10 

5 

0.61-0.66 

0.640 

0.022 

0.009 

2.33 

212 


Whitehead 


Table  14.  Variation  in  pronotal  form  (LP/WP)  in  selected  samples  of  the  seticoll is  subgroup; 
see  Fig.  151. 


Sample 

N 

Males 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

S.  s.  seticollis. 

1 

16 

8 

0.88-0.94 

0.914 

0.028 

0.009 

2.07 

2 

16 

8 

0.90-0.95 

0.918 

0.023 

0.008 

1 .66 

3 

10 

5 

0.89-0.95 

0.917 

0.029 

0.012 

2.12 

4 

18 

9 

0.88-0.96 

0.924 

0.030 

0.009 

2.17 

5 

22 

11 

0.87-0.94 

0.920 

0.026 

0.007 

1.91 

S.  s.  vandykei. 

6 

10 

5 

0.89-0.98 

0.938 

0.039 

0.016 

2.74 

UNITED  STATES 

CALIFORNIA  (9;  ANSP,  CAS,  USNM):  Colorado  Desert  (1;  MCZ);  Alameda  Co.,  Berkeley  (1;  CUNY),  17.5  mi.  s. 
Livermore  (1;  CAS),  Sunol  Valley  (1;  GRNo);  Calaveras  Co.,  Mokelumne  Hill  (9;  CAS);  Colusa  Co.,  Cooks  Springs  (1; 
CAS);  El  Dorado  Co.,  Latrobe  (2;  CAS);  Fresno  Co.,  Camp  Greeley  (5;  CAS,  CNC,  KSUM),  Le  Ferre  Creek  (2;  CAS), 
Trimmer  (1;  CAS),  3.5  mi.  e.  Trimmer  (8;  DRWh,  TLEr,  UASM),  11.6  mi.  s.  Tollhouse  (3;TLEr);  Glenn  Co.,  Elk  Creek  (1; 
CAS);  Humboldt  Co.,  GarberviUe  (1;  CAS);  Kings  Co.,  Stratford  (1;  CAS);  Lake  Co.  (2;  CAS),  Kelsey  Creek  (2;  CAS), 
Middle  Creek  (2;  CAS);  Los  Angeles  Co.  (6;  CAS,  USNM),  Azuza  (1;  CAS),  mi.  ne.  Claremont  (1;  INHS),  La  Canada 
(1;  CAS),  Los  Angeles  (1;  INHS),  Palmdale  (1;  CAS),  Pasadena  (5;  CAS),  Pomona  (3;  MCZ,  RUNB),  Pomona  Mountains 
(5;  CUNY,  MCZ),  San  Gabriel  Mountains  (2;  TCBa),  Saugus  (1;  CAS);  Tujunga  (4;  MCZ);  Mendocino  Co.  (5;  JNeg),  Dry 
Creek  (13;  CAS);  Monterrey  Co.,  Arroyo  Seco  Camp  (1;  TLEr),  Bradley  (3;  MCZ,  UCB);  Napa  Co.  (1;  CNHM),  Monticello 
(1;  UCD),  Pope  Creek  (2;  CAS),  Rutherford  (3;  TLEr),  Santa  Helena  (2;  MCZ,  CAS);  Riverside  Co.,  Riverside  (3;  CAS, 
USNM),  San  Jacinto  Mountains  (1;  CAS);  San  Benito  Co.,  Pinnacles  National  Monument  (1;  CAS),  San  Benito  River  (2; 
CAS);  San  Bemadino  Co.,  0.9  mi.  ne.  Cedar  Springs  (3;  GRNo,  LBSC),  Lytle  Creek  (4;  MCZ),  Mojave  River  (1;  GRNo), 
San  Bemadino  (1;  CAS),  San  Bemadino  Mountains  (3;  MGFT);  San  Diego  Co.,  San  Diego  (5;  MCZ,  CAS),  Warners  (3; 
UKSM);  San  Joaquin  Co.,  Manteca  (4;  CAS);  San  Luis  Obispo  Co.,  Paso  Robles  (3;  CAS,  UKSM),  Santa  Margarita  (4;  CAS); 
San  Mateo  Co.,  Foster  (1;  CAS);  Santa  Barbara  Co.  (1;  CAS),  Santa  Inez  Mountains  (3;  CAS);  Santa  Clara  Co.,  Gilroy  Hot 
Springs  (10;  DJLa,  TLEr),  Mount  Hamilton  (5;  CAS,  TLEr);  Santa  Cruz  Co.  (1;  CNHM);  Shasta  Co.,  Redding  (15;  CAS), 
10  mi.  ne.  Redding  (1;  DHKa);  Sonoma  Co.  (1;  AMNH),  Duncan  Mills  (2;  CAS),  Healdsburg  (3;  CAS),  2 mi.  e.  Healdsburg 
(1;  CAS),  Russian  River  (8;  CAS,  MCZ),  Santa  Rosa  (2;  MCZ),  2.5  mi.  w.  Skaggs  Springs  (8;  CAS),  Sobre  Vista  (2;  CAS), 
Sylvania  (3;  CAS,  MCZ);  Stanislaus  Co.,  22  ml  w.  Patterson  (4;  CAS);  Tehama  Co.  (2;  CAS);  Tulare  Co.,  Kaweah  (1; 
CAS),  Ventura  Co.,  9 mi.  sw.  Stauffer  (1;  UCB);  Yolo  Co.,  Davis  (2;  UCD),  Rumsey  (1;  UCD). 


Collecting  notes.  — Specimens  have  been  collected  from  March  through  October,  most  of 
them  along  gravel  streams.  Some  were  taken  at  lights,  indirect  evidence  that  adults  can  fly. 
I have  not  collected  specimens  of  this  subspecies. 

Taxonomic  notes.  — See  discussion  for  S.  seticollis  vandykei. 

Schizogenius  seticollis  vandykei  new  subspecies 

Type  material.  — Holotype  male  and  allotype  female  labelled  “MEX.  B.Cal.  3 mi.  NW  of 
Miraflores  19—1-1959”  and  “Canon  San  Bemadino.  Boca  de  la  Sierra”  and  “H.  B.  Leech 
Collector”  (CAS).  Nine  additional  specimens  from  various  localities  in  southern  Baja  Cali- 
fornia are  paratypes  (CAS,  DRWh,  UASM). 

Diagnostic  combination.  — All  known  specimens  of  this  subspecies  have  more  than  60 
discal  setae  per  elytron,  and  thus  differ  from  specimens  of  S.  seticollis  seticollis. 

Description.  — As  in  S.  s.  seticollis  except  as  follows.  Base  of  frons  (Fig.  125)  more 


Classification,  phytogeny,  and  zoogeography  of  Schizogenius 


213 


strongly  calloused.  Pronotum  with  10-14  setae  on  each  side.  Elytron  with  18-36  setae  on 
interval  three,  27-35  on  interval  five,  and  22-37  on  interval  seven;  arrangement  of  setae  on 
intervals  three,  five,  and  seven  strongly  biseriate;  discal  setae  about  1.5  times  as  tong  as  the 
maximum  width  of  interval  two.  Male  genitalia  (Fig.  130)  not  distinctive,  two  specimens 
examined. 

Measurements  and  proportions.  See  Table  15.  Of  holotype:  TL,  4.63  mm;  LE,  2.83  mm; 
WH,  0.92  mm;  WP,  1.18  mm;  WE,  1.48  mm;  WF/WH,  0.61;  LP/WP,  0.98;  DP/LP,  0.82; 
LP/WE,  0.78;  Ta/Ti,  0.66;  PS/LP,  0.65.  Of  allotype:  TL,  4.81  mm;LE,  2.96  mm;  WH,  0.95 
mm;  WP,  1.28  mm;  WE,  1.57  mm;  WF/WH,  0.65;  LP/WP,  0.93;  DP/LP,  0.82;  LP/WE,  0.76; 
Ta/Ti,  0.65;  PS/LP,  0.65. 


Table  15.  Descriptive  statistics  for  S.  seticollis  vandykei,  based  on  six  males  and  five  females 
from  southern  Baja  California. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measurerm 

ents,  in  mm. 

TL 

4.14-5.40 

4.625 

0.666 

0.268 

9.60 

LE 

2.54-3.34 

2.856 

0.414 

0.166 

9.67 

WH 

0.83-1.05 

0.915 

0.117 

0.047 

8.52 

WP 

1.06-1.38 

1.199 

0.179 

0.072 

9.97 

WE 

1.34-1.78 

1.505 

0.208 

0.083 

9.20 

B.  Setae  on  k 

:ft  elytron. 

Interval  3 

18-  36 

24.0 

Interval  5 

27-  35 

30.4 

Interval  7 

22-  37 

30.0 

Total 

71-104 

84.4 

16.3 

6.5 

12.86 

C.  Proportion 

is. 

WF/WH 

0.61-0.66 

0.639 

0.022 

0.009 

2.25 

LP/WP 

0.89-0.98 

0.940 

0.038 

0.015 

2.69 

DP/LP 

0.82-0.90 

0.845 

0.041 

0.017 

3.27 

LP/WE 

0.70-0.79 

0.747 

0.040 

0.016 

3.59 

Ta/Ti 

0.61-0.69 

0.659 

0.035 

0.014 

3.49 

PS/LP 

0.61-0.67 

0.637 

0.028 

0.011 

2.90 

Variation.  - See  Tables  11-14. 

Etymology.  — I dedicate  this  subspecies  to  E.  C.  Van  Dyke,  who  provided  much  of  our 
knowledge  about  the  Coleoptera  of  southwestern  United  States. 

Distribution.  — Specimens  of  this  subspecies  are  known  only  from  a few  localities  in 
southern  Baja  California  (Fig.  147).  I studied  11  specimens  from  the  following  localities. 

MEXICO 

BAJA  CALIFORNIA:  Arroyo  Saltito  near  Las  Cruces  (5;  CAS,  DRWh,  UASM),  20  mi.  n.  Comondu  (1;  CAS),  Hamilton 
Ranch  (1;  CAS),  3 mi.  nw.  Miraflores  (2;  CAS). 


214 


Whitehead 


Collecting  notes.  — Adults  of  S.  seticollis  vandykei  are  probably  active  throughout  the 
year,  as  I have  records  for  January,  July,  and  August.  1 know  nothing  about  their  habits, 
but  suspect  they  may  be  found  along  intermittent  streams.  Aside  from  S.  auripennis,  S. 
falli,  and  S.  pygmaeus,  the  subspecies  is  the  only  member  of  the  genus  known  from  Baja 
California. 

Taxonomic  notes.  - I recognize  S.  seticollis  vandykei  as  a distinct  subspecies  because  it 
clearly  is  closely  related  to  S.  seticollis  seticollis , there  is  no  evidence  of  reproductive  isola- 
tion between  them,  and  all  known  specimens  of  each  subspecies  are  distinguished  by  num- 
bers of  discal  elytral  setae.  I suspect  that  the  ranges  of  the  two  subspecies  are  disjunct  in 
central  and  northern  Baja  California,  or  at  least  they  are  joined  only  by  steeply  stepped 
character  dines. 

As  shown  in  Tables  11-14  and  Fig.  148-151,  sample  six,  S.  seticollis  vandykei  does  not 
differ  significantly  in  body  size  from  samples  one  to  five,  S.  seticollis  seticollis ; it  is  more 
different  from  southern  than  northern  samples  of  S.  s.  seticollis  in  eye  size,  but  not  signifi- 
cantly different  from  any  of  them;  and  the  pronotum  is  narrower  than  in  any  of  the  S.  s. 
seticollis  but  not  statistically  significantly  different  from  the  southernmost  samples.  In  the 
number  of  discal  elytral  setae,  however,  despite  a slightly  greater  similarity  of  S.  s.  vandykei 
to  southern  rather  than  northern  samples  of  S.  s.  seticollis,  the  differences  are  large,  indeed 
are  taxonomically  significant  at  1.5  standard  deviations  from  means.  Numbers  of  discal 
elytral  setae  are,  in  general,  directly  correlated  with  body  size.  From  Tables  11  and  12, 
ratios  formed  of  the  mean  values  for  number  of  setae  to  elytral  length  are,  for  samples  one 
to  six,  respectively:  13.6,  12.7,  15.0,  14.6,  17.0,  and  29.6  setae  per  mm.  A definite  increase 
from  north  to  south  is  evident  from  samples  one  to  five,  but  a large  hiatus  still  exists  be- 
tween samples  five  and  six.  I conclude  from  these  observations  that  reproductive  isolation 
between  the  two  subspecies  is  indeterminate,  but  that  they  are  at  least  isolated  geographical- 
ly. Collections  from  central  and  northern  Baja  California  are  needed  to  better  define  geo- 
graphic and  reproductive  relationships. 

Schizogenius  plurisetosus  new  species 

Type  material.  — Holotype  male  and  allotype  female  labelled  “MEX.  Tamaulipas  Rio 
Purificacion  nr.  El  Barretal  Rte.  85  800'  X.  19.  65”  and  “George  E.  Ball  D.  R.  Whitehead 
collectors”  (MCZ).  An  additional  15  specimens  from  various  localities  in  Nuevo  Leon  and 
Tamaulipas  are  paratypes  (BMNH,  CAS,  CNC,  DRWh,  IRSB,  UASM,  USNM). 

Diagnostic  combination.  — Specimens  of  this  species  are  readily  distinguished  from  speci- 
mens of  S.  seticollis  by  having  fewer  accessory  marginal  pronotal  setae,  and  only  one  seta 
in  pit  near  hind  angle.  Also:  apical  portion  of  male  median  lobe  proportionately  longer  and 
more  sharply  deflexed;  front  and  middle  tarsi  less  expanded;  and  range,  in  northeastern 
Mexico,  sharply  disjunct. 

Description.  — As  in  S.  seticollis  seticollis,  except  as  follows.  Legs  paler,  femora  uni- 
formly rufopiceous.  Form  and  sculpture  of  head  and  pronotum,  Fig.  126;  genae  less 
coarsely  sculptured;  pronotum  with  five  to  seven  setae  on  each  side,  usually  with  only  one 
in  a pit  recessed  from  the  hind  angle;  basal  impression  distinctly  punctate;  anterior  trans- 
verse impression  nearly  impunctate.  Elytron  with  18-24  setae  on  interval  three,  17-30 
on  interval  five,  and  17-22  on  interval  seven;  setae  moderately  biseriate;  discal  setae  about 
1.5  times  as  long  as  the  maximum  width  of  interval  two;  striae  indistinctly  punctate. 
Front  and  middle  tarsi  much  narrower.  Male  genitalia  with  apex  of  median  lobe  (Fig.  131, 
slightly  distorted)  sharply  deflexed,  proportionately  long;  endophallus,  Fig.  140;  four  speci- 
mens examined. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


215 


Measurements  and  proportions.  See  Table  16.  Of  holotype:  TL,  3.98  mm;  LE,  2.43  mm; 
WH,  0.80  mm;  WP,  1.06  mm;  WE,  1.24  mm;  WF/WH,  0.61;  LP/WP,  0.93;  DP/LP,  0.81; 
LP/WE,  0.80;  Ta/Ti,  0.60;  PS/LP,  0.63.  Of  allotype:  TL,  4.25  mm;  LE,  2.65  mm;  WH,  0.83 
mm;  WP,  1.11  mm;  WE,  1.35  mm;  WF/WH,  0.62;  LP/WP,  0.92;  DP/LP,  0.83;  LP/WE,  0.76; 
Ta/Ti,  0.66;  PS/LP,  0.66. 


Table  16.  Descriptive  statistics  for  S.  plurisetosus,  based  on  seven  males  from  Nuevo  Leon 
and  Tamaulipas. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measurem 

ents,  in  mm. 

TL 

3.78-4.69 

4.234 

- 

- 

- 

LE 

2.33-2.92 

2.546 

- 

- 

- 

WH 

0.76-0.91 

0.824 

- 

- 

- 

WP 

0.98-1.25 

1.086 

- 

- 

- 

WE 

1.18-1.48 

1.293 

- 

- 

- 

B.  Setae  on  It 

jft  elytron. 

Interval  3 

18-24 

21.0 

Interval  5 

17-30 

21.7 

Interval  7 

17-23 

20.7 

Total 

52-74 

63.4 

C.  Proportior 

is. 

WF/WH 

0.60-0.64 

0.613 

- 

- 

- 

LP/WP 

0.92-0.95 

0.936 

- 

- 

- 

DP/LP 

0.80-0.82 

0.810 

- 

- 

- 

LP/WE 

0.77-0.80 

0.784 

- 

- 

- 

Ta/Ti 

0.59-0.69 

0.633 

- 

- 

- 

PS/LP 

0.63-0.70 

0.661 

- 

- 

- 

Variation.  - See  Table  16.  Two  specimens  have  two  setae  in  a recessed  pit  on  one  side  of 
the  pronotum;  all  others  have  only  one  seta  per  pit. 

Etymology.  — Latin,  pluralis  = more  than  one,  plus  setosus  = bristly,  in  reference  to  the 
large  numbers  of  setae  which  characterize  members  of  this  species. 

Distribution.  — Specimens  of  this  species  have  been  collected  only  in  Nuevo  Leon  and 
Tamaulipas,  from  the  Rio  Salinas,  a tributary  of  the  Rio  Grande,  south  to  the  Rio  Corona, 
a tributary  of  the  Rio  Soto  la  Marina  (Fig.  147).  The  species  is  not  known  to  range  into  the 
United  States,  but  may  do  so.  I studied  17  specimens,  all  collected  by  G.  E.  Ball  or  me 
during  1964  to  1966,  from  the  following  localities. 

MEXICO 

NUEVO  LEON:  Cienega  de  Flores  (5;  UASM),  Linares  (2;  DRWh).  TAMAULIPAS:  El  Barretal  (6;  CAS,  CNC,  MCZ). 
15.2  mi.  n.  Ciudad  Victoria  (1;  BMNH),  21.3  mi.  n.  Ciudad  Victoria  (3;  IRSB;  USNM). 


216 


Whitehead 


Collecting  notes.  - Specimens  of  S.  plurisetosus  were  collected  in  riparian  gravel  bars,  in 
July,  September,  and  October,  at  elevations  ranging  from  500  to  1200'.  At  all  stations  there 
were  fewer  specimens  of  this  species  than  of  others  in  the  genus. 

Taxonomic  notes.  — Despite  accessory  pronotal  setae,  this  species  is  not  closely  related 
to  S.  seticollis,  as  judged  from  form  of  male  genitalia.  It  is  most  closely  related  to  the 
geographically  proximate  S.  multisetosus,  with  similar  male  genitalia.  Even  so,  the  form  of 
the  apex  is  slightly  but  apparently  constantly  more  broadly  rounded.  This  detail,  together 
with  the  apparently  constant  accessory  pronotal  setae,  indicates  that  S.  plurisetosus  and 
S.  multisetosus  are  reproductively  isolated.  Statistical  comparisons  between  them  confirm 
this  distinction.  Thus,  as  shown  in  Tables  18-23  and  Fig.  152-157,  samples  one  to  four,  there 
is  no  important  difference  in  body  size,  eye  size,  or  relative  pronotal  size,  but  numbers  of 
discal  elytral  setae  are  much  greater  in  sample  one,  S.  plurisetosus,  than  in  any  sample  (see 
Fig.  153)  of  S.  multisetosus.  Also,  the  northernmost  sample  of  S.  multisetosus  differs  by 
having  statistically  significantly  broader  pronota  and  statistically  significantly  longer  para- 
median pronotal  sulci. 


Schizogenius  multisetosus  Bates 

Schizogenius  multisetosus  Bates  1891:233.  Type  locality  Huitzo,  Oaxaca;  type  in  BMNH, 

male  specimen  labelled  holotype  here  designated  lectotype  (!). 

Diagnostic  combination.  — The  combination  of  two  pairs  of  marginal  setae  on  the  prono- 
tum,  sharply  deflexed  apical  portion  of  the  median  lobe,  and  lack  of  paramedian  ambulato- 
ry setae  in  females,  distinguishes  this  species  from  other  members  of  the  pluripunctatus 
group.  Also,  numbers  of  setae  on  the  elytral  disc,  in  specimens  examined,  range  from  37  to 
56. 

Description.  - As  in  S.  seticollis  seticollis  except:  legs  paler,  femora  uniformly  rufous  to 
rufopiceous,  tibiae  paler;  abdomen  often  paler  toward  apex.  Form  and  sculpture  of  head 
and  pronotum  as  in  S.  plurisetosus,  Fig.  126,  except  sides  of  pronotum  bisetose  and  basal 
transverse  impression  less  distinctly  punctate.  In  specimens  studied,  elytron  with  12-18 
setae  on  interval  three,  12-21  on  interval  five,  and  12-20  on  interval  seven;  setae  mostly 
adjacent  to  corresponding  inner  striae,  not  markedly  biseriate.  Male  genitalia  with  median 
lobe  (Fig.  132-134)  with  apical  portion  sharply  deflexed,  proportionately  long;  endophallus, 
Fig.  141-142;  24  specimens  examined. 

Measurements  and  proportions.  See  Table  17. 

Variation.  — Samples  of  sufficient  size  for  statistical  analysis  were  available  from  only 
three  localities.  Data  on  variation  in  six  characteristics  in  these  samples,  numbered  two  to 
four  as  in  Fig.  152-157,  are  given  in  Tables  18-23.  All  three  samples  differ  significantly 
from  one  another  in  body  size  (Fig.  152,  Table  18).  Other  statistically  significant  differences 
are:  sample  two,  shorter  paramedian  pronotal  sulci  (Fig.  157,  Table  23);  sample  three, 
proportionately  smaller  thorax  (Fig.  156,  Table  22);  and  sample  four,  broader  pronotum 
(Fig.  155,  Table  21).  These  samples  therefore  represent  populations  which  are  equally  well 
isolated  from  one  another  geographically,  so  no  analysis  of  clinal  relationships  is  feasible. 
The  median  lobe,  especially  at  base,  varies  in  form;  most  specimens  from  the  Rio  Moctezuma 
at  300'  have  a reduced  basal  lobe,  and  those  from  the  Rio  Balsas  usually  have  a less  sharply 
angulate  apical  deflection.  A male  from  Hidalgo,  collected  at  5300'  in  the  Rio  Moctezuma 
drainage  system,  is  in  most  ways  similar  to  the  Rio  Balsas  specimens  but  has  the  large  thorax 
characteristic  of  Tamazunchale  specimens;  it  may  represent  a truly  intermediate  population. 

Etymology.  - Latin,  multus  = most,  plus  setosus  = setose,  in  reference  to  the  numerous 
discal  setae  of  the  elytra. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


217 


Table  17.  Descriptive  statistics  for  S.  multisetosus,  based  on  14  males  from  72.5  miles  south 
of  Valle  Nacional,  Oaxaca. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measurem 

ents,  in  mm. 

TL 

3.84-4.44 

4.093 

0.315 

0.112 

5.13 

LE 

2.34-2.78 

2.546 

0.201 

0.072 

5.27 

WH 

0.75-0.90 

0.816 

0.068 

0.024 

5.51 

WP 

0.95-1.15 

1.049 

0.096 

0.034 

6.07 

WE 

1.20-1.45 

1.304 

0.112 

0.040 

5.72 

B.  Setae  on  le 

ift  elytron. 

Interval  3 

12-17 

14.5 

Interval  5 

15-20 

17.1 

Interval  7 

12-18 

15.9 

Total 

40-54 

47.5 

5.9 

2.1 

8.27 

C.  Proportior 

LS. 

WF/WH 

0.60-0.64 

0.621 

0.020 

0.007 

2.14 

LP/WP 

0.93-0.97 

0.944 

0.020 

0.007 

1.42 

DP/LP 

0.81-0.87 

0.841 

0.027 

0.010 

2.15 

LP/WE 

0.74-0.78 

0.761 

0.024 

0.009 

2.12 

Ta/Ti 

0.62-0.68 

0.649 

0.026 

0.009 

2.67 

PS/LP 

0.62-0.69 

0.654 

0.030 

0.011 

3.09 

Distribution.  — Specimens  of  S.  multisetosus  have  been  collected  at  altitudes  ranging  from 
near  sea  level  in  the  north  to  at  least  5300'.  They  are  from  four  river  systems  in  central 
Mexico  (Fig.  147):  the  Rios  Moctezuma  and  Papaloapan  on  the  Atlantic  slopes,  and  the 
Rios  Balsas  and  Atoyac  on  the  Pacific.  I studied  73  specimens  from  the  following  localities. 

MEXICO 

GUERRERO:  5 mi.  s.  Iguala  (1;CAS).  HIDALGO:  Rio  Tula  near  Tasquillo  5300'(1;  UASM).  MORELOS:  Tetecala  2800' 
(6;  CNC,  MCZ,  UASM),  Yautepec  (1;  BMNH).  OAXACA:  Huitzo  (2;  BMNH),  72.5  mi.  s.  Valle  Nacional  4100'  (25;  UASM). 
PUEBLA:  Tehuitzingo  3700)'  (1;  MGFT),  Tepexco  (22;  UASM).  SAN  LUIS  POTOSL  Tamazunchale  300'  (14;  CAS, 
DRWh,  IRSB,  UASM,  USNM). 

-Collecting  notes.  — Specimens  of  S.  multisetosus  were  collected  during  June,  August, 
September,  and  October.  Some,  including  the  entire  Tepexco  series,  were  collected  at  black 
lights,  having  doubtless  flown  there.  Others  were  hand  collected  in  riparian  gravel  bars,  the 
normal  habitat  for  most  species  in  the  subgenus,  but  on  the  average  in  slightly  drier  places. 
Some  inhabited  streams,  at  least  in  Puebla,  are  dry  during  winter. 

Taxonomic  notes.  — S.  multisetosus , so  far  as  known,  is  allopatric  with  respect  to  all 
other  members  of  the  group.  It  does  share  at  least  one  drainage  system,  the  Rio  Papaloapan, 
with  S.  kulti,  and  may  be  sympatric  with  it.  Lack  of  known  sympatry  and  close  similarity 
of  members  of  the  pluripunctatus  subgroup  provide  no  direct  evidence  of  reproductive 
isolation  between  them,  but  statistical  data  provide  indirect  evidence  that  reproductive  iso- 
lation exists.  Selected  samples  of  S.  plurisetosus,  S.  multisetosus,  S.  pluripunctatus,  and 


218 


Whitehead 


Table  18.  Variation  in  body  size  (LE,  in  mm)  in  selected  samples  of  the  pluripunctatus 
subgroup;  see  Fig.  152. 


Sample 

N 

Males 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

S.  plurisetosus. 

1 

14 

7 

2.33-2.93 

2.625 

0.320 

0.1 14 

8.14 

S.  multisetosus. 

2 

14 

7 

2.50-3.10 

2.734 

0.251 

0.089 

6.12 

3 

22 

11 

2.33-2.90 

2.575 

0.227 

0.064 

5.87 

4 

20 

10 

2.07-2.60 

2.342 

0.232 

0.069 

6.62 

S.  pluripunctatus. 

5 

18 

9 

2.05-2.70 

2.402 

0.251 

0.079 

6.95 

6 

10 

5 

2.15-2.65 

2.382 

0.228 

0.096 

6.39 

7 

18 

9 

2.09-2.68 

2.407 

0.218 

0.069 

6.04 

8 

8 

5 

1.98-2.75 

2.372 

- 

- 

- 

9 

10 

5 

2.15-2.71 

2.445 

0.237 

0.100 

6.47 

S.  kulti. 

10 

10 

5 

2.00-2.55 

2.278 

0.263 

0.111 

7.70 

11 

20 

10 

1.96-2.65 

2.234 

0.237 

0.071 

7.07 

12 

22 

11 

1.75-2.25 

2.050 

0.205 

0.058 

6.65 

13 

12 

6 

2.07-2.44 

2.283 

0.183 

0.070 

5.34 

14 

10 

8 

2.16-2.60 

2.299 

- 

- 

- 

Table  19.  Variation  in  numbers  of  discal  setae  on  left  elytron  in  selected  samples  of  the 
pluripunctatus  subgroup;  see  Fig.  153. 


Sample 

N 

Males 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

S.  plurisetosus. 

1 

14 

7 

52-74 

64.4 

7.4 

2.6 

7.62 

S.  multisetosus. 

2 

14 

7 

42-56 

48.8 

6.0 

2.1 

8.16 

3 

22 

1 1 

42-55 

47.4 

5.6 

1.6 

7.91 

4 

20 

10 

37-54 

47.0 

7.9 

2.4 

11.28 

S.  pluripunctatus. 

5 

18 

9 

39-56 

47.2 

7.4 

2.3 

10.50 

6 

10 

5 

41-57 

47.1 

8.1 

3.4 

11.40 

7 

18 

9 

39-55 

45.3 

7.1 

2.2 

10.51 

8 

8 

5 

35-55 

43.0 

- 

- 

- 

9 

10 

5 

45-61 

52.3 

8.0 

3.4 

10.20 

S.  kulti. 

10 

10 

5 

28-41 

34.6 

6.0 

2.6 

11.66 

11 

20 

10 

18-28 

22.6 

3.6 

1.1 

10.55 

12 

22 

11 

19-26 

21.8 

3.8 

1.1 

11.72 

13 

12 

6 

21-27 

22.9 

3.0 

1.1 

8.62 

14 

10 

8 

23-31 

26.3 

- 

- 

- 

Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


219 


Table  20.  Variation  in  eye  size  (WF/WH)  in  selected  samples  of  the  pluripunctatus  sub- 
group; see  Fig.  154. 


Sample 

N 

Males 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

S.  plurisetosus. 

1 14 

S.  multisetosus. 

7 

0.70-0.66 

0.622 

0.025 

0.009 

2.68 

2 

14 

7 

0.59-0.65 

0.620 

0.027 

0.010 

2.89 

3 

22 

11 

0.60-0.66 

0.625 

0.021 

0.006 

2.25 

4 

20 

10 

0.60-0.64 

0.620 

0.015 

0.004 

1.61 

S.  pluripunctatus. 
5 18 

9 

0.61-0.66 

0.631 

0.020 

0.006 

2.14 

6 

10 

5 

0.59-0.66 

0.627 

0.030 

0.013 

3.19 

7 

18 

9 

0.61-0.66 

0.633 

0.022 

0.007 

2.27 

8 

8 

5 

0.61-0.67 

0.642 

- 

- 

- 

9 

10 

5 

0.60-0.63 

0.614 

0.013 

0.005 

1.37 

S.  kulti. 

10 

10 

5 

0.59-0.63 

0.606 

0.019 

0.008 

2.08 

11 

20 

10 

0.58-0.63 

0.604 

0.020 

0.006 

2.24 

12 

22 

11 

0.60-0.65 

0.617 

0.020 

0.006 

2.20 

13 

12 

6 

0.58-0.61 

0.595 

0.013 

0.005 

1.52 

14 

10 

8 

0.58-0.63 

0.605 

- 

- 

- 

Table  21.  Variation  in  pronotal  form  (LP/WP)  in  selected  samples  of  the  pluripunctatus 
subgroup;  see  Fig.  155. 


Sample 

N 

Males 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

S.  plurisetosus. 

1 

14 

7 

0.90-0.95 

0.932 

0.019 

0.007 

1.34 

S.  multisetosus. 

2 

14 

7 

0.93-1.00 

0.959 

0.032 

0.012 

2.25 

3 

22 

1 1 

0.93-0.98 

0.950 

0.021 

0.006 

1.51 

4 

20 

10 

0.92-0.95 

0.936 

0.013 

0.004 

0.93 

S.  pluripunctatus. 

- 

5 

18 

9 

0.88-0.93 

0.907 

0.017 

0.005 

1.25 

6 

10 

5 

0.91-0.94 

0.919 

0.016 

0.007 

1.19 

7 

18 

9 

0.87-0.93 

0.910 

0.022 

0.007 

1.64 

8 

8 

5 

0.90-0.93 

0.915 

- 

- 

- 

9 

10 

5 

0.90-0.96 

0.931 

0.027 

0.011 

1.92 

S.  kulti. 

10 

10 

5 

0.90-0.93 

0.916 

0.018 

0.007 

1.28 

11 

20 

10 

0.89-0.95 

0.925 

0.024 

0.007 

1.76 

12 

22 

11 

0.90-0.97 

0.932 

0.025 

0.007 

1.80 

13 

12 

6 

0.90-0.95 

0.925 

0.021 

0.008 

1.49 

14 

10 

8 

0.88-0.94 

0.913 

- 

- 

- 

220 


Whitehead 


Table  22.  Variation  in  pronotal  length  relative  to  elytral  width  (LP/WE)  in  selected  samples 
of  the  pluripunctatus  subgroup;  see  Fig.  156. 


Sample 

N 

Males 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

S.  plurisetosus. 

1 

14 

7 

0.74-0.80 

0.775 

0.027 

0.010 

2.30 

S.  multisetosus. 

2 

14 

7 

0.73-0.82 

0.789 

0.050 

0.018 

4.19 

3 

22 

11 

0.73-0.78 

0.752 

0.027 

0.008 

2.42 

4 

20 

10 

0.74-0.80 

0.776 

0.022 

0.007 

1.88 

S.  pluripunctatus. 

5 

18 

9 

0.75-0.83 

0.790 

0.028 

0.009 

2.34 

6 

10 

5 

0.74-0.81 

0.781 

0.039 

0.016 

3.33 

7 

18 

9 

0.71-0.80 

0.764 

0.034 

0.011 

2.95 

8 

8 

5 

0.76-0.81 

0.785 

- 

- 

- 

9 

10 

5 

0.74-0.79 

0.779 

0.036 

0.015 

3.11 

S.  kulti. 

10 

10 

5 

0.74-0.81 

0.784 

0.035 

0.015 

3.01 

11 

20 

10 

0.75-0.83 

0.784 

0.034 

0.010 

2.87 

12 

22 

11 

0.75-0.84 

0.780 

0.035 

0.010 

3.00 

13 

12 

6 

0.77-0.81 

0.792 

0.021 

0.008 

1.79 

14 

10 

8 

0.75-0.82 

0.786 

- 

- 

- 

Table  23.  Variation  in  length  of  paramedian  pronotal  sulcus  relative  to  pronotal  length 
(PS/LP)  in  selected  samples  of  the  pluripunctatus  subgroup;  see  Fig.  157. 


Sample 

N 

Males 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

S.  plurisetosus. 

1 14 

S.  multisetosus. 

7 

0.63-0.70 

0.664 

0.026 

0.009 

2.61 

2 

14 

7 

0.58-0.65 

0.629 

0.028 

0.010 

3.02 

3 

22 

11 

0.62-0.69 

0.658 

0.025 

0.007 

2.51 

4 

20 

10 

0.63-0.70 

0.656 

0.032 

0.009 

3.21 

S.  pluripunctatus. 
5 18 

9 

0.61-0.68 

0.647 

0.030 

0.009 

3.09 

6 

10 

5 

0.63-0.73 

0.663 

0.042 

0.018 

4.27 

7 

18 

9 

0.63-0.71 

0.653 

0.030 

0.010 

3.09 

8 

8 

5 

0.63-0.68 

0.660 

- 

- 

- 

9 

10 

5 

0.63-0.70 

0.673 

0.032 

0.013 

3.14 

S.  kulti. 

10 

10 

5 

0.59-0.68 

0.656 

0.039 

0.017 

4.01 

11 

20 

10 

0.61-0.68 

0.646 

0.029 

0.009 

3.02 

12 

22 

11 

0.59-0.66 

0.628 

0.028 

0.008 

2.93 

13 

12 

6 

0.63-0.70 

0.658 

0.034 

0.013 

3.42 

14 

10 

8 

0.58-0.66 

0.628 

- 

- 

- 

Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


221 


S.  kulti  are  compared  for  six  different  characteristics  in  Tables  18-23  and  Fig.  152-157. 
Satisfactory  comparisons  of  S.  multisetosus  with  the  other  species  are  made  difficult  by 
lack  of  good  information  on  clinal  variation.  Still,  certain  facts  support  the  notion  that  S. 
multisetosus  is  reproductively  isolated  from  the  other  three  forms. 

S.  plurisetosus  is  known  from  rivers  just  north  of  the  Moctezuma  drainage  basin,  and  a 
direct  comparison  between  those  specimens  (sample  one)  and  those  of  S.  multisetosus  from 
Tamazunchale  (sample  two)  is  therefore  relevant.  The  apex  of  the  median  lobe  of  the  male 
genitalia  is  more  broadly  rounded;  accessory  pronotal  setae  are  present  in  all  specimens; 
and  the  number  of  discal  elytral  setae  (Fig.  153,  Table  19)  is  much  greater.  Additional, 
statistically  significant,  differences  are  in  the  form  of  the  pronotum  (Fig.  155,  Table  21) 
and  in  the  length  of  the  paramedian  pronotal  sulci  (Fig.  156,  Table  23).  In  these  two  fea- 
tures, the  S.  plurisetosus  sample  is  similar  to  one  or  both  of  the  two  southern  S.  multiseto- 
sus samples  (samples  three  and  four);  the  Tamazunchale  sample  may  thus  be  considered 
divergent,  indicating  a lack  of  gene  flow  between  it  and  S.  plurisetosus.  Differences  in  these 
five  characteristics,  and  the  geographic  proximity  of  the  two  samples,  indicate  reproductive 
isolation  of  S.  multisetosus  from  S.  plurisetosus. 

Known  geographic  ranges  of  S.  pluripunctatus  (samples  five  to  nine)  and  S.  multisetosus 
are  so  widely  separated  that  statistical  differences  are  equivocal.  However,  constant  differ- 
ences in  form  of  male  genitalia,  constant  presence  of  paramedian  ambulatory  setae  on  ster- 
num seven  in  females  of  S.  pluripunctatus,  and  apparent  geographic  isolation  are  sufficient 
reasons  to  recognize  S.  pluripunctatus  and  S.  multisetosus  as  separate  species. 

S.  kulti  (samples  10  to  14),  particularly  in  the  south,  is  characterized  by  fewer  elytral 
setae  and  smaller  body  size  than  is  S.  multisetosus.  Sample  13  of  S.  kulti,  Valle  Nacional 
and  sample  three  of  S.  multisetosus,  72.5  mi.  s.  Valle  Nacional,  are  from  the  same  river 
system.  Statistically  significant  differences  in  eye  size,  pronotal  form,  and  relative  size  of 
thorax  represent  divergences  in  these  features,  so  there  is  little  doubt  that  the  two  samples 
represent  reproductively  isolated  taxa.  Similar,  though  less  conclusive,  comparisons  between 
other  population  samples  of  the  two  forms  give  similar  results,  even  though  northern  sam- 
ples of  S.  kulti  (samples  10  and  14)  are  characterized  by  large  body  size  and  increased 
numbers  of  elytral  setae.  These  facts,  together  with  constant  differences  in  form  of  male 
median  lobe,  strongly  support  the  conclusion  that  S.  kulti  is  reproductively  isolated  from 
S.  multisetosus. 


Schizogenius  pluripunctatus  LeConte 

Schizogenius  pluripunctatus  LeConte  1852:197.  Type  locality  Colorado  River,  California; 
type  in  MCZ,  specimen  labelled  MCZ  5484  here  designated  lectotype  (!).  LeConte  1857: 
82.  Putzeys  1863:24.  Putzeys  1866:225.  LeConte  1879:34.  Lindroth  1961:165. 
Schizogenius  simplex  LeConte  1852:197.  Type  locality  Colorado  River,  California;  type  in 
MCZ,  specimen  labelled  MCZ  5485  here  designated  lectotype  (!).  LeConte  1857:83, 
established  synonymy. 

Diagnostic  combination.  — Within  the  pluripunctatus  group,  specimens  of  this  species  are 
distinguished  by  the  following  combination  of  characters:  pronotum  without  accessory 
marginal  setae;  females  as  well  as  males  each  with  paramedian  ambulatory  setae  on  sternum 
seven;  apex  of  median  lobe  deflected  at  a weak  angle;  and  elytron  with  35  or  more  discal 
setae. 

Description.  — As  in  S.  seticollis  seticollis,  except:  legs  paler,  femora  uniformly  rufous, 
tibiae  paler;  abdomen  often  paler  toward  apex.  Form  and  sculpture  of  head  and  pronotum 
as  in  S.  plurisetosus  (Fig.  126),  except  sides  of  pronotum  bisetose  and  basal  transverse 


Whitehead 


impression  less  distinctly  punctate.  In  specimens  studied,  elytron  with  12-18  setae  on  in- 
terval three,  12-23  on  interval  five,  and  11-19  on  interval  seven;  setae  mostly  adjacent  to 
corresponding  inner  striae,  not  markedly  biseriate;  total  number  of  setae  per  elytron  35-61; 
striae  usually  indistinctly  punctate.  Females  and  males  each  with  a pair  of  ambulatory 
setae  on  sternum  seven.  Male  genitalia:  median  lobe  (Fig.  135)  with  apical  portion  pro- 
portionately long,  deflected  at  weak  angle;  endophallus,  Fig.  143;  23  specimens  examined. 

Measurements  and  proportions.  See  Table  24. 


Table  24.  Descriptive  statistics  for  S.  pluripunctatus,  based  on  nine  males  from  18.6  miles 
southeast  of  Tonala,  Chiapas. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

nts,  in  mm. 

TL 

3.38-4.26 

3.819 

0.418 

0.186 

7.29 

LE 

2.05-2.60 

2.334 

0.260 

0.116 

7.43 

WH 

0.72-0.89 

0.800 

0.090 

0.040 

7.52 

WP 

0.93-1.19 

1.063 

0.127 

0.057 

7.98 

WE 

1.06-1.33 

1.206 

0.147 

0.065 

8.13 

B.  Setae  on  le 

ft  elytron. 

Interval  3 

13-17 

14.2 

Interval  5 

12-20 

16.1 

Interval  7 

13-19 

14.8 

Total 

39-56 

45.1 

5.5 

2.4 

8.10 

C.  Proportion 

s. 

WF/WH 

0.61-0.65 

0.630 

0.021 

0.009 

2.24 

LP/WP 

0.88-0.91 

0.903 

0.015 

0.007 

1.11 

DP/LP 

0.81-0.85 

0.830 

0.018 

0.008 

1.47 

LP/WE 

0.77-0.83 

0.797 

0.026 

0.012 

2.17 

Ta/Ti 

0.62-0.72 

0.664 

0.054 

0.024 

5.38 

PS/LP 

0.62-0.68 

0.650 

0.031 

0.014 

3.17 

Variation.  — Data  on  variation  in  six  characteristics  among  five  samples  (samples  five  to 
nine)  are  given  in  Tables  18-23  and  Fig.  152-157.  Three  samples  from  Arizona  and  New 
Mexico  (samples  five  to  seven)  are  quite  uniform,  and  are  intermediate  in  most  ways  to  the 
two  Mexican  samples.  I therefore  suspect  that  sample  nine  from  Sinaloa  and  Nayarit  is 
geographically  isolated  from  sample  eight  from  Durango,  and  that  these  two  samples  rep- 
resent populations  near  the  ends  of  a partial  circle  of  races.  This  is  a reasonable  hypothe- 
sis, since  these  areas  are  well  separated  by  high  ranges  of  the  Sierra  Madre  Occidental,  but 
should  be  tested  through  additional  collections. 

One  female  specimen  from  Prescott,  Arizona,  has  only  one  left  paramedian  ambulatory 
seta  on  sternum  seven.  I otherwise  found  this  character  to  be  constant,  and  presume  that  it 
remains  stable  southward  where  the  range  of  S.  pluripunctatus  approaches  that  of  S',  kulti. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


223 


Etymology.  - Latin,  plurimus  = most,  plus  punctum  = small  holes,  in  reference  to  the 
numerous  discal  setae  on  the  elytron. 

Distribution.  - Specimens  of  S.  pluripunctatus  have  been  collected  in  the  Gila  and  Colo- 
rado River  drainages  in  Arizona  and  New  Mexico,  south  to  the  Rio  Acaponeta  drainage 
basin  in  northern  Nayarit,  and  in  the  interior  Rio  Nazas  drainage  in  Durango  (Fig.  147). 
One  specimen  labelled  “N.Y.”  (UKSM)  is  no  doubt  erroneous.  I studied  130  additional 
specimens  from  the  following  localities. 

UNITED  STATES 

ARIZONA  (14;  AMNH,  CAS,  UASM,  USNM):  Cochise  Co.,  Cochise  Stronghold  (1;  UATA),  Fairbanks  (1;  CAS);  Coconino 
Co.,  Bill  Williams  Fork  (9;  KSUM,  MCZ,  UKSM);  Gila  Co.,  Carrizo  Creek  (12;  UASM),  Globe  (1;  CAS),  Roosevelt 
Lake  (1;  AMNH),  Salt  River  Canyon  (3;  ANSP,  MCZ,  UASM),  Verde  River  (1;  MCZ);  Graham  Co.,  Aravaipa  (12; 
BMNH,  CAS,  CNC,  DRWh,  IRSB,  KHSt),  San  Carlos  Reservoir  (1;  MCZ);  Maricopa  Co.,  Phoenix  (2;  LACM);  Pima  Co., 
CoUosal  Cave  Park  (2;  KHSt),  Madera  Canyon  (1;  UASM),  Tucson  (6;  AMNH,  CAS,  JNeg,  KHSt,  UATA);  Santa  Cruz  Co., 
Nogales  (3;  CNHM,  UASM),  5 mi.  n.  Nogales  (1;  MCZ),  Patagonia  (8;  AMNH,  CAS,  UATA),  2 mi.  sw.  Patagonia  (5; 
UATA),  Pena  Blanca  (1;  UASM);  Yavapai  Co.,  Bumble  Bee  (2;  CAS),  Haslampa  District  (2;  CAS),  Prescott  (2;  UASM). 
NEW  MEXICO:  Catron  Co.,  Glenwood  (1;  UASM);  Grant  Co.,  Cliff  (8;  UASM),  26  mi.  n.  Silver  City  (9;  TLEr),  36.4  mi. 
ne.  Silver  City  (1;  MGFT). 


MEXICO 

DURANGO:  12.2  mi.  s.  El  Banco  (8;  UASM).  NAYARIT:  Acaponeta  (1;  UASM),  2.4  mi.  s.  Acaponeta  (6;  UASM),  8 mi. 
nw.  Acaponeta  (1;  CAS).  SINALOA:  26  mi.  ne.  Villa  Union  (3;  LBSC).  SONORA:  7.2  mi.  se.  Alamos  (1;  GRNo). 


Collecting  notes.  — Specimens  of  S.  pluripunctatus  have  been  collected  from  March  to 
September  in  the  United  States,  and  in  January,  July,  and  September  in  Mexico.  Many 
specimens  were  taken  at  lights,  to  which  they  no  doubt  flew.  I have  collected  specimens  in 
gravel  bars  along  streams,  usually  further  from  water  than  specimens  of  most  other  species. 

Taxonomic  notes.  — The  type  of  S.  simplex,  taken  at  the  same  time  and  place  as  that  of 
S.  pluripunctatus,  is  a small  specimen  of  the  same  species  and  was  so  recognized  subsequent 
to  its  description.  Though  the  name  has  line  priority,  it  was  treated  as  a junior  synonym  of 
S.  pluripunctatus  by  LeConte  (1857)  and  by  all  subsequent  authors. 

Though  known  distributions  are  allopatric,  specimens  of  S.  pluripunctatus  and  S.  kulti 
have  been  taken  within  75  miles  of  each  other,  and  the  two  river  systems  involved,  Acapone- 
ta and  Grande  de  Santiago,  approach  each  other  even  more  closely  in  places.  Scarcity  of 
material  from  this  area  precludes  definite  statements  about  reproductive  relationships,  but 
available  evidence  suggests  reproductive  isolation.  In  specimens  seen,  all  females  of  S.  pluri- 
punctatus have  paramedian  ambulatory  setae  on  sternum  seven,  and  males  have  the  apical 
portion  of  the  median  lobe  deflected  at  a slight  but  evident  angle. 

Statistical  evidence  of  distinctness  of  S.  pluripunctatus  and  S.  kulti  is  given  in  Tables  18- 
23  and  Fig.  152-157.  Northern  specimens  of  S.  kulti  (sample  ten)  approach  S.  pluripunctatus 
in  body  size  and  in  numbers  of  elytral  setae,  but  the  Nayarit  sample  of  the  latter  (sample 
nine)  is  divergent  in  these  ways.  Northern  specimens  of  S.  kulti  approach  specimens  of  S. 
pluripunctatus  from  Nayarit,  but  diverge  from  those  from  Durango  (sample  eight),  in  eye 
size.  And,  they  diverge  from  Nayarit  S.  pluripunctatus  in  pronotal  form.  It  is  unlikely  that 
there  is  any  actual  or  potential  gene  flow  between  northern  S.  kulti  and  either  southern 
form  of  S.  pluripunctatus,  and  I therefore  treat  these  taxa  as  distinct  species. 

Schizogenius  kulti  new  species 

Type  material.  — Holotype  male  and  allotype  female  labelled  “MEXICO.  Chiapas.  18.6 
mi.  se.  Tonala.  Rte.  200  100'  III.2.1966”  and  “George  E.  Ball  D.  R.  Whitehead  collectors” 
(MCZ).  Thirty  additional  specimens  from  various  localities  in  Chiapas  are  paratypes  (CNC, 


224 


Whitehead 


DRWh,  UASM.  USNM). 

Diagnostic  combination.  — The  following  characters  combine  to  distinguish  specimens  of 
this  species  from  all  others  in  the  pluripunctatus  group:  median  lobe  arcuate,  apical  deflec- 
tion not  angulate;  females  without  paramedian  ambulatory  setae;  and  pronotum  without 
accessory  marginal  setae.  Most  specimens  have  fewer  than  30  setae  on  the  disc  of  each 
elytron,  a feature  unique  within  the  group,  but  some  specimens  from  northern  parts  of  the 
range  cannot  be  so  distinguished. 

Description.  — As  in  S.  seticollis  seticollis  except:  color  paler,  testaceous  to  rufopiceous; 
front  femora,  coxae,  and  trochanters  ferrugineous;  maxillae,  labial  palpi,  antennae,  front 
tarsi  and  tibiae,  middle  and  hind  legs  except  coxae,  and  apical  margins  of  elytra  and  abdo- 
men rufotestaceous;  no  metallic  luster.  Form  and  sculpture  of  head  and  pronotum  as  in 
S.  plurisetosus  (Fig.  126),  except  sides  of  pronotum  bisetose  and  basal  and  anterior  trans- 
verse impressions  nearly  impunctate.  Elytral  intervals  three,  five,  and  seven  in  specimens 
examined  with  6-14,  6-14,  and  4-14  setae,  respectively;  striae  nearly  impunctate.  Male  geni- 
talia: median  lobe,  Fig.'  136,  nearly  or  quite  arcuate,  apical  portion  proportionately  long 
and  deflexed  at  a slight  angle  or  not  angulate;  endophallus,  Fig.  144;  11  specimens  examined. 

Measurements  and  proportions.  See  Table  25.  Of  holotype:  TL,  3.43  mm;  LE,  2.09  mm; 
WH,  0.71  mm;  WP.  0.92  mm;  WE,  1.04  mm;  WF/WH,  0.63;  LP/WP,  0.93;  DP/LP,  0.83; 
LP/WE.  0.83;  Ta/Ti,  0.65;PS/LP,  0.61 . Of  allotype:  TL,  3.45  mm;LE,  2.10mm;  WH,  0.72 
mm;  WP.  0.92  mm;  WE,  1.08  mm;  WF/WH,  0.60;  LP/WP,  0.95;  DP/LP,  0.83;  LP/WE,  0.82; 
Ta/Ti,  0.65;  PS/LP,  0.59. 


Table  25.  Descriptive  statistics  for  S.  kulti,  based  on  14  males  from  18.6  miles  southeast  of 
Tonala,  Chiapas. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measurerm 

jnts,  in  mm. 

TL 

2.85-3.53 

3.280 

0.349 

0.140 

7.09 

LE 

1.75-2.14 

1.987 

0.205 

0.082 

6.87 

WH 

0.62-0.75 

0.696 

0.066 

0.027 

6.34 

WP 

0.76-0.96 

0.884 

0.097 

0.039 

7.31 

WE 

0.92-1.09 

1.016 

0.090 

0.036 

5.93 

B.  Setae  on  le 

ft  elytron. 

Interval  3 

6-  9 

7.5 

Interval  5 

7-10 

8.0 

Interval  7 

5-  7 

5.9 

Total 

19-25 

21.5 

3.9 

1.6 

11.99 

C.  Proportion 

s. 

WF/WH 

0.60-0.65 

0.620 

0.024 

0.010 

2.60 

LP/WP 

0.90-0.97 

0.935 

0.029 

0.012 

2.04 

DP/LP 

0.81-0.85 

0.832 

0.018 

0.007 

1.41 

LP/WE 

0.75-0.84 

0.807 

0.040 

0.016 

3.32 

Ta/Ti 

0.60-0.71 

0.668 

0.051 

0.020 

5.04 

PS/LP 

0.61-0.65 

0.629 

0.020 

0.008 

2.07 

Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


225 


Variation.  - Date  on  variation  in  six  characteristics  are  given  in  Tables  18-23,  for  samples 
numbered  10  to  14  as  in  Fig.  152-157.  Toward  the  north  in  both  Atlantic  and  Pacific 
drainage  basins,  body  size,  pronotal  width,  and  numbers  of  elytral  setae  increase,  eye  size 
decreases,  and  except  for  the  Veracruz  sample  paramedian  pronotal  sulci  lengthen.  Northern 
populations  are  geographically  isolated  from  one  another,  so  that  their  similarities  may  be 
interpreted  either  as  convergences  or  as  relict  characteristics.  I suspect  the  latter  is  more 
likely,  since  related  species  are  larger  and  more  setose.  I found  no  specimens  with  accessory 
pronotal  setae,  and  no  females  with  paramedian  ambulatory  setae  on  sternum  seven.  The 
form  of  the  median  lobe  is  constantly  arcuate,  the  apical  portion  not  or  hardly  deflected 
at  an  angle. 

Etymology . — The  name  S.  kulti  is  dedicated  to  the  Czechoslovakian  entomologist,  K. 
Kult,  whose  numerous  papers  on  the  Scaritini  contributed  much  to  our  knowledge  of  the 
group.  A specimen  from  Rio  Maria  Linda,  Guatemala,  was  treated  by  Bates  (1881)  as  S. 
tristriatus,  but  was  subsequently  labelled  as  a paratype  of  “S.  brittoni ” Kult,  a pin  label 
name  never  published. 

Distribution.  — Specimens  of  S.  kulti  have  been  collected  at  altitudes  ranging  from  near 
sea  level  to  4100',  from  the  Rio  Grande  de  Santiago  system  in  Nayarit  and  Zacatecas  and 
the  Rio  Atoyac  system  in  Veracruz,  south  to  Guatemala  (Fig.  147).  I studied  122  speci- 
mens of  this  species  from  the  following  localities. 

MEXICO 

CHIAPAS:  3.2  mi.  n.  Arriaga  (1;  USNM),  12.2  mi.  ne.  Chiapa  de  Corzo  (4;  DRWh,  UASM),  Pijijiapan  (4;  CNC),  Tonala 
(1;  USNM),  18.6  mi.  se.  Tonala  (23;  MCZ,  UASM).  GUERRERO:  41.4  mi.  n.  Acapulco  (3;  UASM),  Coyuca  (2;  UASM). 
JALISCO:  0.4  mi.  w.  Cocula  (1;  UASM),  Talpa  de  Allende  (1;  UASM).  NAYARIT:  Jesus  Maria  (2;  UCB),  14  mi.  e.  San 
Bias  (1;  UASM).  OAXACA:  17.7  mi.  w.  El  Camaron  (10;  MGFT,  UASM),  29.4  mi.  e.  El  Coyul  (33;  UASM),  11.1  mi.  n. 
Marias  Romero  (1;  CAS),  Valle  Nacional  (14;  UASM),  Zanatepec  (3;  IRSB).  TABASCO:  Teapa  (3;  BMNH).  VERACRUZ: 
Cordoba  (1;  CAS),  Fortin  de  las  Flores  (9;  DRWh,  FDAG),  Catemaco  (1;  JNeg).  ZACATECAS:  0.9  mi.  n.  Jalpa  (1;  UASM). 

GUATEMALA 

ESCUINTLA:  Rio  Maria  Linda  (1;  BMNH).  IZABAL:  Los  Amates  (1;  MCZ). 


Collecting  notes.  — Adults  of  S.  kulti  are  active  throughout  the  year  in  gravel  bars  along 
streams.  I noted  no  special  habits,  and  found  them  together  with  specimens  of  one  or  more 
other  Schizogenius  species.  Specimens  from  Fortin  de  las  Flores,  Veracruz,  were  taken  at 
black  lights,  and  no  doubt  flew  there.  None  of  the  Fortin  specimens  were  collected  by  hand, 
although  suitable  habitats  are  in  the  vicinity  and  numerous  specimens  of  S.  tristriatus  were 
collected  there. 


The  sallei  group 

Diagnostic  combination.  — Members  of  this  monotypic  group  are  distinguished  by  the 
following  combination  of  characters:  body  strongly  flattened;  paramedian  clypeal  carinae 
extended  to  median  tooth;  clypeal  field  narrow,  no  wider  at  base  than  apex  of  median 
frontal  sulcus;  submentum  without  accessory  setae;  pronotum  without  paralateral  carinae; 
discal  setae  present  on  intervals  three,  five,  and  seven,  total  normally  less  than  30,  average 
length  less  than  1.0  times  maximum  width  of  interval  two;  abdomen  microsculptured  along 
midline,  and  with  microsculpture  in  small  lateral  patches  near  coxal  depressions  of  sternum 
three;  endophallus  with  basal  collar  spines  distinct.  Also:  clypeal  suture  sharply  impressed; 
antennal  articles  five  to  ten  filiform,  elongate;  front  and  middle  tarsi  broadened  and  with 
dense  ventral  pubescence,  especially  in  males;  sternum  seven  with  paramedian  ambulatory 
setae  in  males,  not  in  females;  paramedian  carinae  of  sternum  three  curved  outward  at 


226 


Whitehead 


Fig.  122-123.  Head  and  pronotum,  dorsal  aspect.  122.  S.  truquii  Putzeys,  Cocula,  Jalisco.  123.  S.  brevisetosus  new 
species,  Barrel  Springs  Creek,  Texas.  Fig.  124.  Habitus,  dorsal  aspect,  S.  seticollis  seticollis  Fall,  Mokelumne  Hill,  Cali- 
fornia. Fig.  125-126.  Head  and  pronotum,  dorsal  aspect.  125.  S.  seticollis  vandykei  new  subspecies,  Arroyo  Saltito, 
Baja  California.  126.  S.  plurisetosus  new  species,  Linares,  Nuevo  Leon.  Fig.  127-136.  Male  median  lobe,  lateral  aspect. 
127.  S.  truquii  Putzeys,  Cocula,  Jalisco.  128.  S.  brevisetosus  new  species,  Sanderson,  Texas.  129.  S.  seticollis  seticollis 
Fall,  Dry  Creek,  California.  130.  S.  seticollis  vandykei  new  subspecies,  Arroyo  Saltito,  Baja  California.  131.  S.  plurise- 
tosus new  species,  Cienega  de  Flores,  Nuevo  Leon.  132.  S.  multisetosus  Bates,  Tepexco,  Puebla.  133.  Same,  72.5  mi.  s. 
Valle  Nacional,  Oaxaca.  134.  Same,  Tamazunchale,  San  Luis  Potosi.  135.  S.  pluripunctatus  LeConte,  Patagonia,  Arizona. 
136.  S.  kulti  new  species,  Valle  Nacional,  Oaxaca. 


227 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


137 


145 


147 


LE. 

O 

0 

0 

) 


2.746*2.815 
2 816-2.885 
2.886-2.955 
2.956-3  025 
3.026-3  095 


14 

Fig.  137-144.  Male  endophallus.  137.  5.  truquii  Putzeys,  La  Garita,  Jalisco.  138.  5.  brevisetosus  new  species  Lozi 
Canyon,  Texas.  139.  5.  seticollis  seticollis  Fall,  Trimmer,  California.  140.  5.  plurisetosus  new  species,  El  Barretal  Tan 
ubpas.  141.  5.  multisetosus  Bates,  72.5  mi.  s.  Valle  Nacional,  Oaxaca.  142.  Same,  Tepexco,  Puebla.  143.  S pluripur 
tatus  LeConte,  Acaponeta,  Nayarit.  144.  5.  kulti  new  species,  41.4  mi.  n.  Acapulco,  Guerrero.  Fig.  145-147.  Knoi 
distributions.  145.  S.  truquii  Putzeys.  146.  S.  brevisetosus  new  species.  147.  S.  seticollis  seticollis  Fall,  half-filled  circli 
5.  seticollis  vandykei  new  subspecies,  filled  circles;  S.  pluripunc tatus  LeConte,  filled  triangles;  S.  kulti  new  sped, 
half-filled  triangles;  S.  multisetosus  Bates,  half-filled  squares;  5.  plurisetosus  new  species,  filled  squares.  Fig.  148  Ge 
graphic  variation  in  5.  seticollis  seticollis  Fall,  circles,  and  5.  seticollis  vandykei  new  subspecies,  square;  means  of  bo< 
size.  Table  11.  M 


228 


Whitehead 


30 


20 


152 


LE. 

O 

9 

9 

• 


153 


154 


155 


30 

r'x 

1 > w 

PS 

/ LP 

x.\  i 

20  ° 

0 625-0  634 

vV  \ 

v>  \ i6]  Tfij 

9 

0 G35-0  644 

9 

0.645-0  654 

7a- 

9 

0 655-0.664 

• 

0 665-0 674 

10 

156  t 1000  mi ( 157 


Fig.  149-151.  Geographic  variation  in  S.  seticollis  seticollis  Fall,  circles,  and  S.  seticollis  vandykei  new  subspecies,  square. 
149.  Means  of  numbers  of  discal  setae  on  left  elytron,  Table  12.  150.  Means  of  relative  eye  size,  Table  13.  151.  Means 
of  pronotal  form.  Table  14.  Fig.  152-157.  Geographic  variation  in  S.  plurisetosus  new  species,  hexagon;  S.  multisetosus 
Bates,  diamonds;  S.  pluripunctatus  LeConte,  circles;  and  S.  kulti  new  species,  squares.  152.  Means  of  body  size,  Table 
18.  153.  Means  of  numbers  of  discal  setae  on  left  elytron,  Table  19.  154.  Means  of  relative  eye  size,  Table  20.  155. 
Means  of  pronotal  form,  Table  21.  156.  Means  of  relative  pronotal  size,  Table  22.  157.  Means  of  relative  length  of 
paramedian  pronotal  sulci,  Table  23.  Accompanying  legends  are  for  all  included  taxa. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


229 


apices;  pygidium  crenulate  at  apex  in  females;  color  rufocastaneous;  and  pronotal  hind 
angles  sharply  developed. 

S.  sallei  is  the  only  Schizogenius  species  characterized  by  strong  microsculpture  along 
the  midline  but  not  the  margins  of  the  abdomen. 

Distribution.  — S.  sallei  is  known  from  the  Great  Plains  region  of  North  America,  from 
Kansas  south  to  the  Rio  Grande.  I examined  374  specimens. 

Schizogenius  sallei  Putzeys 

Schizogenius  sallei  Putzeys  1866:228.  Type  locality  Texas;  type  male  in  IRSB,  specimen 

labelled  lectotype  by  me  so  designated  (!).  LeConte  1879:34.  Lindroth  1961:166. 

Diagnostic  combination.  — Abdomen  microsculptured  along  midline,  otherwise  mostly 
unmicrosculptured . 

Description.  — Body  flattened.  Color  castaneous,  unmetallic,  legs  and  palpi  paler. 

Integument.  Conspicuous  microsculpture  on  paramedian  frontal  sulci,  mouthparts,  genae, 
front  legs  except  posterior  surfaces  of  femora  and  trochanters,  middle  legs  except  anterior 
surfaces  of  trochanters,  hind  tibiae  and  posterior  surfaces  of  hind  femora,  elytral  epipleura 
at  base  and  on  apical  two-thirds,  sternum  three  in  coxal  depressions  and  small  paralateral 
patches,  and  middle  of  abdomen  on  sterna  three  to  five  or  six. 

Head.  Fig.  158.  Paramedian  clypeal  carinae  straight,  moderately  elevated  in  basal  half, 
extended  to  median  tooth;  median  field  triangular,  narrow,  no  wider  at  base  than  apex  of 
median  frontal  sulcus.  Clypeal  suture  sharply  defined.  Eye  prominent,  finely  and  uniformly 
faceted.  Neck  densely  punctate.  Gena  coarsely  punctate,  rugose  in  front.  Antennal  articles 
four  to  ten  elongate,  article  five  about  1.7-1. 8 times  longer  than  wide. 

Pronotum.  Fig.  158.  Sides  bisetose;  base  not  rugose;  hind  angles  prominent.  Paramedian 
longitudinal  sulci  moderately  long,  nearly  straight,  strongly  hooked  basally.  Anterior  trans- 
verse impression  strongly  punctate. 

Legs.  Front  and  middle  tarsi  moderately  dilated  and  pubescent  ventrally,  less  so  in  fe- 
males; hind  tarsus  slender,  short.  Paronychia  nearly  as  long  as  tarsal  claws.  Front  tibia 
narrowed  evenly  to  base.  Front  femur  not  strongly  constricted  near  apex. 

Elytra.  Left  elytron  with  about  7-9  setae  on  interval  three,  6-8  on  interval  five,  3-5  on 
interval  seven;  total  17-23  in  specimens  examined.  Striae  deep  and  sharply  engraved,  finely 
punctate  in  basal  two-thirds.  Intervals  one  to  seven  broad  and  flat,  interval  eight  carinate  at 
apex;  intervals  three,  five,  and  seven  broadly  joined  at  apices.  Humeral  denticles  prominent. 

Abdomen.  Sternum  three  with  paramedian  carinae  curved  outward  at  apices.  Sternum 
seven  with  paramedian  ambulatory  setae  in  males  only.  Pygidium  with  apical  margin  crenu- 
late in  females,  entire  in  males. 

Male  genitalia.  Median  lobe,  Fig.  169;  endophallus,  Fig.  178;  8 specimens  examined. 

Measurements  and  proportions.  See  Table  26. 

Variation.  — See  Table  26.  I found  no  significant  geographic  variation  in  relative  eye 
size,  pronotal  form,  body  size,  or  in  numbers  of  elytral  setae. 

Etymology.  — This  species  was  named  after  Auguste  Salle,  who  made  important  early 
collections  of  Coleoptera  in  Mexico. 

Distribution.  — The  known  range  of  this  species  extends  from  Kansas  south  to  the  Rio 
Grande  Valley  in  Texas  (Fig.  185).  This  is  a peculiar  distribution,  especially  since  there  is  no 
evidence  of  restricted  gene  flow  either  between  the  Rio  Grande  and  Colorado  River  systems, 
or  north  and  south  of  the  Red  River.  I have  two  doubtful  records  of  unspecified  localities 
in  Canada  (UKSM)  and  Ohio  (CAS);  both  require  confirmation.  I studied  374  additional 
specimens  from  the  following  localities. 


230 


Whitehead 


UNITED  STATES 

No  locality  (7;  ANSP.  LACM).  KANSAS:  Butler  Co..  Leon  (6;  UKSM);  Riley  Co.  (1;  USNM).  OKLAHOMA:  Carter  Co., 
10.7  mi.  s.  Drake  (2:  TLEr):  Murray  Co..  10.3  mi.  n.  Drake  (6;  TLEr).  TEXAS  (55;  AMNH,  ANSP.  CAS,  CNHM.  INHS, 
IRSB,  KSL'M,  MCZ,  MSUL,  RUNB,  UKSM,  USNM,  ZMLS):  Andrews  Co.,  Fullerton  (3;  CAS,  USNM);  Bexar  Co.,  San 
Antonio  (1;  MCZ);  Blanco  Co.,  Cypress  Mills  (3;  USNM),  Johnson  City  (8;  UASM),  Twin  Sisters  (7;  UASM):  Brown  Co., 
Brownwood  (1;  AMNH);  Comal  Co.,  New  Braunfels  (1;  USNM):  Colorado  Co.,  Columbus  (1;  MSUL):  Cooke  Co.,  4 mi.  sw. 
Era  (3:  BMNH);  Dallas  Co.  (1:  INHS),  Dallas  (7:  INHS.  MCZ,  MSUL,  RTBe):  Erath  Co..  Morgan  Mill  (2;  IRSB):  Guadalupe 
Co.,  Seguin  (3:  UKSM):  Kerr  Co..  Kerrville  (7;  CNC):  Kinney  Co.,  23  mi.  sw.  Brackettville  (107;  AMNH,  ANSP.  CAS, 
CUNY.  DJLa.  FDAG.  HGou,  INHS,  JHen,  JNeg,  MCZ,  MGFT,  MZSP,  TCBa,  UAFA,  UASM,  UATA,  UCB.  ZMLS); 
Lampasas  Co.,  Adamsville  (10;  UASM);  Maverick  Co..  8 mi.  n.  Quemado  (1;  DHKa):  McCulloch  Co.,  16  mi.  s.  Brady  (4: 
CAS):  McLennan  Co..  Waco  (1:  DHKa):  Randall  Co.,  Canyon  (2:  MSUL);  Real  Co.,  Leakey  (1;  UASM),  2 mi.  s.  Leakey 
(34:  UASM;  Taylor  Co.,  25  mi.  sw.  Abilene  (14:  CNHM);  Terrell  Co„  Chandler  Ranch  (7:  UASM);  Travis  Co.,  Austin 
(4:  UASM,  USNM):  Uvalde  Co.,  Gamer  State  Park  (15;  UASM),  Sabinal  (2;  USNM),  17  mi.  nw.  Uvalde  (1;  UASM);  Val 
Verde  Co.,  26  mi.  n.  Comstock  (7:  DRWh).  Devil’s  River  (1;  UATA),  9 mi.  se.  Del  Rio  (8;  UASM),  13  mi.  nw.  Del  Rio 
(24;  UASM),  14  mi.  se.  Del  Rio  (6;  UASM). 


Table  26. 

Descriptive  statistics  for  S. 

sallei,  based  on 

20  males  from 

23  miles  southwest  of 

Brackettville,  Texas. 

Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measurements,  in  mm. 

TL 

4.04-4.50 

4.278 

0.189 

0.056 

2.95 

LE 

to 

l/i 

t j 
oo 
o 

2.660 

0.126 

0.038 

3.16 

WH 

0.82-0.91 

0.866 

0.036 

0.011 

2.79 

WP 

1.05-1.14 

1.096 

0.041 

0.012 

2.52 

WE 

1.29-1.44 

1.372 

0.060 

0.018 

2.90 

1 

B.  Setae  on  left  elytron. 

Interval  3 

7-  9 

7.8 

Interval  5 

6-  8 

7.4 

Interval  7 

3-  5 

4.0 

Total 

17-21 

19.2 

1.9 

0.6 

6.62 

C.  Proportions. 


WF  WH 

0.59-0.64 

0.610 

0.022 

0.007 

i-j 

O 

LP/WP 

0.91-0.96 

0.935 

0.023 

0.007 

1.61 

DP/LP 

0.78-0.84 

0.804 

0.025 

0.007 

I o 

o 

u> 

LP/WE 

0.71-0.77 

0.743 

0.027 

0.008 

2.43 

Ta/Ti 

0.59-0.68 

0.636 

0.033 

0.010 

3.51 

PS  LP 

0.66-0.72 

0.692 

0.030 

0.009 

VO 
00 
r i 

Collecting  notes.  — Specimens  of  this  species  have  been  collected  from  March  to  Septem- 
ber, in  riparian  gravel  bars  and  at  lights.  At  least  in  southern  Texas,  I found  them  abundant. 
The  lack  of  pronounced  geographic  variation  implies  that  S.  sallei  is  a particularly  vagile 
species. 

Taxonomic  notes.  — This  species  is  not  closely  related  to  any  other,  and  I found  no 
problems  in  infraspecific  variation. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


231 


The  tristriatus  group 

Diagnostic  combination.  — Members  of  this  group  are  distinguished  by  the  following 
combination  of  characters:  body  flattened;  paramedian  clypeal  carinae  extended  to  median 
tooth;  bases  of  paramedian  frontal  carinae  not  broadly  fused;  antennal  articles  five  to  ten 
filiform;  discal  setae  present  on  intervals  three,  five,  and  seven;  and  abdomen  extensively 
microsculptured.  Also:  clypeal  suture  sharply  impressed;  clypeal  field  triangular,  less  than 
1.5  apical  width  of  median  frontal  sulcus;  front  and  middle  tarsi  broadened  and  with  dense 
ventral  pubescence,  especially  in  males;  sternum  seven  with  paramedian  ambulatory  setae 
in  males,  not  in  females;  paramedian  carinae  of  sternum  three  curved  at  apices;  pygidium 
crenulate  at  apex  in  most  females  and  some  males;  and  endophallus  with  basal  collar  spines 
distinct. 

Distribution.  — Members  of  this  group  range  from  southeastern  Canada,  through  the 
eastern  and  central  United  States,  south  to  southern  Mexico.  I studied  398  specimens  of  the 
tristriatus  group. 


Schizogenius  tristriatus  Putzeys 

Schizogenius  tristriatus  Putzeys  1846:651.  Type  locality  Mexico;  location  of  type  unknown, 

possibly  Hope  Museum,  Oxford.  Putzeys  1863:24.  Putzeys  1866:227.  Bates  1881:37. 

Kult  1950:140. 

Diagnostic  combination.  - Specimens  of  this  species  are  readily  recognized  by  charac- 
teristics given  in  the  key  to  North  and  Middle  American  species  of  the  subgenus  Schizoge- 
nius. S.  tristriatus  is  the  only  Mexican  member  of  the  tristriatus  group  which  does  not  have 
basally  broadened  front  tibiae. 

Description.  — Body  flattened.  Rufopiceous;  elytral  apices,  abdomen,  and  legs  rufous; 
palpi  testaceous;  no  metallic  luster.  Microsculpture  as  in  S.  dilatus.  Otherwise  as  in  S.  tibialis 
except  as  follows.  Form  and  sculpture  of  head  and  pronotum,  Fig.  159;  eyes  prominent; 
antennal  article  five  1.6- 1.7  times  longer  than  wide.  Front  tibia  (Fig.  164)  strongly  tapered, 
distal  tooth  not  spatulate.  Left  elytron  with  6-8  setae  on  interval  three,  5-7  on  interval  five, 
and  2-4  on  interval  seven,  total  14-18  in  specimens  studied.  Male  genitalia  with  median  lobe, 
Fig.  17-;  endophallus,  Fig.  179;  6 specimens  examined. 

Measurements  and  proportions.  See  Table  27. 

Variation.  — Two  specimens  from  San  Luis  Potosi  and  Queretaro  are  slightly  paler  in 
color,  but  may  simply  be  teneral. 

Etymology.  — Latin,  tres  = three,  plus  stria  = furrow,  in  reference  to  the  median  and 
paramedian  pronotal  sulci. 

Distribution.  — Specimens  of  this  species  have  been  collected  in  eastern  Mexico  from 
San  Luis  Potosi  and  Queretaro  south  to  Veracruz,  at  elevations  ranging  from  2900  to  4100' 
(Fig.  186).  I studied  78  specimens  from  the  following  localities. 

MEXICO 

No  locality  (4;  IRSB,  MCZ).  QUERETARO:  Escanelilla  (1;  UASM).  SAN  LUIS  POTOSf:  Xilitla  (2;  CNC).  VERACRUZ: 
3.2  mi.  sw.  Coscomatepec  (22;  BMNH,  CAS,  CNHM,  DRWh,  IRSB,  MCZ,  MGFT,  UASM,  USNM):  Fortin  de  las  Flores 
(49;  FDAG,  UASM). 

Collecting  notes.  — Specimens  of  S.  tristriatus  have  been  collected  nearly  throughout  the 
year,  from  March  to  November,  either  in  gravel  bars  along  streams  or  at  lights.  Where  I have 
collected  them  in  numbers,  I did  not  find  specimens  of  other  species,  though  at  Fortin  de  las 
Flores  specimens  of  other  species  were  collected  at  lights. 


232 


Whitehead 


Table  27.  Descriptive  statistics  for  S.  tristriatus,  based  on  20  males  from  Fortin  de  las 
Flores,  Veracruz,  Mexico. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

nts,  in  mm. 

TL 

4.16-4.84 

4.574 

0.258 

0.077 

3.76 

LE 

2.60-3.02 

2.854 

0.154 

0.046 

3.59 

WH 

0.84-1.00 

0.928 

0.058 

0.018 

4.20 

WP 

1.10-1.30 

1.215 

0.081 

0.024 

4.44 

WE 

1.43-1.62 

1.531 

0.079 

0.024 

3.43 

B.  Setae  on  le 

ft  elytron. 

Interval  3 

6-  8 

6.7 

Inteval  5 

5-  7 

6.0 

Interval  7 

3-  4 

3.2 

Total 

14-18 

16.0 

1.8 

0.5 

7.31 

C.  Proportion 

s. 

WF/WH 

0.59-0.63 

0.604 

0.018 

0.005 

2.02 

LP/WP 

0.87-0.93 

0.905 

0.024 

0.007 

1.77 

DP/LP 

0.78-0.84 

0.814 

0.020 

0.006 

1.66 

LP/WE 

0.69-0.74 

0.719 

0.024 

0.007 

2.25 

Ta/Ti 

0.60-0.67 

0.625 

0.034 

0.010 

3.62 

PS/LP 

0.62-0.71 

0.668 

0.030 

0.009 

2.95 

Taxonomic  notes.  — There  are  two  specimens  of  S.  tristriatus  in  the  Putzeys  collection 
in  the  Institut  Royal  des  Sciences  Naturelles  de  Belgique  labelled  as  syntypes,  but  I think 
these  probably  are  the  specimens  mentioned  in  a subsequent  paper  (Putzeys,  1863).  How- 
ever, they  were  identified  by  him,  and  do  match  the  original  description,  notably  in  colora- 
tion, and  no  doubt  are  conspecific  with  the  types.  S.  longipennis  Putzeys  was  described  as  a 
variety  of  S.  tristriatus  and  was  considered  as  such  by  Bates  (1881),  but  as  noted  by  Kult 
(1950)  is  not  closely  related. 

Schizogenius  dilatus  new  species 

Type  material.  — Holotype  male  and  allotype  female  labelled  “MEX.  Nuevo  Leon  Rio 
Sabinas  Hidalgo,  4.8  mi.  e.  Sabinas  Hidalgo  800'  X. 22-23.  65”  and  “George  E.  Ball  D.  R. 
Whitehead  collectors”  (MCZ).  An  additional  26  specimens  from  various  localities  in  Nuevo 
Leon  and  Tamaulipas  are  paratypes  (BMNH,  CAS,  CNC,  DRWh,  IRSB,  UASM,  USNM). 

Diagnostic  combination.  — Specimens  of  this  species  are  distinguished  from  others  in  the 
group  by  front  tibia  strongly  broadened  basally  and  femur  distinctly  constricted  near  the 
apex,  but  no  pronounced  microsculpture  on  pro  thoracic  pleura. 

Description.  — Strong  microsculpture  on  paramedian  frontal  sulci,  mouthparts,  genae, 
sides  and  base  of  pronotum,  front  legs,  middle  legs  except  anterior  surfaces  of  trochanters, 
hind  tibiae  and  posterior  surfaces  of  hind  femora,  elytral  epipleura,  and  entire  abdomen. 
Otherwise  as  in  S.  tibialis  except  as  follows.  Eyes  prominent;  antennal  article  five  about 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


233 


1.6- 1.7  times  longer  than  wide.  Front  tibia  (Fig.  165)  slightly  more  tapered,  distal  tooth 
not  spatulate.  Left  elytron  with  8-1 1 setae  on  interval  three,  7-10  on  interval  five,  and  3-5 
on  interval  seven,  total  18-23  in  specimens  studied.  Male  genitalia  with  median  lobe,  Fig. 
171;  and  endophallus,  Fig.  180;  6 specimens  examined. 

Measurements  and  proportions.  See  Table  28.  Of  holotype:  TL,  4.57  mm;  LE,  2.81  mm; 
WH,  0.90  mm;  WP,  1.18  mm;  WE,  1.45  mm;  WF/WH,  0.61;  LP/WP,  0.97;  DP/LP,  0.74; 
LP/WE,  0.79;  Ta/Ti,  0.65;PS/LP,  0.74.  Of  allotype:  TL,  5.05  mm;  LE,  3. 10  mm;  WH,  0.98 
mm;  WP,  1.33  mm;  WE,  1.60  mm;  WF/WH,  0.63;  LP/WP,  0.95;  DP/LP,  0.74;  LP/WE,  0.79; 
Ta/Ti,  0.65;  PS/LP,  0.65. 


Table  28.  Descriptive  statistics  for  S.  dilatus,  based  on  12  males  from  Nuevo  Leon  and 
Tamaulipas,  Mexico. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

nts,  in  mm. 

TL 

4.41-5.08 

4.727 

0.316 

0.122 

4.46 

LE 

2.74-3.11 

2.903 

0.176 

0.068 

4.03 

WH 

0.84-1.00 

0.928 

0.073 

0.028 

5.22 

WP 

1.12-1.33 

1.227 

0.095 

0.037 

5.17 

WE 

1.38-1.62 

1.499 

0.110 

0.042 

4.89 

B.  Setae  on  lei 

ft  elytron. 

Interval  3 

8-  9 

8.7 

Interval  5 

7-  9 

7.5 

Interval  7 

3-  5 

4.1 

Total 

18-22 

20.3 

1.8 

0.7 

6.00 

C.  Proportion; 

WF/WH 

0.60-0.65 

0.624 

0.023 

0.009 

2.51 

LP/WP 

0.94-1.00 

0.967 

0.027 

0.010 

1.84 

DP/LP 

0.74-0.77 

0.755 

0.016 

0.006 

1.44 

LP/WE 

0.76-0.81 

0.789 

0.019 

0.007 

1.57 

Ta/Ti 

0.58-0.65 

0.618 

0.037 

0.014 

3.95 

PS/LP 

0.65-0.74 

0.702 

0.039 

0.015 

3.74 

Etymology.  — Latin,  dilatus  = expanded,  in  reference  to  the  form  of  the  front  tibia. 
Distribution.  — I studied  28  specimens  from  the  following  localities  in  northeastern 
Mexico  (Fig.  187). 

MEXICO 

NUEVO  LEON:  Cienega  de  Flores  (4;  CAS,  IRSB),  Linares  (1;  DRWh),  14.8  mi.  w.  Linares  (2;  USNM),  5 mi.  s.  Monterrey 
(1;  CNC),  4.8  mi.  e.  Sabinas  Hidalgo  (17;  MCZ,  UASM).  TAMAULIPAS:  El  Barretal  (1;  BMNH),  21.3  mi.  n.  Ciudad 
Victoria  (1;  CNC),  39  mi.  s.  Ciudad  Victoria  (1;  DRWh). 


Collecting  notes.  — Specimens  of  S.  dilatus  have  been  collected  from  June  to  October,  in 


234 


Whitehead 


riparian  gravel  bars.  The  specimens  from  14.8  miles  west  of  Linares  were  taken  at  black 
lights.  The  facts  that  one  good  series  was  taken  along  a stream,  and  that  the  eyes  are  not 
reduced,  indicate  epigean  habits. 

Taxonomic  notes.  - The  only  other  species  having  broadened  front  tibiae,  S.  tibialis , is 
well  separated  morphologically,  and  is  also  sympatric  at  least  in  Nuevo  Leon.  S.  dilatus  is 
otherwise  most  similar  to  S.  tristriatus , but  is  clearly  distinct. 

Schizogenius  tibialis  new  species 

Type  material.  - Holotype  male  and  allotype  female  labelled  “MEXICO  S.L.P.  Rte.  85, 
19.3  mi.  n.w.  Tamazunchale  500'  XI.14&20.65”  and  “George  E.  Ball  D.  R.  Whitehead 
collectors”  (MCZ).  Ten  additional  specimens  from  various  localities  in  San  Luis  Potosi, 
Mexico,  are  paratypes  (CAS,  CNC.  DRWh,  IRSB,  UASM,  UKSM). 

Diagnostic  combination.  - Specimens  of  this  species  differ  from  all  others  in  the  genus  by 
the  following  combination  of  characteristics:  prothoracic  pleura  strongly  microsculptured; 
front  tibia  strongly  broadened  basally;  and  front  femur  strongly  constricted  near  apex. 

Description.  — Body  flattened.  Color  castaneous  to  rufopiceous,  without  metallic  luster, 
legs  and  antennae  ferrugineous,  tarsi  and  palpi  testaceous. 

Integument.  Strong  microsculpture  on  paramedian  frontal  sulci,  mouthparts,  genae,  sides 
and  base  of  pronotum,  prothoracic  pleura,  front  legs,  middle  legs  except  anterior  surfaces  of 
trochanters,  hind  legs,  elytral  epipleura,  and  entire  abdomen. 

Head.  Fig.  160,  161.  Generally  as  in  S.  amphibius,  except  eyes  prominent  to  strongly 
flattened,  and  antennal  article  five  about  1.7-1. 8 times  longer  than  wide. 

Pronotum.  Form  and  sculpture,  Fig.  160.  Otherwise  as  in  S.  amphibius  except  anterior 
transverse  impression  strongly  punctate. 

Legs.  Front  and  middle  tarsi  strongly  dilated  and  pubescent  ventrally  in  males,  less  so  in 
females;  hind  tarsus  slender.  Paronychia  conspicuous,  nearly  as  long  as  tarsal  claws.  Front 
tibia  (Fig.  166)  parallel  sided,  not  or  hardly  narrowed  toward  base;  distal  tooth  (Fig.  167- 
168)  varied  in  form.  Front  femur  strongly  constricted  near  apex. 

Elytra.  Left  elytron  with  8-1 1 setae  on  interval  three,  7-10  on  interval  five,  and  3-6  on 
interval  seven.  Striae  deep,  sharply  engraved,  strongly  punctate  in  basal  two-thirds.  Other- 
wise as  in  S.  amphibius. 

Male  genitalia.  Median  lobe,  Fig.  172,  173;  endophallus,  Fig.  1 8 1 ; eight  specimens  exam- 
ined. 

Measurements  and  proportions.  Of  holotype:  TL,  4.56  mm;  LE,  2.77  mm;  WH,  0.87  mm; 
WP.  1.17  mm;  WE,  1.45  mm;  WF/WH,  0.63;  LP/WP,  1.00;DP/LP,  0.75;  LP/WE,  0.81 ; Ta/ 
Ti,  0.65;  PS/LP.  0.65.  Of  allotype:  TL,  4.87  mm;  LE,  2.95  mm;  WH,  0.92  mm;  WP,  1.25 
mm;  WE,  1.54  mm;  WF/WH,  0.66;  LP/WP,  1.01;  DP/LP,  0.74;  LP/WE,  0.82;  Ta/Ti,  0.61; 
PS/LP,  0.72. 

Variation.  — As  S.  tibialis  is  poorly  represented  in  collections,  a statistical  analysis  of 
geographic  variation  is  not  yet  possible.  I do  not  suspect  important  variation  in  body  size, 
body  form,  or  in  numbers  of  elytral  setae.  The  largest  (LE,  3.12  mm)  and  smallest  (LE, 
2.70  mm)  males  are  from  Encarnacion  de  Diaz  and  Saltillo,  respectively,  and  the  largest 
(LE,  3.06  mm)  and  smallest  (LE,  2.73  mm)  females  are  from  Monterrey  and  the  Rio  Balsas, 
respectively. 

Despite  paucity  of  material,  a geographic  pattern  is  evident  from  unusually  pronounced 
variation  in  two  characteristics.  The  distal  tooth  of  the  front  tibia  is  swollen  or  spatulate 
apically  in  specimens  from  San  Luis  Potosi,  Veracruz,  Oaxaca,  and  Chiapas  (Fig.  167),  less  so 
in  specimens  from  Jalisco,  and  hardly  or  not  at  all  in  specimens  from  Coahuila,  Guerrero, 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


235 


Tamaulipas,  and  Texas  (Fig.  168).  Relative  eye  sizes  (WF/WH)  are  summarized  in  Fig. 
188;  eyes  are  prominent  in  specimens  from  San  Luis  Potosi  and  Veracruz  (Fig.  160), 
less  so  in  specimens  from  Oaxaca,  decrease  in  prominence  northward,  and  are  especial- 
ly flattened  in  specimens  from  Indian  Creek  Cave  in  Texas  (Fig.  161).  Only  forms  with 
prominent  eyes  have  been  taken  in  series,  in  San  Luis  Potosi  and  Oaxaca,  and  even 
they  were  not  found  as  abundantly  as  were  specimens  of  other  species.  The  distribu- 
tion of  S.  tibialis  is  probably  correlated  with  the  distribution  of  limestone,  and  per- 
haps of  limestone  caves,  as  all  samples  were  obtained  in  regions  in  or  near  limestone  out- 
croppings. 

Probably  there  are  no  important  barriers  to  gene  flow  along  the  Atlantic  slope  from  San 
Luis  Potosi  south  to  Oaxaca,  then  back  north  along  the  Pacific  slope  to  the  Trans-Volcanic 
Sierra  and  inland  along  its  northern  drainages.  As  there  is  more  similarity  between  Jalisco 
and  San  Luis  Potosi  samples  in  form  of  distal  tooth  of  front  tibia  than  between  the  latter 
and  the  Guerrero  specimen,  there  may  be  some  gene  flow  there.  And  the  eye  size  pattern 
suggests  that  there  is  no  serious  barrier  to  gene  flow  between  Jalisco  and  Atlantic  drainage 
systems  in  Coahuila,  Tamaulipas,  and  Texas.  The  particularly  small  eye  of  Indian  Creek 
Cave  specimens  suggests  a steeply  stepped  cline;  those  specimens  might  be  of  a reproductive 
isolate,  but  the  existence  of  a reasonable  geographic  cline  argues  otherwise.  However,  large- 
eyed populations  from  San  Luis  Potosi  are  structurally  isolated  from  smaller-eyed  forms 
from  Texas  and  Tamaulipas,  and  may  be  reproductively  isolated  and  even  sympatric.  Thus, 
samples  of  S.  tibialis  probably  represent  a circle  of  races,  as  shown  in  Fig.  188.  This  hypo- 
thesis requires  further  investigation.  In  particular,  further  collections  should  be  made  in 
Tamaulipas,  in  the  Sierra  de  Guatemala  and  Sierra  de  Tamaulipas,  to  determine  whether 
sympatry  exists. 

Etymology.  — Latin,  tibialis  = of  the  tibia,  a reference  to  the  peculiar  structure  of  the 
front  tibia. 

Distribution.  - Specimens  of  this  species  have  been  collected  from  the  Nueces  River 
system  in  southern  Texas,  south  to  southeastern  Oaxaca  (Fig.  188)  and  northern  Chiapas, 
in  a pattern  strongly  suggesting  a circle  of  races.  I studied  35  specimens  of  this  species  from 
the  following  localities. 

UNITED  STATES 

TEXAS:  Uvalde  Co.,  Indian  Creek  Cave  (2;  TCBa);  Val  Verde  Co.,  9 mi.  se.  Del  Rio  (1;  USNM). 

MEXICO 

CHIAPAS:  31  mi.  w.  Lazaro  Cardenas  (1;  CNC).  COAHUILA:  14  mi.  n.  Saltillo  (1;  BMNH).  GUERRERO:  Rio  Balsas  (1; 
MCZ).  JALISCO:  9.7  mi.  e.  Encamacion  de  Diaz  (3;  UASM).  NUEVO  LEON:  5 mi.  s.  Monterrey  (2;  CNC).  OAXACA: 
11.1  mi.  n.  Matias  Romero  (1;  UASM),  Tapanatepec  (9;  UKSM).  SAN  LUIS  POTOSI:  El  Salto  de  Agua  (1;  CNC), 
Huichihuayan  (3;  UKSM),  17  ml  n.  Palitla  (1;  DRWh),  Rio  Verde  (1;  IRSB),  2 mi.  e.  Tamazunchale  (1;  UASM),  5. mi.  n. 
Tamazunchale  (1;  CAS),  19.3  mi.  nw.  Tamazunchale  (4;  DRWh,  MCZ,  UASM).  TAMAULIPAS:  73.1  mi.  n.  Manuel  (1; 
UASM).  VERACRUZ:  20  mi.  nw.  Huatusco  (1;  FDAG). 


Collecting  notes.  — Specimens  of  this  species  have  been  collected  nearly  throughout  the 
year.  Some  were  taken  at  lights.  Both  specimens  from  Indian  Creek  Cave  were  taken  in  a 
deep  cave,  and  at  that  locality  the  species  may  be  troglobitic.  Elsewhere  I suspect  individ- 
uals are  found  near  the  surface  only  incidentally,  and  live  at  a much  wider  range  of  depths 
in  gravel  bars  than  do  most  other  species  in  the  genus. 

Taxonomic  notes.  — The  only  other  species  known  to  have  broadened  front  tibiae,  S. 
dilatus,  is  sympatric  with  S.  tibialis  at  least  in  Nuevo  Leon,  and  lacks  marked  subepigean 
adaptations.  Specimens  of  S.  tibialis  have  more  strongly  constricted  front  femora,  and 
coarser  microsculpture  on  the  prothoracic  pleura. 


236 


Whitehead 


Specimens  from  Indian  Creek  Cave  may  represent  a truly  troglobitic  species,  reproduc- 
tively  isolated  from  the  more  southern  epigean  or  subepigean  forms.  Or,  they  may  represent 
a distinctive  subspecies,  since  they  are  visibly  distinct  in  eye  size,  and  geographically  are 
from  an  adjacent  drainage  system,  But  I think  the  eye  size  character  varies  in  a direct, 
though  steeply  stepped,  cline. 

Schizogenius  amphibius  Haldeman 

Clivina  amphibia  Haldeman  1843:299.  Type  locality  southeastern  Pennsylvania;  type  in 

MCZ,  specimen  labelled  “amphibius  2”  here  designated  lectotype  (!).  LeConte  1848: 
215. 

Schizogenius  amphibius,  LeConte  1857:83.  Putzeys  1863:24.  Putzeys  1866:224.  LeConte 

1879:34.  Lindroth  1961:168. 

Clivina  frontalis  LeConte  1848:215.  Type  locality  Westchester  County,  New  York;  type 

male  MCZ,  specimen  labelled  MCZ  5482  here  designated  lectotype  (!). 

Schizogenius  frontalis,  LeConte  1857:83,  suggested  synonymy. 

Diagnostic  combination.  — Specimens  of  this  species  are  easily  distinguished  from  all 
others  in  the  group  by  small  size  and  coarsely  punctate  prothoracic  pleura. 

Description.  — Body  flattened.  Color  testaceous,  without  metallic  luster. 

Integument.  Coarse  microsculpture  in  paramedian  frontal  sulci  and  on  sides  of  abdomen. 
Fine  but  conspicuous  microsculpture  on  mouth-parts,  genae,  front  legs  except  posterior 
surfaces  of  femora  and  trochanters,  middle  legs  except  anterior  surfaces  of  trochanters,  hind 
tibiae  and  posterior  surfaces  of  hind  femora,  elytral  epipleura,  and  middle  of  abdomen. 
Indistinct  microsculpture  sometimes  on  median  portion  of  frons  and  on  prothoracic  pleura. 
Prothoracic  pleura  coarsely  punctate. 

Head.  Fig.  162.  Paramedian  clypeal  carinae  straight,  extended  to  median  tooth,  strongly 
elevated  in  basal  half;  median  field  triangular,  no  wider  at  base  than  apex  of  median  field 
of  frons.  Clypeal  suture  sharply  defined.  Eye  small,  slightly  flattened,  finely  and  uni- 
formly faceted.  Neck  densely  and  coarsely  punctate.  Gena  coarsely  punctate,  rugose  in 
front.  Antennal  articles  four  to  ten  elongate,  filiform,  article  five  about  1.4  times  longer 
than  wide. 

Pronotum.  Form  and  sculpture,  Fig.  162.  Sides  bisetose,  hind  angles  sharp  and  promi- 
nent, base  not  rugose.  Paramedian  longitudinal  sulci  long,  impunctate,  nearly  straight,  deep 
throughout,  in  most  specimens  sharply  hooked  basally.  Anterior  transverse  impression  fine- 
ly punctate. 

Legs.  Front  tarsus  slightly  dilated  and  pubescent  ventrally  in  both  sexes;  middle  and  hind 
tarsi  slender,  short.  Paronychia  nearly  as  long  as  tarsal  claws.  Front  tibia  narrowed  evenly  to 
base.  Front  femur  not  strongly  constricted  near  apex. 

Elytra.  Left  elytron  with  9-14  setae  each  on  intervals  three  and  five,  and  5-10  on  interval 
seven,  adjacent  to  corresponding  inner  striae.  Striae  deep,  sharply  engraved,  finely  punctate 
in  basal  two-thirds.  Intervals  one  to  seven  broad  and  flat,  interval  eight  carinate  at  apex; 
apices  of  intervals  three,  five,  and  seven  broadly  joined.  Humeral  denticles  sharp  and  promi- 
nent. 

Abdomen.  Sternum  three  with  paramedian  carinae  curved  outward  at  apices.  Sternum 
seven  with  paramedian  ambulatory  setae  in  males  only.  Pygidium  with  apical  margin  entire 
in  both  sexes,  or  finely  crenulate  in  a few  females. 

Male  genitalia.  Median  lobe,  Fig.  174,  apex  not  sharply  deflexed;  endophallus  with  elon- 
gate basal  collar  spines;  6 specimens  examined. 

Measurements  and  proportions.  See  Table  29. 


Classification,  phylogeny,  and  zoogeography  of  Schizo genius 


237 


Table  29.  Descriptive  statistics  in  S.  amphibius,  based  on  10  males  from  Ithaca,  New  York. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

nts,  in  mm. 

TL 

3.18-3.38 

3.307 

0.101 

0.042 

2.03 

LE 

1.95-2.10 

2.045 

0.078 

0.033 

2.53 

WH 

0.62-0.68 

0.651 

0.024 

0.010 

2.45 

WP 

0.81-0.88 

0.840 

0.039 

0.016 

3.07 

WE 

0.99-1.05 

1.025 

0.039 

0.016 

2.51 

B.  Setae  on  lei 

ft  elytron. 

Interval  3 

11-14 

12.5 

Interval  5 

10-14 

11.5 

Interval  7 

6-10 

7.7 

Total 

29-36 

31.7 

4.0 

1.7 

8.42 

C.  Proportion; 

WF/WH 

0.68-0.71 

0.698 

0.021 

0.009 

2.00 

LP/WP 

0.93-0.98 

0.952 

0.024 

0.010 

1.70 

DP/LP 

0.76-0.81 

0.781 

0.024 

0.010 

2.05 

LP/WE 

0.76-0.81 

0.781 

0.029 

0.012 

2.45 

Ta/Ti 

0.55-0.65 

0.607 

0.045 

0.019 

4.91 

PS/LP 

0.69-0.78 

0.742 

0.039 

0.017 

3.52 

Variation . — I noted  no  obvious  important  variation,  and  because  the  only  good  popu- 
lation samples  are  clustered  in  a small  geographic  area,  I did  not  study  variation  in  detail. 
The  largest  (LE,  2.80  mm)  and  smallest  (LE,  1.80  mm)  specimens  are  females  from  New 
York. 

Etymology.  — For  S.  amphibius , Greek,  amphibios  = amphibius,  a reference  to  the  habi- 
tat. For  S.  frontalis,  Latin,  frons  = frons,  a reference  to  the  sculpture  of  the  frons. 

Distribution.  — S.  amphibius  is  known  to  range  from  Michigan  and  Quebec  south  to 
Tennessee  and  North  Carolina  (Fig.  190).  An  isolated  form  may  exist  in  Texas,  as  I have 
seen  four  specimens  from  there  but  without  definite  locality  data  (ANSP,  UKSM,  USNM), 
but  confirmation  is  needed;  no  other  specimens  are  known  from  west  of  the  Mississippi 
River.  I studied  188  additional  specimens  from  the  following  localities. 

CANADA 

QUEBEC:  Montreal  (1;  USNM).  See  Lindroth  (1961)  for  additional  records. 

UNITED  STATES 

No  locality  (6;  ANSP,  IRSB,  USNM).  CONNECTICUT:  Hartford  Co.,  West  Hartford  (1;  MCZ).  DISTRICT  OF  COLUM- 
BIA: Washington  (10;  AMNH,  ANSP,  MCZ,  USNM).  ILLINOIS:  Pike  Co.,  Pittsfield  (1;UCD).  INDIANA:  Tippecanoe  Co. 
(1;  UCD).  KENTUCKY:  Ballard  Co.,  Wickliffe  (2;  TCBa);  Jackson  Co.,  Sand  Gap  (1;  TCBa);  Metcalfe  Co.,  Edmonton  (2; 
DRWh);  Powell  Co.,  Slade  (1;  TCBa).  MAINE  (1;  MCZ);  Oxford  Co.,  Paris  (2;  MCZ).  MARYLAND  (4;  INHS,  USNM): 
Frederick  Co.,  Frederick  (1;  RTBe).  MICHIGAN  (2;  CAS).  NEW  HAMPSHIRE:  Grafton  Co.,  Franconia  (4;  AMNH,  MCZ). 
NEW  JERSEY;  Burlington  Co.,  Riverside  (1;  MCZ),  Camden  Co.,  Camden  (1;  MCZ),  Cramer  Hill  (1;  USNM);  Essex  Co., 
Irvington  (1;CAS).  NEW  YORK  (13; CAS,  CNHM,  INHS,  KSUM,  USNM):  New  York  City  vicinity  (5;  AMNH,  CAS,  MCZ); 
Long  Island  (1;  USNM);  Albany  Co.,  Altamont  (1;  CUNY);  Dutchess  Co.,  Hyde  Park  (4;  CAS);  Herkimer  Co.,  Newport  (1; 
MCZ);  Orange  Co.,  West  Point  (21;  USNM);  Tompkins  Co.,  Groton  (4;  CAS),  Ithaca  (20;  CAS,  CUNY,  USNM,  VMKi), 


238 


Whitehead 


Varna  (2;  UASM);  Ulster  Co.,  Esopus  (2;  CUNY),  Phoenicia  (3;  CAS,  CNHM);  Westchester  Co.,  Peekskill  (20;  CAS,  CUNY, 
MCZ,  PSUU),  Tarrytown  (3;  LACM);  Wyoming  Co.,  Pike  (2;  MCZ).  NORTH  CAROLINA:  Buncombe  Co.,  Black  Mount- 
ains (7;  AMNH,  CAS,  MCZ).  OHIO:  Mohican  Point  (1;  UMCG);  Ashtabula  Co.,  Ashtabula  (1 ; MSUL),  Jefferson  ( 1 ; MSUL). 
PENNSYLVANIA  (1;  ANSP):  Allegheny  Co.  (1;  MCZ);  Bucks  Co.,  Parkland  (3;  RUNB);  Philadelphia  Co.,  Frankford  (2; 
USNM);  Pike  Co.,  Milford  (1;  USNM);  Warren  Co.,  Warren  (3;  CAS);  Gray’s  Ferry  (2;  RUNB).  RHODE  ISLAND:  Provi- 
dence Co.,  Providence  (1;  USNM).  TENNESSEE:  Jackson  Co.,  Blackman  Fork  (2;  TCBa);  Smith  Co.,  Lancaster  (l;TCBa), 
Monoville  (1;  TCBa).  Vermont:  Franklin  Co.,  East  Georgia  (1;  RTBe);  Rutland  Co.,  Clarendon  (1;  USNM);  Windham  Co., 
Brattleboro  (2;  RTBe).  VIRGINIA:  Alexandria  Co.  (5;  USNM);  Fairfax  Co.,  Mount  Vernon  (1 ; USNM);  Spotsylvania  Co., 
Fredericksburg  (1;  USNM).  WEST  VIRGINIA:  Greenbrier  Co.,  White  Sulphur  Springs  (3;  CAS,  MCZ). 


Collecting  notes.  — Specimens  of  S.  amphibius  have  been  collected  from  April  until 
September,  generally  in  small  numbers.  I collected  two  specimens  near  Edmonton,  Ken- 
tucky, together  with  specimens  of  S.  sulcifrons,  S.  lineolatus , and  S.  planulatus.  But  a long 
series  of  this  species  only  was  taken  at  West  Point,  New  York,  so  habitat  requirements  of  S. 
amphibius  must  differ  in  some  way  from  those  of  other,  sympatric  species.  Its  range  is  more 
completely  restricted  to  limestone  regions  than  those  of  either  S.  sulcifrons  or  S.  lineolatus , 
and  reduced  eyes  suggest  a more  subterranean  habitat. 

Taxonomic  notes.  — Haldeman’s  types  are  supposed  to  be  in  the  LeConte  Collection,  at 
the  head  of  each  relevant  series.  However,  the  only  specimens  eligible  for  recognition  as 
type  material  of  S.  amphibius  are  specimens  2,  3,  4,  and  5,  each  so  labelled  and  from 
Pennsylvania.  The  first  specimen  in  the  series  is  the  only  specimen  representing  type  mate- 
rial of  S.  frontalis  LeConte,  and  is  the  lectotype.  I selected  specimen  2 as  lectotype  of 
amphibius. 

S.  amphibius  has  no  known  close  relatives,  nor  did  I find  any  interesting  patterns  of 
variation  within  the  species.  If,  however,  specimens  labelled  “Texas”  really  were  found 
there,  they  may  not  be  conspecific.  I doubt  that  S.  amphibius  will  be  found  in  any  area  be- 
tween Texas  and  the  Mississippi  River,  so  that  Texan  populations,  if  they  exist,  would  be 
markedly  disjunct. 

Schizogenius  planulatus  LeConte 

Schizogenius  planulatus  LeConte  1863:5.  Type  area  New  York:  type  in  MCZ;  female  la- 
belled MCZ  5481  here  designated  lectotype  (!).  Putzeys  1866:224.  LeConte  1879:34. 

Lindroth  1961:166. 

Diagnostic  combination.  - Body  strongly  flattened.  Dark  testaceous  to  castaneous,  un- 
metallic. 

Integument.  Conspicuous  microsculpture  on  paramedian  frontal  sulci,  mouthparts,  apex 
of  gula,  genae,  front  legs  except  posterior  surfaces  of  femora  and  trochanters,  middle  legs 
except  anterior  surfaces  of  trochanters,  hind  tibiae  and  posterior  surfaces  of  hind  femora, 
extreme  base  and  sides  of  pronotum,  apical  two-thirds  of  elytral  epipleura,  and  entire  abdo- 
men. Fine  microsculpture  on  basal  third  of  elytral  epipleura,  and  sometimes  on  prothoracic 
pleura  and  median  frontal  sulcus. 

Head.  Fig.  163.  Paramedian  clypeal  carinae  straight,  moderately  elevated  in  basal  half, 
extended  to  median  tooth;  median  field  triangular,  narrow,  no  wider  at  base  than  apex  of 
median  frontal  sulcus.  Clypeal  suture  sharply  defined.  Eye  small,  slightly  flattened,  finely 
and  uniformly  faceted.  Neck  densely  and  coarsely  punctate.  Gena  coarsely  punctate,  rugose 
in  front.  Antennal  articles  four  to  ten  elongate,  article  five  about  2.2  times  longer  than  wide. 

Pronotum.  Fig.  163.  Sides  bisetose;  base  not  rugose;  hind  angles  small.  Paramedian  lon- 
gitudinal sulci  long,  nearly  straight,  deep,  sharply  hooked  basally.  Anterior  transverse  im- 
pression finely  punctate. 

Legs.  Front  and  middle  tarsi  moderately  dilated  and  pubescent  ventrally,  less  so  in 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


239 


females;  hind  tarsus  slender,  short.  Paronychia  nearly  as  long  as  tarsal  claws.  Front  tibia 
narrowed  evenly  to  base.  Front  femur  not  strongly  constricted  near  apex. 

Elytra.  Left  elytron  with  10-14  setae  on  interval  three,  9-12  on  interval  five,  6-9  on  in- 
terval seven;  total  27-34  in  specimens  examined.  Striae  deep  and  sharply  engraved,  finely 
punctate  in  basal  three-fourths.  Intervals  one  to  seven  broad  and  flat,  interval  eight  carinate 
at  apex;  intervals  three,  five,  and  seven  broadly  joined  at  apices.  Humeral  denticles  sharp 
but  not  prominent. 

Abdomen.  Sternum  three  with  paramedian  carinae  curved  outward  at  apices.  Sternum 
seven  with  paired  paramedian  ambulatory  setae  in  males  only.  Pygidium  with  apical  margin 
strongly  serrate  in  females,  entire  in  males. 

Male  genitalia.  Median  lobe,  Fig.  175,  apex  characteristic;  endophallus,  Fig.  182;  5 speci- 
mens examined. 

Measurements  and  proportions.  See  Table  30. 


Table  30.  Descriptive  statistics  for  S.  planulatus,  based  on  1 1 females  plus  four  males  from 
New  York. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

nts,  in  mm. 

TL 

4.85-5.30 

5.133 

0.236 

0.081 

3.06 

LE 

3.00-3.28 

3.190 

0.128 

0.044 

2.67 

WH 

0.97-1.08 

1.042 

0.052 

0.018 

3.34 

WP 

1.28-1.42 

1.357 

0.067 

0.023 

3.30 

WE 

1.55-1.68 

1.629 

0.060 

0.021 

2.44 

B.  Setae  on  le: 

ft  elytron. 

Interval  3 

10-14 

11.6 

Interval  5 

9-12 

10.5 

Interval  7 

6-  9 

7.3 

Total 

27-34 

29.3 

3.0 

1.0 

6.78 

C.  Proportion: 

s. 

WF/WH 

0.66-0.71 

0.687 

0.023 

0.008 

2.25 

LP/WP 

0.88-0.94 

0.909 

0.026 

0.009 

1.88 

DP/LP 

0.74-0.81 

0.777 

0.028 

0.009 

2.37 

LP/WE 

0.71-0.78 

0.755 

0.026 

0.009 

2.29 

Ta/Ti 

0.53-0.65 

0.605 

0.049 

0.017 

5.42 

PS/LP 

0.69-0.74 

0.708 

0.020 

0.007 

1.92 

Etymology.  — Latin,  planus  = flat,  in  reference  to  body  form. 

Distribution.  — I have  seen  specimens  of  this  species  only  from  Kentucky,  New  York, 
and  West  Virginia  (Fig.  189)  but  it  has  been  reported  from  Georgia  (Fattig,  1949)  and 
probably  ranges  throughout  the  Appalachian  region  at  least  in  limestone  areas.  I studied  20 
specimens  from  the  following  localities. 


240 


Whitehead 


UNITED  STATES 

KENTUCKY:  Metcalfe  Co.,  9 mi.  e.  Edmonton  (3;  DRWh,  IRSB).  NEW  YORK:  Erie  Co.,  North  Evans  (2;  CAS,  MCZ); 
Tompkins  Co.,  Ithaca  (13;  CAS,  CUNY,  UASM),  Groton  (1;  JNeg).  WEST  VIRGINIA:  Greenbrier  Co.,  White  Sulphur 
Springs  (1;  MCZ). 


Collecting  notes.  — Specimens  of  this  species  have  been  collected  from  May  to  September. 
I collected  the  Kentucky  specimens  in  a gravel  bar,  along  with  specimens  of  S.  amphibius, 
S.  lineolatus,  and  S.  sulcifrons.  Specimens  of  this  species  probably  tend  to  be  subepigean,  as 
so  few  have  been  collected. 

Taxonomic  notes.  — Specimens  of  this  species  differ  radically  from  the  related  S.  ozarken- 
sis  and  S.  planuloides  in  form  of  male  genitalia,  but  otherwise  probably  are  indistinguisha- 
ble. As  I found  no  important  geographic  variation  in  form  of  male  genitalia  in  any  of  these 
species,  I conclude  that  form  of  genitalia  is  constant  and  that  the  three  geographic  segregates 
truly  represent  distinct  species.  See  discussion  for  S.  planuloides. 

Schizogenius  ozarkensis  new  species 

Type  material.  — Holotype  male  and  allotype  female  labelled  “5  mi.  n.  Stringtown  Atoka 
Co.  OKLAHOMA  7.IX.1964  Awram-Whitehead”  (MCZ).  24  additional  specimens  from  var- 
ious localities  in  Arkansas,  Missouri,  and  Oklahoma  are  paratypes  (BMNH,  CAS,  CNC, 
DRWh,  INHS,  IRSB,  UASM,  USNM). 

Diagnostic  combination.  — Males  of  this  species  differ  strikingly  from  those  of  S.  planu- 
latus  and  S.  planuloides  in  form  of  apex  of  median  lobe.  This  species  is  the  only  member  of 
the  tristriatus  group  known  from  the  Ozark  region,  north  of  the  Red  River  and  west  of  the 
Mississippi  River. 

Description.  — Superficially  as  in  S.  planulatus,  except  frontal  lobes  prominent.  Left 
elytron  with  10-14  setae  on  interval  three,  9-12  on  interval  five,  and  6-9  on  interval  seven; 
total  25-32.  Male  genitalia  with  median  lobe,  Fig.  176,  apex  characteristic;  endophallus, 
Fig.  183;  6 specimens  examined. 

Measurements  and  proportions.  See  Table  31.  Of  holotype:  TL,  4.74  mm;  LE,  2.92  mm; 
WH,  0.95  mm;  WP,  1.24  mm;  WE,  1.48  mm;  WF/WH,  0.68;  DP/LP,  0.74;  LP/WP,  0.93;  LP/ 
WE,  0.78;  PS/LP,  0.76;  Ta/Ti,  0.61.  Of  allotype:  TL,  4.68  mm;  LE,  2.90  mm;  WH,  0.94 
mm;  WP,  1.18  mm;  WE,  1.45  mm;  WF/WH,  0.68;  DP/LP,  0.76;  LP/WP,  0.96;  LP/WE,  0.81 ; 
PS/LP,  0.69;  Ta/Ti,  0.65. 

Etymology.  — I name  this  species  after  the  Ozark  Mountains,  as  this  species  is  known 
only  from  in  or  near  this  area. 

Distribution.  — S.  ozarkensis  is  known  from  only  a small  area  in  eastern  Oklahoma, 
western  Arkansas,  and  southern  Missouri  (Fig.  189).  I studied  27  specimens  from  the  fol- 
lowing localities. 


UNITED  STATES 

ARKANSAS:  Carroll  Co.,  5 mi.  w.  Berryville  (4;  BMNH,  CNC),  Eureka  Springs  (2;  UKSM);  Van  Buren  Co.,  Formosa  (2; 
IRSB);  Washington  Co.  (1;  INHS),  7 mi.  s.  Fayetteville  (8;  UASM).  MISSOURI:  Butler  Co.,  12  mi.  se.  Elsinore  (4;  CAS). 
OKLAHOMA:  Atoka  Co.,  5 mi.  n.  Stringtown  (5;  DRWh,  MCZ,  USNM);  Ottawa  Co.,  Wyandotte  (1;  MCZ). 


Collecting  notes.  — Adults  of  this  species  have  been  found  from  June  to  September,  in 
riparian  gravel  bars. 

Taxonomic  notes.  — Although  adults  of  this  species  cannot  be  distinguished  from  those 
of  S.  planulatus  and  S.  planuloides  by  external  morphology,  details  of  the  male  genitalia 
differ  radically  and  show  no  sign  of  intergradation.  Hence,  I think  that  S.  ozarkensis  is  a 
reproductive  isolate. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


241 


Table  31.  Descriptive  statistics  for  S.  ozarkensis,  based  on  10  females  and  seven  males  from 
northwestern  Arkansas. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

nts,  in  mm. 

TL 

4.50-5.16 

4.848 

0.249 

0.081 

3.43 

LE 

2.80-3.22 

3.006 

0.155 

0.050 

3.44 

WH 

0.90-1.04 

0.978 

0.059 

0.019 

4.03 

WP 

1.10-1.32 

1.225 

0.062 

0.020 

3.40 

WE 

1.38-1.58 

1.502 

0.076 

0.025 

3.37 

B.  Setae  on  left  elytron. 

Interval  3 

10-14 

11.2 

Interval  5 

9-12 

10.4 

Interval  7 

6-  9 

7.2 

Total 

26-32 

28.8 

3.5 

1.1 

8.01 

C.  Proportion: 

WF/WH 

0.64-0.74 

0.671 

0.038 

0.012 

3.76 

LP/WP 

0.92-0.98 

0.948 

0.024 

0.008 

1.69 

DP/LP 

0.73-0.77 

0.751 

0.017 

0.006 

1.52 

LP/WE 

0.74-0.80 

0.771 

0.027 

0.009 

2.29 

Ta/Ti 

0.60-0.68 

0.643 

0.041 

0.014 

4.28 

PS/LP 

0.66-0.74 

0.690 

0.033 

0.011 

3.16 

Schizogenius  planuloides  new  species 

Type  material.  — Holotype  male  labelled  “Cypress  Mills  7/1/88  Texas”  and  “2048” 
(USNM).  An  additional  19  specimens  from  various  localities  in  Texas  are  paratypes  (ANSP, 
CAS,  CNHM,  DHKa,  DRWh,  MSUL,  UASM,  UKSM,  USNM). 

Diagnostic  combination.  — Specimens  of  this  species  are  readily  distinguished  from  those 
of  the  related  S.  planulatus  and  S.  ozarkensis  only  by  form  of  male  genitalia  and  by  the 
allopatric  geographic  distribution. 

Description.  — Superficially  as  in  S.  planulatus,  except  eyes  in  most  specimens  more 
strongly  flattened  and  frontal  lobes  more  prominent.  Left  elytron  with  9-13  setae  on 
interval  three,  9-12  on  interval  five,  and  6-8  on  interval  seven;  total  26-31.  Male  geni- 
talia with  median  lobe,  Fig.  177,  apex  characteristic;  endophallus,  Fig.  184;  3 specimens 
examined. 

Measurements  and  proportions.  See  Table  32.  Of  holotype:  TL,  5.20  mm;  LE,  3.20  mm; 
WH,  1.06  mm;  WP,  1.34  mm;  WE,  1.69  mm;  WF/WH,  0.71;DP/LP,  0.75;  LP/WP,  0.95;  LP/ 
WE,  0.76;  PS/LP,  0.69;  Ta/Ti,  0.70. 

The  claw-bearing  article  of  the  left  hind  tarsus  of  the  type  is  lacking. 

Etymology.  — Latin,  planulatus  + oides  = like  planulatus,  in  reference  to  the  great  exter- 
nal similarity  of  this  species  to  S.  planulatus. 

Distribution.  — This  species  is  known  from  only  six  definite  localities  throughout  much 
of  Texas  (Fig.  189).  Specimens  labelled  simply  “Texas”  may  well  have  been  collected  at  the 


242 


Whitehead 


type  locality,  since  that  was  a favorite  early  collecting  locality  in  Texas.  I studied  22  speci- 
mens of  this  species  from  the  following  localities. 

UNITED  STATES 

TEXAS  (15;  ANSP,  CAS,  MSUL,  UKSM,  USNM):  Blanco  Co.,  Cypress  Mills  (1;  USNM);  Coryell  Co.  (1;  MCZ);  Dallas  Co., 
Dallas  (1;  MCZ);  Kinney  Co.,  23  mi.  sw.  Brackettville  (1;  DRWh);  McLennon  Co.,  Waco  (1;  DHKa);  Taylor  Co.,  25  mi. 
sw.  Abilene  (1;  CNHM);  Terrell  Co.,  Independence  Creek  (1;  UASM). 


Table  32.  Descriptive  statistics  for  S.  planuloides , based  on  six  females  and  12  males  from 
Texas. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

nts,  in  mm. 

TL 

4.94-5.54 

5.194 

0.244 

0.077 

3.13 

LE 

3.05-3.45 

3.204 

0.160 

0.050 

3.33 

WH 

1.02-1.15 

1.084 

0.050 

0.016 

3.07 

WP 

1.22-1.40 

1.322 

0.076 

0.024 

3.82 

WE 

1.50-1.75 

0.628 

0.095 

0.030 

3.88 

B.  Setae  on  lei 

~t  elytron. 

Interval  3 

9-13 

11.0 

Interval  5 

9-12 

10.4 

Interval  7 

6-  8 

6.9 

Total 

26-31 

28.3 

2.3 

0.7 

5.25 

C.  Proportions 

WF/WH 

0.68-0.73 

0.703 

0.022 

0.007 

2.13 

LP/WP 

0.93-0.98 

0.956 

0.021 

0.007 

1.49 

DP/LP 

0.72-0.78 

0.752 

0.024 

0.008 

2.16 

LP/WE 

0.75-0.79 

0.776 

0.019 

0.006 

1.67 

Ta/Ti 

0.63-0.68 

0.645 

0.021 

0.007 

2.19 

PS/LP 

0.64-0.73 

0.675 

0.033 

0.011 

3.30 

Collecting  notes.  — Specimens  of  this  species  have  been  collected  from  March  through 
July,  some  at  lights  distant  from  water.  I collected  one  specimen  near  Brackettville  in  a 
riparian  gravel  bar,  along  with  several  specimens  of  S.  scopaeus  and  over  100  of  S.  sallei. 
Scarcity  of  specimens,  and  more  strongly  reduced  eyes,  suggest  that  this  species  is  more 
strongly  subepigean  than  either  S.  planulatus  or  S.  ozarkensis.  As  in  those  species,  I suspect 
that  geographic  distribution  depends  on  surface  and  subsurface  limestone. 

Taxonomic  notes.  — The  well  differentiated  male  genitalia  of  this  species  indicates  re- 
productive isolation  from  S.  planulatus  and  S.  ozarkensis.  The  geographically  intermediate 
S.  ozarkensis  has  less  specialized  male  genitalia,  but  the  genitalia  of  S.  planulatus  and  S. 
planuloides  are  specialized  in  different  ways.  As  shown  in  Tables  30-32,  the  sample  of  S. 
planulatus  is  characterized  by  statistically  significantly  broader  pronota,  the  sample  of  S. 
ozarkensis  by  statistically  significantly  smaller  body  size,  and  the  sample  of  S.  planuloides 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


243 


Fig.  158-163.  Head  and  pronotum,  dorsal  aspect.  158.  S.  sallei  Putzeys,  Comstock,  Texas.  159.  S.  tristriatus  Putzeys, 
Coscomatepec,  Veracruz.  160.  S.  tibialis  new  species,  Tamazunchale,  San  Luis  Potosi.  161.  Same,  Indian  Creek  Cave, 
Texas.  162.  S.  amphibius  Haldeman,  Edmonton,  Kentucky.  163.  S.  planulatus  LeConte,  Edmonton,  Kentucky.  Fig.  164- 
166.  Left  front  tibia,  posterior  aspect.  164.  S.  tristriatus  Putzeys,  Coscomatepec,  Veracruz.  165.  S.  dilatus  new  species, 
Sabinas  Hidalgo,  Nuevo  Leon.  166.  S.  tibialis  new  species,  Tamazunchale,  San  Luis  Potosi.  Fig.  167-168.  Distal  portion 
of  left  front  tibia,  posterior  aspect.  167.  S.  tibialis  new  species,  Tamazunchale,  San  Luis  Potosi.  168.  Same,  Indian 
Creek  Cave,  Texas.  Fig.  169-173.  Male  median  lobe,  lateral  aspect.  169.  S.  sallei  Putzeys,  Austin,  Texas.  170.  S.  tristri- 
atus Putzeys,  Fortin  de  las  Flores,  Veracruz.  171.  S.  dilatus  new  species,  Sabinas  Hidalgo,  Nuevo  Leon.  172.  S.  tibialis 
new  species,  Encarnacion  de  Diaz,  Jalisco.  173.  Same,  Indian  Creek  Cave,  Texas. 


244 


Whitehead 


Fig.  174-177.  Male  median  lobe,  lateral  aspect.  174.  S.  amphibius  Haldeman,  West  Point,  New  York.  175.  S.  planulatus 
LeConte,  North  Evans,  New  York.  176.  S.  ozarkensis  new  species,  Elsinore,  Missouri.  177.  S.  planuloides  new  species, 
Texas.  Fig.  178-184.  Male  endophallus.  178.  S.  sallei  Putzeys,  Sabinal,  Texas.  179.  S.  tristriatus  Putzeys,  Fortin  de  las 
Flores,  Veracruz.  180.  S.  dilatus  new  species,  Sabinas  Hidalgo,  Nuevo  Leon.  181.  S.  tibialis  new  species,  Tapanatepec, 
Oaxaca.  182.  S.  planulatus  LeConte,  Ithaca,  New  York.  183.  S.  ozarkensis  new  species,  Stringtown,  Oklahoma.  184. 
S.  planuloides  new  species,  Cypress  Mills,  Texas. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


245 


185 


188 


WF  / WH 
O 0.63-0.65 
0 0.66-0.68 
3 0.69-0.71 

• 0.76-0.78 


1000  mi 


\ ,1  , disuibu«0"s-  185.  S.  zallei  Putzeys.  186.  S.  Mstriams  Putzeys.  187.  5.  dilatus  new  species.  188 

' b'al'S  "eW  spec,es  (with  cUnal  ''ariation  in  relative  eye  size).  189.  S.  plcmutatus  LeConte,  circles;  S.  ozarkensis  new 
species,  triangles;  S.  planulmdes  new  species,  squares.  190.  S.  amphibius  Haldeman;  open  symbol  represents  state  record 


246 


Whitehead 


by  statistically  significantly  smaller  eye  size.  As  judged  from  comparisons  with  other  mem- 
bers of  the  tristriatus  group,  the  ancestor  of  the  planulatus  subgroup  probably  had  larger 
eyes  as  in  S.  planulatus  and  S.  ozarkensis,  smaller  size  as  in  S.  ozarkensis,  and  narrower 
pronota  as  in  S.  ozarkensis  and  S.  planuloides.  In  all  of  these  ways,  S.  ozarkensis  is  most 
like  the  probable  ancestor  of  the  subgroup. 

Since  I have  no  females  from  the  type  locality,  I designate  no  allotype  for  S.  planu- 
loides. 


The  lineolatus  group 

Diagnostic  combination.  — From  other  members  of  the  genus  with  antennae  elongate, 
pronotal  hind  angles  prominent,  discal  setae  on  intervals  three,  five,  and  seven,  and  para- 
median sternal  carinae  curved  at  apices,  members  of  this  group  are  distinguished  by  the 
following  combination  of  characters:  body  strongly  flattened;  elytra  aeneopiceous,  or  rufo- 
castaneous  with  pale  apices;  paramedian  pronotal  sulci  not  extended  forward  nearly  to 
anterior  transverse  impression;  elytron  with  total  discal  setae  fewer  than  30,  their  average 
length  less  than  1.0  times  maximum  width  of  interval  two;  and  abdomen  unmicrosculp- 
tured  except  in  small  patches  near  coxal  depressions  of  sternum  three.  Also  paramedian 
clypeal  carinae  extended  to  median  tooth;  clypeal  field  narrow,  no  wider  at  base  than  apex 
of  median  frontal  sulcus;  clypeal  suture  sharply  impressed;  frontal  carinae  not  confused  at 
base;  submentum  without  accessory  setae;  pronotum  without  paralateral  carinae,  without 
accessory  marginal  setae;  front  and  middle  tarsi  broadened  and  with  dense  ventral  pubes- 
cence, especially  in  males;  sternum  seven  with  paramedian  ambulatory  setae  in  males,  not 
in  females;  pygidium  crenulate  at  apex  or  not  in  females;  and  endophallus  with  basal  collar 
spines  distinct. 

Distribution.  — The  range  of  the  single  included  species,  S.  lineolatus , covers  much  of 
eastern  North  America  south  to  the  Rio  Grande.  I examined  897  specimens. 

Schizogenius  lineolatus  Say 

Clivina  lineolata  Say  1823:22.  Type  locality  Allegheny,  Pennsylvania  (designated  by  Lin- 

droth,  1961);  neotype  male  designated  by  Lindroth  and  Freitag  (1969).  LeConte  1848: 
214. 

Schizogenius  lineolatus , LeConte  1857:82.  Putzeys  1863:24.  Putzeys  1866:228.  LeConte 

1879:34.  Lindroth  1961:166. 

Diagnostic  combination.  — Specimens  of  this  species  are  best  distinguished  by  character- 
istics given  in  the  key.  In  eastern  North  America,  the  only  other  dark  species  of  Schizogenius 
with  setae  on  elytral  interval  seven  is  S.  sulcifrons.  Specimens  of  that  species  generally  have 
less  sharply  developed  pronotal  hind  angles,  more  elytral  setae,  and  concolorous  elytra  and 
pronotum. 

Description.  - Body  flattened.  Color  light  to  dark  castaneous;  legs  rufous;  tarsi  and  palpi 
testaceous; elytra  rufopiceous  and  in  most  specimens  strongly  aeneous,  except  in  Rio  Grande 
Valley  where  castaneous,  paler  toward  apices,  and  weakly  aeneous. 

Integument.  Conspicuous  microsculpture  on  paramedian  frontal  sulci,  mouthparts,  genae, 
front  legs  except  posterior  surfaces  of  femora  and  trochanters,  middle  legs  except  anterior 
surfaces  of  trochanters,  hind  tibiae  and  posterior  surfaces  of  hind  femora,  elytral  epipleura 
on  apical  two-thirds  and  in  most  eastern  specimens  on  shoulder  and  on  basal  third,  and 
sternum  three  in  coxal  depressions  and  small  paralateral  patches. 

Head.  Fig.  191.  Paramedian  clypeal  carinae  straight,  moderately  elevated  in  basal  half, 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


247 


extended  to  median  tooth;  median  field  triangular,  narrow,  no  wider  at  base  than  apex  of 
median  frontal  sulcus.  Clypeal  suture  sharply  defined.  Eye  prominent,  finely  and  uniformly 
faceted.  Neck  densely  punctate.  Gena  coarsely  punctate,  rugose  in  front.  Antennal  articles 
four  to  ten  elongate,  article  five  about  1.7-1 .8  times  longer  than  wide. 

Pronotum.  Fig.  191.  Sides  bisetose;  base  not  rugose;  hind  angles  prominent.  Paramedian 
longitudinal  sulci  quite  long,  nearly  straight,  strongly  hooked  basally.  Paralateral  carinae 
absent.  Anterior  transverse  impression  strongly  punctate  in  most  specimens. 

Legs.  Front  and  middle  tarsi  moderately  dilated  and  pubescent  ventrally,  less  so  in  fe- 
males; hind  tarsus  slender,  short.  Paronychia  nearly  as  long  as  tarsal  claws.  Front  tibia 
narrowed  evenly  to  base.  Front  femur  not  strongly  constricted  near  apex. 

Elytra.  Left  elytron  with  about  six  to  nine  setae  on  interval  three,  five  to  eight  on  interval 
five,  two  to  five  on  interval  seven;  total  13-20  in  specimens  examined.  Striae  deep  and 
sharply  engraved,  finely  punctate  in  basal  two-thirds.  Intervals  one  to  seven  broad  and  flat, 
interval  eight  carinate  at  apex;  intervals  three,  five,  and  seven  broadly  joined  at  apices. 
Humeral  denticles  prominent. 

Abdomen.  Sternum  three  with  paramedian  carinae  curved  outward  at  apices.  Sternum 
seven  with  paramedian  ambulatory  setae  in  males  only.  Pygidium  with  apical  margin  crenu- 
late  in  some  females,  entire  in  males. 

Male  genitalia.  Median  lobe,  Fig.  196;  endophallus,  Fig.  201,  virga  small;  20  specimens 
examined. 

Measurements  and  proportions.  See  Table  33. 


Table  33.  Descriptive  statistics  for  S.  lineolatus,  based  on  20  males  from  5 miles  north  of 
Stringtown,  Oklahoma. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

mts,  in  mm. 

TL 

3.80-4.44 

4.412 

0.288 

0.077 

4.15 

LE 

2.35-2.74 

2.558 

0.153 

0.046 

4.00 

WH 

0.81-0.95 

0.875 

0.052 

0.016 

3.97 

WP 

1.00-1.20 

1.092 

0.083 

0.025 

5.08 

WE 

1.20-1.42 

1.324 

0.084 

0.025 

4.22 

B.  Setae  on  le 

ft  elytron. 

Interval  3 

6-  8 

6.8 

Interval  5 

5-  8 

6.2 

Interval  7 

2-  4 

3.1 

Total 

14-19 

16.1 

1.9 

0.6 

7.78 

C.  Proportion 

s. 

WF/WH 

0.55-0.59 

0.567 

0.017 

0.005 

1.99 

LP/WP 

0.87-0.94 

0.910 

0.021 

0.006 

1.55 

DP/LP 

0.80-0.86 

0.830 

0.024 

0.007 

1.93 

LP/WE 

0.73-0.77 

0.749 

0.019 

0.006 

1.73 

Ta/Ti 

0.60-0.69 

0.643 

0.037 

0.011 

3.85 

PS/LP 

0.76-0.82 

0.786 

0.025 

0.007 

2.08 

248 


Whitehead 


Variation.  — Statistical  data  on  variation  in  body  size,  numbers  of  elytral  setae,  relative 
eye  size,  and  pronotal  form  are  given  in  Tables  34-37  for  17  samples  of  S.  lineolatus  (Fig. 
21 1-214).  Body  size  (Table  34,  Fig.  211)  decreases  quite  regularly  from  northeast  to  south- 
west, with  no  geographically  proximate  samples  statistically  significantly  distinct  from  one 
another.  Numbers  of  elytral  setae  (Table  35,  Fig.  212)  tend  to  increase  from  northeast  to 
southwest,  and  the  Rio  Grande  sample  (number  17)  barely  is  statistically  significantly  dis- 
tinct from  the  Guadalupe  River  sample  (number  16).  Eye  size  (Table  36,  Fig.  213)  increases 
generally  southwest  to  the  Guadalupe  River  (sample  16),  with  a statistically  significant  in- 
crease in  the  east  between  samples  four  and  five,  but  is  sharply  reduced  between  samples  16 
and  17.  The  pronotum  (Table  37,  Fig.  214)  narrows  from  northeast  to  southwest,  with 
statistically  significant  gaps  between  the  Red  River  (sample  14),  the  Guadalupe  River  (sam- 
ple 16),  and  the  Rio  Grande  (sample  17). 

Variation  in  extent  of  evident  microsculpture  on  basal  third  of  elytral  epipleura  and  on 
shoulder  follows  a pattern  of  gradual  reduction  from  east  to  west.  Microsculpture  on  the 
basal  third  of  the  epipleura  is  present  on  most  specimens  from  east  of  the  Appalachians, 
absent  from  most  specimens  from  west  of  the  Mississippi  River,  and  varied  in  specimens 
from  intermediate  areas.  Microsculpture  is  evident  on  the  shoulder  of  most  specimens  from 
all  areas  except  in  many  from  the  most  western  localities  such  as  in  western  Oklahoma  and 
eastern  Wyoming,  but  is  most  strongly  developed  in  eastern  specimens. 

Specimens  from  the  Rio  Grande  Valley  (sample  17)  differ  from  all  others,  except  some 
teneral  specimens,  in  elytral  coloration:  reddish  rather  than  blackish,  and  paler  toward 
apices.  In  this  characteristic,  these  specimens  do  not  intergrade  with  those  from  the  nearest 
localities  where  S.  lineolatus  is  known  to  exist,  in  the  Guadalupe  River  system  (sample  16) 
some  125-150  miles  distant.  These  two  samples  also  are  statistically  distinct  in  the  number 
of  elytral  setae,  relative  eye  size,  and  pronotal  form,  but  are  not  distinct  in  epipleural  micro- 
sculpture  or  in  body  size.  Lack  of  differentiation  in  some  characteristics,  evidence  of  clinal 
continuity  in  others,  and  in  particular  the  fine  clinal  sequence  in  variation  in  pronotal  form 
shown  by  samples  14-16-17  strongly  indicate  that  the  Rio  Grande  population  is  not  repro- 
ductively  isolated.  But  the  difference  in  coloration  is  constant,  statistically  significant  dif- 
ferences exist  for  numbers  of  elytral  setae,  relative  eye  size,  and  pronotal  form,  and  the 
direction  of  clinal  variation  is  reversed  in  the  relative  eye  size  characteristic.  I conclude  from 
these  facts  that  the  Rio  Grande  form  is  geographically  isolated  from  populations  to  the 
north,  and  predict  that  truly  intermediate  populations  do  not  exist. 

Etymology.  — Latin,  lineola  = fine  line,  probably  in  reference  to  the  presence  of  para- 
median pronotal  sulci. 

Distribution.  — Four  specimens  are  labelled  “Fla.”  I doubt  that  S.  lineolatus  really  occurs 
in  Florida,  as  I think  there  is  no  suitable  habitat  there.  Otherwise,  the  known  range  of  this 
species  extends  west  from  southern  Canada  and  eastern  United  States  to  at  least  eastern 
Wyoming,  and  south  to  the  Rio  Grande  Valley  (Fig.  206).  I studied  893  additional  speci- 
mens from  the  following  localities. 


CANADA 

No  locality  (5;  ANSP,  CNHM,  MCZ,  UKSM).  ONTARIO:  16  mi.  w.  Bondhead  (1;  UASM),  London  (20;CNHM,  UASM), 
Ottawa  (1;  TLEr),  Port  Credit  (1;  CUNY),  Port  Stanley  (1;  UASM),  Saint  Catherines  (3;  JNeg),  Toronto  (5;  MCZ,  UASM), 
Wheatley  (1;  KHSt).  QUEBEC:  Montreal  (1;  MCZ),  Potton  Springs  (10;  MCZ).  For  other  Canadian  records,  see  Lindroth 
(1961). 


UNITED  STATES 

No  locality  (8;  ANSP,  CUNY.  IRSB,  USNM).  ALABAMA:  Tuscaloosa  Co.,  15  mi.  n.  Tuscaloosa  (1 1 ; DJLa).  ARKANSAS: 
Carroll  Co.,  Berryville  (1;  MCZ),  4 mi.  w.  Berryville  (27;  UASM);  Newton  Co.,  Jasper  (1;  BMNH);  Sevier  Co.,  6 mi.  n. 
Lockesburg  (1;  MCZ);  Van  Buren  Co.,  Formosa  (2;  IRSB);  Washington  Co.  (2;  INHS),  7 mi.  s.  Fayetteville  (7;  UASM). 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


249 


Table  34.  Variation  in  body  size  (LE,  in  mm)  in  selected  samples  of  S.  lineolatus',  see  Fig. 
211. 


Sample 

N 

Males 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

1 

10 

5 

2.82-3.08 

2.902 

0.127 

0.053 

2.91 

2 

10 

5 

2.65-3.08 

2.844 

0.218 

0.092 

5.12 

3 

18 

9 

2.55-3.15 

2.828 

0.197 

0.062 

4.68 

4 

12 

6 

2.58-2.98 

2.813 

0.212 

0.082 

5.03 

5 

28 

14 

2.40-3.04 

2.735 

0.252 

0.064 

6.14 

6 

16 

8 

2.55-2.95 

2.756 

0.178 

0.069 

4.30 

7 

12 

6 

2.62-3.02 

2.802 

0.178 

0.069 

4.24 

8 

12 

6 

2.55-3.00 

2.750 

0.184 

0.071 

4.46 

9 

10 

5 

2.40-2.92 

2.663 

0.237 

0.100 

5.93 

10 

10 

5 

2.55-2.98 

2.766 

0.199 

0.084 

4.81 

11 

14 

7 

2.45-2.90 

2.643 

0.189 

0.067 

4.76 

12 

14 

7 

2.48-2.88 

2.677 

0.156 

0.056 

3.90 

13 

22 

11 

2.42-3.00 

2.703 

0.200 

0.057 

4.94 

14 

26 

13 

2.35-2.85 

2.629 

0.197 

0.052 

5.00 

15 

16 

8 

2.38-2.98 

2.619 

0.273 

0.091 

6.96 

16 

18 

9 

2.38-2.85 

2.616 

0.206 

0.065 

5.26 

17 

20 

10 

2.32-2.88 

2.630 

0.195 

0.058 

4.94 

Table  35.  Variation  in  numbers  of  discal  setae  on  left  elytron  in  selected  samples  of  S. 
lineolatus',  see  Fig.  212. 


Sample 

N 

Males 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

1 

10 

5 

13-17 

15.3 

1.7 

0.7 

7.58 

2 

10 

5 

16-19 

16.6 

1.6 

0.7 

6.48 

3 

18 

9 

15-19 

15.9 

1.6 

0.5 

6.62 

4 

12 

6 

15-17 

16.2 

1.3 

0.5 

5.16 

5 

28 

14 

14-17 

15.6 

1.2 

0.3 

5.04 

6 

16 

8 

15-17 

15.6 

1.2 

0.4 

5.23 

7 

12 

6 

15-18 

15.9 

1.5 

0.7 

6.26 

8 

12 

6 

13-17 

15.5 

1.9 

0.7 

8.02 

9 

10 

5 

14-17 

15.6 

1.3 

0.5 

5.41 

10 

10 

5 

15-18 

16.0 

1.6 

0.7 

6.58 

11 

14 

7 

15-20 

17.1 

1.5 

0.9 

9.60 

12 

14 

7 

14-17 

16.1 

1.1 

0.4 

4.77 

13 

22 

11 

14-19 

16.3 

1.7 

0.5 

7.17 

14 

26 

13 

14-18 

16.2 

1.5 

0.4 

6.30 

15 

16 

8 

15-19 

16.5 

1.5 

0.5 

6.26 

16 

18 

9 

14-18 

16.0 

1.8 

0.6 

7.43 

17 

20 

10 

16-19 

17.2 

1.4 

0.4 

5.28 

250 


Whitehead 


Table  36.  Variation  in  eye  size  (WF/WH)  in  selected  samples  of  S.  lineolatus ; see  Fig. 
213. 


Sample 

N 

Males 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

1 

10 

5 

0.59-0.63 

0.614 

0.023 

0.010 

2.57 

2 

10 

5 

0.60-0.62 

0.612 

0.012 

0.005 

1.29 

3 

18 

9 

0.58-0.63 

0.607 

0.021 

0.006 

2.26 

4 

12 

6 

0.58-0.62 

0.601 

0.016 

0.006 

1.80 

5 

28 

14 

0.56-0.61 

0.590 

0.016 

0.004 

1.87 

6 

16 

8 

0.58-0.61 

0.592 

0.017 

0.006 

1.87 

7 

12 

6 

0.57-0.60 

0.582 

0.015 

0.006 

1.77 

8 

12 

6 

0.56-0.60 

0.579 

0.020 

0.008 

2.26 

9 

10 

5 

0.56-0.60 

0.579 

0.022 

0.009 

2.50 

10 

10 

5 

0.56-0.59 

0.577 

0.014 

0.006 

1.64 

11 

14 

7 

0.54-0.59 

0.579 

0.023 

0.008 

2.70 

12 

14 

7 

0.55-0.59 

0.574 

0.018 

0.006 

2.12 

13 

22 

11 

0.56-0.59 

0.576 

0.015 

0.004 

1.75 

14 

26 

13 

0.55-0.59 

0.569 

0.017 

0.004 

1.95 

15 

16 

8 

0.55-0.58 

0.565 

0.015 

0.005 

1.83 

16 

18 

9 

0.54-0.60 

0.564 

0.020 

0.006 

2.37 

17 

20 

10 

0.57-0.60 

0.579 

0.015 

0.004 

1.67 

Table  37.  Variation  in  pronotal  form  (LP/WP)  in  selected  samples  of  S.  lineolatus ; see  Fig. 
214. 


Sample 

N 

Males 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

1 

10 

5 

0.86-0.90 

0.880 

0.022 

0.009 

1.69 

9 

10 

5 

0.84-0.91 

0.880 

0.030 

0.012 

2.14 

3 

18 

9 

0.85-0.91 

0.882 

0.025 

0.008 

1.92 

4 

12 

6 

0.84-0.94 

0.883 

0.041 

0.016 

3.07 

5 

28 

14 

0.85-0.93 

0.888 

0.015 

0.006 

1.87 

6 

16 

8 

0.86-0.93 

0.886 

0.023 

0.008 

1.74 

7 

12 

6 

0.86-0.94 

0.898 

0.033 

0.013 

2.46 

8 

12 

6 

0.84-0.92 

0.882 

0.037 

0.014 

2.82 

9 

10 

5 

0.87-0.94 

0.899 

0.035 

0.015 

2.59 

10 

10 

5 

0.87-0.91 

0.885 

0.019 

0.008 

1.43 

11 

14 

V 

0.88-0.93 

0.901 

0.023 

0.008 

1.73 

12 

14 

7 

0.88-0.94 

0.909 

0.026 

0.009 

1.90 

13 

22 

! 11 

0.86-0.93 

0.903 

0.030 

0.008 

2.19 

14 

26 

13 

0.87-0.93 

0.902 

0.025 

0.007 

1.85 

15 

16 

8 

0.89-0.95 

0.912 

0.022 

0.007 

1.63 

16 

18 

9 

0.88-0.94 

0.922 

0.023 

0.007 

1.64 

17 

20 

10 

0.90-0.98 

0.940 

0.031 

0.009 

2.18 

Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


251 


DISTRICT  OF  COLUMBIA:  Washington  (16;  AMNH,  CAS,  MCZ,  MSUL,  USNM).  ILLINOIS  (8;  INHS,  MCZ,  USNM). 
Cook  Co.,  Palos  Park  (1;CNHM);  La  Salle  Co.,  Ottawa  (1 ; RTBe);  McLean  Co.,  Normal  (1;  INHS);  Peoria  Co.,  East  Peoria 
(2;  CNHM);  Vermilion  Co.,  Fairmount  (1;  RTBe).  INDIANA  (1;  MSUL):  Parke  Co.  (9;  UCD),  The  Shades  State  Park  (3; 
RTBe);  Tippicanoe  Co.  (2;  MCZ),  1 mi.  ne.  Lafayette  (1;  GRNo),  Wabash  River  (2;  CAS).  IOWA:  Herrold  (1;  USNM); 
Boone  Co.,  Boone  (6;  UASM);  Cedar  Co.  (1;  USNM);  Johnson  Co.,  Iowa  City  (1;  USNM).  KANSAS:  Bourbon  Co.,  Fort 
Scott  (2;  USNM);  Douglas  Co.,  Lawrence  (1;  DJLa);  Pottawatomie  Co.,  Onaga  (5;  KSUM,  USNM);  Riley  Co.,  Manhattan 
(1;  KSUM);  Wilson  Co.,  Benadict  (2;  CAS).  KENTUCKY:  Ballard  Co.,  Wickliffe  (9;  TCBa);  Cumberland  Co.,  Marrowbone 
Creek  (7;  TCBa);  Green  Co.,  Greasy  Creek  (1;  TCBa);  Jackson  Co.,  Sand  Gap  (2;  TCBa);  Jessamine  Co.,  Indian  Falls  (2; 
TCBa);  Metcalfe  Co.,  8 mi.  e.  Edmonton  (20;  UASM);  Powell  Co.,  Slade  (2;  TCBa);  Hockcastle  Co.,  Crooked  Creek  (1; 
TCBa).  MARYLAND  (3;  CAS,  MSUL,  USNM):  Difficult  (1;  MCZ);  Baltimore  Co.,  Baltimore  (2;  CAS);  Frederick  Co., 
Frederick  (7;  DRWh,  RTBe);  Harford  Co.,  Edgewood  (7;  DRWh);  Prince  Georges  Co.,  Bladensburg  (1;  CUNY).  MISSOURI: 
Barry  Co.,  Cassville  (1;  USNM);  Boone  Co.,  Columbia  (3;  USNM);  Crawford  Co.,  Meramec  River  (2;  USNM);  Laclede  Co., 
Bennet  Springs  (7;  USNM);  McDonald  Co.,  3 mi.  n.  Noel  (3;  CAS);  Ozark  Co.,  Gainesville  (18;  UASM);  Reynolds  Co., 
Bunker  (1;  USNM);  Ripley  Co.,  Buffalo  Creek  (1;  CAS),  Doniphan  (2;  USNM);  Saint  Louis  Co.,  Saint  Louis  (2;  UMCG); 
Taney  Co.,  5 mi.  e.  Forsythe  (2;  MGFT);  Wright  Co.,  Mountain  Grove  (14;  TCBa).  MONTANA:  Roosevelt  Co.,  Wolf  Point 
(1;  CAS).  NEW  JERSEY  (2;  ANSP):  Bergen  Co.,  Westwood  (1;  CAS);  Camden  Co.,  Camden  (9;  CAS,  LACM,  RUNB), 
Cramer  Hill  (1;  USNM);  Essex  Co.,  Irvington  (2;  AMNH);  Gloucester  Co.,  Westville  (2;  MCZ);  Middlesex  Co.,  Jamesburg 
(3;  CUNY);  Morris  Co.,  Boonton  (6;  USNM);  Somerset  Co.  (1;  USNM);  Warren  Co.,  Phillipsburg  (11;  CAS,  MSUL).  NEW 
YORK  (5;  CAS,  CNHM,  INHS,  USNM);  New  York  City  (2;  CAS);  West  Hebron  (3;  CAS);  Albany  Co.,  Altamont  (1; 
CUNY);  Broome  Co.,  Chenango  Valley  State  Park  (13;  GRNo);  Cattaraugus  Co.,  Allegany  State  Park  (1;  USNM);  Dutchess 
Co.,  Fishkill  (6;  CAS);  Erie  Co.,  Buffalo  (2;  CAS),  North  Evans  (3;  CAS);  Greene  Co.,  Ashland  (4;  CUNY,  MCZ);  Rockland 
Co.,  Hillbum  (2;  USNM);  Tompkins  Co.,  Groton  (7;  JNeg,  UATA),  Ithaca  (28;  CAS,  CUNY,  FDAG,  USNM,  VMKi); 
Ulster  Co.,  Phoenicia  (1;  CAS);  Wyoming  Co.,  Pike  (2;  MCZ).  NORTH  CAROLINA:  (2;  MCZ)  Buncombe  Co.,  Black 
Mountains  (1;  CAS).  OHIO:  Mohican  Point  (3;  UMCG);  Ashtabula  Co.,  Ashtabula  (2;  MSUL),  Conneaut  (1;  MSUL),  Rock 
Creek  (1;  MSUL);  Athens  Co.,  Athens  (1;  UATA);  Crawford  Co.,  Plankton  (1;  CAS);  Cuyahoga  Co.,  Cleveland  (1;  FDAG); 
Knox  Co.,  Gambier  (1;  UMCG);  Preble  Co.,  West  Alexandria  (1;  RTBe);  Warren  Co.,  Twenty  Mile  Strand  (19;  DJLa). 
OKLAHOMA:  Atoka  Co.,  Atoka  (6;  CAS,  MCZ,  USNM),  5 mi.  n.  Stringtown  (80;  UASM);  Carter  Co.,  10.7  mi.  s.  Drake 
(7;  TLEr);  Cherokee  Co.,  15  mi.  sw.  Talequah  (1;  BMNH);  Comanche  Co.,  Wichita  National  Forest  (13;  CAS);  Cotton  Co. 
(2;  CAS);  Craig  Co.,  Grand  Lake  (6;  DHKa);  McCurtain  Co.,  7 mi.  sw.  Smithville  (10;  UASM);  Murray  Co.,  10.3  mi.  n. 
Drake  (6;  TLEr);  Washita  Co.  (1;  CAS).  PENNSYLVANIA  (15;  AMNH,  ANSP,  CAS,  MCZ,  MSUL):  Willow  MiUs  (1; 
VMKi);  Allegheny  Co.,  Allegheny  (1;  MCZ);  Bucks  Co.  (4;  CAS,  RUNB),  Point  Pleasant  (1;  CAS);  Cumberland  Co., 
Lemoyne  (3;  CAS),  New  Cumberland  (12;  CAS,  CUNY,  MCZ,  VMKi),  West  Fairview  (10;  CAS);  Monroe  Co.,  Delaware 
Water  Gap  (12;  AMNH,  MCZ);  Northampton  Co.,  Bethlehem  (4;  CNHM),  Easton  (8;  CAS,  CNHM);  Philadelphia  Co., 
Frankford  (3;  USNM),  Mount  Airy  (1;  ANSP),  Philadelphia  (5;  MCZ),  Wyoming  (1;  USNM);  Pike  Co.,  Milford  (6;  USNM); 
York  Co.,  5 mi.  w.  Davidsburg  (1;  GRNo).  SOUTH  CAROLINA  (1;  MCZ).  SOUTH  DAKOTA:  Mellete  Co.,  White  River 
(1;  UASM).  TENNESSEE:  Carter  Co.,  Elizabethan  (1 ; CNHM);  Davidson  Co.,  Nashville  (4;  CAS,  CUNY,  LACM);  Jackson 
Co.,  Cummins  Mill  (7;  TCBa);  Lincoln  Co.,  2 mi.  n.  Howell  (5;  UASM);  Smith  Co.,  Lancaster  (2;  TCBa);  Warren  Co., 
Cardwell  Mountain  (1;  TCBa);  White  Co.,  Caney  Fork  River  (2;  TCBa),  Sparta  (1;  TCBa).  TEXAS  (21 ; ANSP,  CAS,  MCZ, 
RUNB,  UKSM,  USNM);  Blanco  Co.,  Cypress  Mills  (2;  USNM),  Johnson  City  (22;  UASM);  Colorado  Co.,  Columbus  (9; 
USNM);  Cooke  Co.,  4 mi.  sw.  Era  (1;  CNC);  Erath  Co.,  Morgan  Mill  (2;  INHS);  Hamilton  Co.,  6 mi.  n.  Hamilton  (6;  ANSP); 
Kerr  Co.,  Kerrville  (1;  CNC);  Kinney  Co.,  23  mi.  sw.  Brackettville  (3;  DRWh);  Lampasas  Co.,  Adamsville  (7;  AMNH); 
McCulloch  Co.,  16  mi.  s.  Brady  (3;  CAS);  Maverick  Co.,  8 mi.  n.  Quemado  (5;  UASM);  Parker  Co.,  5 mi.  sw.  Weatherford 
(3;  CNHM);  Val  Verde  Co.,  13  mi.  nw.  Del  Rio  (17;  UASM).  VERMONT:  Clarendon  (3;  RTBe);  South  Alberg  (1;  RTBe); 
Addison  Co.,  North  Ferrisburg  (5;  RTBe);  Bennington  Co.,  Pownall  (3;  RTBe);  Chittenden  Co.,  Charlotte  (2;  RTBe), 
Shelburne  (1;  CAS);  Franklin  Co.,  East  Georgia  (13;  RTBe);  Rutland  Co.,  Fair  Haven  (9;  RTBe);  Windham  Co.,  Brattleboro 
(1;  USNM),  Newfane  (1;  RTBe),  Townshend  (4;  RTBe);  Windsor  Co.,  (1;  RTBe).  VIRGINIA:  Arlington  Co.,  Rosslyn  (4; 
MCZ,  USNM);  Fairfax  Co.  (1;  USNM),  Alexandria  (13;  AMNH,  CAS,  USNM),  Black  Pond  (1;  USNM),  Glencarlyn  (1; 
USNM),  Great  Falls  (2;  AMNH,  USNM),  Mount  Vemon  (4;  USNM);  Loudon  Co.  (2;  AMNH);  Roanoke  Co.,  Buffalo'Creek 
(6; CUNY), Roanoke  (1;MZSP).  WEST  VIRGINIA:  Greenbrier  Co.,  White  Sulphur  Springs  (1;  CAS).  WISCONSIN  (6;  CAS, 
CNHM,  USNM):  Milwaukee  Co.,  Milwaukee  (1;  MCZ),  Wauwatosa  (1;  CNHM).  WYOMING:  Niobrara  Co.,  37  mi.  n.  Lusk 
(8;  UASM). 

MEXICO 

TAMAULIPAS:  34.9  mi.  s.  Nuevo  Laredo  (2;  UASM). 

Collecting  notes.  — Specimens  of  this  abundant  species  have  been  collected  from  April  to 
October,  most  of  them  in  gravel  bars  along  streams. 

Taxonomic  notes.  — This  species  and  S.  sulcifrons  were  long  confused  in  the  literature, 
but  Lindroth  (1961)  recognized  their  distinctness.  They  are  not  closely  related.  A distinctive 
form  of  S.  lineolatus  in  the  Rio  Grande  Valley  does  not  meet  my  criteria  for  recognition  as 
a subspecies,  but  is  no  doubt  geographically  isolated. 


252 


Whitehead 


The  longipennis  group 

Diagnostic  combination.  — Members  of  this  group  are  distinguished  by  the  following 
combination  of  characters:  body  moderately  to  strongly  flattened;  paramedian  clypeal  cari- 
nae  extended  to  median  tooth;  clypeal  field  narrow,  no  wider  at  base  than  apex  of  median 
frontal  sulcus;  submentum  without  accessory  setae;  pronotum  without  paralateral  carinae; 
discal  setae  present  on  intervals  three,  five,  and  seven,  total  less  than  20,  average  length  less 
than  1.0  times  maximum  width  of  interval  two;  abdomen  without  extensive  microsculpture, 
and  without  small  lateral  patches  near  coxal  depressions  of  sternum  three;  endophallus  with 
enlarged  basal  collar  spines.  Also:  clypeal  suture  sharply  impressed  in  most  specimens;  an- 
tennal articles  five  to  ten  filiform;  front  and  middle  tarsi  broadened  and  with  dense  ventral 
pubescence,  especially  in  males;  sternum  seven  with  paramedian  ambulatory  setae  in  males, 
not  in  females;  paramedian  carinae  of  sternum  three  not  or  weakly  curved  outward  at  apices; 
and  pygidium  crenulate  or  not  at  apex  in  females. 

Distribution.  — Members  of  this  group  range  from  the  Colorado  River  system  in  Arizona 
and  New  Mexico  in  the  west  and  from  Tamaulipas  and  Nuevo  Leon  in  the  east  south  to 
Costa  Rica.  I examined  1406  specimens  of  the  longipennis  group. 

Taxonomic  notes.  - I have  not  critically  studied  geographic  variation  in  members  of  this 
group,  as  three  species  are  neither  widespread  nor  represented  by  sufficient  material.  Four 
species  are  distinguished  by  male  genitalic  characteristics  or  by  superficial  characters.  Varia- 
tion in  at  least  S.  longipennis  is  extensive  and  no  doubt  is  worthy  of  study,  but  such  study 
is  not  required  for  species  recognition. 

Schizogenius  neovalidus  new  species 

Type  material.  — Holotype  male  and  allotype  female  labelled  “Gila  River,  nr.  Cliff,  Grant 
Co.  NEW  MEXICO  26.VIII.1964  Awram-Whitehead”  (MCZ).  An  additional  107  specimens 
from  various  localities  in  Arizona  and  New  Mexico  are  paratypes  (AMNH,  ANSP,  BMNH, 
CAS,  CNC,  CNHM,  CUNY,  DJLa,  DRWh,  IRSB,  JNeg,  MGFT,  TLEr,  UASM,  UATA, 
USNM). 

Diagnostic  combination.  — Within  the  longipennis  group,  the  following  characteristics  of 
the  male  genitalia  are  diagnostic  of  this  species:  apex  of  median  lobe  (Fig.  197)  elongate, 
deflexed  at  sharp  angle,  right  margin  strongly  flanged;  basal  collar  spines  of  endophallus 
slender,  elongate.  In  Arizona,  most  matured  specimens  may  be  distinguished  from  matured 
specimens  of  S.  longipennis  by  rufopiceous  rather  than  rufous  femora,  and  by  form  of 
frontal  carinae  (Fig.  192).  From  matured  specimens  of  S',  chiricahuanus  matured  specimens 
of  S.  neovalidus  are  distinguished  by  piceous  or  dark  rufopiceous  rather  than  rufous  or  light 
rufopiceous  elytra. 

Description.  — Body  flattened.  Color  piceous,  elytra  not  to  weakly  aeneous;  femora  in 
most  specimens  dark  rufopiceous;  tibiae,  tarsi,  and  antennae  rufous;  palpi  testaceous. 

Integument.  Conspicuous  microsculpture  on  paramedian  frontal  sulci,  mouthparts,  genae, 
front  legs  except  posterior  surfaces  of  femora  and  trochanters,  middle  legs  except  anterior 
surfaces  of  trochanters,  and  hind  tibiae  and  posterior  surfaces  of  hind  femora.  Sternum  three 
without  conspicuous  paralateral  patches  of  microsculpture. 

Head.  Fig.  192,  frontal  carinae  three  abbreviated  in  most  specimens.  Paramedian  clypeal 
carinae  straight,  not  markedly  elevated  in  basal  half,  extended  to  median  tooth;  median 
field  triangular,  narrow,  no  wider  at  base  than  apex  of  median  frontal  sulcus.  Clypeal  suture 
weakly  to  strongly  engraved.  Eye  prominent,  finely  and  uniformly  faceted.  Neck  densely 
punctate.  Gena  coarsely  punctate,  rugose  in  front.  Antennal  articles  four  to  ten  elongate, 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


253 


article  five  about  1.6-1. 7 times  longer  than  wide. 

Pronotum.  Fig.  192.  Sides  bisetose;  base  not  rugose;  hind  angles  reduced.  Paramedian 
longitudinal  sulci  moderately  long,  nearly  straight  to  strongly  bent  near  middle,  strongly 
hooked  basally.  Anterior  transverse  impression  not  or  weakly  punctate;  basal  impression 
impunctate. 

Legs.  Front  tarsus  strongly  dilated  in  males,  moderately  in  females;  middle  tarsus  mod- 
erately dilated  in  both  sexes;  hind  tarsus  slender,  short.  Paronychia  nearly  as  long  as 
tarsal  claws.  Front  tibia  narrowed  evenly  to  base.  Front  femur  not  strongly  constricted  near 
apex. 

Elytra.  Left  elytron  with  about  6-8  setae  on  interval  three,  5-8  on  interval  five,  3-5  on 
interval  seven;  total  14-19  in  specimens  examined.  Striae  deep  and  sharply  engraved,  finely 
punctate  in  basal  three-fourths.  Intervals  one  to  seven  broad  and  weakly  convex,  interval 
eight  carinate  at  apex;  intervals  three,  five,  and  seven  broadly  joined  at  apices.  Humeral 
denticles  moderately  prominent. 

Abdomen.  Sternum  three  with  paramedian  carinae  not  or  weakly  curved  outward  at 
apices.  Sternum  seven  with  paramedian  ambulatory  setae  in  males  only.  Pygidium  with 
apical  margin  entire  in  males  and  females. 

Male  genitalia.  Median  lobe,  Fig.  197,  apex  elongate,  deflexed  at  a 50-60  degree  angle, 
right  ventral  margin  strongly  flanged;  endophallus,  Fig.  202,  virga  0.42-0.48  length  of  me- 
dian lobe,  basal  collar  spines  six  to  eight  times  longer  than  wide;  13  specimens  examined. 

Measurements  and  proportions.  See  Table  38.  Of  holotype:  TL,  4.94  mm;  LE,  3.09  mm; 
WH,  1.02  mm;  WP,  1.32  mm;  WE,  1.64  mm;  WF/WH,  0.65;  LP/WP,  0.87;  DP/LP,  0.85; 
LP/WE,  0.70;  Ta/Ti,  0.63;  PS/LP,  0.64.  Of  allotype:  TL,  5.29  mm;  LE,  3.35  mm;  WH,  1.02 
mm;  WP,  1.38  mm;  WE,  1.78  mm;  WF/WH,  0.65;  LP/WP,  0.88;  DP/LP,  0.86;  LP/WE,  0.69; 
Ta/Ti,  0.64;  PS/LP,  0.63. 

Variation.  - See  Table  38.  In  some  specimens,  the  front  femora  may  be  rufous  or  the 
third  frontal  carina  not  abbreviated  basally;  I otherwise  observed  no  important  variation. 

Etymology.  — Latin,  neo  = new,  plus  validus  = strong.  Since  the  Arizona  form  of  S. 
longipennis  was  named  S.  validus  by  Fall,  I here  name  this  similar  looking  new  species  by 
the  similar  name,  S.  neovalidus. 

Distribution.  — The  known  range  of  this  species,  except  for  one  female  specimen  collect- 
ed recently  by  D.  J.  Larson  on  the  Verde  River,  is  restricted  to  upper  reaches  of  the  Gila 
River  system  in  southeastern  Arizona  and  southwestern  New  Mexico  (Fig.  207).  I examined 
1 10  specimens  from  the  following  localities. 

UNITED  STATES 

ARIZONA:  Gila  Co.,  nr.  Carrizo  (3;  UASM),  6 mi.  n.  Payson  (1;  DJLa);  Graham  Co.,  nr.  Aravaipa  (35;  AMNH,  ANSP, 
BMNH,  CAS,  CNC,  CNHM,  CUNY,  DRWh,  IRSB,  JNeg,  MGFT,  UASM,  UATA,  USNM);  Pinal  Co.,  9 mi.  nw.  Payson  (1; 
CAS).  NEW  MEXICO:  Catron  Co.,  Glenwood  (11;  UASM);  Grant  Co.,  Cliff  (33;  MCZ,  UASM),  Gila  (1;  UASM),  26  mi.  n. 
Silver  City  (15;  TLEr),  36.4  mi.  ne.  Silver  City  (7;  UASM),  71.6  mi.  ne.  Silver  City  (3;  UASM). 


Collecting  notes.  — Specimens  of  this  species  have  been  collected  from  late  May  through 
late  August,  in  riparian  gravel  bars. 

Taxonomic  notes.  — Until  now,  specimens  of  the  longipennis  group  from  Arizona  have 
all  been  named  S.  validus  Fall.  But  these  specimens  clearly  represent  three  species,  and 
further,  the  name  S.  validus  falls  as  a synonym  of  S.  longipennis  Putzeys.  Males  of  all  three 
species  are  well  characterized  by  details  of  the  male  median  lobe  and  endophallus,  but  fe- 
males cannot  be  distinguished  with  certainty.  Two  females  from  Aravaipa  are  identified  as 
S.  longipennis , and  a female  and  male  of  S.  neovalidus  from  Payson  are  well  within  the  range 
of  S.  longipennis . Thus,  these  two  species  are  sympatric. 


254 


Whitehead 


Table  38.  Descriptive  statistics  for  S.  neovalidus,  based  on  16  males  from  Cliff,  New  Mexico. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

nts,  in  mm. 

TL 

4.79-5.49 

5.224 

0.366 

0.122 

4.65 

LE 

3.00-3.41 

3.248 

0.212 

0.071 

4.34 

WH 

0.96-1.14 

1.059 

0.076 

0.025 

4.78 

WP 

1.22-1.50 

1.401 

0.128 

0.043 

6.09 

WE 

1.57-1.86 

1.736 

0.131 

0.044 

5.04 

B.  Setae  on  le 

ft  elytron. 

Interval  3 

6-  8 

6.7 

Interval  5 

6-  7 

6.2 

Interval  7 

3-  5 

3.7 

Total 

15-18 

16.6 

1.4 

0.5 

5.76 

C.  Proportion 

s. 

WF/WH 

0.61-0.65 

0.632 

0.019 

0.006 

1.96 

LP/WP 

0.84-0.92 

0.881 

0.037 

0.012 

2.80 

DP/LP 

0.82-0.89 

0.848 

0.026 

0.009 

2.03 

LP/WE 

0.69-0.73 

0.711 

0.015 

0.005 

1.40 

Ta/Ti 

0.61-0.69 

0.654 

0.033 

0.011 

3.40 

PS/LP 

0.58-0.67 

0.614 

0.033 

0.011 

3.56 

Schizogenius  longipennis  Putzeys 

Schizogenius  tristriatus  longipennis  Putzeys  1866:227.  Type  locality  “Mexique,”  here  re- 
stricted to  Fortin  de  las  Flores,  Veracruz;  lectotype  female  labelled  “longipennis.  Chd. 
Mex.  (C.  Chd.)”  on  green  paper  in  Putzeys’  script,  in  IRSB  (!),  specimen  labelled  lecto- 
type here  designated.  Bates  1881:37. 

Schizogenius  longipennis , Kult  1950:140. 

Schizogenius  validus  Fall  1901:210.  Type  locality  Rio  Verde,  central  Arizona;  type  male  in 
MCZ  (!),  specimen  labelled  “M.C.Z.  Type  23860”  here  designated  lectotype.  Lindroth 
1961:166.  NEW  SYNONYMY. 

Diag}iostic  combination.  — Males  of  this  species  are  distinguished  from  males  of  other 
members  of  the  longipennis  group  by  the  following  combination  of  characters  of  the  male 
genitalia;  apex  of  median  lobe  short,  deflected  at  weak  angle,  ventral  margin  in  most  speci- 
mens weakly  swollen  near  angulation,  not  flanged  (Fig.  198);  endophallus  with  virga  en- 
larged, about  0.42-0.48  length  of  median  lobe;  basal  collar  spines  slender,  elongate,  about  10 
times  longer  than  wide.  In  Arizona,  most  matured  specimens  of  S.  longipennis  are  distin- 
guished from  matured  specimens  of  S.  chiricahuanus  by  darker  body  color,  and  from  those 
of  S.  neovalidus  by  reddish  front  femora.  In  western  Mexico,  most  specimens  of  S.  longi- 
pennis are  distinguished  from  most  specimens  of  S.  pacificus  by  larger  body  size,  relatively 
larger  eyes,  less  convex  body,  indistinct  micro  sculpture  on  sternum  two  and  basal  margin  of 
pronotum,  pronotum  proportionately  smaller  in  relation  to  rest  of  body,  elytra  less  strongly 
aeneous,  and  paramedian  pronotal  sulci  shorter,  less  sharply  terminated  at  apices. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


255 


Description.  - As  in  S.  neovalidus  except  as  follows.  Color  of  legs  variable,  front  femora 
rufous  to  piceous.  Form  and  sculpture  of  head  and  pronotum,  Fig.  193;  base  of  frontal 
carina  three  not  abbreviated  in  most  specimens;  paramedian  pronotal  sulci  in  most  speci- 
mens sharply  terminated  at  apices.  Left  elytron  with  13-21  discal  setae  in  specimens  exam- 
ined. Pygidium  with  apical  margin  crenulate  in  some  females. 

Male  genitalia.  Median  lobe,  Fig.  198;  apex  short,  deflexed  at  a 30-40  degree  angle,  right 
ventral  margin  not  flanged,  ventral  margin  in  most  specimens  distinctly  swollen  near  angula- 
tion; endophallus,  Fig.  203,  virga  0.42-0.48  length  median  lobe,  basal  collar  spines  about 
10  times  longer  than  wide;  34  specimens  examined. 

Measurements  and  proportions.  See  Table  39. 


Table  39.  Descriptive  statistics  for  S.  longipennis,  based  on  20  males  from  Paso  de  Ovejas, 
Veracruz,  Mexico. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

nts,  in  mm. 

TL 

4.87-5.54 

5.270 

0.305 

0.091 

3.85 

LE 

3.04-3.50 

3.319 

0.192 

0.057 

3.85 

WH 

0.97-1.10 

1.046 

0.058 

0.017 

3.72 

WP 

1.26-1.47 

1.390 

0.095 

0.028 

4.57 

WE 

1.61-1.82 

1.742 

0.098 

0.029 

3.76 

B.  Setae  on  le 

ft  elytron. 

Interval  3 

5-  7 

6.1 

Interval  5 

5-  7 

5.6 

Interval  7 

2-  4 

3.3 

Total 

12-17 

15.0 

1.5 

0.4 

6.49 

C.  Proportion: 

s. 

WF/WH 

0.58-0.63 

0.605 

0.017 

0.005 

1.90 

LP/WP 

0.86-0.93 

0.887 

0.028 

0.008 

2.07 

DP/LP 

0.83-0.90 

0.862 

0.026 

0.008 

2.03 

LP/WE 

0.68-0.73 

0.708 

0.022 

0.007 

2.06 

Ta/Ti 

0.59-0.69 

0.645 

0.036 

0.011 

3.71 

PS/LP 

0.56-0.66 

0.618 

0.037 

0.011 

4.02 

Variation.  — Geographic  variation  not  studied  in  detail.  In  general,  there  is  a trend  toward 
size  reduction  and  reduced  numbers  of  discal  elytral  setae  from  north  to  south.  The  fre- 
quency of  crenulate  pygidial  margins  in  females  is  higher  from  Arizona  to  Sinaloa  than  else- 
where. The  front  femur  tends  to  be  reddish  from  Arizona  to  Sinaloa,  reddish  in  the  apical 
fourth  from  Jalisco  to  Guerrero,  and  piceous  elsewhere.  And  the  tibiae  tend  to  be  testaceous 
to  rufous  from  Arizona  to  Guerrero,  and  dusky  or  piceous  elsewhere. 

Etymology.  - Latin,  longus  = long,  plus  penna  = wing,  in  reference  to  the  elongate 
elytra. 


256 


Whitehead 


Distribution.  — The  known  range  of  this  species  extends  from  southern  Arizona  in  the 
west  and  from  Nuevo  Leon  and  Tamaulipas  in  the  east  south  at  least  to  Costa  Rica  (Fig. 
208).  I studied  1125  specimens  from  the  following  localities. 

UNITED  STATES 

ARIZONA  (17;  AMNH,  ANSP,  CAS,  INHS,  USNM):  Hot  Springs  (2;  CAS),  Riverside  (6;  ANSP,  CAS,  USNM);  Cochise 
Co.,  2.8  mi.  s.  Portal  (1;  TLEr);  Gila  Co.,  33  mi.  s.  Globe  (1;  DJLa),  Pinal  Mountains  (1;  CUNY),  Roosevelt  Lake  (2;  CAS, 
UKSM),  Sierra  Ancha  Mountains  (1;  UASM);  Graham  Co.,  Aravaipa  (2;  CUNY,  DRWh);  Maricopa  Co.,  Phoenix  (1;  MCZ); 
Santa  Cruz  Co.,  Pena  Blanca  (2;  UASM),  Sycamore  Canyon  (2;  CAS);  Yavapai  Co.,  Bumble  Bee  (1;  CAS). 

MEXICO 

No  locality  (5;  IRSB,  MCZ);  Matamoros  (1;  CAS).  CHIAPAS:  3.2  mi.  n.  Arriaga  (1;  CNC),  18.4  mi.  ne.  Chiapa  de  Corzo 
(2;  UASM),  32.5  mi.  e.  Comitan  (1;  UASM),  Huehuetan  (10;  UASM),  Huixtla  (2;  DRWh),  12.8  mi.  nw.  Huixtla  (1;  CNC), 
38.2  mi  nw.  Huixtla  (1;  CNC),  Pijijiapan  (18;  UASM),  Solosuchiapa  (1;  UASM),  4.0  mi.  s.  Solosuchiapa  (4;  UASM),  18.6 
mi.  se.  Tonala  (45;  UASM).  COLIMA:  3.4  mi.  se.  Colima(54;  UASM),  8 mi.  sw.  Colima  (3;  UASM).  GUERRERO:  41.4  mi. 
n.  Acapulco  (42;  UASM).  JALISCO:  4 mi.  s.  Atenquique  (7;  UASM),  Cocula  (3;  IRSB),  Ixtapa  (34;  UASM),  8.5  mi.  n. 
Juchitlan  (37;  UASM),  La  Garita  (1;  CAS),  17.9  mi.  w.  Magdalena  (8;  UASM),  Pitillal  (48;  UASM),  Puerto  Vallarta  (1; 
UASM),  Talpa  de  Allende  (76;  CNHM,  CUNY,  INHS,  JNeg,  MCZ,  MZSP,  UASM,  UATA,  USNM).  MICHOACAN:  8.5  ml 
n.  Nueva  Italia  (10;  UASM).  MORELOS:  Puente  de  Ixtla  (1;  CBoP),  Tejalpa  (1;  FDAG),  Tetecala  (9;  UASM).  NAYARIT: 
Acaponeta  (5;  UASM),  2.4  mi.  s.  Acaponeta  (9;  UASM),  8 mi.  nw.  Acaponeta  (11;  CAS),  13.8  mi.  e.  San  Bias  (20;  UASM). 
NUEVO  LEON:  Linares  (1;  DRWh),  32.9  ml  n.  Montemorelos  (2;  UASM).  OAXACA:  5 mi.  w.  El  Camaron  (l;HFHo), 
17.7  ml  w.  El  Camaron  (1;  BMNH),  25  mi.  e.  El  Camaron  (28;  UASM),  29.4  mi.  e.  El  Coyul  (25;  UASM),  Juchatengo 
(3;  AMNH),  11.1  mi.  n.  Matias  Romero  (1;  BMNH),  9.9  mi.  n.  Pochutla  (7;  UASM),  18.6  mi.  n.  Pochutla  (3;  UASM),  19.1 
mi.  s.  Suchixtepec  (1;  ANSP),  5 mi.  e.  Tapanatepec  (1;  ANSP),  Tehuantepec  (2;  BMNH),  Valle  Nacional  (50;  UASM),  72.5 
mi.  s.  Valle  Nacional  (16;  UASM),  97.3  mi.  s.  Valle  Nacional  (1;  CBoP),  Zanatepec  (20;  UASM),  76  mi.  w.  Zanatepec  (1; 
UASM),  18.4  mi.  w.  Zanatepec  (4;  UASM).  PUEBLA:  Tehuitzingo  (2;  CNHM),  Tepexco  (23;  UASM).  QUERETARO: 
Escanelilla  (9;  UASM),  Jalpan  (6;  UASM),  6.4  mi.  e.  Pinal  de  Amoles  (1;  JHeS).  SAN  LUIS  POTOSf:  17  mi.  n.  Palitla  (36; 
UASM),  2.7  ml  w.  Santa  Catarina  (8;  UASM),  Tamazunchale  (26;  UASM).  SINALOA:  Alamosa  (1;  CAS),  Choix  (2;  UCD), 
5.5  mi.  nw.  Choix  (1;UCD),  Chupaderos  (2;  UCD),  Concordia  (19;  UASM),  11.2  mi.  ne.  Concordia  (1;  JHeS),  30.6  mi.  s. 
Culiacan  (6;  UASM),  8 mi.  w.  El  Palmito  (1;  CNC),  Mazatlan  (2;  MCZ),  7 mi.  n.  Mazatlan  (2;  LBSC),  Rio  Piaxtla  (3; 
UASM),  Rosario  (11;  UASM),  21-33  mi.  e.  Villa  Union  (13;  CNC,  LBSC,  UCB).  SONORA:  Alamos  (1;CAS),  7.2  mi.  se. 
Alamos  (5;  GRNo),  10  mi.  s.  Alamos  (1;  UCD).  TABASCO:  Teapa  (54;  MGFT,  UASM).  TAMAULIPAS:  El  Barretal  (46; 
UASM),  Encino  (4;  UASM),  15.2  ml  n.  Ciudad  Victoria  (6;  UASM),  21.3  mi.  n.  Ciudad  Victoria  (30;  UASM),  39  mi.  s. 
Ciudad  Victoria  (25;  UASM).  VERACRUZ:  Coatzacoalcos  (2;  USNM),  Cordova  (4;  CAS,  USNM),  Coyame  (1;  UASM), 
Fortin  de  las  Flores  (7;  FDAG,  UASM),  20  mi.  nw.  Huatusco  (1;  FDAG),  Jalapa  (3;  MCZ,  USNM),  21.8  mi.  e.  Jalapa  (1; 
MCZ),  Paso  de  Ovejas  (52;  UASM). 


GUATEMALA 

EL  QUICHE:  Sacapulas  (1;  AMNH). 

HONDURAS 

CORTES:  La  Lima  (2;  FDAG). 

COSTA  RICA 

CARTAGO:  Turrialba  (1;  UAFA).  LIMON:  Guapiles  (1;  USNM).  PUNTARENAS:  Palmar  Sur  (2;  UAFA),  6 mi.  n.  Palmar 
Sur  (22;  UAFA),  3 mi.  s.  Palmar  Sur  (1;  UAFA). 


Collecting  notes.  — Adult  specimens  of  S.  longipennis  have  been  collected  throughout  the 
year,  in  riparian  gravel  bars  or  at  lights.  In  material  at  hand,  teneral  adults  are  most  abun- 
dant in  the  months  just  before  and  just  after  the  wet  season,  which,  in  most  of  Mexico, 
extends  from  June  through  August. 

Taxonomic  notes.  — Putzeys  (1866)  indicated  that  his  specimens  of  S.  longipennis  were 
collected  together  with  his  specimens  of  S.  tristriatus.  As  these  must  have  come  from  Vera- 
cruz, and  most  likely  from  the  Cordova  area,  and  as  I have  seen  specimens  of  both  species 
from  Fortin  de  las  Flores,  I have  restricted  the  type  locality  to  that  locality.  Despite  the 
treatment  of  S.  longipennis  as  a form  of  S.  tristriatus  by  both  Putzeys  (1866)  and  Bates 
(1881),  these  names  clearly  refer  to  unrelated  species. 

The  range  of  S.  longipennis  is  sympatric  with  those  of  the  other  members  of  the  longi- 


Classification,  phytogeny,  and  zoogeography  of  Schizogenius 


257 


pennis  group,  and  there  hence  is  no  question  that  S.  longipennis  is  a distinct  species.  I have 
seen  the  types  of  both  S.  longipennis  and  S.  validus.  Specimens  of  S.  longipennis  from 
Arizona  do  differ  in  various  ways  from  Veracruz  specimens,  as  noted  in  my  discussion  of 
variation,  but  I have  no  evidence  to  suggest  that  variation  in  any  characteristic  is  disjunct. 
Thus,  I think  the  names  S.  longipennis  and  S.  validus  are  synonyms. 

Schizogenius  chiricahuanus  new  species 

Type  material.  — Holotype  male  and  allotype  female  labelled  “Cave  Ck.,  Cochise  Co. 
ChiricahuaMts.  Ariz.  ca.600(y  Aug.  17,  1960”  and  “G.E.  Ball  family  & R.B.  Madge  collec- 
tors” (MCZ).  An  additional  30  specimens  from  various  localities  in  Arizona  are  paratypes 
(AMNH,  CUNY,  DJLa,  DRWh,  IRSB,  KHSt,  UASM,  UKSM,  USNM). 

Diagnostic  combination.  — Within  the  longipennis  group,  matured  specimens  of  this  spe- 
cies are  distinguished  by  rufous  coloration.  Also,  characteristics  of  the  male  genitalia  (Fig. 
199)  are  diagnostic. 

Description.  - As  in  S.  neovalidus  except  as  follows.  Average  size  smaller.  Color  light  to 
dark  rufous,  not  piceous  or  dark  rufopiceous.  Form  and  sculpture  of  head  and  pronotum, 
Fig.  194.  Left  elytron  with  about  6-8  setae  on  interval  three,  6-8  on  interval  five,  and  3-5  on 
interval  seven;  total  15-20  in  specimens  examined.  Pygidium  with  apex  crenulate  in  some 
females. 

Male  genitalia.  Median  lobe,  Fig.  199,  apex  short,  deflexed  at  40-45  degree  angle,  right 
ventral  margin  not-  flanged,  ventral  margin  not  swollen  near  angulation;  endophallus,  Fig. 
204,  virga  0.32-0.38  length  median  lobe,  basal  collar  spines  about  five  times  longer  than 
wide  or  less;  four  specimens  examined. 

Measurements  and  proportions.  See  Table  40.  Of  holotype:  TL,  4.52  mm;  LE,  2.89  mm; 
WH,  0.90  mm;  WP,  1.20  mm;  WE,  1.53  mm;  WF/WH,  0.64;  LP/WP,  0.85;  DP/LP,  0.89;  LP / 
WE,  0.67;  Ta/Ti,  0.61;  PS/LP,  0.66.  Of  allotype:  TL,  4.91  mm;  LE,  3.12  mm;  WH,  0.98 
mm;  WP,  1.30  mm;  WE,  1.63  mm;  WF/WH,  0.62;  LP/WP,  0.86;  DP/LP,  0.86;  LP/WE,  0.69; 
Ta/Ti,  0.65;  PS/LP,  0.67. 

Etymology.  — I name  S.  chiricahuanus  after  the  Chiricahua  Mountains  of  southern  Arizo- 
na, since  most  specimens  known  to  me  were  collected  there. 

Distribution.  - Specimens  of  this  species  are  known  only  from  a few  localities  in  south- 
ern Arizona  (Fig.  209).  I studied  35  specimens  from  the  following  localities. 

UNITED  STATES 

ARIZONA:  Cochise  Co.,  Cave  Creek  (19;  MCZ,  UASM),  Chiricahua  Mountains  (4;  UKSM,  USNM),  Huachuca  Mountains 
(1;  KHSt)  Portal  (2;  KHSt),  5 mi.  w.  Portal  (4;  AMNH,  CUNY,  UCB),  Rucker  Lake  (1;  IRSB);  Gila  Co.,  Roosevelt  Lake 
(1;  DJLa),  Sierra  Ancha  Mountains  (3;  DRWh,  MCZ). 


Collecting  notes.  — Specimens  of  this  species  have  been  collected  in  riparian  gravel  bars, 
from  June  through  August. 

Taxonomic  notes.  — S.  chiricahuanus  is  sympatric  with  both  S.  longipennis  and  S.  neova- 
lidus in  southern  Arizona,  or  at  least  nearly  so.  This  fact,  and  well  marked  differences  in 
structures  of  male  genitalia  and  in  other  features,  indicate  that  S.  chiricahuanus  is  repro- 
ductively  isolated  from  those  species.  Known  ranges  of  S.  chiricahuanus  and  S.  pacificus  do 
not  overlap,  and  differences  in  structures  of  male  genitalia  are  smaller,  but  well  marked 
differences  in  body  form  and  in  other  characteristics  indicate  that  these  two  forms  are  also 
reproductively  isolated.  Differences  in  habitus  are  clearly  indicated  in  Tables  38-41  for  the 
four  species  of  the  longipennis  group. 


258 


Whitehead 


Table  40.  Descriptive  statistics  for  S.  chiricahuanus,  based  on  16  males  from  Cochise 
County,  Arizona. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

mts,  in  mm. 

TL 

4.23-4.71 

4.426 

0.210 

0.070 

3.17 

LE 

2.68-2.97 

2.812 

0.119 

0.040 

2.83 

WH 

0.85-0.97 

0.904 

0.046 

0.015 

3.42 

WP 

1.08-1.26 

1.164 

0.068 

0.023 

3.92 

WE 

1.43-1.57 

1.491 

0.070 

0.023 

3.12 

B.  Setae  on  le 

ft  elytron. 

Interval  3 

6-  8 

6.9 

Interval  5 

6-  7 

6.6 

Interval  7 

3-  5 

3.7 

Total 

16-19 

17.1 

1.6 

0.5 

6.35 

C.  Proportion 

s. 

WF/WH 

0.59-0.65 

0.617 

0.026 

0.009 

2.78 

LP/WP 

0.83-0.90 

0.866 

0.030 

0.010 

2.35 

DP/LP 

0.84-0.89 

0.868 

0.028 

0.009 

2.16 

LP/WE 

0.63-0.71 

0.678 

0.030 

0.010 

2.91 

Ta/Ti 

0.58-0.69 

0.642 

0.040 

0.013 

4.19 

PS/LP 

0.62-0.69 

0.654 

0.026 

0.009 

2.67 

Schizogenius  pacificus  new  species 

Type  material.  - Holotype  male  and  allotype  female  labelled  “Acapulco  Gro.,  Mex.  Aug. 
18,  1938  Lipovsky”  (MCZ).  An  additional  5 1 specimens  from  various  localities  in  Guerrero, 
Jalisco,  and  Sinaloa  are  paratypes  (CAS,  DRWh,  MCZ,  UASM,  USNM). 

Diagnostic  combination.  — Within  the  longipennis  group,  specimens  of  S.  pacificus  are 
distinguished  from  those  of  S.  longipennis  and  S.  neovalidus  by  numerous  details  of  male 
genitalia  (Fig.  200),  and  from  specimens  of  S.  chiricahuanus  by  dark  coloration.  In  addition, 
specimens  of  S.  pacificus  differ  from  specimens  of  the  sympatric  S.  longipennis  by  distinct 
microsculpture  on  base  of  pronotum  and  on  sternum  two,  and  by  a more  convex  body. 

Description.  — As  in  S.  neovalidus  except  as  follows.  Body  convex.  Color  piceous,  elytra 
in  most  specimens  distinctly  aeneous;  femora  in  most  specimens  partly  or  wholly  piceous  or 
dark  rufopiceous;  front  tibia  rufous  to  rufopiceous,  hind  tibiae  rufotestaceous.  Additional 
conspicuous  microsculpture  at  base  of  pronotum  and  on  sternum  two  in  most  specimens. 
Head  and  pronotum,  Fig.  195.  Left  elytron  with  about  5-7  setae  on  interval  three,  5-7  on 
interval  five,  2-4  on  interval  seven;  total  13-17  in  specimens  examined.  Pygidium  apex  cre- 
nulate  in  females  or  not. 

Male  genitalia.  Median  lobe,  Fig.  200,  apex  moderate,  deflexed  at  a 40-45  degree  angle, 
right  ventral  margin  not  flanged,  ventral  margin  not  swollen  near  angulation;  endophallus, 
Fig.  205,  virga  0.32-0.38  length  median  lobe,  basal  collar  spines  about  five  times  longer  than 
wide;  five  specimens  examined. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


259 


Measurements  and  proportions.  See  Table  41.  Of  holotype:  TL,  4.27  mm;  LE,  2.62  mm; 
WH,  0.94  mm;  WP,  1.25  mm;  WE,  1.45  mm;WF/WH,  0.61;LP/WP,  0.84;  DP/LP,  0.91;  LP / 
WE,  0.72;  Ta/Ti,  0.65;  PS/LP,  0.57.  Of  allotype:  TL,  4.67  mm;  LE,  2.90  mm;  WH,  0.99 
mm;  WP,  1.31  mm;  WE,  1.57  mm;  WF/WH,  0.62;  LP/WP,  0.86;  DP/LP,  0.93;  LP/WE,  0.72; 
Ta/Ti,  0.71;  PS/LP,  0.60. 


Table  41.  Descriptive  statistics  for  S.  pacificus,  based  on  16  males  from  San  Juan  Abajo, 
Jalisco,  Mexico. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

nts,  in  mm. 

TL 

4.01-4.92 

4.327 

0.415 

0.138 

6.32 

LE 

2.45-2.99 

2.676 

0.244 

0.081 

6.08 

WH 

0.89-1.02 

0.995 

0.068 

0.023 

4.56 

WP 

1.13-1.38 

1.25 

0.110 

0.037 

5.85 

WE 

1.39-1.63 

1.491 

0.121 

0.040 

5.41 

B.  Setae  on  le 

ft  elytron. 

Interval  3 

6-  7 

6.2 

Interval  5 

5-  7 

5.8 

Interval  7 

2-  4 

3.0 

Total 

14-17 

15.0 

1.1 

0.4 

4.87 

C.  Proportion: 

s. 

WF/WH 

0.61-0.64 

0.626 

0.013 

0.004 

1.43 

LP/WP 

0.83-0.91 

0.862 

0.032 

0.011 

2.49 

DP/LP 

0.88-0.94 

0.902 

0.026 

0.009 

1.95 

LP/WE 

0.68-0.77 

0.728 

0.036 

0.012 

3.26 

Ta/Ti 

0.63-0.71 

0.685 

0.037 

0.012 

3.58 

PS/LP 

0.55-0.63 

0.597 

0.034 

0.011 

3.75 

Etymology.  — I name  this  species  after  the  Pacific  Ocean,  since  most  known  specimens 
were  collected  along  Pacific  drainage  systems. 

Distribution.  — Specimens  of  S.  pacificus  have  been  collected  along  various  rivers  of  the 
Pacific  slope  from  southern  Sinaloa  to  southern  Oaxaca,  and  along  one  river  of  the  Atlantic 
slope  in  the  Isthmus  of  Tehuantepec  in  Oaxaca  (Fig.  210).  I studied  136  specimens  from  the 
following  localities. 


MEXICO 

GUERRERO:  Acapulco  (7;  MCZ),  24.8  mi.  e.  Acapulco  (1;  CAS),  C'oyuca  (2;  IRSB).  JALISCO:  4 mi.  s.  Atenquique  (4; 
UASM),  San  Juan  Abajo  (33;  UASM).  OAXACA:  22.2  mi.  n.  Matias  Romero  (12;  UASM).  18.4  mi.  w.  Zanatepec  (70; 
AMNH,  ANSP,  BMNH,  CNC,  CNHM,  CUNY,  JHeS,  JNeg,  MGFT,  MZSP,  UASM).  SINALOA:  Rosario  (4;  CAS,  DRWh, 
USNM),  Villa  Union  (2;  UASM),  26  mi.  ne.  Villa  Union  (1;  LBSC). 


Collecting  notes.  — Specimens  of  this  species  have  been  collected  in  riparian  gravel  bars, 


260 


Whitehead 


Fig.  191-195.  Head  and  pronotum,  dorsal  aspect.  191.  S.  lineolatus  Say,  Frederick,  Maryland.  192.  S.  neovalidus  new 
species,  Aravaipa,  Arizona.  193.  S.  longipennis  Putzeys,  Linares,  Nuevo  Leon.  194.  S.  chiricahuanus  new  species,  Portal, 
Arizona.  195.  S.  pacificus  new  species,  Rosario,  Sinaloa.  Fig.  196-200.  Male  median  lobe,  lateral  aspect.  196.  S.  lineo- 
latus Say,  Edgewood,  Maryland.  197.  S.  neovalidus  new  species,  Glenwood,  New  Mexico.  198.  S.  longipennis  Putzeys, 
Palitla,  Veracruz.  199.  S.  chiricahuanus  new  species,  Sierra  Ancha  Mountains,  Arizona.  200.  S.  pacificus  new  species. 
Villa  Union,  Sinaloa. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


261 


207  208 


201-205.  Male  endophallus.  201.  S.  lineolatus  Say,  Berryville,  Arkansas.  202.  S.  neovalidus  new  species,  Carrizo,  Arizona. 
203.  S.  longipennis  Putzeys,  Alamos,  Sonora.  204.  S.  chiricahuanus  new  species,  Portal,  Arizona.  205.  S.  pacificus  new 
species,  Acapulco,  Guerrero.  Fig.  206-208.  Known  distributions.  206.  S.  lineolatus  Say;  open  symbol  represents  state 
record  only.  207.  S.  neovalidus  new  species  208.  S.  longipennis  Putzeys. 


262 


Whitehead 


210 


1000  mi 


211 


213 


lrig.  209-210.  Known  distributions.  209.  S.  chiricahuanus  new  species.  210.  S.  pacificus  new  species.  Fig.  211-214. 
Geographic  variation  in  S.  lineolatus  Say.  211.  Means  of  body  size,  Table  34.  212.  Means  of  numbers  of  discal  setae  on 
left  elytron,  Table  35.  213.  Means  of  relative  eye  size,  Table  36.  214.  Means  of  pronotal  form.  Table  37. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


263 


or  at  lights,  in  January,  March,  June,  August,  and  December.  I suspect  that  its  habi- 
tat differs  somewhat  from  that  of  S.  longipennis,  since  specimens  of  the  two  species 
have  been  taken  together  in  only  four  places,  and  then  in  quite  unequal  samples.  The 
more  convex  body  form  of  S.  pacificus  suggests  a sandier  habitat  than  that  of  S.  longi- 
pennis. 

Taxonomic  notes.  - Specimens  of  this  species  are  well  differentiated  from  specimens  of 
other  members  of  the  longipennis  group  in  characters  of  male  genitalia  or  habitus  or  both, 
and  are  unquestionably  reproductively  isolated  from  them. 

The  depressus  group 

Diagnostic  combination.  — Members  of  this  group  are  distinguished  by  the  following  com- 
bination of  characters:  body  moderately  to  strongly  flattened;  paramedian  clypeal  carinae 
extended  to  median  tooth;  clypeal  field  narrow,  no  wider  at  base  than  apex  of  median 
frontal  sulcus;  submentum  without  accessory  setae;  pronotum  with  paralateral  carinae  not 
or  weakly  developed;  discal  setae  present  on  intervals  three,  five,  and  seven,  total  norm- 
ally less  than  30,  average  length  less  than  1.0  times  maximum  width  of  interval  two; 
elytra  concolorous  with  pronotum  if  aeneopiceous,  not  pale  at  apices  if  rufocastaneous; 
abdomen  without  extensive  microsculpture  except  in  small  lateral  patches  near  coxal 
depressions  of  sternum  three,  or,  if  extensively  microsculptured  then  pronotal  hind  angles 
not  prominent;  endophallus  with  basal  collar  spines  distinct  but  small.  Also;  clypeal 
suture  sharply  impressed  in  most  species;  antennal  articles  five  to  ten  filiform;  front 
and  middle  tarsi  broadened  and  with  dense  ventral  pubescence,  especially  in  males;  stern- 
um seven  with  paramedian  ambulatory  setae  in  males,  not  in  females;  paramedian  cari- 
nae of  sternum  three  curved  outward  at  apices;  and  pygidium  crenulate  or  not  at  apex 
in  females. 

Distribution.  - The  composite  range  of  members  of  this  group  is  that  of  the  whole  genus. 
I examined  5114  specimens  of  the  depressus  group. 

Taxonomic  notes.  — I made  no  attempt  to  classify  South  American  members  of  this 
large  and  difficult  group,  as  I studied  insufficient  material  to  make  a worthwhile  con- 
tribution. The  only  species  of  the  group  known  to  extend  into  South  America  from 
Middle  America  is  S.  pygmaeus,  which  ranges  at  least  to  Colombia.  Colombian  speci- 
mens of  that  species  may  be  distinguished  from  specimens  of  other  Colombian  species 
of  the  depressus  group  by  details  of  the  male  genitalia.  Also,  they  lack  distinct  micro- 
sculpture on  the  median  field  of  the  frons,  have  little  or  no  microsculpture  on  the  sides 
of  the  pronotum,  and  have  a relatively  convex  pronotum  with  no  indication  of  paralat- 
eral ridges. 

In  addition  to  the  Colombian  specimens  of  S.  pygmaeus,  I examined  513  specimens  of 
one  or  perhaps  several  closely  related  South  American  species  of  the  depressus  group,  from 
a few  localities  in  Argentina,  Bolivia,  Brazil,  and  Colombia  (AMNH,  CAS,  DRWh,  GRNo, 
IRSB,  JNeg,  MCZ,  MGFT,  MZSP,  UASM).  All  of  these  have  piceous  body  color,  as  do 
southern  specimens  of  S.  pygmaeus  and  the  Middle  American  S.  emdeni.  I here  refer  to 
these  specimens  as  a single  taxon,  S.  “apicalis.”  Putzeys  (1863:24)  described  S.  apicalis , 
based  on  12  specimens  from  the  Amazon  River.  One  male  from  the  type  series  (IRSB)  is 
labelled  “Amaz.”  and  “S.  apicalis  Ptz.”  on  green  paper.  As  this  specimen  does  not  agree 
with  the  original  description  in  various  diagnostic  characteristics,  I make  no  lectotype  desig- 
nation until  other  type  material  is  seen.  I have  seen  type  material  of  no  other  described 
South  American  species  of  the  depressus  group,  and  am  unable  to  place  other  species  in  that 
group  from  original  or  subsequent  descriptions. 


264 


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Schizogenius  arimao  Darlington 

Schizogenius  arimao  Darlington  1934:71.  Type  locality  Soledad,  Cuba;  holotype  male  in 

MCZ  (!). 

Diagnostic  combination.  — Specimens  of  this  species  differ  from  others  of  the  depressus 
group  by  uniformly  piceous  elytra,  abdomen  microsculptured,  and  frontal  carinae  basally 
confused. 

Description.  - Body  moderately  flattened.  Color  piceous,  not  aeneous;  front  femora  ru- 
fopiceous,  legs  otherwise  rufous;  palpi  testaceous. 

Integument.  Strong  microsculpture  on  paramedian  frontal  sulci,  mouthparts,  genae,  front 
legs  except  posterior  surfaces  of  femora  and  trochanters,  middle  legs  except  anterior  surfaces 
of  trochanters,  hind  tibiae  and  posterior  surfaces  of  hind  femora,  elytral  epipleura  at  base 
and  on  apical  two-thirds,  and  abdomen. 

Head.  Fig.  215.  Paramedian  clypeal  carinae  straight,  extended  to  median  tooth,  strongly 
elevated  in  basal  half;  median  field  triangular,  no  wider  at  base  than  apex  of  median  frontal 
sulcus.  Clypeal  suture  sharply  defined.  Eye  prominent,  Finely  and  uniformly  faceted.  Neck 
densely  punctate.  Gena  coarsely  punctate,  rugose  in  front.  Antennal  articles  four  to  ten 
elongate,  filiform,  article  five  about  1.4  times  longer  than  wide. 

Pronotum.  Fig.  215.  Sides  bisetose,  hind  angles  obsolete,  base  not  rugose.  Paralateral 
carinae  weakly  developed.  Paramedian  longitudinal  sulci  elongate,  hooked  basally.  Anterior 
transverse  impression  strongly  punctate. 

Legs.  Front  and  middle  tarsi  slightly  dilated  and  pubescent  ventrally  in  both  sexes;  hind 
tarsus  slender.  Paronychia  conspicuous,  nearly  as  long  as  tarsal  claws.  Front  tibia  narrowed 
evenly  to  base.  Front  femur  not  strongly  constricted  near  apex. 

Elytra.  Left  elytron  with  about  seven  setae  each  on  intervals  three  and  five,  and  about 
four  on  interval  seven  (Darlington,  1934).  Striae  deep,  sharply  engraved,  strongly  punc- 
tate in  basal  two-thirds.  Intervals  one  to  seven  broad  and  flat,  interval  eight  carinate  at 
apex;  apices  of  intervals  three,  five,  and  seven  broadly  joined.  Humeral  denticles  promi- 
nent. 

Abdomen.  Sternum  three  with  paramedian  carinae  curved  outward  at  apices.  Sternum 
seven  with  a pair  of  paramedian  ambulatory  setae  in  males  only  (Darlington,  1934).  Pygi- 
dium  not  examined. 

Male  genitalia.  Median  lobe,  Fig.  222;  1 specimen  studied. 

Measurements  and  proportions.  Based  on  one  male  from  the  type  locality.  TL,  3.61  mm; 
LE,  2.23  mm;  WH.  0.74  mm;  WP,  0.94  mm;  WE,  1.19  mm:  WF/WH,  0.61;  LP/WP,  0.94;  DP/ 
LP,  0.81;  LP/WE,  0.75;  Ta/Ti,  0.65;  PS/LP,  0.66. 

Etymology.  — S.  arimao  was  named  after  the  Arimao  River. 

Distribution.  - S.  arimao  is  known  definitely  only  from  two  localities  in  Cuba  (Darling- 
ton, 1934)  (Fig.  235);  I did  not  confirm  a Jamaican  record  (Darlington,  1941).  Although  I 
saw  type  material  in  the  Museum  of  Comparative  Zoology,  my  description  is  based  on  just 
one  topotypic  male  given  to  me  by  Darlington. 

Taxonomic  notes.  — This  species  has  no  known  close  relatives.  Darlington  suggested  a 
relationship  with  S.  tristriatus,  but  I think  it  most  closely  related  to  the  South  American 
S.  “apicalis”  and  the  Middle  American  S.  emdeni.  In  particular,  the  weak  but  evident 
paralateral  pronotal  carinae  suggest  a relationship  with  S.  “ apicalis ”,  a suggestion  sup- 
ported by  zoogeographic  considerations,  reduced  hind  angles,  short  antennae,  small  size, 
dark  color,  and  comparatively  elongate  tarsi.  Abdominal  microsculpture  of  this  species  is  a 
characteristic  convergent  in  the  taxon  S.  ochthocephalus  and  in  members  of  the  tristriatus 
group. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


265 


Schizogenius  emdeni  new  species 

Type  material.  — Holotype  female  labelled  “S.  Geronimo.  Guatemala.  Champion.”,  “B. 
C.A.  Col.  I.  1.  Schizogenius  tristriatus,  Putz.”,  “TYPE”,  and  “Schizogenius  Emdeni  Kt. 
57  det.  K.  Kult”  (BMNH).  Four  males  and  two  females  from  Palmar  Sur,  Costa  Rica  are 
paratypes  (DRWh,  UAFA,  UASM). 

Diagnostic  combination.  — Best  distinguished  from  S.  pygmaeus,  the  only  other  known 
piceous  Middle  American  member  of  the  depressus  group  without  abdominal  microsculp- 
ture, by  details  of  male  genitalia.  Also,  most  specimens  have  fewer  than  15  elytral  setae, 
and  reduced  paralateral  patches  of  microsculpture  on  sternum  three. 

Description.  — As  in  S.  pygmaeus  except  as  follows.  Body  color  piceous,  elytra  distinctly 
aeneous.  Patches  of  microsculpture  in  coxal  depressions  of  sternum  three  smaller,  less  dis- 
tinct. Head  and  pronotum,  Fig.  216.  Left  elytron  with  five  or  six  setae  on  interval  three, 
five  on  interval  five,  and  two  or  three  on  interval  seven;  total  12-14  in  specimens  examined. 
Male  genitalia  with  median  lobe,  Fig.  223;  one  specimen  examined. 

Measurements  and  proportions.  Of  holotype:  TL,  4.04  mm;  LE,  2.48  mm;  WH,  0.80  mm; 
WP,  1.11  mm;  WE,  1.29  mm;  WF/WH,  0.61;  LP/WP,  0.92;  DP/LP,  0.83;  LP/WE,  0.79; 
Ta/Ti,  0.62;  PS/LP,  0.66.  Of  type  series:  TL,  3.31-3.751-4.04  mm;  LE,  2.05-2.316-2.48 
mm;  WH,  0.65-0.739-0.80  mm;WP,  0.86-1.009-1.11  mm;  WE,  1.06-1.196-1 .29  mm;  WF/WH, 
0.60-0.616-0.64;  LP/WP,  0.90-0.941-0.98;  DP/LP,  0.79-0.814-0.84;  LP/WE,  0.76-0.784-0.80; 
Ta/Ti,  0.62-0.657-0.68;  PS/LP,  0.62-0.651-0.70. 

Etymology.  — K.  Kult  recognized  this  species  as  distinct,  and  planned  to  name  it  after  the 
well  known  coleopterist  Fritz  van  Emden.  As  he  never  published  the  name,  I do  so  here. 

Distribution.  — S.  emdeni  is  so  far  known  only  from  two  localities  in  Middle  America 
(Fig.  235).  I studied  seven  specimens  from  the  following  localities. 

GUATEMALA 

BAJA  VERAPAZ:  San  Geronimo  (1;  BMNH). 

COSTA  RICA 

PUNTARENAS:  Palmar  Sur  (6;  DRWh,  UAFA,  UASM). 

Collecting  notes.  — In  Costa  Rica,  specimens  of  S.  emdeni  were  collected  at  black  lights 
by  R.  T.  Allen,  in  August. 

Taxonomic  notes.  — I selected  as  holotype  the  specimen  earlier  selected  by  Kult,  in  order 
to  avoid  future  confusion.  There  is  no  doubt  that  the  Costa  Rican  specimens  are  conspecific 
with  it. 

Reduced  numbers  of  elytral  setae  and  reduced  paralateral  patches  of  microsculpture  on 
sternum  three  superficially  suggest  placement  in  the  longipennis  group.  I place  S.  emdeni  in 
the  depressus  group  because  the  patches  of  microsculpture  are  present,  and  because  of  simi- 
larities in  some  statistical  characteristics  with  S.  arimao  and  S.  “apicalis”. 

Schizogenius  sulcifrons  Putzeys 

Schizogenius  sulcifrons  Putzeys  1846:652.  Type  locality  “Amerique  boreale,”  restricted  to 
Rumney,  New  Hampshire  by  Lindroth  (1961);  lectotype  female  designated  by  Lindroth 
(1961),  in  Hope  Museum  at  Oxford,  not  seen  by  me,  Lindroth  1961:167. 

Schizogenius  lineolatus,  LeConte  1857:83.  Putzeys  1863:24.  Putzeys  1866:228.  Gemminger 
and  Harold  1868:206.  LeConte  1879:34.  Leng  1920:48.  Csiki  1927:551. 

Diagnostic  combination.  — Specimens  of  S.  sulcifrons  are  distinguished  from  those  of 


266 


Whitehead 


other  piceous  species  of  Schizogenius  found  in  eastern  North  America  by  having  in  combi- 
nation: more  than  20  discal  setae  per  elytron  in  most  specimens;  weakly  developed  pronotal 
hind  angles;  and  uniformly  piceous  pronotum  and  elytra.  Most  specimens  differ  from  most 
of  the  western  S.  litigiosus  by  abdomen  paler  than  elytra. 

Description.  — Body  flattened.  Color  piceous  above,  rufopiceous  below;  legs  and  antennae 
rufous;  palpi  testaceous;  elytra  in  most  specimens  strongly  aeneous. 

Integument.  Conspicuous  microsculpture  on  paramedian  frontal  sulci,  mouthparts,  genae, 
front  legs  except  posterior  surfaces  of  femora  and  trochanters,  middle  legs  except  anterior 
surfaces  of  trochanters,  hind  tibiae  and  posterior  surfaces  of  hind  femora,  elytral  epipleura 
on  apical  two-thirds,  and  sternum  three  in  coxal  depressions  and  small  paralateral  patches. 

Head.  Fig.  217.  Paramedian  clypeal  carinae  straight,  moderately  elevated  in  basal  half, 
extended  to  median  tooth;  median  field  triangular,  narrow,  no  wider  at  base  than  apex  of 
median  frontal  sulcus.  Clypeal  suture  sharply  defined.  Eye  prominent,  finely  and  uniformly 
faceted.  Neck  densely  punctate.  Gena  coarsely  punctate,  rugose  in  front.  Antennal  articles 
four  to  ten  elongate,  article  five  about  1.6- 1.8  times  longer  than  wide. 

Pronotum.  Fig.  217.  Sides  bisetose;  base  not  rugose;  hind  angles  weakly  developed.  Para- 
median longitudinal  sulci  moderately  elongate,  nearly  straight,  strongly  hooked  basally. 
Paralateral  carinae  absent.  Anterior  transverse  impression  not  or  weakly  punctate. 

Legs.  Front  and  middle  tarsi  moderately  dilated  and  pubescent  ventrally,  less  so  in  fe- 
males; hind  tarsus  slender,  short.  Paronychia  nearly  as  long  as  tarsal  claws.  Front  tibia 
narrowed  evenly  to  base.  Front  femur  not  strongly  constricted  near  apex. 

Elytra.  Left  elytron  with  about  seven  to  eleven  setae  each  on  intervals  three  and  five, 
five  to  eight  on  interval  seven;  total  19-30  in  specimens  examined.  Striae  deep  and  sharply 
engraved,  finely  punctate  in  basal  two-thirds.  Intervals  one  to  seven  broad  and  flat,  interval 
eight  carinate  at  apex;  intervals  three,  five,  and  seven  broadly  joined  at  apices.  Humeral 
denticles  moderate. 

Abdomen.  Sternum  three  with  paramedian  carinae  curved  outward  at  apices.  Sternum 
seven  with  paramedian  ambulatory  setae  in  males  only.  Pygidium  with  apical  margin  nor- 
mally crenulate  in  females,  entire  in  males. 

Male  genitalia.  Median  lobe,  Fig.  224;  six  specimens  examined. 

Measurements  and  proportions.  See  Table  42.  As  this  sample  includes  males  and  females, 
these  data  are  not  directly  comparable  with  data  given  for  other  members  of  the  depressus 
group.  Males  tend  to  be  smaller  than  females;  the  mean  LE  of  six  males  from  Rumney,  New 
Hampshire,  is  2.505  mm. 

Variation.  - I had  insufficient  material  of  this  species  for  a useful  statistical  analysis  of 
variation.  I noticed  no  important  geographic  variation  in  either  pronotal  form  or  in  numbers 
of  elytral  setae.  But,  from  the  midwestern  states,  through  the  Washington.  D.  C.  area, 
through  New  York  state,  to  New  Hampshire,  there  are  definite  trends  toward  larger  body 
size  and  reduced  eyes.  These  trends  agree  with  those  observed  for  S.  lineolatus  in  the  same 
general  areas. 

Etymology.  — Latin,  sulcus  = groove  + frons  = front,  in  reference  to  the  plurisulcate 
frons. 

Distribution.  — The  known  range  of  S.  sulcifrons  includes  much  of  North  America  east 
of  the  Mississippi  River,  from  New  Brunswick  (Lindroth,  1961)  to  Georgia  in  the  east,  and 
from  Wisconsin  to  Mississippi  in  the  west  (Fig.  236).  I studied  131  specimens  from  the 
following  localities. 

CANADA 

No  locality  (1;  ANSP).  ONTARIO:  London  (6;  CNHM,  UASM),  Saint  Augustine  (1;  USNM),  Toronto  (3;  CUNY,  MCZ). 
See  Lindroth  (1961)  for  additional  Canadian  records. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


267 


UNITED  STATES 

No  locality  (9;  ANSP,  CAS,  PSUU,  UKSM).  ALABAMA:  Jackson  Co.,  Big  Coon  Creek  (l;TCBa).  DISTRICT  OF  COLUM- 
BIA: Washington  (7;  CAS,  USNM).  GEORGIA  (1;  ANSP).  ILLINOIS:  Vermilion  Co.,  Kickapoo  State  Park  (2;  RTBe). 
INDIANA  (1 ; CNHM):  Parke  Co.,  The  Shades  State  Park  (1 ; RTBe);  Tippecanoe  Co.  (3;  CNC,  MCZ,  UATA).  KENTUCKY: 
Metcalfe  Co.,  8 mi.  e.  Edmonton  (2;  DRWh).  MAINE:  Oxford  Co.,  Paris  (1 ; MCZ).  MARYLAND:  Frederick  Co.,  Frederick 
(1;  RTBe);  Harford  Co.,  Edgewood  (3;  CUNY);  Prince  Georges  Co.,  Bladensburg  (1;  USNM).  MASSACHUSETTS:  Franklin 
Co.,  Northfield  (2;  MCZ).  MISSISSIPPI:  Ireland  (1;  UKSM).  NEW  HAMPSHIRE:  Carroll  Co.,  North  Conway  (2;  MCZ); 
Grafton  Co.,  Rumney  (15;  CNHM,  CNC,  MCZ,  UASM).  NEW  JERSEY  (1 ; CAS);  Warren  Co.,  Phillipsburg  (1 ; CAS).  NEW 
YORK  (7;  CAS,  UKSM,  USNM);  New  Windsor  (2;  USNM);  Erie  Co.,  Lancaster  (1;  CAS);  Schuyler  Co.,  Watkins  Glen  (3; 
AMNH,  MCZ);  Suffolk  Co.,  Riverhead  (1;  CUNY);  Tompkins  Co.,  Groton  (5;  JNeg),  Ithaca  (15;  CAS,  CUNY,  USNM); 
Ulster  Co.,  Esopus  (5;  CUNY,  MCZ).  NORTH  CAROLINA:  Buncombe  Co.,  Black  Mountains  (1;  CAS).  OHIO:  Knox  Co., 
Gambier  (3;  UMCG).  PENNSYLVANIA  (2;  CAS);  Allegheny  Co.,  Allegheny  (1;  MCZ);  Monroe  Co.,  Delaware  Water  Gap 
(2;  AMNH);  Philadelphia  Co.,  Philadelphia  (2;  MCZ);  Pike  Co.,  Milford  (1;  USNM).  SOUTH  CAROLINA  (1;  MCZ). 
TENNESSEE:  Davidson  Co.,  Nashville  (2;  CAS,  USNM);  Monroe  Co.,  Sweetwater  (3;  AMNH).  VERMONT  (1;  MCZ): 
Chittenden  Co.,  Milton  (1;  RTBe);  Rutland  Co.,  Poultney  (1;  RTBe).  VIRGINIA:  Loudon  Co.  (4;  AMNH).  WISCONSIN 
(1;  CUNY). 


Table  42.  Descriptive  statistics  for  S.  sulcifrons,  based  on  six  males  and  six  females  from 
Rumney,  New  Hampshire. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measurements,  in  mm. 

TL 

3.69-4.50 

4.128 

0.314 

0.121 

5.08 

LE 

2.32-2.85 

2.601 

0.215 

0.083 

5.50 

WH 

0.75-0.85 

0.808 

0.039 

0.015 

3.20 

WP 

0.97-1.17 

1.086 

0.087 

0.033 

5.33 

WE 

1.19-1.43 

1.329 

0.104 

0.040 

5.24 

B.  Setae  on  lei 

ft  elytron. 

Interval  3 

8-  9 

8.5 

Interval  5 

7-10 

8.4 

Interval  7 

5-  7 

6.0 

Total 

20-25 

22.9 

2.8 

1.1 

8.00 

C.  Proportion? 

WF/WH 

0.62-0.68 

0.658 

0.026 

0.010 

2.58 

LP/WP 

0.85-0.91 

0.892 

0.023 

0.009 

1.73 

DP/LP 

0.80-0.85 

0.828 

0.024 

0.009 

r.9i 

LP/WE 

0.70-0.75 

0.727 

0.021 

0.008 

1.97 

Ta/Ti 

0.59-0.66 

0.621 

0.035 

0.013 

3.73 

PS/LP 

0.68-0.76 

0.707 

0.039 

0.015 

3.68 

Collecting  notes.  — Specimens  of  S.  sulcifrons  have  been  collected  from  late  March 
through  September,  generally  in  riparian  gravel  bars.  I found  specimens  of  this  species,  S. 
lineolatus,  S.  amphibius,  and  S.  planulatus  along  a small  stream  near  Edmonton,  Kentucky. 

Taxonomic  notes.  - See  discussion  by  Lindroth  (1961).  As  I did  not  see  the  lectotype, 
I accept  Lindroth’s  concept  of  the  species.  Though  S.  sulcifrons  and  S.  lineolatus  have  been 


268 


Whitehead 


confused  in  the  past,  they  are  not  really  closely  related.  S.  sulcifrons  is  closely  related  to 
S.  litigiosus,  but  ranges  of  the  two  forms  are  widely  separated  and  there  is  no  evidence  of 
intergradation  between  them. 


Schizogenius  litigiosus  Fall 

Schizogenius  litigiosus  Fall  1901:210.  Type  locality  Sylvania,  California;  holotype  female 
in  MCZ  (!).  Lindroth  1961:168. 

Schizogenius  depressus,  Hatch  1949:118.  Hatch  1953:69. 

Diagnostic  combination.  — Fully  colored  specimens  of  this  species  are  distinguished  from 
other  members  of  the  depressus  group  in  western  North  America  by  their  piceous  colora- 
tion. Most  specimens  are  distinguished  from  most  specimens  of  the  eastern  S.  sulcifrons  by 
having  concolorous  abdomen  and  elytra. 

Description.  - Body  flat.  Color  piceous;  front  femora  rufous  to  rufopiceous;  antennae, 
tibiae,  tarsi,  and  middle  and  hind  femora  rufous;  palpi  testaceous;  elytra  not  or  weakly 
aeneous,  rufescent  toward  apices  in  many  specimens. 

Integument,  head,  pronotum,  legs,  elytra,  and  abdomen  as  in  S.  sulcifrons  except  as 
follows.  Head  and  pronotum,  Fig.  218.  Left  elytron  with  about  six  to  nine  setae  each  on 
intervals  three  and  five,  three  to  five  on  interval  seven;  total  16-22  in  specimens  examined. 
Apex  of  pygidium  normally  entire  in  females  and  males. 

Male  genitalia.  Median  lobe,  Fig.  225;  six  specimens  examined. 

Measurements  and  proportions.  See  Table  43. 


Table  43.  Descriptive  statistics  for  S.  litigiosus,  based  on  18  males  from  Clear  Lake,  Cali- 
fornia. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

ints,  in  mm. 

TL 

3.43-4.34 

4.033 

0.315 

0.099 

5.21 

LE 

2.13-2.70 

2.495 

0.191 

0.060 

5.11 

WH 

0.71-0.88 

0.822 

0.061 

0.019 

4.92 

WP 

0.87-1.15 

1.054 

0.107 

0.034 

6.76 

WE 

1.10-1.38 

1.284 

0.098 

0.031 

5.08 

B.  Setae  on  le 

ft  elytron. 

Interval  3 

6-  8 

7.2 

Interval  5 

6-  8 

7.0 

Interval  7 

3-  5 

4.4 

Total 

16-21 

18.6 

2.1 

0.6 

7.40 

C.  Proportions. 


WF/WH 

0.60-0.65 

0.617 

0.022 

0.007 

2.35 

LP/WP 

0.90-0.97 

0.935 

0.024 

0.007 

1.69 

DP/LP 

0.78-0.82 

0.801 

0.020 

0.006 

1.63 

LP/WE 

0.74-0.79 

0.768 

0.022 

0.007 

1.92 

Ta/Ti 

0.60-0.70 

0.648 

0.040 

0.013 

4.12 

PS/LP 

0.65-0.73 

0.688 

0.034 

0.011 

3.25 

Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


269 


Variation.  — There  is  a definite  trend  toward  reduced  eye  size  from  northern  California 
to  southern  British  Columbia,  and  there  may  also  be  a trend  toward  increased  body  size 
from  south  to  north.  I otherwise  found  no  conspicuous  geographic  variation.  I had  in- 
sufficient material  from  most  areas  for  useful  statistical  analysis  of  geographic  variation. 

Etymology.  - Latin,  litigiosus  = contentious  or  quarrelsome;  Fall  probably  suggested  this 
name  because  this  species  had  previously  been  confused  with  S.  pluripunctatus  and  S.  de- 
pressus. 

Distribution.  - The  known  range  of  S.  litigiosus  extends  from  southern  British  Columbia 
to  southern  California,  west  of  the  Rocky  Mountains  (Fig.  237).  I studied  629  specimens 
from  the  following  localities. 


CANADA 

BRITISH  COLUMBIA:  Duncan  (13;  CAS,  KSUM,  MCZ),  Keremeos  (8;  BMNH,  DRWh,  IRSB,  MGFT).  For  other  Canadian 
records,  see  Lindroth  (1961). 


UNITED  STATES 

CALIFORNIA  (13;  AMNH,  CAS,  KSUM,  MCZ,  USNM);  La  Panza  (1;  CAS);  Alameda  Co.,  Alameda  Creek  (1;  CAS),  20  mi. 
s.  Livermore  (1;  UCB);  Amador  Co.,  Cold  Creek  (2;  CAS);  Butte  Co.,  Oroville  (7;  CAS);  Paradise  (2;  CAS);  Calaveras  Co., 
Mokelumne  Hill  (23;  CAS,  MCZ);  Colusa  Co.,  Cooks  Springs  (2;  CAS);  Del  Norte  Co.  (3;  CNHM);  Eldorado  Co.  (7;  CNHM, 
MGFT),  Coloma  (6;  GRNo);  Fresno  Co.,  Camp  Greeley  (1;  CAS);  Glenn  Co.,  Elk  Creek  (1;  CAS),  25  mi.  w.  Elk  Creek  (1; 
CAS),  Hamilton  City  (2;  UCD);  Humboldt  Co.  (2;  CAS),  6 mi.  e.  Bridgeville  (1;  TLEr),  Garberville  (2;  CAS),  Hoopa  (2; 
CAS,  UASM),  Larabee  Creek  (2;  CAS),  Mad  River  (1;  CAS),  North  Dobbyn  Creek  (1;  CAS),  Redwood  Creek  (1;  UASM), 
Shively  (3;  UCD),  Willow  Creek  (2;  CAS);  Kings  Co.,  Stratford  (2;  CAS);  Lake  Co.  (2;  CAS),  Clear  Lake  (28;  CAS), 
Grossford  (1;  CAS),  Kelseyville  (3;  CAS),  Middle  Creek  (23;  CAS),  Middletown  (5;  CAS),  North  Fork  Cache  Creek  (10; 
UCD),  Scott  Creek  (1;  CAS);  Los  Angeles  Co.,  Cole  (1;  CAS),  Pomona  (1;  USNM),  3 mi.  s.  Valyermo  (l;GRNo);  Madera 
Co.,  North  Fork  (1;  CUNY);  Marin  Co.  (1;  CAS),  Fairfax  (15;  CAS);  Mendocino  Co.  (3;  CNHM),  Black  Butte  River  (18; 
CAS),  Bloody  Run  Creek  (1;  CAS),  Dry  Creek  (4;  CAS),  Eel  River  (3;  CAS),  Longvale  (13;  CAS),  Mailliard  (2;  CAS),  8 mi. 
w.  Navarro  (1;  CAS),  2 mi.  nw.  Philo  (1;  CAS),  Twin  Rocks  (1;  CAS),  Williams  Creek  (13;  CAS),  2 mi.  s.  Yorkville  (3; 
CAS);  Modoc  Co.,  9.5  mi.  s.  Cedarville  (35;  CAS),  Lake  City  (2;  CAS);  Napa  Co.  (6; CAS,  USNM),  Monticello  (10;  CAS, 
UCD,  MCZ),  Pope  Valley  (6;  CAS),  Saint  Helena  (6;  CAS,  KSUM,  MCZ,  UASM);  Placer  Co.,  Auburn  (3;  UCD),  Penryn 
(1;  UCB);  Plumas  Co.,  Clio  (4;  CAS);  Sacramento  Co.,  Cosumes  River  (2;  CAS),  Fair  Oaks  (2;  UCD);  San  Bemadino  Co., 
Hesperia  (1;  CAS),  Mojave  River  (5;  GRNo,  TLEr);  San  Diego  Co.  (2;  LACM);  San  Francisco  Co.  (3;  CAS);  San  Joaquin 
Co.,  San  Joaquin  River  (1;  CAS),  Santa  Clara  Co.,  Anderson  Reservoir  (1;  DHKa),  Gilroy  Hot  Springs  (11;  CAS,  DJLa, 
TLEr),  Los  Gatos  (1;  CAS),  Mount  Hamilton  (3;  CAS);  Shasta  Co.,  Redding  (9;  CAS);  Siskiyou  Co.  (1;  USNM),  2.3  mi. 
nw.  Callahan  (9;  CAS),  Dillon  Creek  (6;  UASM),  Scott  River  (1;  CAS),  Yreka  (1;  MCZ);  Sonoma  Co.  (6;  CAS,  CUNY, 
LACM,  MCZ),  Agua  Caliente  (2;  CNHM),  Annapolis  (2;  CAS),  Cloverdale  (1;  CAS),  Duncan  Mills  (3;  CAS),  Guemeville 
(12;  CAS),  Healdsburg  (2;  CAS),  Preston  (1;  CAS),  Rio  Nido  (1;  CAS),  Russian  River  (6;  CAS,  MCZ),  Santa  Rosa  (10; 
CAS,  MCZ,  MSUL),  2.5  mi.  w.  Skaggs  Springs  (2;  CAS),  Sylvania  (5;  CAS,  MCZ,  LACM);  Stanislaus  Co.,  22  mi.  w. 
Patterson  (4;  CAS);  Tehama  Co.,  Red  Bluff  (29;  CNHM);  Trinity  Co.,  2 mi.  e.  Burnt  Ranch  (6;  CAS,  RTBe),  Canyon  City 
(1;  CAS);  Clear  Creek  (1;  TLEr),  Douglas  City  (3;  UASM),  Hayfork  Creek  (9;  CAS),  Hyampton  (3;  CAS),  Mad  River  (3; 
CAS,  UASM),  4 mi.  se.  Ruth  (22;  CAS),  6 mi.  s.  Ruth  (5;  CAS),  Ruth  Dam  (3;  GRNo,  TLEr),  Trinity  Center  (2;  CAS), 
Weaverville  (1;  CAS);  Tulare  Co.,  Fairview  (1;  UCB),  Kaweah  (1;  CAS);  Yolo  Co.,  Davis  (13;  UCD),  Putah  Canyon  (2; 
UCD),  Rumsey  (1;  UCD).  IDAHO:  Nez  Perce  Co.,  6 mi.  n.  Lenore  (1;  DHKa);  Owahee  Co.,  Hot  Creek  Falls  (3;  CNHM). 
NEVADA  (2;  ANSP):  Lyon  Co.,  Weeks  (3;  UASM);  Washoe  Co.,  Pyramid  Lake  (1;  UCD).  OREGON  (1;  ANSP):  Jackson 
Co.,  Cow  Creek  (3;  CAS,  MCZ),  Eagle  Point  (1;  CNHM),  Medford  (17;  CAS,  MCZ,  UCD,  USNM),  Talent  (2;  UCD); 
Josephine  Co.  (7;  CNHM);  Lane  Co.,  Eugene  (3;  CNHM),  Middle  Fork  of  Willamette  River  (1;  CNHM);  Malhuer  Co., 
Sucker  Creek  Canyon  (3;  CNHM);  Umatilla  Co.,  McKay  Reservoir  ( 1 ; CNHM) ; Wasco  Co.,  The  Dalles  (1 ; USNM);  Wheeler 
Co.,  John  Day  Gorge  (1;  CNHM);  Yamhill  Co.,  Dayton  (2;  MCZ).  WASHINGTON:  Yakima  Co.,  Toppenish  (12;  KSUM, 
MCZ,  USNM). 

Collecting  notes.  — In  the  north,  specimens  of  this  species  have  been  collected  from  May 
to  September.  In  central  California,  adults  may  be  found  throughout  the  year.  Adults  are 
found  in  riparian  gravel  bars. 

Taxonomic  notes.  — This  species  was  confused  with  S',  depressus  by  Hatch  (1949,  1953), 
but  well  distinguished  by  Lindroth  (1961).  The  range  of  S.  litigiosus  is  entirely  included 
within  that  of  S.  depressus , and  specimens  of  both  species  have  often  been  found  at  the 
same  locality. 


270 


Whitehead 


Schizogenius  pygmaeus  Van  Dyke 

Schizogenius  pygmaeus  Van  Dyke  1925:12.  Type  locality  Clear  Lake,  California;  holotype 

in  CAS,  not  seen  by  me. 

Schizogenius  championi  Kult  1950:142.  Type  locality  Pantaleon,  Escuintla,  Guatemala; 

holotype  in  BMNH,  not  studied.  NEW  SYNONYMY. 

Diagnostic  combination.  — Within  the  depressus  group,  all  dark  or  bicolored  specimens 
seen  from  Mexico  belong  to  S.  pygmaeus,  and  none  from  the  United  States  or  Canada  do. 
In  Middle  America  south  of  Mexico,  dark  forms  of  this  species  are  best  distinguished  from 
specimens  of  S.  emdeni  by  characteristics  given  in  the  key.  In  South  America,  specimens  of 
S.  pygmaeus  are  distinguished  from  other  members  of  the  depressus  group  by  details  of 
male  genitalia.  Specimens  of  pale  forms  of  S.  pygmaeus  are  best  distinguished  from  speci- 
mens of  pale  forms  of  S.  falli , S.  ochthocephalus,  and  S.  depressus  by  characteristics  given 
in  the  key.  Ranges  of  S.  pygmaeus  and  S.  scopaeus  are  largely  allopatric.  The  latter  species 
is  distributed  along  Atlantic  drainage  systems;  its  range  extends  south  of  the  Rio  Grande 
only  in  Nuevo  Leon  and  Tamaulipas  where  it  is  sympatric  with  black  forms  of  S.  pygmaeus. 
The  range  of  S.  pygmaeus  reaches  the  Rio  Grande  drainage  basin  only  in  southern  Chihua- 
hua, where  S.  scopaeus  is  not  known  to  occur.  Where  the  two  species  approach  one  another 
or  overlap  in  range,  specimens  are  distinguishable  by  conspicuous  differences  in  the  male 
genitalia  and,  in  northeastern  Mexico,  by  color. 

Description.  - Body  weakly  convex.  Color  testaceous  or  ferrugineous  to  brunneous  to 
piceous,  some  specimens  distinctly  bicolored  with  head  and  pronotum  darker  than  elytra; 
some  dark  specimens  slightly  aeneous;  legs  and  antennae  ferrugineous  or  testaceous;  palpi 
testaceous. 

Integument.  Distinct  microsculpture  on  paramedian  frontal  sulci,  mouthparts,  genae, 
front  legs  except  posterior  surfaces  of  femora  and  trochanters,  middle  legs  except  anterior 
surfaces  of  trochanters,  hind  tibiae  and  posterior  surfaces  of  hind  femora,  shoulders  and 
apical  two-thirds  of  elytral  epipleura,  and  small  areas  in  coxal  depressions  of  sternum  three. 

Head.  Fig.  219.  Clypeus  with  paramedian  carinae  straight,  extended  to  median  tooth, 
strongly  elevated  in  basal  half;  median  field  narrow,  no  wider  at  base  than  apex  of  median 
frontal  sulcus.  Clypeal  suture  sharply  defined.  Eye  prominent,  finely  and  uniformly  faceted. 
Neck  finely,  densely  punctate.  Gena  coarsely  punctate,  rugose  in  front.  Antennal  articles 
four  to  ten  elongate,  article  five  about  1.4-1. 5 times  longer  than  wide. 

Pronotum.  Fig.  219.  Sides  bisetose;  base  not  rugose;  hind  angles  weakly  developed.  Para- 
median longitudinal  sulci  moderately  long,  nearly  straight,  strongly  hooked  basally.  Ante- 
rior transverse  impression  weakly  to  strongly  punctate. 

Legs.  Front  and  middle  tarsi  moderately  dilated  and  pubescent  ventrally,  less  so  in  fe- 
males; hind  tarsus  slender,  short.  Paronychia  nearly  as  long  as  tarsal  claws.  Front  tibia  nar- 
rowed evenly  to  base.  Front  femur  not  strongly  constricted  near  apex. 

Elytra.  Left  elytron  with  about  six  to  ten  setae  on  interval  three,  five  to  nine  on  interval 
five,  three  to  six  on  interval  seven;  total  14-24  in  specimens  examined.  Striae  sharply  en- 
graved, finely  punctate  in  basal  two-thirds.  Intervals  one  to  seven  broad  and  flat,  interval 
eight  carinate  at  apex;  intervals  three,  five,  and  seven  broadly  joined  at  apices.  Humeral 
denticles  weakly  developed. 

Abdomen.  Sternum  three  with  paramedian  carinae  curved  outward  at  apices.  Sternum 
seven  with  paramedian  ambulatory  setae  in  males  only.  Pygidium  with  apical  margin  finely 
crenulate  in  some  females. 

Male  genitalia.  Median  lobe,  Fig.  226,  227,  form  of  apex  variable  (Fig.  234);  20  specimens 
examined. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


271 


Measurements  and  proportions.  See  Table  44. 


Table  44.  Descriptive  statistics  for  S.  pygmaeus,  based  on  20  males  from  Clear  Lake, 
California. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

1 

A.  Measurements,  in  mm. 

TL 

2.80-3.22 

3.032 

0.152 

0.045 

3.35 

LE 

1.72-1.98 

1.861 

0.091 

0.027 

3.25 

WH 

0.57-0.66 

0.629 

0.038 

0.011 

4.03 

WP 

0.72-0.84 

0.792 

0.045 

0.013 

3.79 

WE 

0.89-1.02 

0.958 

0.043 

0.013 

2.98 

B.  Setae  on  lei 

ft  elytron. 

Interval  3 

7-  9 

8.0 

Interval  5 

7-  9 

7.6 

Interval  7 

3-  6 

4.6 

Total 

17-23 

20.2 

2.3 

0.7 

7.46 

C.  Proportions 

WF/WH 

0.64-0.70 

0.660 

0.021 

0.006 

2.11 

LP/WP 

0.91-0.98 

0.936 

0.023 

0.007 

1.67 

DP/LP 

0.77-0.83 

0.802 

0.023 

0.007 

1.91 

LP/WE 

0.74-0.80 

0.771 

0.022 

0.006 

1.88 

Ta/Ti 

0.55-0.70 

0.646 

0.052 

0.016 

5.40 

PS/LP 

0.66-0.73 

0.690 

0.027 

0.008 

2.57 

Variation.  — As  shown  in  Fig.  242,  specimens  from  interior  and  Pacific  drainage  basins 
south  to  and  including  the  Rio  Balsas  are  reddish,  without  conspicuous  variation.  Mature 
specimens  from  coastal  lowland  Atlantic  drainage  basins  in  Mexico,  and  from  most  localities 
south  of  Mexico,  are  dark  piceous.  In  Pacific  drainage  systems  in  Chiapas  and  Oaxaca,  most 
specimens  are  strongly  bicolored,  but  others  are  piceous  or  intermediate.  Two  specimens 
from  La  Lima,  Honduras  are  bicolored.  In  upland  areas  in  Atlantic  drainage  systems  in  both 
Oaxaca  and  Chiapas,  dark  and  bicolored  forms  are  commonly  found  together.  In  Guerrero 
south  of  the  Rio  Balsas,  and  in  upland  Atlantic  drainage  areas  in  San  Luis  Potosi,  most 
specimens  are  dark  red  and  many  are  slightly  bicolored.  Specimens  from  5000'  on  the  Rio 
Tula  in  Hidalgo  are  brown.  In  short,  there  is  a gradation  from  red  forms  ( S . pygmaeus ) to 
dark  forms  ( S . championi)  in  Mexico,  both  eastward  and  southward. 

Statistical  data  on  variation  in  body  size,  numbers  of  elytral  setae,  relative  eye  size,  and 
pronotal  form  are  given  in  Tables  45-48.  Major  discontinuities,  representing  strong  statis- 
tically significant  differences,  are  shown  in  Fig.  243-246  by  heavy  broken  lines.  Two  or 
more  such  gaps  are  found  in  central  California  (setae,  LP/WP),  between  southern  California 
and  Arizona  (setae,  WF/WH),  along  the  Sierra  Madre  Occidental  (LE,  setae,  WF/WH),  and 
along  the  Sierra  Madre  del  Sur  (LE,  setae,  LP/WP).  These  gaps  strongly  suggest  that  gene 


a tic 

fei 

N 

22 

16 

14 

16 

24 

12 

18 

24 

12 

18 

26 

16 

18 

10 

12 

20 

16 

12 

12 

10 

12 

26 

20 

16 

22 

12 

12 

24 

9 

6 

6 


Whitehead 


n body  size  (LE,  in  mm)  in  selected  samples  of  S.  pygmaeus ; see  Fig. 
es  each  50%  in  samples  1-28. 


Range 

Mean 

1.5SD 

2SE 

CV(%) 

1.77-2.00 

1.878 

0.091 

0.029 

3.58 

1.70-2.00 

1.907 

0.130 

0.044 

4.56 

1.71-2.16 

1.907 

0.152 

0.054 

5.32 

1.62-2.03 

1.841 

0.197 

0.066 

7.14 

1.87-2.15 

2.028 

0.128 

0.035 

4.20 

1.75-2.15 

1.970 

0.170 

0.065 

5.74 

1.80-2.16 

1.988 

0.142 

0.047 

4.76 

1.81-2.12 

1.954 

0.125 

0.034 

4.26 

1.83-2.13 

1.962 

0.164 

0.063 

5.57 

1.88-2.20 

2.019 

0.126 

0.040 

4.16 

1.80-2.23 

1.984 

0.164 

0.043 

5.51 

1.87-2.30 

2.078 

0.158 

0.053 

5.08 

1.68-1.95 

1.839 

0.106 

0.033 

3.83 

1.70-1.93 

1.817 

0.115 

0.050 

4.32 

1.75-2.07 

1.957 

0.140 

0.054 

4.77 

1.69-2.02 

1.860 

0.143 

0.043 

5.13 

1.82-2.12 

1.938 

0.115 

0.038 

3.96 

1.90-2.24 

2.027 

0.139 

0.054 

4.58 

1.73-1.95 

1.818 

0.101 

0.039 

3.70 

1.80-2.03 

1.910 

0.132 

0.055 

4.57 

1.75-2.08 

1.951 

0.153 

0.059 

5.24 

1.91-2.25 

2.114 

0.127 

0.033 

4.01 

1.86-2.14 

2.002 

0.114 

0.034 

3.78 

1.98-2.26 

2.103 

0.131 

0.044 

4.14 

1.88-2.25 

2.090 

0.154 

0.044 

4.91 

1.99-2.30 

2.113 

0.149 

0.057 

4.70 

1.99-2.29 

2.170 

0.132 

0.051 

4.04 

1.95-2.28 

2.092 

0.142 

0.039 

4.53 

1.96-2.21 

2.101 

2.00-2.21 

2.135 

1.99-2.12 

2.038 

Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


273 


Table  46. 
Fig.  244. 

Sample 


1 

2 

3 

4 

5 

6 

7 

8 
9 

10 

11 

12 

13 

14 

15 

16 

17 

18 

19 

20 

21 

22 

23 

24 

25 

26 

27 

28 

29 

30 

31 


Variation  in  numbers  of  elytral  setae  in  selected  samples  of  S.  pygmaeus ; see 
Males  and  females  each  50%  in  samples  1-28. 


N 

Range 

Mean 

1.5SD 

2SE 

CV(%] 

22 

18-23 

20.6 

1.7 

0.6 

6.30 

16 

19-22 

20.5 

1.5 

0.5 

4.94 

14 

16-19 

18.4 

1.3 

0.5 

4.62 

16 

19-22 

20.2 

1.3 

0.4 

4.23 

24 

17-23 

19.6 

2.0 

0.5 

6.85 

12 

18-23 

19.4 

2.3 

0.9 

8.06 

18 

15-22 

19.2 

2.6 

0.8 

9.02 

24 

17-22 

19.0 

2.1 

0.6 

7.54 

12 

17-21 

19.0 

1.6 

0.6 

5.50 

18 

16-20 

18.6 

1.7 

0.5 

6.19 

26 

15-21 

18.7 

2.1 

0.6 

7.68 

16 

17-21 

19.2 

2.1 

0.7 

7.18 

18 

14-18 

16.3 

1.5 

0.5 

6.30 

10 

15-17 

16.2 

0.9 

0.4 

3.90 

12 

15-19 

16.8 

1.7 

0.6 

6.62 

20 

15-18 

16.8 

1.5 

0.4 

5.44 

16 

14-18 

16.1 

2.0 

0.7 

8.44 

12 

16-20 

17.5 

1.5 

0.6 

5.71 

12 

14-18 

16.0 

1.7 

0.7 

7.05 

10 

15-18 

16.9 

1.7 

0.7 

6.51 

12 

16-20 

17.5 

1.8 

0.7 

6.67 

26 

16-20 

17.4 

1.6 

0.4 

6.11 

20 

16-20 

17.8 

1.6 

0.5 

5.93 

16 

18-21 

19.1 

1.6 

0.5 

5.57 

22 

16-22 

18.6 

2.3 

0.6 

8.07 

12 

18-21 

19.5 

1.5 

0.6 

5.13 

12 

19-24 

21.9 

2.1 

0.8 

6.29 

24 

19-25 

21.8 

2.0 

0.6 

6.20 

9 

19-24 

21.4 

6 

18-21 

19.5 

6 

17-19 

18.5 

274 


Whitehead 


Table  47. 
Fig.  245. 

Sample 


1 

2 

3 

4 

5 

6 

7 

8 
9 

10 

11 

12 

13 

14 

15 

16 

17 

18 

19 

20 

21 

22 

23 

24 

25 

26 

27 

28 

29 

30 

31 


Variation  in  relative  eye  size  (WF/WH)  in  selected  samples  of  S.  pygmaeus ; see 
Males  and  females  each  50%  in  samples  1-28. 


N 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

22 

0.65-0.67 

0.658 

0.013 

0.004 

1.30 

16 

0.64-0.67 

0.655 

0.017 

0.006 

1.76 

14 

0.64-0.68 

0.659 

0.017 

0.006 

1.73 

16 

0.60-0.66 

0.637 

0.022 

0.007 

2.27 

24 

0.60-0.65 

0.628 

0.018 

0.005 

1.97 

12 

0.61-0.65 

0.631 

0.017 

0.007 

1.85 

18 

0.60-0.64 

0.624 

0.015 

0.005 

1.58 

24 

0.60-0.65 

0.622 

0.019 

0.005 

2.08 

12 

0.60-0.62 

0.610 

0.01 1 

0.004 

1.21 

18 

0.60-0.64 

0.618 

0.016 

0.005 

1.72 

26 

0.59-0.65 

0.619 

0.024 

0.006 

2.60 

16 

0.60-0.64 

0.623 

0.015 

0.005 

1.63 

18 

0.59-0.63 

0.604 

0.016 

0.005 

1.81 

10 

0.59-0.64 

0.616 

0.025 

0.010 

2.67 

12 

0.59-0.63 

0.608 

0.021 

0.008 

2.31 

20 

0.58-0.62 

0.604 

0.020 

0.006 

2.23 

16 

0.59-0.63 

0.615 

0.015 

0.005 

1.68 

12 

0.59-0.62 

0.603 

0.016 

0.006 

1.78 

12 

0.58-0.62 

0.598 

0.020 

0.008 

2.23 

10 

0.58-0.62 

0.599 

0.019 

0.008 

2.15 

12 

0.57-0.61 

0.595 

0.022 

0.008 

2.43 

26 

0.58-0.61 

0.599 

0.015 

0.004 

1.52 

20 

0.58-0.62 

0.598 

0.018 

0.005 

1.90 

16 

0.58-0.63 

0.606 

0.020 

0.007 

2.16 

22 

0.60-0.64 

0.619 

0.018 

0.005 

1.92 

12 

0.62-0.65 

0.638 

0.016 

0.006 

1.66 

12 

0.61-0.65 

0.634 

0.020 

0.008 

2.07 

24 

0.60-0.66 

0.633 

0.023 

0.006 

2.40 

9 

0.60-0.65 

0.628 

6 

0.60-0.63 

0.615 

6 

0.58-0.63 

0.600 

Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


275 


Table  48.  Variation  in  pronotal  form  (LP/WP)  in  selected  samples  of  S.  pygmaeus',  see  Fig. 
246.  Males  and  females  each  50%  in  samples  1-28. 


Sample 

N 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

1 

22 

0.90-0.98 

0.934 

0.026 

0.007 

1.86 

2 

16 

0.92-0.97 

0.941 

0.020 

0.007 

1.42 

3 

14 

0.89-0.94 

0.913 

0.022 

0.008 

1.63 

4 

16 

0.89-0.95 

0.922 

0.025 

0.008 

1.84 

5 

24 

0.86-0.94 

0.907 

0.030 

0.008 

2.15 

6 

12 

0.87-0.95 

0.913 

0.036 

0.014 

2.61 

7 

18 

0.88-0.94 

0.909 

0.025 

0.008 

1.83 

8 

24 

0.88-0.93 

0.908 

0.023 

0.006 

1.71 

9 

12 

0.88-0.93 

0.899 

0.022 

0.008 

1.61 

10 

18 

0.88-0.93 

0.906 

0.021 

0.007 

1.53 

11 

26 

0.88-0.91 

0.919 

0.022 

0.006 

1.61 

12 

16 

0.89-0.95 

0.910 

0.028 

0.009 

2.08 

13 

18 

0.88-0.93 

0.904 

0.019 

0.006 

1.38 

14 

10 

0.87-0.94 

0.898 

0.032 

0.014 

2.39 

15 

12 

0.88-0.94 

0.916 

0.023 

0.009 

1.64 

16 

20 

0.88-0.94 

0.906 

0.021 

0.006 

1.54 

17 

16 

0.88-0.93 

0.897 

0.023 

0.008 

1.71 

18 

12 

0.89-0.93 

0.910 

0.019 

0.007 

1.41 

19 

12  ! 

0.88-0.93 

0.908 

0.023 

0.009 

1.70 

20 

,0  j 

0.88-0.91 

0.901 

0.016 

0.007 

1.22 

21 

12 

0.87-0.94 

0.914 

0.027 

0.011 

2.00 

22 

26 

0.87-0.93 

0.902 

0.023 

0.006 

1.73 

23 

20 

0.88-0.93 

0.905 

0.023 

0.007 

1.70 

24 

16 

0.88-0.93 

0.908 

0.023 

0.008 

1.67 

25 

22 

0.88-0.93 

0.904 

0.022 

0.006 

1.66 

26 

12 

0.89-0.95 

0.919 

0.030 

0.012 

2.20 

27 

. 12 

0.88-0.95 

0.912 

0.031 

0.012 

2.24 

28 

24  j 

0.89-0.94 

0.920 

0.022 

0.006 

1.60 

29 

9 

0.90-0.96 

0.927 

30 

6 

0.88-0.92 

0.905 

31 

6 

0.88-0.93 

0.910 

276 


Whitehead 


flow  is  impeded  across  these  regions. 

Fig.  234  portrays  observed  variation  in  form  of  male  genitalia  in  S.  pygmaeus.  Because  of 
considerable  individual  variation,  analysis  of  geographic  variation  is  difficult.  I found  no 
important  clinal  variation  except  for  a sudden  increase  in  size  of  apex  toward  northern 
California.  Specimens  from  this  area  cannot  be  distinguished  by  form  of  apex  of  male  geni- 
talia from  specimens  of  S.  scopaeus  from  northeastern  Mexico  and  the  lower  Rio  Grande  in 
Texas. 

I conclude  that  some  gene  flow  exists  between  narrow  bodied  forms  of  S.  pygmaeus  in 
central  and  northern  California  and  wider  bodied  forms  further  south,  and  between  small 
eyed  forms  in  California  and  large  eyed  forms  in  Arizona.  There  is  little  or  no  gene  flow 
across  the  higher  parts  of  the  Sierra  Madre  Occidental,  but  gene  flow  is  unrestricted  from 
north  to  south  on  both  sides  of  this  mountain  range.  South  of  the  Rio  Balsas  and  across  the 
Sierra  Madre  del  Sur,  gene  flow  is  limited.  Across  northern  Mexico  from  the  eastern  side  of 
the  Sierra  Madre  Occidental  to  the  Atlantic  coast,  there  is  no  evidence  that  gene  flow  is 
restricted.  Nor  is  gene  flow  restricted  along  the  Atlantic  coast  to  Middle  America,  across  to 
the  Pacific  in  Chiapas,  and  west  from  there  to  the  southern  side  of  the  Sierra  Madre  del  Sur. 
No  geographically  proximate  forms  are  reproductively  isolated,  and  no  forms  warrant  formal 
recognition  as  subspecies. 

Etymology.  — Latin,  pygmaeus  = dwarf,  a reference  to  the  small  size  of  these  beetles. 

Distribution.  — Specimens  of  S.  pygmaeus  have  been  found  at  elevations  ranging  from 
near  sea  level  to  about  6500'.  S.  pygmaeus  ranges  from  northern  California,  Arizona,  ex- 
treme western  New  Mexico,  southern  Chihuahua,  Durango,  Zacatecas,  San  Luis  Potosi, 
Nuevo  Leon,  and  Tamaulipas,  southward  at  least  to  Colombia  (Fig.  239).  I studied  1143 
specimens  of  this  species  from  the  following  localities. 

UNITED  STATES 

ARIZONA  (9;  ANSP,  CUNY,  MCZ,  UKSM,  USNM):  10  mi.  e.  Continental  (1;  UATA),  Hot  Springs  (2;  USNM),  Riverside 
(3;  CAS,  USNM);  Cochise  Co.,  Chiricahua  Mountains  (1;  UKSM),  Cochise  Stronghold  (4;  TLEr,  UATA),  Palominas  (1; 
UASM),  Portal  (1;  UCB),  5 mi.  w.  Portal  (4;  AMNH,  CNC),  Rucker  Lake  (1;  UASM);  Coconino  Co.,  Bill  Williams  Fork 
(1;  UKSM);  Gila  Co.,  Globe  (3;  MCZ,  UASM),  33  mi.  s.  Globe  (1;  DJLa),  Pinal  Mountains  (2;  UASM),  Rice  (3;  MCZ, 
UASM),  Salt  River  (2;  USNM),  San  Carlos  Lake  (1 ; UASM);  Graham  Co.,  Aravaipa  (19;  KHSt,  UASM),  Galiuro  Mountains 
(2;  USNM);  Graham  Mountains  (2;  UATA),  Power’s  Garden  (2;  UASM),  Thatcher  (1;  UCD):  Greenlee  Co.,  Gila  River 
(1;  KHSt);  Pima  Co.,  Ajo  Mountains  (1;  CAS),  Arivaca  (11;  CAS,  MCZ),  Baboquivari  Mountains  (6;  CAS,  MCZ),  Organ 
Pipe  National  Monument  (1;  CAS),  Quitobaquito  (17;  UASM),  Sabino  Canyon  (38;  AMNH,  CNC,  TCBa,  TLEr,  UATA, 
UCD,  UKSM),  Sahuarita  (1;  MCZ),  Santa  Catalina  Mountains  (4;  CAS,  KHSt),  Tanque  Verde  (14;  UATA),  Tucson  (8; 
CAS,  KHSt,  MCZ,  UATA,  USNM);  Pinal  Co.,  14  mi.  e.  Oracle  (1;  CAS),  Superior  (1;  CAS);  Santa  Cruz  Co.,  Madera  Canyon 
(4;  UATA,  UCD),  Nogales  (6;  CAS,  CNHM),  Patagonia  (12;  CAS,  CNHM,  CUNY,  KHSt,  TLEr,  UATA),  Pena  Blanca  (10; 
CNC,  KHSt,  UASM,  UATA),  Santa  Rita  Mountains  (1;  KHSt),  Tumacacori  Mountains  (17;  CAS);  Yuma  Co.,  Fort  Yuma 
(2;  USNM).  CALIFORNIA  (12;  ANSP,  CAS,  MCZ):  Alameda  Co.,  20  mi.  s.  Livermore  (1;  TLEr);  Butte  Co.,  Oroville 
(1;  CAS);  Calaveras  Co.,  Mokelumne  Hill  (6;  CAS),  El  Dorado  Co.  (1;  CNHM);  Fresno  Co.,  La  Fevre  Creek  (3;  CAS),  11.6 
mi.  s.  Tollhouse  (5;  TLEr);  Lake  Co.  (3;  CAS),  Clear  Lake  (59;  CAS),  North  Fork  Cache  Creek  (2;  UCD);  Los  Angeles  Co., 
Pasadena  (1;  CAS),  Pomona  (1 ; RUNB),  San  Gabriel  Canyon  (2;  TCBa),  Tujunga  Canyon  (4;  LACM,  MSUL);  Mendocino 
Co.,  Longvale  (1;  CAS);  Orange  Co.,  San  Juan  Guard  Station  (3;  GRNo);  Riverside  Co.,  Palm  Springs  (15;  AMNH,  CAS, 
CNC);  San  Bemadino  Co.  (1;  MCZ),  10  mi.  ne.  Earp  (2;  UCB),  Mojave  River  (1;  TLEr),  San  Bemadino  Mountains  (1; 
MGFT);  San  Diego  Co.,  Elsinore  Lake  (4;  CAS),  Julian  (4;  KHSt),  Poway  (4;  CAS),  Warners  Hot  Springs  (1;  MSUL);  Santa 
Barbara  Co.,  Santa  Barbara  (1;  CAS);  Santa  Clara  Co.,  East  Palo  Alto  (1;  CAS),  Gilroy  Hot  Springs  (3;  TLEr),  Mount 
Hamilton  (29;  CAS);  Sonoma  Co.,  Rio  Nido  (1;  CAS);  Stanislaus  Co.,  Del  Puerto  Creek  (1;  DHKa),  Patterson  (2;  CAS), 
20  mi.  w.  Patterson  (1;  TLEr);  Yolo  Co.,  Davis  (9;  UCD).  NEW  MEXICO:  Grant  Co.,  Cliff  (3;  DRWh),  21.9  mi.  ne.  Pinos 
Altos  (1;  UASM). 


MEXICO 

AGUASCALIENTES:  2.8  ml  s.  Aguascalientes  (1;  UASM).  BAJA  CALIFORNIA  (4;  CAS):  Catavinia  (4;  CAS),  20  mi.  n. 
Comondu  (1;  CAS),  Ensenada  (1;  UKSM),  12.4  mi  e.  La  Paz  (1;  CAS),  Las  Cruces  (7;  CAS),  5 mi.  nw.  Miraflores  (1; 
CAS),  5 mi.  s.  Miraflores  (1;  CAS),  5 mi.  w.  San  Bartolo  (11;  CAS),  Santa  Rosa  (2;  CNHM,  MCZ),  Santiago  (1;  CAS),  6 
mi.  sw.  Santiago  (4;  UATA),  Triunfo  (8;  CAS).  CHIAPAS:  3.2  mi.  n.  Arriaga  (15;  UASM),  20.9  mi.  n.  Arriaga  (19;  UASM), 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


277 


5.9  mi.  e.  Chiapa  de  Corzo  (1;  UASM),  12.2  mi.  ne.  Chiapa  de  Corzo  (13;  HFHo,  UASM),  32.5  mi.  e.  Comitan  (3;  BMNH), 
Huehuetan  (2;  IRSB),  Huixtla  (12;  UASM),  Macuilapa  (6;  FDAG),  San  Quintih  (38;  UASM),  Tonala  (2;  UASM),  18.6  mi. 
se.  Tonala  (9;  UASM).  CHIHUAHUA:  Catarinas  (7;  AMNH),  Parral  (1;  AMNH),  15  mi.  e.  Parral  (1;  AMNH).  COLIMA: 
3.4  mi.  se.  Colima  (5;  UASM),  8 mi.  sw.  Colima  (1;  UASM).  DURANGO:  12.2  mi.  s.  El  Banco  (30;  UASM),  4.2  mi.  w. 
Vicente  Guerrero  (8;  UASM).  GUANAJUATO:  9.8  mi.  s.  Silao  (2;  UASM).  GUERRERO:  Acapulco  (4;  MCZ),  24.8  mi.  e. 
Acapulco  (3;  UASM),  41.4  mi.  n.  Acapulco  (8;  UASM),  Coyuca  (4;  UASM),  23.7  mi.  n.  Zumpango  del  Rio  (1;  UASM), 
30.8  mi.  n.  Zumpango  del  Rio  (1;  UASM).  HIDALGO:  Tasquillo  (6;  UASM).  JALISCO:  Ajijic  (1;  CAS),  Cocula  (19; 
UASM),  9.7  mi.  e.  Encamacion  de  Diaz  (8;  UASM),  8.5  mi.  n.  Juchitlan  (10;  UASM),  10.6  mi.  s.  La  Huerta  (1;  UASM), 
17.7  mi.  nw.  Los  Volcanes  (1;  UASM),  17.9  mi.  w.  Magdalena  (2;  UASM),  Pitallal  (31;  UASM),  Puerto  Vallarta  (5;  UASM), 
Talpa  de  Allende  (4;  UASM).  MEXICO:  Tejupilco  (3;  MCZ).  MICHOACAN:  8.5  mi.  n.  Nueva  Italia  (32;  UASM),  20  mi.  n. 
Nueva  Italia  (3;  UASM).  MORELOS:  Tetecala  (2;  UASM).  NAYARIT:  2.4  mi.  s.  Acaponeta  (1;  UASM),  20.3  mi.  w. 
Compostela  (1;  GRNo),  14  mi.  e.  San  Bias  (3;  UASM).  NUEVO  LEON:  14.8  mi.  w.  Linares  (3;  UASM).  OAXACA:  17.7 
mi.  w.  El  Camaron  (2;  UASM),  25  mi.  w.  El  Camaron  (11;  UASM),  29.4  mi.  e.  El  Coyul  (1;  UASM),  Huitzo  (2;  BMNH), 

11.1  mi.  n.  Matias  Romero  (7;  UASM),  19  mi.  s.  Matias  Romero  (3;  CNC),  22.5  mi.  w.  Oaxaca  (2;  MGFT),  9.9  mi.  n. 
Pochutla  (15;  UASM),  Rio  Jalpan  (1;  FDAG),  Salina  Cruz  (3;  AMNH),  Tehuantepec  (2;  BMNH),  Totolapan  (1;  UKSM), 
Valle  Nacional  (1;  UASM),  Zanatepec  (25;  UASM),  18.4  mi.  w.  Zanatepec  (17;  UASM).  PUEBLA:  Acatlan  (2;  UCB),  9 mi. 
n.  Amatitlan  (2;  CAS),  Petlalcingo  (17;  UASM),  Tehuitzingo  (4;  UASM),  Tepexco  (13;  UASM).  QUERETARO:  Escanelilla 
(1;  UASM),  Jalpan  (3;  UASM).  SAN  LUIS  POTOSI:  14  mi.  e.  Ciudad  del  Maiz  (1;  CAS),  Ciudad  del  Valles  (1;  CBoP),  El 
Naranjo  (1;  CAS),  Huichihuayan  (1;  UKSM),  7.5  mi.  nw.  Mexquitic  (1;  UASM),  2.7  mi.  w.  Santa  Catarina  (23;  UASM), 
Tamazunchale  (1;  UASM),  19.3  mi.  nw.  Tamazunchale  (1;  UASM),  Vergel  (1 ; CBoP).  SINALOA:  Concordia  (18;  UASM), 

11.2  mi.  ne.  Concordia  (1;  UASM),  12  mi.  s.  Mazatlan  (5;  CAS,  TLEr,  UCB),  Rosario  (1;  UASM),  28  mi.  e.  Villa  Union. 
SONORA  (4;  CNC):  10  mi.  s.  Agua  Prieta  (1;  KHSt),  Alamos  (16;  CAS),  7 mi.  s.  Alamos  (10;  UCB,  UCD),  10  mi.  s. 
Alamos  (1;  UCD),  10  mi.  w.  Alamos  (5;  AMNH),  5 mi.  w.  Alamos  (1;  UATA),  16  mi.  ne.  Ciudad  Obregon  (1;  CNC), 
Hermosillo  (4;  CAS),  10  mi.  e.  Navajoa  (4;  UATA),  San  Carlos  Bay  (3;  CAS).  TAMAULIPAS:  Ciudad  Victoria  (1;  USNM), 

15.2  mi.  n.  Ciudad  Victoria  (1;  UASM),  21.3  mi.  n.  Ciudad  Victoria  (13;  UASM),  Encino  (2;  UASM).  VERACRUZ:  Bobo 
(1;  BMNH),  Catemaco  (2;  JNeg),  Fortin  de  las  Flores  (1;  FDAG),  20  mi.  nw.  Huatusco  (5;  FDAG),  21.8  mi.  e.  Jalapa  (9; 
UASM),  Paso  de  Ovejas  (12;  UASM).  ZACATECAS:  Jalpa  (6;  UASM),  Sain  Alto  (4;  UASM). 

GUATEMALA 

ALTA  VERAPAZ:  Trece  Aguas  (I;  USNM).  CHIQUIMULA:  Chiquimula  (5;  AMNH).  GUATEMALA:  Chinautla  (1; 
BMNH).  QUICHE:  Sacapulas  (1;  AMNH). 


EL  SALVADOR 

LA  PAZ:  La  Herradura  (4;  JNeg).  SAN  SALVADOR:  Guzapa  (4;  JNeg),  San  Salvador  (4;  JNeg). 

HONDURAS 

COMAYAGUA:  Rancho  Chiquito  (4;  FDAG).  CORTES:  La  Lima  (2;  FDAG).  El  Zamarano  (1;  OSUC). 

NICARAGUA 

RIVAS:  10  km.  nw.  Sapoa  (1;  FDAG). 

COSTA  RICA 

GUANACASTE:  5 km.  n.  Canas  (1;  LACM).  LIMON:  Los  Diamantes  (1;FDAG).  PUNTARENAS:  Palmar  Sur  (1;  UAFA), 
6 mi.  n.  Palmar  Sur  (12;  UAFA).  7 mi.  nw.  Palmar  Sur  (1;  UAFA),  Villa  Neilly  (1;  FDAG). 

COLOMBIA 

MAGDALENA:  Aracataca  (1;  MCZ),  Rio  Frio  (7;  MCZ). 


Collecting  notes.  — Specimens  of  this  species  have  been  taken  throughout  the  year,  at 
lights  or  in  riparian  gravel  bars.  At  most  localities  where  specimens  of  S.  pygmaeus  and  S. 
falli  have  been  collected  together,  one  of  the  two  species  was  strongly  numerically  domi- 
nant. 

Taxonomic  notes.  — Although  I studied  holotypes  of  neither  S.  pygmaeus  nor  S.  cham- 
pioni,  I did  study  paratypes  of  both  and  have  no  doubt  that  my  association  of  the  names  is 
correct.  As  shown  in  my  analysis  of  geographic  variation  above,  these  two  names  clearly 
refer  to  a single  species.  Lindroth  (1961)  suggested  that  the  form  in  southern  California 
might  be  a distinct  species,  but  though  most  specimens  are  distinguishable  from  topotypic 
specimens  of  S.  pygmaeus,  there  is  no  evidence  for  reproductive  isolation. 


278 


Whitehead 


Schizogenius  scopaeus  new  species 

Type  material.  — Holotype  male  and  allotype  female  labelled  “Limpia  Canyon,  2 mi.  n.w. 
Fort  Davis,  Texas.  3 .VIII .63  D.R.  Whitehead”  (MCZ).  An  additional  98  specimens  from 
Jeff  Davis  County,  Texas  are  paratypes  (AMNH,  ANSP.  BMNH.  CAS,  CNC,  CNHM.  DJLa, 
DRWh,  IRSB,  MGFT,  UASM,  UCD.  USNM). 

Diagnostic  combination.  — Specimens  of  this  species  are  reliably  distinguished  from  red 
specimens  of  S.  pygmaeus  only  by  form  of  apex  of  male  median  lobe,  and  by  geographic 
distribution. 

Description.  - As  in  S.  pygmaeus  except  as  follows.  Body  color  testaceous  to  ferrugi- 
neous,  not  brunneous,  piceous,  bicolored,  or  aeneous.  Left  elytron  with  about  eight  to 
eleven  setae  on  interval  three,  seven  to  ten  on  interval  five,  four  to  seven  on  interval  seven; 
total  18-27  in  specimens  examined.  Male  genitalia  with  median  lobe,  Fig.  228-229,  apex  in 
most  specimens,  particularly  western  specimens,  broader  than  in  specimens  of  S.  pygmaeus 
(Fig.  234);  20  specimens  examined. 

Measurements  and  proportions.  See  Table  49.  Of  holotype:  TL,  3.25  mm;  LE,  2.00  mm; 
WH,  0.66  mm;  WP.  0.84  mm;  WE,  1.04  mm;  WF/WH,  0.64;  LP/WP,  0.93;  DP/LP,  0.81; 
LP/WE,  0.75;  Ta/Ti,  0.60;  PS/LP.  0.68.  Of  allotype:  TL,  3.50  mm;  LE,  2.18  mm;  WH,  0.71 
mm;  WP,  0.92  mm;  WE,  1.12  mm;  WF/WH,  0.63;  LP/WP,  0.90;  DP/LP,  0.78;  LP/WE,  0.74; 
Ta/Ti,  0.64;  PS/LP,  0.72. 


Table  49.  Descriptive  statistics  for  S.  scopaeus , based  on  20  males  from  Limpia  Canyon, 
Texas. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

nts,  in  mm. 

TL 

3.04-3.60 

3.265 

0.199 

0.059 

4.07 

LE 

1.86-2.22 

2.010 

0.144 

0.043 

4.78 

WH 

0.64-0.75 

0.685 

0.040 

0.012 

3.89 

WP 

0.79-0.93 

0.848 

0.050 

0.015 

3.91 

WE 

0.97-1.18 

1.054 

0.078 

0.023 

4.93 

B.  Setae  on  lei 

4 elytron. 

Interval  3 

9-1 1 

9.6 

Interval  5 

7-10 

8.6 

Interval  7 

4-  7 

5.4 

Total 

20-26 

23.5 

2.3 

0.7 

6.55 

C.  Proportions 

WF/WH 

0.60-0.66 

0.628 

0.023 

0.007 

2.47 

LP/WP 

0.88-0.95 

0.924 

0.026 

0.008 

1.86 

DP/LP 

0.78-0.84 

0.804 

0.022 

0.007 

1.86 

LP/WE 

0.72-0.77 

0.747 

0.023 

0.007 

2.04 

Ta/Ti 

0.60-0.70 

0.658 

0.044 

0.013 

4.48 

PS/LP 

0.64-0.77 

0.694 

0.038 

0.01 1 

3.67 

Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


279 


Variation.  — Statistical  data  on  variation  in  body  size,  numbers  of  elytral  setae,  relative 
eye  size,  and  pronotal  form  are  given  in  Tables  50-53  and  summarized  in  Fig.  243-246. 
From  north  to  south,  numbers  of  elytral  setae  increase,  eyes  increase  in  size,  and  pronota 
narrow.  Specimens  from  central  Texas  tend  to  be  smaller  than  specimens  from  the  Rio 
Grande  Valley  to  the  South  and  from  Arkansas  and  Missouri  to  the  north.  The  form  of  apex 
of  median  lobe  of  male  genitalia  (Fig.  234)  shows  considerable  variation;  in  particular,  the 
apex  is  much  larger  in  specimens  from  Colorado  and  western  Texas. 

Etymology.  - Greek,  skopaios  = dwarf.  This  word  is  an  etymological  synonym  of  the 
Latin  pygmaeus,  chosen  to  indicate  the  great  similarity  of  these  two  species. 

Distribution.  - S.  scopaeus  ranges  over  an  area  limited  in  the  north  by  the  Missouri  River, 
in  the  east  by  the  Mississippi  River,  in  the  west  by  the  Rocky  Mountains,  and  in  the  south 
by  the  Rio  Grande  except  for  a small  area  in  northeastern  Mexico  (Fig.  238).  I studied  375 
specimens  of  this  species  from  the  following  localities. 

UNITED  STATES 

No  locality  (1;  ANSP).  ARKANSAS:  Carroll  Co.,  5 mi.  w.  Berryville  (15;  UASM);  Washington  Co.,  7 mi.  s.  Fayetteville 
(7;  UASM).  COLORADO:  Baca  Co.,  Regnier  (1 ; AMNH);  Pueblo  Co.,  38  mi.  e.  Walsenburg  (6;  UASM).  MISSOURI:  Butler 
Co.,  12  mi.  se.  Elsinore  (5;  CAS);  Crawford  Co.,  Meramec  River  (1;  USNM);  McDonald  Co.,  3 mi.  n.  Noel  (1;  CAS); 
Reynolds  Co.,  Ellington  (2;  USNM);  Ripley  Co.,  5.5  mi.  n.  Briar  (8;  CAS);  Wayne  Co.,  Williamsville  (1;  CNC).  OKLAHO- 
MA: Atoka  Co.,  5 mi.  n.  Stringtown  (4;  UASM);  Carter  Co.,  10.7  mi.  s.  Drake  (1 ; TLEr);  Cimarron  Co.,  Black  Mesa  State 
Park  (1;  CAS);  Comanche  Co.,  Wichita  National  Forest  (13;  CAS);  Cotton  Co.  (1;  CAS);  Murray  Co.,  10.3  mi.  n.  Drake  (4; 
TLEr).  TEXAS  (11;  AMNH,  CAS,  CNHM,  INHS,  MCZ,  MSUL,  UKSM,  USNM):  Austin  Co.,  Austin  (8;  CAS,  USNM). 
Bexar  Co.,  San  Antonio  (1;  CAS);  Blanco  Co.,  Cypress  Mills  (2;  USNM),  Johnson  City  (12;  UASM),  Twin  Sisters  (2; 
UASM);  Brewster  Co.,  Alpine  (1;  MCZ),  Big  Bend  National  Park  (1;  CNC),  Green  Valley  (1;  CAS),  22  mi.  s.  Marathon 
(3;  UCD);  Culberson  Co.,  2.5  mi.  e.  Nickle  Creek  Station  (1;  CNHM);  Jeff  Davis  Co.,  Barrel  Springs  Creek  (8;  BMNH, 
DRWh,  IRSB,  MGFT),  Davis  Mountains  State  Park  (4;  CAS,  DJLa),  Fort  Davis  (21;  AMNH,  ANSP,  CAS,  CNC,  MCZ, 
UCD),  4 mi.  w.  Fort  Davis  (4;  CNC),  Limpia  Canyon  (64;  AMNH,  CNC,  CNHM,  MCZ,  UASM,  USNM);  Kinney  Co.,  23  mi. 
sw.  Brackettville  (6;  UASM);  Lampasas  Co.,  Adamsville  (2;  JNeg);  Llano  Co.,  Enchanted  Rock  (13;  CNC);  Maverick  Co., 
8 mi.  n.  Quemado  (9;  UASM);  McCulloch  Co.,  Brady  (2;  KSUM),  16  mi.  s.  Brady  (33;  CAS),  San  Saba  Camp  (1;  KSUM); 
Terrell  Co.,  Chandler  Ranch  (6;  UASM),  Independence  Creek  (14;  UASM);  Val  Verde  Co.,  Del  Rio  (4;  CNC,  USNM),  13 
mi.  nw.  Del  Rio  (17;  UASM). 

MEXICO 

NUEVO  LEON:  Cienega  de  Flores  (8;  UASM),  Linares  (1;  JHeS),  Montemorelos  (11;  UASM),  32.9  mi.  n.  Montemorelos 
(1;  UASM),  5 mi.  s.  Monterrey  (16;  CNC,  HFHo),  6 mi.  s.  Monterrey  (1;  FDAG),  4.8  mi.  e.  Sabinas  Hidalgo  (1;  UASM). 
TAMAULIPAS:  39  mi.  s.  Ciudad  Victoria  (1;  CBoP),  Villagran  (11;  UASM). 


Collecting  notes.  — Specimens  of  S.  scopaeus  are  abundant  in  gravel  bars  along  both 
permanent  and  intermittent  streams.  In  northeastern  Mexico  they  are  found  in  less  protected 
places  than  are  specimens  of  S.  pygmaeus , such  as  along  the  intermittent  Arroyo  Villagran 
in  Tamaulipas.  All  specimens  were  collected  between  April  and  October.  On  repeated  occa- 
sions, specimens  have  been  taken  at  lights  in  the  vicinity  of  Fort  Davis  and  Limpia  Canyon, 
Texas. 

Taxonomic  notes.  — I recognize  S.  scopaeus  as  a new  species  distinct  from  S.  pygmaeus 
because  these  taxa  overlap  in  range  in  northeastern  Mexico  and  because  I cannot  show  that 
they  form  a continuous  circle  of  races  through  Durango,  Chihuahua,  and  western  Texas. 
Such  a circle  of  races  is  quite  possible,  and  is  strongly  suggested  in  statistical  comparisons 
summarized  in  Fig.  243-246.  But  male  genitalia  in  this  area  are  divergent  (Fig.  234);  western 
forms  of  S.  scopaeus , so  far  as  studied,  have  broad  apices  while  those  of  S.  pygmaeus  in 
Durango  and  Chihuahua  have  narrow  apices.  A detailed  study  of  material  from  the  Rio 
Conchos  system  in  Chihuahua  will  be  required  to  finally  decide  whether  the  forms  here 
separated  as  S.  scopaeus  and  S.  pygmaeus  are  conspecific  or  not.  A stepped  cline  in  the 
genitalic  characteristic  is  surely  a possibility.  However,  S.  scopaeus  is  largely  replaced  by  the 
related  S.  falli  in  the  Big  Bend  region  of  Texas,  and  Chihuahua  S.  pygmaeus  and  Texas  S. 
scopaeus  may  therefore  be  geographically  isolated  along  the  lower  parts  of  the  Rio  Conchos. 


280 


Whitehead 


Table  50.  Variation  in  body  size  (LE,  in  mm)  in  selected  samples  of  S.  scopaeus ; see  Fig. 
243.  Males  and  females  each  50%. 


Sample 

N 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

1 

22 

1.77-2.10 

1.940 

0.122 

0.035 

4.19 

2 

26 

1.83-2.15 

1.975 

0.122 

0.032 

4.10 

3 

14 

1.82-2.05 

1.964 

0.096 

0.034 

3.24 

4 

22 

1.89-2.28 

2.068 

0.151 

0.043 

4.86 

5 

22 

1.71-2.01 

1.879 

0.122 

0.035 

4.34 

6 

20 

1.71-2.02 

1.888 

0.140 

0.042 

4.95 

7 

20 

1.85-2.14 

1.971 

0.139 

0.041 

4.71 

8 

12 

1.87-2.18 

1.975 

0.121 

0.047 

4.09 

Table  51.  Variation  in  numbers  of  elytral  setae  in  selected  samples  of  S.  scopaeus',  see  Fig. 
244.  Males  and  females  each  50%. 


Sample 

N 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

1 

22 

21-27 

23.0 

2.6 

0.7 

7.51 

2 

26 

20-27 

23.2 

2.4 

0.6 

7.01 

3 

14 

21-27 

22.9 

2.8 

1.0 

8.20 

4 

22 

20-28 

23.6 

2.4 

0.7 

6.85 

c 

22 

18-26 

21.6 

2.6 

0.7 

8.01 

6 

20 

19-25 

21.6 

2.0 

0.6 

6.27 

7 

20 

18-23 

20.6 

1.7 

0.5 

5.50 

8 

12 

19-24 

21.0 

1.9 

0.7 

6.09 

Table  52.  Variation  in  relative  eye  size  (WF/WH)  in  selected  samples  of  S.  scopaeus',  see 
Fig.  245.  Males  and  females  each  50%. 


Sample 

N 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

1 

22 

0.59-0.65 

0.616 

0.019 

0.005 

2.05 

2 

26 

0.60-0.65 

0.624 

0.020 

0.005 

2.18 

3 

14 

0.61-0.64 

0.629 

0.015 

0.005 

1.58 

4 

22 

0.60-0.65 

0.629 

0.023 

0.006 

2.42 

5 

22 

0.63-0.66 

0.640 

0.012 

0.003 

1.23 

6 

20 

0.62-0.67 

0.643 

0.025 

0.008 

2.63 

7 

20 

0.63-0.68 

0.650 

0.020 

0.006 

2.09 

8 

12 

0.64-0.68 

0.654 

0.020 

0.008 

2.00 

Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


281 


Table  53.  Variation  in  pronotal  form  (LP/WP)  in  selected  samples  of  S.  scopaeus ; see  Fig. 
246.  Males  and  females  each  50%. 


Sample 

N 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

1 

22 

0.89-0.96 

0.922 

0.030 

0.009 

2.19 

2 

26 

0.88-0.97 

0.925 

0.028 

0.007 

1.99 

3 

14 

0.90-0.96 

0.931 

0.031 

0.011 

2.22 

4 

22 

0.89-0.96 

0.922 

0.026 

0.007 

1.88 

5 

22 

0.87-0.95 

0.911 

0.030 

0.009 

2.23 

6 

20 

0.87-0.94 

0.914 

0.025 

0.007 

1.82 

7 

20 

0.89-0.93 

0.904 

0.015 

0.004 

1.10 

8 

12 

0.87-0.93 

0.902 

0.026 

0.010 

1.96 

Schizogenius  falli  new  species 

Type  material  — Holotype  male  and  allotype  female  labelled  “MEX.  Nuevo  Leon  Rio 
Sabinas  Hidalgo,  4.8  mi.  e.  Sabinas  Hidalgo  800'  X.22-23.  65”  and  “George  E.  Ball  D. 
R.  Whitehead  collectors”  (MCZ).  An  additional  34  specimens  from  various  localities  in 
Nuevo  Leon  are  paratypes  (BMNH,  CAS,  CNC,  CNHM,  DRWh,  IRSB,  MGFT,  UASM, 
USNM). 

Diagnostic  combination.  — Within  the  depressus  group,  specimens  of  this  species  are 
recognized  by  the  following  combination  of  characters:  pale  color;  small  size;  abdomen 
without  extensive  microsculpture;  and  frontal  carinae  strongly  fused  basally. 

Description.  — As  in  S.  pygmaeus  except  as  follows.  Body  color  testaceous  to  ferrugi- 
neous,  not  brunneous,  piceous,  bicolored,  or  aeneous.  Head  and  pronotum,  Fig.  220;  fron- 
tal carinae  confused  basally;  pronotal  hind  angles  more  prominent.  Left  elytron  with  about 
seven  to  eleven  setae  on  interval  three,  six  to  ten  on  interval  five,  four  to  six  on  interval 
seven;  total  17-27  in  specimens  examined.  Male  genitalia  with  median  lobe,  Fig.  230;  10 
specimens  examined. 

Measurements  and  proportions.  See  Table  54.  Of  holotype:  TL,  3.80  mm;  LE,  2.30  mm; 
WH,  0.80  mm;  WP,  1.02  mm;  WE,  1 .20  mm;  WF/WH,  0.66;  LP/WP,  0.94;  DP/LP,  0.76;  LP/ 
WE,  0.79;  Ta/Ti,  0.59;  PS/LP,  0.71.  Of  allotype:  TL,  3.78  mm;  LE,  2.34  mm;  WH,  0.80 
mm;  WP,  0.99  mm;  WE,  1.18  mm;  WF/WH,  0.66;  LP/WP,  0.93;  DP/LP,  0.77;  LP/WE,  0.78; 
Ta/Ti,  0.56;  PS/LP,  0.70. 

Variation.  — Data  on  variation  in  body  size,  numbers  of  elytral  setae,  relative  eye  size, 
and  pronotal  form  are  given  in  Tables  55-58,  and  summarized  in  Fig.  247-250.  In  general, 
body  size  increases  from  west  to  east,  with  no  important  gaps  between  geographically  proxi- 
mate samples  (Fig.  247).  Specimens  from  interior  parts  of  the  range  tend  to  have  more  setae 
than  do  those  from  peripheral  areas,  and  there  is  a statistically  significant  difference  be- 
tween the  Chihuahua  and  Durango  samples  (16  and  17)  (Fig.  248).  Relative  eye  size  (Fig. 
249)  tends  to  decrease  from  west  to  east,  and  there  is  a statistically  significant  difference 
between  the  Boquillas  and  Marathon  samples  (9  and  10).  Specimens  from  the  northeast 
and  southwest  of  the  range  (Fig.  250)  tend  to  have  wider  pronota  than  those  from  else- 
where, and  the  samples  from  Cochise  Stronghold  and  Portal  (5  and  6)  are  statistically 
significantly  different  in  this  characteristic.  For  a more  extended  discussion,  see  the  taxo- 
nomic notes  section  under  S.  ochthocephalus. 


282 


Whitehead 


Table  54.  Descriptive  statistics  for  S.  falli,  based  on  20  males  from  4.8  miles  east  of  Sabinas 
Hidalgo,  Nuevo  Leon. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

nts,  in  mm. 

TL 

3.52-4.18 

3.858 

0.237 

0.071 

4.10 

LE 

2.15-2.58 

2.358 

0.150 

0.045 

4.25 

WH 

0.74-0.85 

0.794 

0.041 

0.012 

3.40 

WP 

0.93-1.09 

1.019 

0.065 

0.019 

4.27 

WE 

1.10-1.29 

1.203 

0.071 

0.021 

3.91 

B.  Setae  on  lei 

ft  elytron. 

Interval  3 

7-  9 

8.2 

Interval  5 

6-  9 

7.5 

Interval  7 

4-  6 

5.3 

Total 

19-24 

21.0 

1.7 

0.5 

5.44 

C.  Proportion: 

WF/WH 

0.65-0.70 

0.671 

0.020 

0.006 

1.99 

LP/WP 

0.92-0.97 

0.942 

0.020 

0.006 

1.40 

DP/LP 

0.75-0.78 

0.762 

0.013 

0.004 

1.09 

LP/WE 

0.78-0.83 

0.801 

0.022 

0.007 

1.85 

Ta/Ti 

0.56-0.62 

0.594 

0.024 

0.007 

2.75 

PS/LP 

0.67-0.76 

0.709 

0.032 

0.009 

3.00 

Etymology.  — I dedicate  the  name  S.  falli  in  recognition  of  the  many  important  contri- 
butions to  our  knowledge  of  North  American  Coleoptera  made  by  the  late  H.  C.  Fall. 

Distribution.  - The  known  range  of  S.  falli  extends  west  from  southern  Illinois  to  ex- 
treme southern  California  and  south  to  Colima  (Fig.  240).  I studied  1069  specimens  from 
the  following  localities. 


UNITED  STATES 

No  locality  (2;  ANSP).  ARIZONA  (13;  AMNH,  ANSP,  INHS,  MCZ,  USNM):  Cochise  Co.,  Bear  Canyon  (1;CUNY),  Benson 
(1;  UATA),  Carr  Canyon  (1;  CAS),  Cave  Creek  (4;  UASM),  Chiricahua  Mountains  (2;  CAS,  USNM),  Cochise  Stronghold 
(42;  DJLa,  TLEr,  UATA),  Douglas  (2;  CNHM,  KHSt),  Guadalupe  Canyon  (1;  KHSt),  Portal  (4;  UCB),  5 mi.  w.  Portal 
(7;  AMNH,  CUNY,  FDAG,  UCB,  UCD),  Texas  Pass  (15;  CUNY),  Whetstone  Mountains  (2;  CAS),  Wilcox  (1;CAS),  8 mi. 
e.  Wilcox  (3;  GRNo);  Gila  Co.,  Globe  (32;  CAS,  CNC,  MCZ,  UASM,  USNM),  33  mi.  s.  Globe  (2;  DJLa),  Rice  (4;  MCZ, 
UASM),  San  Carlos  Lake  (16;  CUNY,  MCZ,  UASM);  Graham  Co.,  Aravaipa  (2;  CAS,  UASM),  Galiuro  Mountains  (1; 
USNM),  Marijilda  Canyon  (2;  CAS,  UATA),  Power’s  Garden  (8;  UASM),  Thatcher  (5;  UCD);  Maricopa  Co.,  Haslampa 
District  (1;  CAS),  Phoenix  (6;  MCZ,  USNM),  Salt  River  (2;  MCZ),  Tempe  (1;  USNM),  Verde  River  (4;  LACM,  MCZ), 
Wickenburg  (1;  LACM);  Mojave  Co.,  Kingman  (1;  UKSM);  Pima  Co.,  Alamo  Canyon  (3;  CAS),  Arivaca  (8;  CAS,  MCZ, 
UKSM),  Azo  (3;  CAS),  Baboquivari  Canyon  (6;  CAS),  Brown  Canyon  (4;  CAS,  UATA),  Catalina  Springs  (2;  USNM), 
Collosal  Cave  Park  (4;  KHSt),  Elkhom  Ranch  (1;  CAS),  Kits  Peak  Rincon  (2;  AMNH),  Organ  Pipe  National  Monument 
(10;  UATA,  UCD),  Pantano  (3;  CUNY),  Pepper  Sauce  Canyon  (1;  CAS),  17  mi.  w.  Quijotoa  (1;  UASM),  Quitobaquito 
(3;  AMNH,  UASM),  Robles  Ranch  (3;  UCD),  Sabino  Canyon  (15;  CAS,  RTBe,  TCBa,  TLEr,  UCD,  UKSM,  USNM),  Santa 
Catalina  Mountains  (11;  CAS,  KHSt,  UATA),  Staghorn  Ranch  (1;  CNC),  Tanque  Verde  (4;  UATA),  Tucson  (101;  ANSP, 
CAS,  JNeg,  KHSt,  UATA,  UCB,  USNM);  Pinal  Co.,  7 mi.  ne.  Apache  Junction  (3;  GRNo),  10  mi.  w.  Casa  Grande  (1; 
CAS),  Oracle  (2;  USNM),  14  mi.  e.  Oracle  (1;  CAS),  Superior  (3;  UATA);  Santa  Cruz  Co.,  11.5  mi.  se.  Arivaca  (1;  CAS), 
Black  Dike  Prospect  (2;  AMNH),  Calabasas  Canyon  (3;  UASM),  Canelo  (6;  UATA),  Madera  Canyon  (30;  DHKa,  UATA, 
UCD),  Nogales  (32;  AMNH,  CAS,  CNHM,  UASM,  UCD),  Patagonia  (10;  CAS,  CNHM,  TLEr,  UASM,  UATA),  Pena  Blanca 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


283 


Table  55.  Variation  in  body  size  (LE,  in  mm)  in  selected  samples  of  S.  falli  and  S.  ochtho- 
cephalus  (sample  one  from  Davis,  California);  see  Fig.  247.  Males  and  females  each  50%. 


Sample 

N 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

1 

12 

2.19-2.50 

2.292 

0.136 

0.053 

3.97 

2 

12 

2.09-2.47 

2.247 

0.167 

0.064 

4.95 

3 

24 

2.08-2.42 

2.211 

0.166 

0.045 

4.99 

4 

20 

2.00-2.38 

2.182 

0.136 

0.041 

4.16 

5 

20 

2.05-2.45 

2.208 

0.145 

0.043 

4.38 

6 

14 

2.00-2.40 

2.229 

0.155 

0.055 

4.64 

7 

14 

2.14-2.40 

2.281 

0.096 

0.042 

3.43 

8 

14 

2.17-2.52 

2.340 

0.136 

0.048 

3.87 

9 

20 

2.00-2.59 

2.309 

0.234 

0.070 

6.75 

10 

20 

2.17-2.47 

2.320 

0.158 

0.047 

4.54 

11 

10 

2.10-2.50 

2.295 

0.192 

0.081 

5.57 

12 

18 

2.20-2.58 

2.368 

0.165 

0.052 

4.64 

13 

20 

1.95-2.35 

2.186 

0.198 

0.059 

6.05 

14 

20 

1.87-2.39 

2.080 

0.201 

0.060 

6.44 

15 

20 

1.90-2.33 

2.170 

0.195 

0.058 

5.97 

16 

20 

2.04-2.55 

2.304 

0.215 

0.064 

6.23 

17 

18 

2.15-2.50 

2.326 

0.140 

0.044 

4.02 

18 

20 

2.20-2.58 

2.396 

0.183 

0.055 

5.09 

Table  56.  Variation  in  numbers  of  < 

slytral  setae  in  selected  samples 

of  S.  falli ; 

and  S.  ochtho- 

cephalus  (sample  one 

from  Davis, 

California);  see  Fig. 

248.  Males  and  females  each  50%. 

Sample 

N 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

1 

12 

18-22 

19.9 

1.7 

0.7 

5.85 

2 

12 

19-23 

21.0 

1.8 

0.7 

5.74 

3 

24 

19-25 

21.2 

2.0 

0.6 

6.37 

4 

20 

19-24 

21.0 

2.0 

0.6 

6.29 

5 

20 

19-25 

21.5 

2.3 

0.7 

7.16 

6 

14 

20-25 

21.9 

1.8 

0.6 

5.50 

7 

14 

20-25 

21.7 

2.2 

0.8 

-6.62 

8 

14 

20-23 

21.6 

1.5 

0.5 

4.71 

9 

20 

19-24 

21.8 

2.1 

0.6 

6.48 

10 

20 

19-24 

21.1 

2.4 

0.7 

7.52 

11 

10 

20-24 

21.3 

2.0 

0.8 

6.28 

12 

18 

18-25 

21.2 

2.3 

0.7 

7.29 

13 

20 

18-24 

21.4 

2.6 

0.8 

8.08 

14 

20 

17-26 

21.0 

3.4 

1.0 

10.73 

15 

20 

18-24 

21.1 

2.2 

0.6 

6.86 

16 

20 

17-25 

20.4 

2.8 

0.8 

9.06 

17 

18 

19-27 

22.1 

2.6 

0.8 

7.85 

18 

20 

19-24 

21.4 

2.1 

0.6 

6.68 

284 


Whitehead 


Table  57.  Variation  in  relative  eye  size  (WF/WH)  in  selected  samples  of  S.  falli  and  S.  och- 
thocephalus  (sample  one  from  Davis,  California);  see  Fig.  249.  Males  and  females  each  50%. 


Sample 

N 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

1 

12 

0.65-0.69 

0.671 

0.017 

0.007 

1.74 

2 

12 

0.62-0.66 

0.636 

0.016 

0.006 

1.70 

3 

24 

0.62-0.66 

0.645 

0.015 

0.004 

1.58 

4 

20 

0.63-0.67 

0.651 

0.015 

0.004 

1.49 

5 

20 

0.63-0.66 

0.644 

0.014 

0.004 

1.47 

6 

14 

0.62-0.66 

0.641 

0.017 

0.006 

1.78 

7 

14 

0.63-0.68 

0.654 

0.025 

0.009 

2.52 

8 

14 

0.63-0.68 

0.653 

0.019 

0.007 

1.94 

9 

20 

0.63-0.68 

0.666 

0.021 

0.006 

2.14 

10 

20 

0.63-0.66 

0.648 

0.019 

0.006 

1.91 

11 

10 

0.65-0.69 

0.669 

0.024 

0.010 

2.38 

12 

18 

0.64-0.68 

0.661 

0.014 

0.004 

1.42 

13 

20 

0.62-0.66 

0.647 

0.016 

0.005 

1.67 

14 

20 

0.61-0.67 

0.650 

0.022 

0.007 

2.26 

15 

20 

0.63-0.67 

0.652 

0.019 

0.006 

1.98 

16 

20 

0.65-0.70 

0.669 

0.026 

0.008 

2.61 

17 

18 

0.64-0.68 

0.662 

0.021 

0.007 

2.09 

18 

20 

0.65-0.70 

0.673 

0.026 

0.008 

2.56 

Table  58.  Variation  in  pronotal  form  (LP/WP)  in 

selected  samples 

of  S.  falli  i 

ind  S.  ochtho- 

cephalus  (sample  one 

from  Davis,  < 

California);  see  Fig.  250.  Males  and  females  each  50%. 

Sample 

N 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

1 

12 

0.93-0.96 

0.952 

0.014 

0.005 

0.98 

2 

12 

0.94-0.98 

0.958 

0.017 

0.006 

1.16 

3 

24 

0.90-0.99 

0.950 

0.035 

0.010 

2.47 

4 

20 

0.92-0.98 

0.952 

0.024 

0.007 

1.71 

5 

20 

0.93-0.98 

0.956 

0.028 

0.008 

1.96 

6 

14 

0.90-0.97 

0.937 

0.025 

0.009 

1.80 

7 

14 

0.92-0.98 

0.944 

0.023 

0.008 

1.65 

8 

14 

0.92-0.99 

0.950 

0.030 

0.011 

2.11 

9 

20 

0.90-1.00 

0.950 

0.030 

0.009 

2.11 

10 

20 

0.94-0.98 

0.952 

0.015 

0.004 

1.06 

11 

10 

0.90-0.96 

0.936 

0.030 

0.013 

2.15 

12 

18 

0.90-0.96 

0.931 

0.024 

0.008 

1.72 

13 

20 

0.93-0.98 

0.935 

0.017 

0.005 

1.18 

14 

20 

0.92-0.99 

0.947 

0.032 

0.010 

2.25 

15 

20 

0.92-0.97 

0.946 

0.018 

0.006 

1.31 

16 

20 

0.91-0.98 

0.944 

0.025 

0.008 

1.80 

17 

18 

0.90-0.99 

0.944 

0.036 

0.011 

2.57 

18 

20 

0.92-0.98 

0.948 

0.022 

0.007 

1.56 

Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


285 


(30;  CNC,  CUNY,  UASM);  Santa  Rita  Mountains  (9;  UKSM,  USNM),  Sycamore  Canyon  (12;  CAS,  KHSt,  UATA);  Yavapai 
Co.,’  Bumble  Bee  (3;  CAS),  Congress  (1;  UATA),  Jerome  (4;  CAS),  Mayer  (1;  GRNo);  Yuma  Co.,  Yuma  (10;  UKSM, 
USNM).  ARKANSAS:  Caroll  Co.,  Berry  town  (7;  UKSM),  Eureka  Springs  (7;  UKSM);  Madison  Co.,  Patrick  (1;  UKSM); 
Washington  Co.  (1;  INHS),  7 mi.  s.  Fayetteville  (5;  UASM).  CALIFORNIA  (2;  MCZ,  USNM):  Imperial  Co.,  Potholes  (2; 
CAS);  Riverside  Co.,  Palm  Springs  (1;  USNM);  San  Bernadino  Co.,  10  mi.  ne.  Earp  (3;  UCB).  COLORADO:  Montrose  Co., 
Cimarron  (1;  CUNY).  ILLINOIS:  Pike  Co.,  Pittsfield  (1;  UCD).  MISSOURI:  Crawford  Co.,  Meramec  River  (1;  USNM); 
Ripley  Co.,  5.5  mi.  n.  Briar  (1;  CAS);  Wright  Co.,  Mountain  Grove  (l;TCBa).  NEBRASKA:  Douglas  Co.,  Omaha  (1;  CAS). 
NEW  MEXICO:  Dona  Ana  Co.,  Agustin  Pass  (2;  CAS),  Las  Cruces  (1;  CNC),  White  Sands  (1;  UKSM);  Grant  Co.,  Cliff 
(1;  DRWh),  Silver  City  (1;  CNC);  Hidalgo  Co.,  Lordsburg  (1;  CNC);  Luna  Co.,  Deming  (1;  UCD),  7.5  mi.  nw.  Florida  (7; 
CAS);  Rio  Arriba  Co.,  San  Juan  Pueblo  (1;  DRWh);  Torrance  Co.,  4 mi.  w.  Abo  (4;  GRNo).  OKLAHOMA:  Cimarron  Co., 
Black  Mesa  State  Park  (1;  CAS).  TEXAS  (20;  AMNH,  ANSP,  CAS,  INHS,  MCZ,  USNM):  Blanco  Co.,  Cypress  Mills  (3; 
USNM),  Round  Mountain  (1;  MCZ);  Brewster  Co.  (1;  MCZ),  Black  Gap  Refuge  (1;  CNC),  Boquillas  (62;  AMNH,  CNC, 
UCB),  Green  Valley  (1;  CAS),  Hot  Springs  (21;  AMNH,  CNC),  Lajitas  (2;  CNC),  22  mi.  s.  Marathon  (39;  UCD),  Maverick 
(3;  CNC),  Nine  Point  Draw  (1;  CNC),  Panther  Junction  (1;  CNC),  Rio  Grande  Village  (4;  CAS,  UASM),  Terlingua  (2; 
CNC),  Tomillo  Flat  (3;  CNC);  Jeff  Davis  Co.,  Davis  Mountains  State  Park  (10;  CAS,  DHKa,  DJLa),  Fort  Davis  (7;  CAS, 
CNC);  Llano  Co.,  Enchanted  Rock  (7; CNC);  Presidio  Co.,  4 mi.  w.  Lajitas  (1;  JHeS),  6 mi.  e.  Presidio  (21 ; UASM);  Terrell 
Co.,  10  mi.  e.  Dryden  (1;  DJLa),  Lozier  Canyon  (1;  MCZ);  Val  Verde  Co.,  Del  Rio  (3;  CAS,  USNM).  UTAH:  San  Juan  Co., 
8 mi.  e.  Bluff  (1;  UATA). 


MEXICO 

BAJA  CALIFORNIA:  10  mi.  sw.  Canipole  (3;  UATA),  20  mi.  n.  Comondu  (5;  CAS),  Conception  Bay  (1;  CAS),  8 mi.  nw. 
El  Progreso  (1;  CAS),  La  Paz  (2;  CAS),  25  mi.  w.  La  Paz  (1;  CAS),  Purissima  (2;  UASM),  5 mi.  w.  San  Bartolo  (7;  CAS), 
12  mi.  nw.  San  Bartolo  (3;  CAS),  10  mi.  sw.  San  Jose  del  Cabo  (8;  UATA),  25  mi.  s.  Santa  Rosalia  (1;  CAS),  6 mi.  sw. 
Santiago  (12;  UATA),  Triunfo  (7;  CAS).  CHIHUAHUA:  25  mi.  sw.  Camargo  (2;  AMNH),  Catarinas  (5;  AMNH),  70-75  mi. 
n.  Chihuahua  (2; GRNo,  UCB),  33  mi.  s.  Ciudad  Jimenez  (3;  DJLa),  Parral  (1;  UCB),  15  mi.  e.  Parral  (3;  AMNH),  40  mi.  ne. 
Parral  (3;  UASM),  5 mi.  w.  Parrita  (3;  UCB),  63  mi.  w.  Santa  Barbara  (4;  AMNH),  Valle  de  Olivos  (2;  AMNH).  COAHUI- 
LA:  Boquillas  del  Carmen  (3;  CNC),  15  mi.  n.  Saltillo  (1;  UASM).  COLIMA:  8 mi.  sw.  Colima  (1;  UASM).  DURANGO: 
12.2  mi.  s.  El  Banco  (25;  UASM),  Las  Nieves  (3;  UASM),  4.2  mi.  w.  Vicente  Guerrero  (1;  UASM).  NAYARIT:  Acaponeta 
(1;  CAS),  2.4  mi.  s.  Acaponeta  (2;  UASM).  NUEVO  LEON:  Cienega  de  Flores  (2;  CAS),  14.8  mi.  w.  Linares  (2;  BMNH), 
5 mi.  s.  Monterrey  (2;  CNC),  4.8  mi.  e.  Sabinas  Hidalgo  (30;  CNHM,  DRWh,  IRSB,  MCZ,  MGFT,  UASM,  USNM). 
SINALOA:  30.6  mi.  s.  Culiacan  (1;  UASM),  26  mi.  n.  Pericos  (1;  CAS),  4 mi.  s.  Villa  Union  (1;  UCB).  SONORA  (2; 
BMNH):  Alamos  (8;  CAS),  5 mi.  w.  Alamos  (3;  UATA),  10  mi.  w.  Alamos  (5;  AMNH),  Hermosillo  (40;  CAS,  UATA), 
40  mi.  n.  Hermosillo  (13;  CAS),  10  mi.  e.  Navajoa  (21;  UATA),  San  Bernadino  (1;  CAS),  San  Carlos  Bay  (1;  CAS),  San 
Lorenzo  (1;  CNHM),  40  mi.  nw.  Santa  Ana  (2;  GRNo).  TAMAULIPAS:  Nuevo  Laredo  (1;  USNM).  ZACATECAS:  25  mi. 
w.  Fresnillo  (1;  CAS),  1.3  mi.  se.  Sain  Alto  (1;  UASM). 


Collecting  notes.  — Specimens  of  this  species  have  been  collected  from  April  to  October, 
and,  in  Mexico,  in  January.  Most  were  found  in  riparian  gravel  bars,  or  were  taken  at  lights. 

Taxonomic  notes.  — Within  the  depressus  group,  S.  falli  is  similar  to  S.  pygmaeus,  S. 
scopaeus,  and  S.  depressus , and  has  in  the  past  been  confused  with  them.  But  these  species 
are  sympatric,  and  specimens  are  constantly  distinguishable  by  form  of  frontal  carinae  and 
by  details  of  male  genitalia.  That  S.  falli  is  specifically  distinct  is  further  confirmed  by  the 
fact  that  sympatric  population  samples  of  these  species  are  statistically  significantly  different 
in  various  ways. 


Schizogenius  ochthocephalus  new  species 

Type  material.  — Holotype  male  and  allotype  female  labelled  “Davis  Cal  1.17  1956,” 
“Taken  in  flood  debris,”  and  “E.  A.  Kurtz  Collector”  (UCD).  An  additional  16  specimens 
from  various  localities  in  central  and  northern  California  are  paratypes  (CAS,  DRWh,  UASM, 
UCD). 

Diagnostic  combination.  — Within  the  depressus  group,  this  is  the  only  known  pale  spe- 
cies with  extensive  abdominal  microsculpture. 

Description.  — As  in  S.  falli  except  as  follows.  Left  elytron  with  about  seven  to  eight 
setae  on  interval  three,  six  to  eight  on  interval  five,  four  to  six  on  interval  seven;  total  18-21 
in  specimens  examined.  Abdomen  extensively  microsculptured.  Male  genitalia  with  median 
lobe,  Fig.  231;  four  specimens  examined 


286 


Whitehead 


Measurements  and  proportions.  See  Table  59.  Of  holotype:  TL,  3.64  mm;  LE,  2.25  mm; 
WH,  0.75  mm;  WP,  0.93  mm;  WE,  1.12  mm;  WF/WH,  0.65;  LP/WP,  0.96;  DP/LP,  0.78; 
LP/WE,  0.79;  Ta/Ti,  0.56;  PS/LP,  0.70.  Of  allotype:  TL,  3.88  mm;  LE,  2.40  mm;  WH,  0.80 
mm;  WP.  1.02  mm;  WE.  1.25  mm;  WF/WH,  0.67;  LP/WP,  0.94;  DP/LP,  0.78;  LP/WE,  0.77; 
Ta/Ti,  0.61;  PS/LP,  0.71. 


Table  59.  Descriptive  statistics  for  S.  ochthocephalus,  based  on  1 1 males  from  various 
localities  in  California. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

nts,  in  mm. 

TL 

3.51-3.77 

3.633 

0.122 

0.049 

2.23 

LE 

2.18-2.34 

2.247 

0.074 

0.030 

2.18 

WH 

0.71-0.77 

0.743 

0.026 

0.010 

2.33 

WP 

0.89-0.97 

0.928 

0.035 

0.014 

2.54 

WE 

1.10-1.17 

1.127 

0.034 

0.014 

2.03 

B.  Setae  on  le 

ft  elytron. 

Interval  3 

7-  8 

7.4 

Interval  5 

6-  8 

7.1 

Interval  7 

5-  6 

5.2 

Total 

18-21 

19.6 

1.8 

0.7 

6.14 

C.  Proportion: 

s. 

WF/WH 

0.65-0.69 

0.670 

0.020 

0.008 

2.00 

LP/WP 

0.94-0.97 

0.955 

0.021 

0.009 

1.49 

DP/LP 

0.76-0.81 

0.784 

0.020 

0.008 

1.73 

LP/WE 

0.77-0.80 

0.784 

0.013 

0.005 

1.11 

Ta/Ti 

0.55-0.60 

0.580 

0.034 

0.014 

3.93 

PS/LP 

0.69-0.73 

0.706 

0.019 

0.008 

1.78 

Etymology.  - Greek,  ochthos  = mound,  plus  kephalos  = head,  in  reference  to  lumpy 
appearance  of  head  caused  by  basal  coalescence  of  frontal  sulci. 

Distribution.  — S.  ochthocephalus  is  known  from  only  a few  localities  in  central  and 
northern  California  (Fig.  240).  I studied  the  following  18  specimens. 

UNITED  STATES 

CALIFORNIA:  Calaveras  Co.,  Mokelumne  Hill  (1;  CAS);  Humboldt  Co.,  Fembridge  (1;  UASM);  Lake  Co.,  Clear  Lake  (1; 
CAS);  Sacramento  Co.,  Fair  Oaks  (1;  UCD);  Sonoma  Co.,  Guemeville  (1;  CAS);  Yolo  Co.,  Davis  (13;  DRWh,  UCD). 


Collecting  notes.  - I have  not  collected  specimens  of  this  species,  but  presume  they  live  in 
riparian  gravel  bars.  Available  specimens  were  collected  in  January,  March,  April,  and  June. 

Taxonomic  notes.  — This  species  is  sympatric  with  S.  pygmaeus  and  S.  depressus,  and  dif- 
fers constantly  from  them  by  having  abdominal  micro  sculpture  and  basally  confused  frontal 
carinae.  S.  ochthocephalus  clearly  shares  a common  ancestry  with  S.falli,  and  may  be  only  a 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


287 


geographic  isolate  of  that  species,  but  I think  it  is  reproductively  isolated  for  the  following 
reasons.  There  is  evidently  at  least  one  complete  morphological  discontinuity  between  the 
two  taxa,  as  all  specimens  of  the  former  have  the  abdomen  completely  microsculptured, 
while  none  of  the  latter  do.  Mean  eye  size  in  S.  ochthocephalus  (sample  one)  is  taxonomi- 
cally  significantly  smaller  (Table  57)  than  in  the  geographically  nearest  sample  of  S.  falli 
(sample  two,  Yuma),  and  in  this  characteristic  an  otherwise  continuous  cline  is  interrupted 
(Fig.  249).  These  discontinuities  indicate  that  S.  ochthocephalus  is  more  likely  reproductive- 
ly isolated  from  S.  falli  than  not,  and  justify  my  recognition  of  them  as  distinct  species.  The 
S.  ochthocephalus  sample  is  further  characterized  by  fewer  elytral  setae  than  the  Yuma 
sample  of  S.  falli  (Table  56,  Fig.  248),  but  the  difference  is  not  statistically  significant. 

Since  no  related  species  in  the  depressus  group  have  extensive  abdominal  microsculpture, 
presence  of  such  microsculpture  in  S.  ochthocephalus  must  be  a specialization.  The  four 
characteristics  studied  by  statistical  analysis  exemplify  central-peripheral  patterns  of  vari- 
ation (Brown,  1958;  Ball  1960),  in  which  peripheral  character  states  are  best  regarded  as 
ancestral.  I judge  that  large  body  size  (Fig.  247)  and  small  numbers  of  elytral  setae  (Fig. 
248)  are  peripheral  character  states,  and  hence  probably  ancestral.  The  small  body  size 
character  state  is  evidently  spreading  outward  from  the  Sonoran  Region,  but  has  reached 
neither  S.  ochthocephalus  nor  eastern  populations  of  S.  falli.  Increased  numbers  of  elytral 
setae  may  be  a character  state  now  spreading  outward  from  the  Chihuahuan  Region,  with 
impeded  gene  flow  between  the  Chihuahua  and  Durango  samples,  and  no  gene  flow  between 
S.  falli  and  S.  ochthocephalus . The  evidence  for  central-peripheral  variation  in  relative  eye 
size  (Fig.  249)  is  stronger;  the  small  eye  character  state  is  definitely  peripheral,  and  hence 
most  probably  is  ancestral.  The  enlarged  eye  character  state  apparently  is  spreading  outward 
from  the  Sonoran  Region,  but  does  not  affect  S.  ochthocephalus , and,  curiously,  is  only 
weakly  evident  in  the  Big  Bend  part  of  Texas. 

Even  more  interesting  is  the  pattern  of  variation  in  pronotal  form  (Fig.  250).  The  wide 
I pronotum,  as  in  samples  12  and  13,  is  a peripheral  character  state  which  I think  must  be 
| ancestral.  Further  support  for  this  notion  comes  from  a comparison  of  two  samples  from  the 
Dragoon  Mountains  in  Arizona.  One  sample  from  Texas  Pass,  not  tabulated  or  figured,  was 
collected  in  1917;  the  second  sample,  collected  in  1965,  is  from  Cochise  Stronghold  (sample 
five).  Data  for  the  Texas  Pass  sample  are  as  follows:  N,  12;  Range,  0.89-0.98;  Mean,  0.936; 
1.5SD,  0.030;  2SE,  0.01 1;  CV(%),  2.1 1.  If  the  statistical  data  for  these  two  samples  proper- 
ly reflect  their  respective  populations,  there  must  have  been  a shift  in  time  from  broad  to 
narrow  pronotal  form.  The  difference  is  statistically  significant,  and  I therefore  interpret  it 
as  biologically  significant;  there  has,  in  the  region  of  the  Dragoon  Mountains,  been  a shift 
from  the  ancestral  to  derived  character  state  within  a period  of  less  than  50  years.  I suggest 
that  in  S.  falli  the  narrow  pronotum  character  state  is  spreading  from  two  foci,  in  the  Sono- 
ran Region  and  along  the  Rio  Grande.  If  this  assertion  is  correct,  then  the  narrow  pronotum 
in  S.  ochthocephalus  must  be  an  independently  acquired  specialization.  However,  my  inter- 
i pretation  may  be  overly  simplified,  in  view  of  the  complex  pattern  of  variation  in  pronotal 
form.  A test  of  my  hypothesis  of  central-peripheral  variation  may  be  made  by  searching 
for  other  shifts  in  time,  from  narrow  to  broad  or  broad  to  narrow  pronotal  form. 

Schizogenius  depressus  LeConte 

| Schizogenius  depressus  LeConte  1852:197.  Type  locality  Colorado  River,  California;  type 
in  MCZ,male  labelled  MCZ  5843  here  designated  lectotype  (!).  LeConte  1857:83.  Putzeys 
1863:24.  Putzeys  1866:223.  LeConte  1879:34.  Hatch  1949:118.  Hatch  1953:69.  Lind- 
roth  1961:168. 


288 


Whitehead 


Diagnostic  combination.  — Among  pale  members  of  the  depressus  group  without  ven- 
tral microsculpture,  specimens  of  S.  depressus  are  distinguished  by  prominent  pronotal 
hind  angles,  frontal  sulci  not  or  weakly  confused  basally,  and  elytral  length  over  2.0 
mm. 

Description.  — As  in  S.  pygmaeus  except  as  follows.  Larger,  elytral  length  2.05  mm  or 
more.  Body  color  testaceous  to  ferrugineous  or  castaneous,  not  piceous,  bicolored,  or 
aeneous.  Head  and  pronotum,  Fig.  221;  eyes  uniformly  faceted  except  in  some  Mexican 
specimens,  where  inner  facets  enlarged;  neck  densely  to  sparsely  punctate;  antennal  article 
five  about  1.4  to  1.6  times  longer  than  wide;  pronotal  hind  angles  prominent  in  most  speci- 
mens; anterior  transverse  impression  finely  punctate.  Left  elytron  with  six  to  twelve  setae 
on  interval  three,  six  to  1 1 on  interval  five,  four  to  seven  on  interval  seven;  total  16-30  in 
specimens  examined;  humeral  denticles  moderately  sharp  and  prominent.  Male  genitalia 
with  median  lobe,  Fig.  232;  endophallus,  Fig.  233;  20  specimens  examined. 

Measurements  and  proportions.  See  Table  60. 

Variation.  — Specimens  from  the  Rio  Balsas  drainage  in  the  state  of  Mexico  have  much 
larger  inner  than  outer  eye  facets.  Specimens  from  Zacatecas  also  have  large  inner  eye 
facets,  though  less  pronouncedly  so.  Eyes  in  all  other  specimens  are  uniformly  faceted,  in- 
cluding those  from  the  Rio  Tula  drainage  in  the  state  of  Mexico.  The  Zacatecas  locality  is, 
in  a sense,  geographically  intermediate  between  the  two  state  of  Mexico  localities,  as  it  is 
from  an  interior  drainage  system,  while  the  Rio  Balsas  drains  to  the  Pacific  and  the  Rio  Tula 
to  the  Atlantic. 


Table  60.  Descriptive  statistics  for  S.  depressus , based  on  20  males  from  Clear  Lake, 
California. 


Character 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

A.  Measureme 

nts,  in  mm. 

TL 

3.39-3.92 

3.706 

0.194 

0.058 

3.49 

LE 

2.06-2.40 

2.263 

0.123 

0.037 

3.61 

WH 

0.70-0.81 

0.764 

0.039 

0.012 

3.41 

WP 

0.91-1.05 

0.990 

0.052 

0.016 

3.50 

WE 

1.06-1.24 

1.165 

0.069 

0.020 

3.92 

B.  Setae  on  lei 

ft  elytron. 

Interval  3 

6-  9 

7.0 

Interval  5 

6-  8 

7.2 

Interval  7 

4-  6 

4.6 

Total 

16-22 

18.8 

2.0 

0.6 

7.15 

C.  Proportion; 

WF/WH 

0.63-0.68 

0.650 

0.020 

0.006 

2.08 

LP/WP 

0.90-0.96 

0.930 

0.024 

0.007 

1.69 

DP/LP 

0.76-0.81 

0.788 

0.023 

0.007 

1.98 

LP/WE 

0.77-0.83 

0.790 

0.024 

0.007 

1.99 

Ta/Ti 

0.59-0.68 

0.634 

0.041 

0.012 

4.31 

PS/LP 

0.70-0.78 

0.735 

0.032 

0.009 

2.88 

Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


289 


Statistical  data  on  variation  in  body  size,  numbers  of  elytral  setae,  relative  eye  size,  and 
pronotal  form  are  given  in  Tables  61-64,  and  summarized  in  Fig.  251-254.  The  main  fea- 
tures of  variation  in  body  size  (Fig.  251)  are  the  following.  Western  samples  are  varied,  but 
all  show  clear  evidence  of  clinal  relationships  with  geographically  related  samples.  Eastern 
samples  decrease  in  mean  body  size  south  to  the  Colorado  River,  and  increase  from  there 
south  into  Mexico.  Northern  samples  are  not  statistically  significantly  different  from  one 
another,  but  seem  not  to  be  clinally  continuous  across  the  Rocky  Mountains. 

Variation  in  numbers  of  elytral  setae  (Fig.  252)  may  be  summarized  as  follows.  In  the 
United  States,  samples  from  the  Rio  Papigochic,  Colorado  River,  Great  Basin,  Rio  Grande, 
Canadian  River,  and  Missouri  River  drainage  basins  have  statistically  significantly  more  ely- 
tral setae  than  any  of  the  western  samples.  All  eastern  and  all  western  samples  are  evidently 
clinally  related  to  one  another.  Northern  samples  do  not  appear  to  be  clinally  continuous 
across  the  Rocky  Mountains.  The  Chihuahua  sample  (number  22)  is  probably  related  to  the 
Arizona  sample  (number  21)  via  a steep  cline.  Additional  collections  are  needed  to  determine 
relationships  between  the  quite  different  Chihuahua  and  Zacatecas  samples  (22  and  23). 


Table  61.  Variation  in  body  size  (LE,  in  mm)  in  selected  samples  of  S.  depressus  (Fig.  251). 
Males  and  females  each  50%. 


Sample 

N 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

1 

4 

2.22-2.50 

2.378 

2 

14 

2.32-2.92 

2.563 

0.221 

0.079 

5.74 

3 

20 

2.16-2.71 

2.442 

0.226 

0.067 

6.17 

4 

12 

2.22-2.58 

2.432 

0.184 

0.071 

5.09 

5 

30 

2.25-2.78 

2.560 

0.183 

0.044 

4.76 

6 

16 

2.32-2.66 

2.487 

0.165 

0.055 

4.43 

7 

4 

2.39-2.58 

2.482 

8 

40 

2.05-2.60 

2.321 

0.154 

0.035 

4.71 

9 

16 

2.31-2.69 

2.501 

0.152 

0.050 

4.04 

10 

16 

2.12-2.56 

2.325 

0.203 

0.068 

5.82 

11 

12 

2.29-2.69 

2.514 

0.181 

0.070 

4.80 

12 

14 

2.26-2.72 

2.461 

0.166 

0.059 

4.50 

13 

14 

2.35-2.75 

2.520 

0.213 

0.076 

5.62 

14 

20 

2.28-2.72 

2.502 

0.169 

0.050 

4.50 

15 

6 

2.32-2.70 

2.558 

16 

18 

2.30-2.70 

2.529 

0.167 

0.053 

4.41 

17 

16 

2.15-2.78 

2.530 

0.241 

0.080 

6.34 

18 

26 

2.35-2.69 

2.494 

0.143 

0.037 

3.83 

19 

24 

2.25-2.75 

2.508 

0.200 

0.054 

5.31 

20 

48 

2.19-2.72 

2.449 

0.187 

0.036 

5.10 

21 

40 

2.25-2.78 

2.520 

0.164 

0.035 

4.36 

22 

4 

2.65-2.85 

2.775 

23 

24 

2.50-2.92 

2.676 

0.161 

0.044 

4.01 

290 


Whitehead 


Table  62.  Variation  in  numbers  of  elytral  setae  in  selected  samples  of  S.  depressus  (Fig. 
252).  Males  and  females  each  50%. 


Sample 

N 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

1 

4 

18-21 

19.5 

2 

14 

18-22 

20.4 

1.7 

0.6 

5.65 

3 

20 

17-22 

19.8 

2.3 

0.7 

7.61 

4 

12 

18-21 

19.5 

1.3 

0.5 

4.09 

5 

30 

18-23 

20.4 

1.9 

0.5 

6.12 

6 

16 

18-23 

20.1 

1.9 

0.6 

6.43 

7 

4 

17-21 

18.8 

8 

40 

16-22 

19.2 

1.8 

0.4 

6.15 

9 

16 

17-21 

19.2 

1.6 

0.5 

5.53 

10 

16 

18-23 

19.9 

1.9 

0.6 

6.33 

11 

12 

18-22 

19.8 

1.7 

0.7 

6.02 

12 

14 

18-22 

19.8 

1.6 

0.6 

5.31 

13 

14 

19-22 

20.3 

1.8 

0.6 

5.94 

14 

20 

17-22 

19.8 

2.0 

0.6 

6.68 

15 

6 

20-23 

21.8 

16 

18 

21-25 

23.0 

2.3 

0.7 

6.67 

17 

16 

20-25 

22.1 

2.1 

0.7 

6.37 

18 

26 

20-24 

22.0 

1.7 

0.4 

5.05 

19 

24 

20-28 

23.0 

3.2 

0.9 

9.16 

20 

48 

20-30 

22  2 

2.9 

0.6 

8.66 

21 

40 

19-25 

22.6 

2.0 

0.5 

6.93 

22 

4 

24-27 

25.5 

23 

24 

19-23 

20.5 

2.0 

0.5 

6.44 

Variation  in  relative  eye  size  (Fig.  253)  has  the  following  main  features.  In  both  east 
and  west,  there  is  a general  trend  toward  increased  eye  size  from  north  to  south,  with 
particularly  large  eyes  characteristic  of  Mexican  samples.  In  the  United  States,  eye  size 
in  eastern  samples  is  quite  uniform  but  in  the  west  a discordant  pattern  of  variation  is 
evident  in  the  north.  Northern  samples  do  not  appear  to  be  clinally  continuous  across  the 
Rocky  Mountains. 

The  chief  features  of  variation  in  pronotal  form  (Fig.  254)  are  as  follows.  The  pattern 
of  variation  in  western  samples  is  complex,  and  no  definite  trends  are  evident,  but  geo- 
graphically related  samples  are  probably  clinally  continuous.  Among  eastern  samples  there 
is  a definite  cline  toward  narrower  pronotal  form  from  north  to  south.  Across  the  Rocky 
Mountains  in  the  north,  there  is  strong  evidence  of  clinal  discontinuity  in  this  character- 
istic. 

Among  Mexican  specimens,  only  those  from  Zacatecas  were  numerous  enough  to  form 
a good  sample  for  statistical  analysis;  thus,  no  detailed  account  of  variation  is  yet  possible. 
The  Chihuahua  specimens  are  evidently  related  by  steep  character  dines  to  specimens 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


291 


Table  63.  Variation  in  relative  eye  size  (WF/WH)  in  selected  samples  of  S.  depressus  (Fig. 
253).  Males  and  females  each  50%. 


Sample 

N 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

1 

4 

0.66-0.69 

0.675 

2 

14 

0.63-0.68 

0.658 

0.021 

0.008 

2.13 

3 

20 

0.63-0.69 

0.670 

0.023 

0.007 

2.24 

4 

12 

0.66-0.69 

0.676 

0.016 

0.006 

1.47 

5 

30 

0.63-0.67 

0.654 

0.017 

0.004 

1.73 

6 

16 

0.66-0.70 

0.674 

0.015 

0.005 

1.63 

7 

4 

0.65-0.68 

0.670 

8 

24 

0.62-0.68 

0.655 

0.020 

0.005 

2.01 

9 

16 

0.63-0.68 

0.659 

0.017 

0.006 

1.70 

10 

16 

0.65-0.69 

0.666 

0.016 

0.005 

1.63 

11 

12 

0.63-0.67 

0.650 

0.022 

0.009 

2.27 

12 

14 

0.63-0.67 

0.652 

0.020 

0.007 

2.01 

13 

14 

0.65-0.67 

0.663 

0.012 

0.004 

1.25 

14 

20 

0.63-0.68 

0.657 

0.018 

0.005 

1.79 

15 

6 

0.64-0.67 

0.657 

16 

18 

0.64-0.68 

0.661 

0.016 

0.005 

1.63 

17 

16 

0.64-0.69 

0.659 

0.024 

0.008 

2.44 

18 

26 

0.62-0.70 

0.654 

0.026 

0.007 

2.67 

19 

24 

0.63-0.68 

0.664 

0.022 

0.006 

2.26 

20 

48 

0.62-0.68 

0.653 

0.025 

0.005 

2.51 

21 

24 

0.63-0.68 

0.655 

0.022 

0.006 

2.70 

22 

4 

0.62-0.67 

0.642 

23 

24 

0.62-0.66 

0.640 

0.018 

0.005 

1.93 

from  the  Chiricuahua  Mountains  of  Arizona.  Specimens  from  Zacatecas  differ  markedly 
from  those  from  Chihuahua,  particularly  in  numbers  of  elytral  setae,  but  the  differences 
probably  reflect  mainly  a lack  of  intermediate  collections.  Specimens  from  Atlantic  and 
Pacific  drainages  in  the  state  of  Mexico  are  strongly  differentiated.  Rio  Tula  and  Guana- 
juato specimens  differ  from  Rio  Balsas  specimens  by  narrower  pronota,  and  by  uniformly 
faceted  and  much  smaller  eyes.  Zacatecas  specimens  are  intermediate  in  these  character- 
istics, and  may  also  be  considered  geographically  intermediate  since  they  are  from  an 
interior  drainage  system.  I suspect  that  the  Rio  Balsas  and  Rio  Tula  populations  are  geo- 
graphically quite  isolated  from  one  another  despite  their  proximity,  and  that  they  may 
even  be  reproductively  isolated. 

In  America  north  of  Mexico,  sample  means  for  numbers  of  elytral  setae  in  eastern  samples 
j of  S.  depressus  are  statistically  significantly  greater  than  in  western  samples.  Eastern  samples 
I are  similar  in  all  ways  measured,  and  are  linked  to  one  another  by  definite  character  dines. 
Western  samples  are  more  varied,  but  are  similarly  related  by  character  dines.  There  is  no 
evidence  of  reproductive  isolation  between  eastern  and  western  forms  in  the  south,  but 


292 


Whitehead 


Table  64.  Variation  in  pronotal  form  (LP/WP)  in  selected  samples  of  S.  depressus  (Fig.  254). 
Males  and  females  each  50%. 


Sample 

N 

Range 

Mean 

1.5SD 

2SE 

CV(%) 

1 

4 

0.91-0.94 

0.925 

2 

14 

0.91-0.96 

0.939 

0.023 

0.008 

1.64 

3 

20 

0.90-0.95 

0.932 

0.019 

0.006 

1.76 

4 

12 

0.91-0.96 

0.932 

0.026 

0.010 

1.84 

5 

30 

0.89-0.95 

0.928 

0.021 

0.005 

1.53 

6 

16 

0.90-0.96 

0.927 

0.025 

0.008 

1.79 

7 

4 

0.93-0.97 

0.948 

8 

24 

0.89-0.98 

0.933 

0.032 

0.009 

2.29 

9 

16 

0.89-0.96 

0.928 

0.029 

0.010 

2.10 

10 

16 

0.89-0.96 

0.929 

0.032 

0.011 

2.27 

11 

12 

0.91-0.98 

0.938 

0.034 

0.013 

2.40 

12 

14 

0.92-0.96 

0.941 

0.017 

0.006 

1.21 

13 

14 

0.87-0.96 

0.936 

0.038 

0.013 

2.67 

14 

20 

0.91-0.95 

0.932 

0.020 

0.006 

1.46 

15 

6 

0.88-0.94 

0.915 

16 

18 

0.89-0.95 

0.923 

0.025 

0.008 

1.82 

17 

16 

0.89-0.95 

0.923 

0.022 

0.007 

1.62 

18 

26 

0.89-0.95 

0.923 

0.024 

0.006 

1.76 

19 

24 

0.88-0.96 

0.930 

0.028 

0.008 

2.04 

20 

48 

0.89-0.98 

0.928 

0.024 

0.005 

1.75 

21 

24 

0.89-0.98 

0.939 

0.028 

0.008 

2.06 

22 

4 

0.90-0.97 

0.942 

23 

24 

0.88-0.96 

0.928 

0.031 

0.008 

2.23 

differences  in  mean  numbers  of  elytral  setae  do  indicate  geographic  isolation.  In  the  north, 
the  Montana  sample  (sample  15)  is  similar  in  most  ways  to  some  of  the  more  distant 
western  samples,  especially  samples  two  and  five,  but  is  divergent  from  them  in  pronotal 
form.  Western  samples  geographically  closer  to  the  Montana  sample,  particularly  samples 
three  and  four,  are  more  differentiated  in  most  or  all  characteristics  measured.  In  the 
Pacific  Northwest,  these  characteristics  vary  in  definite  geographic  patterns,  apparently 
independently  of  one  another  but  not  randomly.  Most  notably,  small  eyed  forms  extend 
southwest  from  the  Snake  River  to  northern  California,  thus  separating  large  eyed  forms 
to  the  north  and  south  (Fig.  253).  These  facts  suggest  that  the  northern  populations  were 
once  clinally  continuous  across  the  Rocky  Mountains,  but  later  became  geographically 
and  perhaps  even  reproductively  isolated.  The  eastern  form  may  relatively  recently  have 
crossed  the  mountain  barrier  to  interact  with  the  western  form,  resulting  in  character 
displacement  (Brown  and  Wilson,  1956).  This  hypothesis  of  character  displacement  may 
be  tested  by  searching  for  bimodal  character  state  distributions  in  large  samples  collected 
along  the  western  slopes  of  the  Rocky  Mountains.  Insufficient  material  is  available  at 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


293 


present  for  such  an  analysis. 

Although  I have  no  direct  evidence  of  character  displacement  in  the  northwest,  I cannot 
otherwise  explain  how  such  a complex  pattern  of  variation  and  differentiation  may  have 
originated.  I assume  that  my  hypotheses  are  correct,  and  postulate  the  following  sequence 
of  events.  S.  depressus  was  once  widespread  in  western  North  America,  with  clinal  continui- 
ty in  the  south  via  the  Colorado  River,  in  the  north  via  the  Columbia  and  Missouri  Rivers, 
and  perhaps  also  across  the  Great  Basin.  With  deteriorating  climatic  conditions  in  the 
Pleistocene,  and  particularly  with  advances  of  glaciation  southward  along  the  Rocky  Moun- 
tains, clinal  continuity  in  the  north  was  repeatedly  broken  by  geographic  isolation.  During 
subsequent  interglaciation,  geographic  isolation  may  have  been  partially  maintained.  Much 
later,  the  retreat  of  the  Wisconsin  glaciation  was  accompanied  by  a rapid  spread  of  desert 
conditions  in  the  south  (Martin  and  Mehringer,  1965),  and  resulted  in  a break  in  clinal  con- 
tinuity there.  As  climates  in  the  north  moderated,  renewed  contact  between  eastern  and 
western  forms  was  established,  but  too  late  for  renewed  clinal  continuity.  Possible  datings 
for  Late  Pleistocene  events  (Martin  and  Mehringer,  1965)  are:  glacial  advance,  70,000  years 
ago;  glacial  retreat,  12,000  years  ago;  and  reestablishment  of  contact  between  eastern  and 
western  forms  in  the  north,  less  than  12,000  years  ago. 

Etymology.  — Latin,  depressus  = flattened,  in  reference  to  the  flattened  body  form. 

Distribution.  — S.  depressus  ranges  further  north,  and  reaches  higher  elevations,  than  any 
other  species  in  the  genus  (Fig.  241).  It  ranges  in  the  east  along  upper  reaches  of  the  Rocky 
Mountain  drainage  systems  as  far  north  as  Montana,  and  in  the  west  it  occurs  in  coastal  and 
montane  regions  north  to  southern  British  Columbia.  To  the  south,  S.  depressus  extends  to 
both  Atlantic  and  Pacific  drainage  basins  in  the  state  of  Mexico,  where  specimens  have  been 
found  as  high  as  8000'  above  sea  level.  One  specimen  labelled  “D.  C.”  belongs  to  this  species 
and  is  no  doubt  incorrectly  labelled.  I examined  1230  additional  specimens  of  this  species 
from  the  following  localities. 


CANADA 

BRITISH  COLUMBIA:  Salmon  Arm  (2;  CAS,  MCZ),  Vaseaux  Lake  (1;  UASM),  Vernon  (6;  CAS,  KSUM);  see  Lindroth 
(1961)  for  additional  Canadian  records. 


UNITED  STATES 

No  locality  (4;  IRSB,  MGFT,  USNM).  ARIZONA  (29;  AMNH,  ANSP,  CAS,  CUNY,  INHS,  USNM):  Cochise  Co.,  Bear 
Canyon  (1;  CUNY),  Bisbee  (1;  CAS),  Cave  Creek  (22;  AMNH,  CAS,  TCBa,  UASM),  Chiricahua  Mountains  (12;  CAS, 
UKSM,  USNM),  Cochise  Stronghold  (2;  TLEr,  UATA),  Huachuca  Mountains  (1;  KHSt),  Montezuma  Pass  (1;  CNC), 
Palmerlee  (1;  USNM),  Pinery  Creek  (2;  CNHM),  5 mi.  w.  Portal  (32;  AMNH,  CNC,  CUNY,  LBSC,  MSUL,  TLEr,  UCB, 
UCD),  Ramsey  Canyon  (1;  UATA),  Rucker  Lake  (1;  UASM),  Sierra  Vista  (1;  CNC),  South  Fork  Forest  Camp  (1;  UATA), 
Texas  Pass  (1;  CAS),  Turkey  Creek  (1;  UATA);  Coconino  Co.  (1;  CNHM),  Flagstaff  (6;  CAS,  UKSM),  Oak  Creek  Canyon 
(1;  UASM);  Gila  Co.,  Globe  (4;  CAS,  CNC,  KSUM),  6 mi.  n.  Payson  (1;  DJLa),  Rice  (1;  UASM),  Roosevelt  Lake  (1; 
UASM),  San  Carlos  Lake  (1;  CUNY);  Graham  Co.,  Arcadia  Forest  Camp  (1;  UATA),  Camp  Geronimo  (1;  UATA),  Noon 
Creek  (2;  UATA),  Power’s  Garden  (1;  UASM);  Greenlee  Co.,  Clifton  (1;  CAS),  Diamond  Creek  (3;  UASM);  Pima  Co., 
Rincon  Mountains  (4;  UASM),  Sahuarita  (3;  MCZ);  Santa  Cruz  Co.,  Madera  Canyon  (6;  DHKa,  UATA,  UCD),  Nogales  (2; 
CAS,  MGFT),  Pena  Blanca  (1;  UCB),  Santa  Rita  Mountains  (3;  UKSM),  Sycamore  Canyon  (3;  CAS,  KHSt);  Yavapai  Co., 
Crown  King  (3;  CAS),  Prescott  (12;  CAS,  MCZ,  UASM).  CALIFORNIA  (29;  AMNH,  ANSP,  CNHM,  INHS,  KSUM,  MCZ, 
RUNB,  UKSM,  USNM):  Alameda  Co.,  Arroyo  Mocho  (7;  TLEr,  UCB),  Berkeley  (2;  CUNY),  Livermore  (1;  CAS),  Sunol 
(1;  UCB);  Alpine  Co.,  Markleville  (1;  CAS);  Calaveras  Co.,  Mokelumne  Hill  (3;  CAS);  Colusa  Co.,  Cooks  Springs  (2;  CAS); 
Contra  Costa  Co.,  March  Springs  Creek  (3;  CAS,  MCZ),  Moraga  (1;  CAS),  Mount  Diablo  (2;  CAS),  San  Pablo  Valley  (1; 
UCB);  Del  Norte  Co.  (2;  CNHM);  El  Dorado  Co.  (11;  CNHM),  Pollock  Pines  (1;  UCD);  Fresno  Co.,  Kings  River  (1;  CAS), 
Stevenson  Creek  (3;  CAS),  Trimmer  (6;  CAS);  Humboldt  Co.,  6 mi.  e.  Bridgeville  (l;GRNo),  Fernbridge  (1;  UASM), 
Frenchman  Creek  (2;  CAS);  Inyo  Co.,  Panamint  Mountains  (1;  USNM);  Kern  Co.  (4;  USNM);  Lake  Co.,  Adams  Springs  (1 ; 
CAS),  Clear  Lake  (90;  CAS),  Lower  Lake  (2;  CAS),  North  Fork  Cache  Creek  (33;  UCD),  Middletown  (1;  CAS);  Los 
Angeles  Co.,  Arroyo  del  Valle  (1;  CAS),  Crystal  Lake  (9;  TCBa),  Fish  Canyon  (1;  GRNo),  Pasadena  (4;  CAS),  San  Antonio 
Canyon  (1;  GRNo),  Santa  Monica  (3;  CAS,  INHS),  Tanbark  Flat  (1;  UCD),  Tapia  Park  (1;  LACM),  Tujunga  Canyon  (7; 
LACM);  Marin  Co.  (1;  KSUM),  Camp  Taylor  (2;  CAS,  CNHM),  Fairfax  (5;  CAS),  Lake  Lagunitas  (1;  CAS),  San  Anselmo 
(1;  CAS);  Mariposa  Co.  (1;  CNHM),  Miami  Ranger  Station  (1;  CAS),  Yosemite  Valley  (1;  INHS);  Mendocino  Co.  (2; 


294 


Whitehead 


CNHM,  MCZ),  Bloody  Run  Creek  (1 ; CAS),  8 mi.  w.  Navarro  (3;  CAS),  2 mi.  nw.  Philo  (1 ; CAS);  Modoc  Co.  (9;  CNHM, 
MGFT),  Davis  Creek  (2;  CAS),  4.5  mi.  n.  Fort  Bidwell  (3;  CAS),  3 mi.  s.  Lake  City  (1;  CAS);  Mono  Co.,  Fales  Hot  Springs 
(1;  CAS);  Napa  Co.  (1;  CNHM),  Mount  Saint  Helena  (1;  CAS),  Rutherford  (5;  TLEr);  Nevada  Co.  (4;  CNHM,  MGFT), 
Truckee  (9;  CAS,  MCZ,  USNM);  Orange  Co.,  El  Toro  (1;  CAS),  Silverado  Canyon  (l;GRNo);  Placer  Co.  (1;  USNM),  Bear 
Valley  (2;  CAS),  Lake  Tahoe  (20;  CAS,  MCZ,  UASM,  UCD),  Tahoe  City  (3;  CAS);  Riverside  Co.,  Banning  (1;  CAS), 
Idyllwild  (2;  CAS,  UCB),  Pinon  Flat  (1;  UCB),  15  mi.  e.  Redlands  (1;  LBSC),  Riverside  (2;  CAS),  San  Jacinto  Mountains 
(1;  CAS);  San  Benito  Co.,  Pinnacles  National  Monument  (1;  CAS);  San  Bernadino  Co.,  5.5  mi.  sw.  Big  Pine  Flat  (1 ; GRNo), 
Cedar  Springs  (11;  GRNo,  LBSC),  9 mi.  nw.  Fawnskin  (1;  TLEr),  San  Bernadino  Mountains  (1;  CAS),  South  Fork  Santa 
Ana  River  (1;  CAS);  San  Diego  Co.,  Poway  (2;  CAS,  GRNo),  Warners  Springs  (2;  CAS);  San  Luis  Obispo  Co.,  La  Panza 
(2;  CAS),  San  Luis  Obispo  (2;  CAS);  San  Mateo  Co.  (2;  CAS,  MCZ);  Santa  Barbara  Co.  (1;  CAS),  Santa  Cruz  Island  (3; 
CAS,  UCD),  Santa  Inez  Mountains  ( 1 ; CAS) ; Santa  Clara  Co.,  Alum  Rock  Park  ( 1 ; TLEr),  Arroyo  Bayo  ( 1 ; TLEr),  Gilroy 
Hot  Springs  (11;  CAS,  TLEr),  Los  Gatos  (2;  CAS),  San  Martin  (23;  CAS),  Stanford  (1;  CAS);  Siskiyou  Co.  (2;  CAS, 
USNM),  2.3  mi.  nw.  Callahan  (1;  CAS),  Dunsmuir  (1;  USNM),  1.3  mi.  e.  Grenada  (1;  CAS),  McCloud  (1 ; CAS);  Sonoma 
Co.  (4;  CAS,  CUNY,  LACM),  Cazadero  (1;  CAS),  GuerneviUe  (1;  CAS),  Healdsburg  (1;  CAS),  Santa  Rosa  (1;  MSUL); 
Stanislaus  Co.,  Del  Puerto  Creek  (1;  TLEr);  Trinity  Co.,  10  mi.  sw.  Big  Bar  (3;  CAS),  Hyampon  (1;  CAS),  4 mi.  se.  Ruth 
(2;  CAS);  Tulare  Co.,  Gray  Meadow  (1;  CAS),  Kaweah  (1;  CAS),  Sequoia  National  Park  (2;  CAS,  UCD);  Tuolumne  Co., 
Hardin  Flat  (1;  CAS).  COLORADO  (6;  KSUM,  USNM):  Baca  Co.,  Regnier  (1;  AMNH);  Boulder  Co.,  6.9  mi.  n.  Golden 
(14;  TLEr),  Lyons  (2;  UASM),  South  Boulder  Creek  (3;  DHKa),  2 mi.  e.  Wonderview  (1;  DHKa);  El  Paso  Co.,  Colorado 
Springs  (1 ; USNM);  Fremont  Co.,  Canon  City  (7;  ANSP,  CAS,  MCZ,  USNM);  Jefferson  Co.,  Coal  Creek  Canyon  (1;  CAS), 
Golden  (3;  DHKa),  2 mi.  n.  Golden  (3;  TLEr),  2 mi.  e.  Morrison  (10;  DHKa),  Waterton  (5;  DHKa);  Huerfano  Co.,  La  Veta 
(1;  MCZ);  La  Plata  Co.,  Durango  (1;  MCZ);  Larimer  Co.,  Fort  Collins  (2;  CAS).  IDAHO:  Idaho  Co.,  39  mi.  e.  Lowell  (1; 
HGou);  Kootenai  Co.,  Hayden’s  Lake  (1;  CAS);  Nez  Perce  Co.,  Juliaetta  (2;  CUNY,  USNM),  6 mi.  w.  Lenore  (12;  DHKa, 
HGou),  Waha  Lake  (19;  AMNH,  CAS,  KSUM,  MCZ,  USNM).  MONTANA:  Blaine  Co.,  Bear  Paw  Mountains  (2;  USNM); 
Cascade  Co.,  Armington  (3;  DJLa,  UASM);  Lewis  and  Clark  Co.,  Helena  (1;  UASM),  4 mi.  s.  Wolf  Creek  (1;  UASM). 
NEW  MEXICO  (3;  UKSM):  Bernalillo  Co.,  Albuquerque  (1;  USNM);  Catron  Co.,  3 mi.  s.  Beaverhead  (28;  TLEr),  12.3  mi. 
n.  Glenwood  (4;  UASM);  Grant  Co.,  4 mi.  e.  Hillsboro  (1;  CNHM),  23  mi.  n.  Mimbres  (5;  TLEr),  Silver  City  (1;  MCZ),  14 
mi.  n.  Silver  City  (1;MSUL),  26  mi.  n.  Silver  City  (16;  TLEr),  36.4  mi.  ne.  Silver  City  (3;  UASM),  71.6  mi.  ne.  Silver  City 
(1;  UASM);  Rio  Arriba  Co.,  San  Juan  Pueblo  (1;  DRWh);  Sandoval  Co.,  Jemez  Mountains  (5;  CAS),  Jemez  Springs  (2; 
TCBa),  6 mi.  sw.  Jemez  Springs  (1 ; GRNo);  San  Miguel  Co.,  Las  Vegas  (2;  INHS),  Porvenir  (1;  CAS),  Sapello  (15;  UASM); 
Taos  Co.,  Taos  (2;  CAS).  OREGON:  Columbia  Co.,  Portland  (1;  CUNY);  Gilliam  Co.,  Rock  Creek  (1;  CAS);  Grant  Co., 
Blue  Mountains  (2;  CAS);  John  Day  Gorge  (2;  CNHM);  Jackson  Co.,  Medford  (14;  CAS,  UCD),  Talent  (48;  UCD),  Tolo 
(1;  UCD);  Lake  Co.,  Crooked  Creek  (5;  MSUL),  Lakeview  (1;  CNHM),  Paisley  (2;CNHM);  Malhuer  Co.,  Sucker  Creek  (5; 
CAS,  CNHM);  Marion  Co.,  Detroit  (1;  CAS);  Yamhill  Co.,  Dayton  (1;  MCZ).  SOUTH  DAKOTA:  Lawrence  Co.,  Spearfish 
(1;  DRWh).  TEXAS:  Randall  Co.,  Canyon  (5;  MSUL).  UTAH  (2;  JNeg):  Beaver  Co.,  Beaver  Creek  Hills  (2;  USNM),  South 
Creek  (1;  USNM);  Juab  Co.,  Nephi  (1;  USNM);  San  Juan  Co.,  Arch  Canyon  (2;  MCZ);  Utah  Co.,  American  Fork  (30; 
ANSP,  CAS,  USNM);  Provo  (30;  CAS,  MCZ,  USNM);  Weber  Co.,  Ogden  (23;  CNHM,  MCZ,  USNM);  Chad’s  Ranch  (2; 
MCZ,  USNM).  WASHINGTON:  Seattle  (4;  KSUM);  Franklin  Co.,  Kahlotus  (1;  USNM);  Walla  Walla  Co.,  College  Place 
(1;  CNC),  Walla  Walla  (21;  CAS).  WYOMING:  Laramie  Co.,  11  mi.  n.  Cheyenne  (22;  UASM). 

MEXICO 

CHIHUAHUA:  Catarinas  (2;  AMNH),  23.0  mi.  s.  Minaca  (1;  UASM),  Santa  Barbara  (3;  AMNH),  63  mi.  w.  Santa  Barbara 
(1;  AMNH).  GUANAJUATO  (1;  BMNH).  MEXICO:  Temascaltepec  (3;  MCZ),  Villa  Carbon  (3;  JHeS).  ZACATECAS: 
General  Enrique  Estrada  (69;  UASM,  IRSB,  BMNH). 


Collecting  notes.  — I have  collected  specimens  of  this  species  at  several  localities,  all  at 
comparatively  high  altitudes  or  latitudes,  in  gravel  bars  along  unshaded  streams.  The  only 
locality  where  I collected  a good  series  was  near  Laramie,  Wyoming,  where  I found  speci- 
mens of  no  other  species  of  the  genus.  At  lower  elevations  elsewhere,  however,  specimens 
have  been  taken  together  with  specimens  of  one  or  more  other  species.  Specimens  of  S. 
depressus  have  been  taken  throughout  the  year  in  the  more  southern  parts  of  the  United 
States. 

Taxonomic  notes.  — S.  depressus  is  closely  related  to  S.  pygmaeus,  S.  scopaeus,  S.  ochtho- 
cephalus,  and  S.  falli,  but  is  sympatric  with  all  of  them  and  clearly  is  reproductively  isolated. 
As  noted  by  Lindroth(1961),  Hatch  (1949,  1953)  erred  in  placing  S.  litigiosus  as  a synonym 
of  S.  depressus. 

Eastern  and  western  populations  of  S.  depressus,  in  both  the  northern  and  southern  ex- 
tremes of  its  range,  may  be  reproductively  isolated.  Here  is  a fertile  field  for  future  investi- 
gations at  the  population  level. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


295 


Fig.  215-221.  Head  and  pronotum,  dorsal  aspect.  215.  S.  arimao  Darlington,  Soledad,  Cuba.  216.  S.  emdeni  new  spe- 
| cies,  San  Geronimo,  Guatemala.  217.  S.  sulcifrons  Putzeys,  Edmonton,  Kentucky.  218.  S.  litigiosus  Fall,  Gilroy,  Cali- 

i fomia.  219.  S.  pygmaeus  Van  Dyke,  Cliff,  New  Mexico.  220.  S.  falli  new  species,  Sabinas  Hidalgo,  Nuevo  Leon.  221. 
S.  depressus  LeConte,  San  Juan  Pueblo,  New  Mexico.  Fig.  222-232.  Male  median  lobe,  lateral  aspect.  222.  S.  arimao 
Darlington,  Soledad,  Cuba.  223.  S.  emdeni  new  species,  Palmar  Sur,  Costa  Rica.  224.  S.  sulcifrons  Putzeys,  Rumney, 
New  Hampshire.  225.  S.  litigiosus  Fall,  Willow  Creek,  California.  226.  S.  pygmaeus  Van  Dyke,  Tollhouse,  California, 
j 227.  Same,  Linares,  Nuevo  Leon.  228.  S.  scopaeus  new  species,  Montemorelos,  Nuevo  Leon.  229. Same,  Brackettville, 
| Texas.  230.  S.  falli  new  species,  Power’s  Garden,  Arizona.  231.  S.  ochthocephalus  new  species,  Fembridge,  California. 

232.  S.  depressus  LeConte,  Power’s  Garden,  Arizona.  Fig.  233.  Male  endophallus,  S.  depressus  LeConte,  Oliver,  British 
! Columbia. 


296 


Whitehead 


Fig.  234.  Geographic  variation  in  form  of  apex  of  male  median  lobe  in  S.  pygmaeus  Van  Dyke,  circles,  and  S.  scopaeus 
new  species,  squares. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


297 


235 


1000  mi 


40 


30 


237 


Fig.  235-239.  Known  distributions.  235.  S.  arimao  Darlington,  circles,  and  S.  emdeni  new  species,  triangles.  236.  S.  sul- 
cifrons  Putzeys;  open  symbols  represent  state  records  only.  237.  S.  litigiosus  Fall.  238.  S.  scopaeus  new  species.  239. 
S.  pygmaeus  Van  Dyke. 


298 


Whitehead 


240 


1000  mi 


Fig.  240-241.  Known  distributions.  240.  S.  falli  new  species,  circles,  and  S.  ochthocephalus  new  species,  triangles.  241. 
S.  depressus  LeConte.  Fig.  242.  Geographic  variation  in  body  color  of  S.  pygmaeus  Van  Dyke,  circles,  and  S.  scopaeus 
new  species,  squares;  legend  applies  to  both  tax  a. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


299 


1000  mi 


Fig.  243-244.  Geographic  variation  in  S.  pygmaeus  Van  Dyke,  circles,  and  S.  scopaeus  new  species,  squares;  broken 
lines  indicate  probable  major  bariers  to  gene  flow.  243.  Means  of  body  size,  Tables  45  and  50.  244.  Means  of  numbers 
of  discal  setae  on  left  elytron,  Tables  46  and  51. 


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245 


1000  mi 


Fig.  245-246.  Geographic  variation  in  S.  pygmaeus  Van  Dyke,  circles,  and  S.  scopaeus  new  species,  squares;  broken 
lines  indicate  probable  major  barriers  to  gene  flow.  245.  Means  of  relative  eye  size.  Tables  47  and  52.  246.  Means  of 
pronotal  form,  Tables  48  and  53.  Fig.  247-248.  Geographic  variation  in  S.  ochthocephalus  new  species,  Sample  1,  and 
S.  falli  new  species,  Samples  2-18.  247.  Means  of  body  size,  Table  55.  248.  Means  of  numbers  of  discal  setae  on  left 
elytron,  Table  56. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


301 


W 

249 


1000  mi 


250 


253  254 

Fig.  249-250.  Geographic  variation  in  S.  ochthocephalus  new  species,  Sample  1,  and  S.  falli  new  species,  Samples  2-18. 
249.  Means  of  relative  eye  size,  Table  57.  250.  Means  of  pronotal  form,  Table  58.  Fig.  251-254.  Geographic  variation 
in  S.  depressus  LeConte.  251.  Means  of  body  size,  Table  61.  252.  Means  of  numbers  of  discal  setae  on  left  elytron, 
Table  62.  253.  Means  of  relative  eye  size.  Table  63.  254.  Means  of  pronotal  form,  Table  64. 


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PHYLOGENY 


Introduction 

A taxonomic  study  normally  includes  aids  for  identification  of  specimens,  and  organizes 
information  for  storage  and  retrieval.  I believe  it  should  do  much  more  than  this.  It  should 
be  based  on  evolutionary  theory,  and  thus  furnish  sufficient  background  to  predict  relation- 
ships or  similarities  in  characteristics  not  yet  studied,  and  to  make  whatever  critical  com- 
parisons are  needed  to  test  those  predictions.  Therefore,  phylogenetic  and  biogeographic 
analyses  are  important  adjuncts  to  taxonomic  work. 

I here  give  a general  summary  of  my  philosophical  and  procedural  approaches,  which 
derive  from  a blending  of  patristic,  cladistic,  and  even  some  phenetic  philosophies  and  pro- 
cedures. I do  not  claim  originality  for  ideas  aired  in  this  section.  All  have  at  least  been 
touched  upon,  and  most  have  been  explored  in  detail,  by  one  or  more  of  the  following: 
Blackwelder  (1962);  Brundin  (1966);  Crowson  (1970);  Darlington  (1970);  Ehrlich  and 
Raven  (1969);  Hennig  (1966);  Hull  (1964,  1970);  Mayr  (1969);  Sokal  and  Crovello  (1970); 
and  Tuomikoski  (1967).  See  also  Darlington  (1971)  for  an  able  review  of  “modem  tax- 
onomy, reality,  and  usefulness”.  The  following  discussion  amplifies  those  of  my  ideas  about 
phylogeny  that  do  not  completely  agree  with  those  of  Darlington  and  Mayr,  with  whom  I 
am  in  closest  general  agreement;  it  closes  with  a series  of  examples,  some  from  the  literature 
and  some  from  my  own  work. 

Classification,  phylogeny,  and  zoogeography 

The  evolutionary  history  of  a taxon  relates  closely  to  its  past  and  present  distribution 
patterns  and  to  environmental  factors  responsible  for  those  patterns.  Thus,  to  classify  taxa 
in  accordance  both  with  evolutionary  theory  and  with  all  of  the  facts  known  about  the 
organisms,  one  must  determine  probable  evolutionary  pathways  and  correlate  these  with 
probable  historical  zoogeography.  The  phylogeny,  zoogeography,  and  classification  of  a 
group  must  be  compatible  with  one  another,  and  a classification  is  therefore  likely  to  be 
most  useful  if  completed  after  a careful  consideration  of  phylogenetic  and  zoogeographic 
evidence. 

Discussions  on  phylogeny  and  zoogeography  provide  indications  of  the  taxonomist’s 
logic,  and  may  even  suggest  compatibility  tests  for  evaluating  the  integrity  of  his  work. 
Without  these  discussions,  only  experience  permits  users  to  evaluate  a taxonomic  work. 
Thus,  the  user  is  entitled  not  only  to  know  something  about  the  taxonomist’s  philosoph- 
ical and  procedural  approaches,  but  also  to  know  where  compatibility  tests  may  most 
appropriately  be  made.  In  this  study  of  Schizogenius  I did  my  best  to  ensure  compati- 
bility. Phylogeny  was  a major  consideration  in  developing  the  entire  classification,  and 
my  original  classification  was  modified  repeatedly  until  all  incompatibilities  obvious  to 
me  were  eliminated.  Zoogeography  was  considered  in  working  out  relationships  at  the 
species  level,  and  among  many  of  the  species  groups.  Relationships  among  the  truquii, 
lineolatus,  longipennis,  depressus,  brevisetosus,  pluripunctatus,  sallei,  tristriatus,  capitalis, 
lindrothi,  darlingtoni,  ocellatus,  and  quadripunctatus  groups  were  analyzed  with  zoogeo- 
graphic considerations  in  mind.  But  relationships  of  all  other  species  groups  to  these  groups 
and  to  one  another  were  analyzed  independently  of  zoogeographic  considerations,  so  the 
user  may,  if  he  so  desires,  test  for  compatibility  between  zoogeography  and  phylogeny 
among  these  groups.  I give  examples  of  compatibility  between  phylogeny  and  zoogeo- 
graphy in  the  carabid  genus  Evarthrus  and  the  carabid  division  Brachinida,  and  an  ex- 
ample of  noncompatibility  in  the  milliped  genus  Cleptoria. 


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303 


Phylogenetic  methods 

In  general,  my  methods  for  reconstructing  a phylogeny  conform  with  Hennig’s  (1966) 
holomorphological  and  chorological  methods.  No  data  are  available  to  permit  use  of  paleon- 
tological and  parasitological  methods,  so  these  require  no  further  discussion  here.  Hennig’s 
principles  for  working  out  phylogenetic  relationships  may  be  summarized  as  follows:  recog- 
nize and  define  a monophyletic  group;  determine  its  sister  group;  and  repeat  these  processes 
until  the  phylogeny  of  a group  is  completely  reconstructed.  I apply  these  principles  by  using 
four  closely  related  procedures.  I first  develop  from  morphological  data  a probable  phylo- 
geny which  best  fits  these  data.  Then  I compare  this  phylogeny  with  chorological  data,  and 
modify  as  required.  From  the  modified  phylogeny,  I estimate  the  most  probable  morpho- 
logical and  chorological  characteristics  of  the  common  ancestors  of  sister  groups,  and  if 
necessary  modify  the  phylogeny  accordingly.  Finally,  the  revised  phylogeny  is  correlated 
with  major  paleogeological  events,  and  again  revised  if  necessary. 

In  general,  the  simplest  system  for  explanation  of  all  available  morphological  and  choro- 
logical data  is  the  system  of  choice.  I believe  further  that  a phylogeny  should  be  recon- 
structed as  completely  as  possible,  so  long  as  sufficient  data  are  available  and  so  long  as 
these  data  are  not  to  be  overruled  by  rigid  a priori  decisions.  Darlington  (1970)  rightly 
criticizes  Brundin  (1966)  for  gross  oversimplifications  in  interpretation  of  certain  of  Hennig’s 
(1966)  phylogenetic  and  zoogeographic  procedures  and  interpretations.  But  I think  cladistic 
methods  are  useful  as  analytical  tools,  at  least  to  develop  broad  generalizations. 

Monophyly,  paraphyly,  and  polyphyly.  — According  to  Darlington  (1970),  “the  central 
thesis  of  cladism  is  that  taxonomic  categories  [i.e.,  taxa]  must  be  monophyletic  in  a special, 
rigid  sense,  and  that  their  rank  must  be  determined  by  time  of  origin  rather  than  by  degree 
of  difference  or  extent  of  diversification.”  Tuomikoski  (1967),  who  otherwise  basically 
agrees  with  Hennig  (1966)  and  Brundin  (1966),  suggests  that  paraphyletic  taxa  may  be 
acceptable  under  certain  conditions.  I (1966b)  treated  the  genus  Halocoryza  as  a paraphy- 
letic taxon  in  disregard  of  the  sister  group  rule,  the  rule  of  monophyly,  and  the  rule  of 
ranking  by  time  of  origin.  This  was  done  to  account  for  the  relative  divergence  of  Schizoge- 
nius from  New  World  Halocoryza ; all  species  of  Halocoryza  are  more  similar  to  one  another 
than  to  any  species  of  Schizogenius  in  known  characteristics,  and  probably  also  in  character- 
istics not  yet  studied.  This  is  consistent  with  my  premise  that  a classification  should  be 
based  on  hypotheses  about  patristic  relationships,  rather  than  cladistic  relationships  alone. 
My  treatment  of  the  genera  Halocoryza  and  Schizogenius  parallels  classifications  in  which 
birds  are  segregated  from  crocodiles  and  other  reptiles  (Darlington,  1970;  Hennig,  1966). 

Taxa  are  not  always  monophyletic  in  the  “special,  rigid  sense”  of  Hennig  (1966),  Brundin 
(1966),  and  Tuomikoski  (1967).  This  is  clear  in  botanical  diploid-tetraploid  systems  in 
which  progeny  are  reproductively  isolated  from  their  parents.  If  tetraploidy  may  appear 
once,  then  it  may  reappear;  obviously,  the  parent  species  is  then  paraphyletic,  and  the 
daughter  species  is  polyphyletic.  Further,  as  suggested  by  Crowson  (1970),  the  tetraploid 
progeny  might  in  turn  spontaneously  produce  diploid  progeny,  so  that  both  the  diploid  and 
the  tetraploid  forms  are  at  once  paraphyletic  and  polyphyletic.  It  is  not  my  intention  here 
to  discuss  plant  speciation,  but  Crowson  does  suggest  that  diploid-tetraploid  shifts  may  have 
also  played  some  part  in  animal  evolution.  If  so,  and  if  such  diploid-tetraploid  pairs  are 
accepted  as  biological  species,  then  it  follows  that  strictly  monophyletic  species  are  not 
always  definable;  Hennig’s  techniques  provide  no  practical  way  to  distinguish  stem  species 
from  daughter  species  in  such  systems. 

That  the  stem  species  concept,  a theoretical  element  of  the  cladist’s  phylogenetic  system, 
should  not  be  confused  with  the  biological  species  concept  is  clear.  Indeed,  the  very  point 
of  the  stem  species  concept  is  to  make  a monophyletic  definition  of  extant  biological  species 


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possible.  But,  as  in  diploid-tetraploid  sister  species,  the  concept  fails  in  practice.  And  an 
even  more  radical  departure  from  the  pattern  of  a stem  species  resolved  into  two  mono- 
phyletic  daughter  species  results  when  two  parent  species  hybridize  successfully  to  produce 
an  unquestionably  polyphyletic  hybrid  species  (Hull,  1964). 

What  really  is  wrong  with  the  stem  species  as  a rigid  concept,  and  with  the  concept  of 
rigidly  monophyletic  taxa,  however,  is  to  be  found  where  species  differentiation  is  gradual, 
as  it  must  ordinarily  be.  If  geographic  isolation  develops  such  that  each  isolate  initially 
includes  two  or  more  populations,  those  populations  of  each  isolate  that  once  shared  the 
most  gene  flow  are  genetically  most  closely  related  to  one  another.  If  species  differentiation 
follows,  daughter  species  are  paraphyletic;  and  evidence  of  paraphyletic  relationships  may 
remain  for  some  time  after  completion  of  reproductive  isolation.  If  geographic  isolation 
results  from  establishment  of  a founder  population,  the  parental  isolate  is  paraphyletic 
because  initially  some  individuals  or  populations  are  genetically  more  closely  related  to  the 
founder  than  are  other  individuals  or  populations.  And  if  limited  gene  flow  continues  in 
one  or  both  directions  even  while  species  differentiation  takes  place,  one  or  both  isolates 
are  polyphyletic  in  the  early  stages  of  species  differentiation.  Thus,  strict  monophyly  can- 
not be  required  of  species  level  taxa,  since  paraphyly  and  even  polyphyly  may  be  inherent 
in  their  evolutionary  histories.  And  if  this  is  so,  then  by  logical  extension  absolute  require- 
ment of  strict  monophyly  even  for  taxa  of  higher  categories  may  be  voided. 

I do  not  accept  as  species  taxa  geographic  isolates  which  are  evidently  paraphyletic,  as 
judged  from  character  dines,  and  probably  would  not  do  so  even  if  these  geographic  isolates 
were  known  to  be  reproductively  isolated.  For  instance,  in  Schizogenius  tibialis , I suspect 
that  the  Indian  Creek  Cave  form  is  reproductively  isolated  from  its  geographically  nearest 
neighbors;  but  it  is  more  similar  and  doubtless  more  closely  related  genetically  to  them  than 
to  any  geographically  more  distant  populations  of  that  species.  I look  for  evidence  of  pro- 
pinquity of  ancestry  at  the  population  level,  and,  if  I find  such  evidence,  I regard  the  forms 
in  question  as  conspecific.  At  some  indeterminate  time  after  reproductive  isolation  is  a- 
chieved,  such  evidence  of  propinquity  of  ancestry  should  disappear;  I would  then  treat  the 
forms  in  question  as  distinct  species,  or,  where  genetic  relationships  are  in  doubt,  as  sub- 
species. This  propinquity  of  ancestry  criterion  depends  on  whether  paraphyly  remains  evi- 
dent, or  whether  such  evidence  is  lost.  Thus,  at  species  level,  I in  fact  approach  the  rigidly 
monophyletic  taxa  required  by  Hennig.  But  having  decided  that  there  is  no  theoretical 
basis  for  rigidly  monophyletic  taxa,  I do  not  insist  that  higher  taxa  always  be  monophyletic, 
though  I think  such  a criterion  is  normally  desirable. 

The  important  point  here  is  not  whether  or  not  taxa  should  be  rigidly  monophyletic.  The 
real  question  is  how  to  devise  classifications  and  reconstructed  phylogenies  that  best  show 
relationships  and  best  suggest  hypotheses  to  be  tested  by  other  workers.  If  they  are  intended 
to  have  maximum  heuristic  value,  their  construction  cannot  be  rigidly  bound  by  any  con- 
cepts other  than  evolutionary  theory.  The  simplest  possible  models  should  be  used  initially, 
but  should  freely  be  modified  for  particular  situations  as  complexities  in  these  situations 
become  evident.  I accept  cladistic  concepts  as  working  models,  but  do  not  insist  that  all 
situations  fit  a model  in  all  ways.  My  views  on  this  subject  are  analogous  to  my  use  of  the 
biological  species  concept  as  a model. 

Secondary  cladistic  concepts.  — Three  of  Darlington’s  (1970)  criticisms  of  Hennig’s  ( 1 966) 
and  Brundin’s  (1966)  methodologies  are:  that  there  is  no  fundamental  reason  why  species 
should  invariably  split  in  a simple,  dichotomous  fashion;  that  phylogenies  need  not  be 
strictly  dichotomous;  and  that  rates  of  evolution  and  divergence  need  be  neither  con- 
stant nor  the  same  for  different  groups.  If  the  range  of  a species  is  divided  into  three 
or  more  approximately  equal  parts  by  the  same  or  related  phenomena,  and  if  rates  of 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


305 


evolution  and  divergence  are  constant,  then  reproductive  isolation  must  develop  at  nearly 
equal  rates  in  all  geographic  isolates.  If  speciation  results  from  a gradual  process,  there  should 
be  no  clearly  definable  branching  points.  And  if  these  isolates  form  a pattern  in  which  the 
intermediate  isolate  has  undergone  the  least  amount  of  evident  evolutionary  change,  then  it 
might  be  difficult  or  even  impossible  to  determine  sister  species  relationships  even  if  their 
phylogeny  truly  was  dichotomous.  Nevertheless,  it  may  still  be  possible  to  realistically  sim- 
plify most  such  situations,  and,  as  I prefer  a dichotomous  system  I attempt  to  do  so;  an  ex- 
ample is  given  for  the  species  taxa  S.  planulatus,  S.  ozarkensis,  and  S.  planuloides. 

That  rates  of  evolutionary  change  and  divergence  are  neither  always  constant  nor  the 
same  in  all  groups  is  well  known.  But,  at  least  for  balanced  continental  faunas,  I think  it 
worthwhile  to  assume  an  average  rate  of  speciation  in  the  group  studied,  because  on  that 
basis  its  phylogeny  can  be  tested  against  known  paleogeological  events.  This  assumption,  in 
other  words,  makes  available  an  analytical  method  for  testing  a reconstructed  phylogeny  for 
compatibility  with  historical  zoogeography.  Some  correlated  assumptions  are  the  following. 
Changes  in  evolutionary  rates  imposed  by  gradual  environmental  change  are  negligible.  In 
a major  taxon  whose  phylogeny  has  been  worked  out,  at  least  one  major  lineage  is  sufficient- 
ly diverse  and  complete  that  extinction  patterns  may  be  disregarded.  And  apparent  large 
gaps  between  related  genus-group  taxa  are  associated  with  rapid  rates  of  evolutionary  change 
and  divergence  resulting  from  exploitation  of  new  habitats  and  resulting  major  new  adapta- 
tions. Analytical  methods  required  to  test  a reconstructed  phylogeny  for  compatibility  with 
historical  zoogeography  are  the  following.  The  average  time  required  for  speciation  is  de- 
termined from  what  is  known  about  living  species  taxa  of  the  group  in  question,  and  from 
probable  historical  factors  involved  in  their  evolution.  In  the  phylogeny  to  be  tested,  this 
average  time  is  treated  as  equal  to  the  time  between  successive  dichotomies  in  the  lineage 
with  the  maximum  number  of  known  dichotomies.  The  age  of  any  desired  common  ances- 
try is  thus  the  maximum  number  of  dichotomies  between  that  ancestry  and  the  present, 
multiplied  by  the  average  time  span  required  for  speciation.  These  assumptions  and  methods 
are  generalizations,  and  perhaps  even  gross  oversimplifications,  but  the  interesting  point  is 
that  where  I have  tested  them  they  seem  to  work.  Examples  are  given  for  the  carabid  divi- 
sion Brachinida  and  the  carabid  genus  Evarthrus.  I used  these  techniques  to  help  place  such 
species  taxa  as  S.  arimao  in  both  classification  and  reconstructed  phylogeny,  whenever  mor- 
phological data  suggested  two  or  more  equally  plausible  evolutionary  pathways. 

Phylogeny,  classification,  and  the  biological  species 

Various  relationships  among  phylogeny,  classification,  and  biological  species  were  dis- 
cussed by  Hull  (1970).  I here  summarize  my  views  about  relationships  and  compatibilities 
among  these,  with  particular  emphasis  on  the  biological  species  concept  as  a unifying  model 
of  great  evolutionary  importance.  I also  discuss  how  I apply  the  biological  species  concept 
in  certain  situations  in  which  reproductive  relationships  are  discordant  with  suspected  phy- 
logenetic relationships,  and  explain  my  rationale  for  interpreting  these  situations.  In  this 
discussion,  I start  from  Hull’s  (1970)  restricted  definition  of  “classification,”  Hennig’s 
(1966)  definition  of  “phylogenetic  relationships,”  and  Mayr’s  (1969)  definition  of  “species”. 

A concise  statement  of  my  interpretation  of  relationships  among  phylogeny,  classifica- 
tion, and  biological  species  is  the  following.  Organisms  are  classified  into  taxa,  each  of  which 
is  based  on  the  hypothesis  that  included  organisms  are  more  closely  related  to  one  another 
than  to  excluded  organisms,  at  least  at  any  one  time.  A reconstructed  phylogeny  is  based  on 
hypotheses  about  how  taxa  are  related  to  one  another.  In  this  system,  the  species  is  not  a 
necessarily  fundamental  evolutionary  unit;  but  the  classification  and  reconstructed  phylo- 
geny are  unified  by  the  biological  species  concept,  which  consequently  is  a fundamental 


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evolutionary  model. 

Compatibility  of  patristic  classification  and  reconstructed  phytogeny.  - I prefer  patristic 
classifications  over  cladistic  classifications,  because  the  cladistic  relationships  implied  in  the 
latter  may  be  adequately  expressed  in  a reconstructed  phylogeny.  Inherent  in  a patristic 
classification  is  the  hypothesis  that  extant  elements  of  a taxon  share  closer  patristic  relation- 
ships with  one  another  than  with  extant  elements  of  excluded  taxa.  A reconstructed  phylo- 
geny, on  the  other  hand,  must  reflect  evolutionary  history,  and  is  based  on  hypotheses  about 
cladistic  relationships  between  taxa.  The  patristic  classification  and  the  reconstructed  phylo- 
geny are  therefore  statements  based  on  different  sets  of  hypotheses.  For  heuristic  purposes, 
however,  they  are  interdependent,  and  must  therefore  be  compatible  with  one  another. 

Compatibility  between  an  evolutionary  classification  and  a reconstructed  phylogeny  is 
achieved  if  units  of  at  least  one  level  are  common  to  both.  Taxa  of  species  rank  are  such 
units.  Higher  taxa  in  an  evolutionary  classification  may  not  be  compatible  with  the  recon- 
structed phylogeny,  and  units  below  species  rank  do  not  normally  appear  in  the  recon- 
structed phylogeny.  Taxa  of  species  rank  are  the  units  best  suited  to  insure  compatibility 
between  the  evolutionary  classification  and  the  reconstructed  phylogeny  for  the  following 
additional  reasons. 

Hennig  (1966)  discussed  differences  between  ontogenetic,  tokogenetic,  and  phylogenetic 
relationships,  which  apply  respectively  to  semophoronts,  individuals,  and  reproductively  iso- 
lated groups  of  individuals  or  populations.  An  evolutionary  classification  is  possible  because 
semophoronts,  individuals,  and  populations  cluster  to  form  more  or  less  discrete  groups 
which  are  linked  by  phylogenetic  relationships.  Semophoronts,  individuals,  and  populations 
that  share  ontogenetic  and  tokogenetic  relationships  are  conspecific,  but  these  relationships 
are  not  sufficient  to  define  a species.  Ontogenetic  and  tokogenetic  relationships  between 
geographic  isolates  are  replaced  by  phylogenetic  relationships,  but  are  renewed  if  gene  ex- 
change occurs  before  reproductive  isolation  is  achieved.  This  potential  instability  of  phylo- 
genetic relationships  is  sufficient  reason  for  units  of  compatibility  between  an  evolutionary 
classification  and  a reconstructed  phylogeny  to  be  taxa  of  at  least  species  rank. 

A species  is  a unit  of  classification  based  on  known  or  hypothetical  relationships  among 
included  semophoronts,  individuals,  and  populations;  these  relationships  are  interpreted  in 
terms  of  a model,  the  biological  species  concept.  Since  only  for  taxa  of  species  rank  is  there 
such  a model,  species  level  taxa  are  better  suited  as  units  of  compatibility  between  an  evo- 
lutionary classification  and  a reconstructed  phylogeny  than  are  taxa  of  higher  rank.  Thus, 
species  have  the  important  function  of  unifying  facts,  hypotheses,  and  theories,  and  this 
is  why  I consider  the  biological  species  concept  as  a fundamental  evolutionary  model. 

Some  problems  in  applying  the  biological  species  concept.  — Semophoronts  and  individ- 
uals may  be  classified  (Blackwelder,  1962;  Hennig,  1966),  but  since  they  are  characterized 
by  having  ontogenetic  or  tokogenetic  relationships  they  do  not  evolve  independently  from 
one  another  and  therefore  are  not  fundamental  evolutionary  units.  Some  authors  believe 
that  species  are  the  real  units  of  evolution  (Mayr,  1969).  Others  (see  Hull,  1970)  suggest 
that  species  are  not  fundamental  evolutionary  units,  and  that  populations  may  be  such  units 
(Ehrlich  and  Raven,  1969;  Sokal  and  Crovello,  1970).  Their  basic  arguments  are  that  selec- 
tion is  the  prime  factor  in  evolution  and  that  gene  flow  normally  is  negligible.  But  how  can 
one  explain  patterns  of  continuous  or  nearly  continuous  geographic  variation  if  gene  flow 
is  negligible?  One  might  argue  that  similarities  between  geographically  proximate  popula- 
tions result  exclusively  from  similar  selective  regimes.  But  such  an  argument  is  superficial; 
it  does  not  explain,  for  example,  the  circular  pattern  of  variation  in  relative  eye  size  in  S. 
tibialis  (Fig.  188).  I think  that  all  character  dines  in  Schizogenius  species  studied  by  me 
indicate  genetic  relationships  between  at  least  some  geographically  proximate  population 


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307 


samples,  and  that  these  genetic  relationships  result  from  relatively  recent  gene  flow.  How- 
ever, though  populations  must  interact  with  some  frequency  if  there  are  non-random  pat- 
terns of  geographic  variation,  they  must  be  sufficiently  independent  of  one  another  for 
selection  to  act  to  produce  this  variation.  Except  in  such  species  that  include  only  one 
population,  species  normally  are  not  really  fully  integrated  evolutionary  units  since  included 
populations  are  independent  of  one  another  to  varied  degrees. 

A problem  raised  by  the  notion  that  species  are  not  necessarily  fundamental  evolutionary 
units  is  that  reproductive  relationships  may  not  be  concordant  with  suspected  phylogenetic 
relationships.  This  problem  arises  as  a consequence  of  loss  of  tokogenetic  relationships  be- 
tween populations  or  groups  of  populations  (i.e.,  geographic  isolation)  before  reproductive 
isolation  is  attained.  These  populations  or  groups  of  populations  have  phylogenetic  relation- 
ships, but  according  to  the  biological  species  concept  they  are  conspecific.  If  the  evolu- 
tionary classification  and  reconstructed  phylogeny  are  to  have  maximum  heuristic  value, 
interpretations  of  discordant  reproductive  and  phylogenetic  relationships  must  be  flexible 
and  must  be  made  at  the  population  level.  That  some  species  taxa  may  not  perfectly  fit  the 
biological  species  concept  does  not  diminish  the  importance  or  validity  of  that  model.  Data 
compared  with  a model  either  fit  it  or  fail  to  fit  it,  and  models  in  biology  are  not  exception- 
less; why  would  they  be,  when  life  itself  has  no  rigorous,  universally  accepted  definition? 
Indeed,  from  evolutionary  theory  the  existence  of  exceptions  is  to  be  predicted.  The  bio- 
logical species  concept  is  a good  model  because  most  data  do  fit  it,  and  its  importance  is 
that  it  provides  a reasonably  consistent  basis  for  species  recognition. 

Reproductive  isolation  is  a criterion  for  defining  species  in  terms  of  discrete  evolutionary 
potential.  However,  before  reproductive  isolation  is  attained,  isolates  may  have  a marked 
evolutionary  potential  even  if  that  potential  is  not  discrete.  An  example  is  found  in  the  taxa 
S.  pygmaeus  and  S.  scopaeus.  Alternatively,  if  possible  reproductive  isolates  retain  evidence 
of  paraphyletic  relationships,  their  evolutionary  potential,  though  discrete,  may  be  of  rela- 
tively recent  origin,  and  may  be  limited.  An  example  of  this  alternative  is  the  taxon  S. 
tibialis.  I base  species  taxa  on  hypotheses  about  past  or  present  genetic  relationships  between 
populations  in  addition  to  known  or  suspected  relationships  among  semophoronts  and 
individuals.  Recognition  of  these  taxa  depends  on  evidence  about  how  or  perhaps  whether 
populations  most  probably  interact  or  have  interacted  with  one  another.  Thus,  some  species 
taxa  in  Schizogenius  such  as  S.  tibialis , S.  pygmaeus,  and  S.  scopaeus  do  not,  or  may  not, 
exactly  fit  the  biological  species  concept.  But  even  if  they  are  not  fundamental  evolutionary 
units,  these  taxa  seem  more  consistent  with  the  differing  hypotheses  on  which  I define  spe- 
cies, base  classifications,  and  reconstruct  phytogenies  than  they  would  if  redefined  in  terms 
of  discrete  evolutionary  potential.  I think  they  should  be  reclassified  if,  and  only  if,  under- 
lying hypotheses  fail,  regardless  of  additional  data  on  reproductive  isolation. 

Some  examples  of  phylogenetic  analyses 

In  this  section  I illustrate  with  examples,  some  from  the  literature  and  some  from  my 
study  of  Schizogenius , certain  points  raised  in  preceding  sections.  These  examples  are 
keyed  to  the  pertinent  section  of  discussion.  A further  reason  for  including  these  examples 
is  to  give  the  reader  insight  into  details  of  my  phylogenetic  methods.  Not  all  of  the  Schizo- 
genius phylogeny  is  discussed  in  detail,  but  these  examples  are  representative  of  my  methods 
throughout. 

Compatibility  of  phylogeny  and  zoogeography.  - The  examples  given  here,  both  from 
the  literature,  are  chosen  to  show  how  compatibility  tests  between  phylogeny  and  zoogeo- 
graphy can  be  made.  To  test  techniques  outlined  above,  I examined  the  phylogeny  and 
historical  zoogeography  proposed  for  Brachinus  (Erwin,  1970)  and  Evarthrus  (Ball  and 


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Freitag,  in  Freitag,  1969).  For  various  reasons,  the  average  time  required  for  speciation  in 
the  genus  Schizogenius  is  considered  to  be  approximately  3,000,000  years.  The  Brachinus 
and  Evarthrus  examples  were  selected  to  find  if  this  time  span  might  be  common  among 
North  American  carabid  beetles. 

Erwin’s  proposed  phylogeny  suggests  a maximum  of  23  dichotomies  to  reach  the  common 
ancestor  of  American  Neobrachinus  species.  The  true  number,  as  determined  from  known 
species  taxa,  may  be  somewhat  less,  since  Erwin’s  data  suggest  that  the  following  taxa  form 
monophyletic  groups:  B.  oaxacensis,  B.  patruelis,  and  B.  conformis\  B.  ovipennis  and  B. 
medius ; and  B.  tenuicollis  and  B.  cyanipennis.  This  would  reduce  the  required  minimum 
number  of  dichotomies  to  19.  If  the  3,000,000  year  time  span  is  correct,  ancestral  Neobra- 
chinus entered  North  America  some  57,000,000  to  69,000,000  years  ago.  This  fits  Erwin’s 
hypothesis  that  entry  into  North  America  occured  at  some  time  before  Eocene,  probably 
during  Paleocene  some  58,000,000  to  63,000,000  years  ago.  Erwin  further  suggested  that 
the  primitive  ancestor  of  the  bombardier  beetles  of  the  Division  Brachinida  evolved  in 
Middle  to  Late  Cretaceous.  His  phylogeny  indicates  a minimum  of  seven  and  a maximum 
of  ten  additional  dichotomies  to  reach  back  to  this  brachinine  ancestor.  At  3,000,000  years 
per  dichotomy,  this  ancestor  evolved  between  78,000,000  and  99,000,000  years  ago,  a 
period  well  within  Erwin’s  suggested  limits.  The  only  fault  I found  with  Erwin’s  work  is 
that  his  phylogeny  is  incompletely  reconstructed.  It  is  sufficiently  complete,  however,  that 
it  can  be  tested  for  compatibility  and  correlation  with  the  zoogeography,  and  I find  that 
there  is  indeed  a strong,  positive  correlation.  I therefore  suggest  that  Erwin’s  work  may  be 
used  with  confidence,  for  heuristic  purposes. 

There  are  limits  to  the  applicability  of  these  procedures.  Ball  and  Freitag  suggest  that 
the  ancestor  of  the  pterostichine  genus  Evarthrus  entered  North  America  via  the  Behring 
land  bridge  in  Early  Tertiary.  No  dating  of  this  early  event  by  my  techniques  is  possible 
from  the  Evarthrus  phylogeny,  because  all  western  lineages,  if  there  were  any,  have  become 
extinct.  Whether  this  dating  can  be  done  from  examination  of  the  phylogeny  of  the  Old 
World  sister  group  cannot  be  judged  until  such  a phylogeny  is  reconstructed.  However, 
suggestions  about  subsequent  events  can  be  tested  and  dated  according  to  the  suggested 
Evarthrus  phylogeny.  Members  of  the  genus  spread  across  temperate  Arcto-Tertiary  forests 
in  the  north,  but  in  Middle  Tertiary  receded  eastward  as  did  those  forests.  The  common 
ancestor  of  living  Evarthrus  species  evolved  after  this  eastward  recession.  Depending  on  how 
the  peculiar  taxa  E.  gravesi  and  E.  hypherpiformis  fit  the  phylogeny,  there  may  be  as  few 
as  nine  to  as  many  as  eleven  dichotomies  needed  to  reach  that  common  ancestor.  Its  age 
thus  falls  between  27,000,000  and  33,000,000  years,  which  corresponds  to  Oligocene,  or 
Middle  Tertiary,  in  support  of  Ball  and  Freitag’s  ideas.  Moreover,  members  of  only  the  sub- 
striatus  and  torvus  groups  extend  much  westward  of  southeastern  forested  regions.  Just 
four  dichotomies  are  required  to  reach  their  common  ancestor,  which  may  still  have  been  a 
component  of  the  eastern  forest  fauna;  the  timing  is  about  12,000,000  years  ago,  in  Early 
Pliocene.  This  supports  Ball  and  Freitag’s  idea  that  Gulf  Coast  grassland  habitats  were  not 
entered  by  Evarthrus  species  before  post-Miocene  time,  because  if  they  were  the  genus 
should  today  be  represented  in  northeastern  Mexico.  My  conclusions  are  that  Ball  and 
Freitag’s  phylogenetic  and  zoogeographic  analyses  are  compatible,  and  so  probably  correct, 
and  that  therefore  their  work  may  confidently  be  used  for  heuristic  purposes. 

The  data  required  for  these  compatibility  tests  are  the  authors’  suggested  phylogenies 
and  suggested  timings  for  zoogeographic  events.  For  both  Brachinida  and  Evarthrus , com- 
patibilities were  found  at  two  different  points  in  time.  General  conclusions  from  these 
analyses  are  that  for  each  the  phylogeny  and  zoogeography  are  compatible,  and  that  both 
of  these  taxonomic  works  are  therefore  probably  highly  reliable.  The  taxonomy  must  have 


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309 


been  carefully  done  and  well  thought  out  for  these  compatibilities  to  be  evident.  An  addi- 
tional conclusion  is  that  the  3,000,000  year  time  span  between  dichotomies  may  be  a useful 
generalization  for  North  American  Carabidae,  although  in  Evarthrus  this  time  span  may  be 
somewhat  shorter  as  my  datings  are  at  upper  limits  of  acceptability. 

Incompatibility  of  taxonomy,  phylogeny,  and  zoogeography.  - Hoffman  (1967),  in  his 
review  of  the  milliped  genus  Cleptoria,  proposed  aesthetically  pleasing  phylogenetic  and 
zoogeographic  patterns  but  gave  no  indication  about  timing.  Thus,  my  test  for  compatibility 
between  phylogeny  and  zoogeography,  illustrated  above  for  Evarthrus  and  Brachinus,  is 
inapplicable.  A user  interested  in  studying  relationships  between  species  must  assess  the 
reliability  of  the  taxonomy,  phylogeny,  and  zoogeography  by  other  means,  if  given  no  firm 
basis  to  believe  the  phylogeny  and  zoogeography  have  good  heuristic  value.  Without  per- 
sonal experience  with  the  organisms,  the  only  way  he  can  judge  the  soundess  of  their  taxo- 
nomy is  to  carefully  read  and  compare  all  descriptive  material,  including  information  about 
geographic  variation.  Based  on  his  conclusions  about  taxonomy,  he  must  then  decide  for 
himself  about  phylogeny  and  zoogeography. 

Hoffman’s  data  indicate  that  all  five  species  taxa  included  in  Cleptoria,  except  perhaps 
C.  rileyi  and  C.  bipraesidens , have  long  been  reproductively  isolated.  These  data  also  sup- 
port his  ideas  that  the  taxon  C.  divergens  is  the  most  primitive  in  the  genus,  and  that  the 
origin  of  the  genus  was  in  the  southern  Appalachians.  But,  though  I think  the  taxonomy  is 
sound,  I think  Hoffman’s  explanation  of  the  phylogeny  and  zoogeography  (Fig.  259)  in 
Cleptoria  is  improbable.  He  suggests  that  the  five  allopatric  species  taxa  evolved  in  a “hop- 
scotch” pattern,  from  the  southern  Appalachians  southward  into  the  piedmont  region.  Such 
a pattern  would  probably  be  comparatively  recent  in  development,  and  some  evidence  of 
gene  flow  patterns  should  remain;  I would  expect  to  find  evidence  of  paraphyletic  relation- 
ships, as  discussed  for  Schizogenius  tibialis  later  in  this  section.  However,  no  gene  flow 
patterns  are  evident  between  species  taxa  in  Cleptoria.  I think  an  orderly  “hop-scotch” 
pattern  is  unlikely,  if  species  taxa  really  are  reproductive  isolates,  since  if  isolates  are  so 
readily  founded  they  should  also  be  susceptible  to  replacement  or  displacement  by  subse- 
quent founders.  The  pattern  suggests  much  greater  vagility  than  is  reflected  in  known  facts 
of  distribution  and  morphology.  An  essentially  linear  sequence  among  reproductive  isolates 
would  more  probably  result  from  a “taxon  cycle”  system  of  displacement  (Wilson,  1961); 
but  then  the  taxon  C.  divergens  should  be  the  most  apomorphic  of  the  genus,  not  the  most 
plesiomorphic. 

A review  of  Hoffman’s  data  suggests  that  a more  plausible  system  of  evolutionary  path- 
ways in  the  milliped  genus  Cleptoria  (Fig.  260)  is  the  following.  The  first  dichotomy  in  the 
phylogeny  of  Cleptoria  was  a split  of  a piedmont  form,  with  telopodite  shortened  and 
thickened  and  gonopods  not  interlocked,  from  the  more  conservative  montane  C.  divergens. 
The  piedmont  form  developed  isolates  north  and  south  of  the  Savannah  River,  with  the 
peculiarly  specialized  C.  abbot ti  representing  the  latter.  A subsequent  extension  by  the 
northern  isolate  southward  across  the  Savannah  River  resulted  in  the  separation  of  the 
northern  C.  macra  from  the  southern  ancestor  of  C.  rileyi  and  C.  bipraesidens,  which  lost 
the  prefemoral  process.  These  forms  diverged  somewhere  near  their  present  ranges,  and 
displaced  C.  abbotti  from  the  western  part  of  its  range.  I thus  do  not  think  that  C.  abbotti 
is  closest  to  C.  rileyi,  and  in  fact  suspect  that  C.  macra  is  closer  to  C.  rileyi  and  C.  biprae- 
sidens than  is  C.  abbotti.  Support  for  these  suggestions  is  evident  in  Hoffman’s  diagrams  of 
the  process  of  sternum  three,  which  show  C.  rileyi  and  C.  macra  as  intermediate  between  C. 
divergens  and  C.  abbotti',  I doubt  that  this  similarity  is  a coincidence.  Thus,  though  I have 
not  personally  studied  specimens  of  Cleptoria , my  reinterpretations  from  Hoffman’s  data 
seem  to  explain  evolutionary  pathways  at  least  as  well  as  do  Hoffman’s  interpretations. 


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When  phylogenetic  and  zoogeographic  analyses  are  published,  they  should  allow  a user 
to  verify  for  himself  the  probable  correctness  of  suggested  relationships.  If  an  author’s  own 
interpretations  are  compatible  with  morphological  and  chorological  data,  no  reinterpreta- 
tions should  be  needed.  But  the  heuristic  value  of  an  author’s  interpretations  are  suspect 
is  his  published  data  suggest  more  plausible  interpretations. 

Phytogeny:  dichotomous  or  trichotomous?  - The  taxa  Schizogenius  planulatus,  S.  ozar- 
kensis,  and  S.  planuloides  have  ranges  now  separated  by  the  Mississippi  and  Red  Rivers  (Fig. 
189).  Perhaps  these  rivers  were  at  one  time  not  effective  barriers,  but  became  so  because  of 
some  major  change  in  climate  which  increased  water  loads  and  thus  eliminated  suitable 
habitats  by  deposition  of  silt  or  mud.  The  three  isolates  evolved  independently,  and  male 
genitalia  became  more  specialized  in  S.  planulatus  (Fig.  175)  and  S.  planuloides  (Fig.  177) 
than  in  the  geographically  intermediate  S.  ozarkensis  (Fig.  176).  This  sequence  of  events 
suggests  a trichotomous  phylogeny.  But  other  zoogeographic  factors  suggest  that  a dicho- 
tomous phylogeny  is  more  probable. 

The  Mississippi  River,  near  its  confluence  with  the  Ohio  River,  should  pose  no  major 
barrier  to  these  organisms,  since  limestone  uplands  in  southern  Illinois  nearly  link  those  of 
western  Kentucky  and  eastern  Missouri.  Drainage  systems  from  uplands  of  this  entire  region 
should  provide  suitable  habitats  for  members  of  this  complex.  Indeed,  there  may  be  extant 
populations  of  one  or  both  of  S.  planulatus  and  S.  ozarkensis  in  southern  Illinois.  Further, 
these  species  range  primarily  in  regions  of  deciduous  forests  and  may  therefore  be  adapted 
to  cooler  conditions  than  is  S.  planuloides,  a species  of  warmer,  more  arid  regions.  I think 
the  ancestor  of  these  taxa  evolved  in  subtropical  Texas  or  northeastern  Mexico,  since  that 
is  the  region  of  greatest  diversity  among  their  relatives.  The  first  dichotomy  in  their  phylo- 
geny followed  establishment  of  a northern  isolate,  which  became  cool  adapted  and  spread 
across  temperate  forested  areas  in  the  Ozark  and  Appalachian  regions.  Meanwhile,  the  male 
genitalia  of  the  southern  isolate  evolved  its  specialized  characteristics.  The  second  dichotomy 
developed  when  Early  Pleistocene  climatic  deterioration  caused  the  northern  isolate  to  sub- 
divide into  isolates  restricted  to  the  southern  Ozark  and  southern  Appalachian  regions.  Then 
the  Appalachian  isolate  diverged  in  characteristics  of  the  male  genitalia  from  the  more  con- 
servative Ozark  isolate.  Following  retreating  Pleistocene  glaciations,  the  ranges  of  Ozark  and 
Appalachian  isolates  expanded  northward,  and  may  now  approach  or  even  overlap  one 
another  in  the  region  near  the  confluence  of  the  Mississippi  and  Ohio  Rivers. 

Problem  species.  — Some  species  of  Schizogenius  may  not  fit  precisely  the  biological  spe- 
cies concept.  Examples  are  the  taxa  S.  tibialis,  S.  pygmaeus,  and  S.  scopaeus.  Where  evi- 
dence from  geographic  variation  indicates  relatively  recent  gene  flow  between  geographical- 
ly proximate  populations,  I regard  them  as  conspecific.  If  no  evidence  of  such  gene  flow 
between  these  populations  exists,  either  directly  or  through  some  alternative  geographic 
sequence,  and  if  morphological  and  chorological  evidence  suggests  reproductive  isolation, 
then  I recognize  them  as  distinct  species. 

Lindsay  and  Vickery  (1967)  found  from  studies  of  Mimulus  “a  picture  of  the  frequent 
emergence,  change,  and  disappearance  of  distinctive  populations  with  only  the  rare  forma- 
tion of  one  sufficiently  distinct  to  be  on  its  own  evolutionary  path.”  I suspect  the  same  is 
true  in  Schizogenius. 

In  the  taxon  S.  tibialis  (Fig.  188),  the  Indian  Cave  Creek  form  may  or  may  not  be  a repro- 
ductive isolate,  but  regardless  shows  evidence  of  past  genetic  continuity  with  geographically 
proximate  populations  to  the  south.  It  is  restricted  and  peripheral  in  known  distribution,  is 
undoubtedly  of  comparatively  recent  origin,  and  probably  has  a low  potential  for  survival. 
Suspected  reproductive  and  phylogenetic  relationships  are  discordant;  if  this  form  is  recog- 
nized as  a separate  species  because  of  reproductive  isolation,  the  southern  form  is  reduced  to 


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311 


an  evidently  paraphyletic  taxon.  I regard  the  Indian  Cave  Creek  isolate  as  conspecific  with 
other  populations  of  S.  tibialis  because  it  is  patristically  more  closely  related  to  geographi- 
cally proximate  populations  than  to  more  distant  ones.  I do  not  here  regard  possible  repro- 
ductive isolation  as  a useful  criterion  for  species  recognition,  because  though  evolutionary 
potential  may  be  discrete  I do  not  think  it  is  great.  I would  not  recognize  the  Indian  Cave 
Creek  form  as  a distinct  species  even  if  known  to  be  reproductively  isolated,  unless  hypothe- 
ses about  patristic  relationships  were  found  to  fail.  One  such  failure  would  be  to  find  the 
two  forms  sympatric;  the  hypothesis  of  paraphyletic  relationships  would  then  be  falsified. 

In  contrast,  certain  geographically  distant  populations  of  the  taxa  S.  pygmaeus  (Fig.  239) 
and  S.  scopaeus  (Fig.  238)  may  well  not  be  reproductively  isolated  from  one  another,  but 
geographically  proximate  populations  evidently  are.  These  taxa,  both  of  which  are  abundant 
and  widespread,  are  sympatric  in  northeastern  Mexico  where  they  are  doubtless  reproduc- 
tively isolated.  No  certain  evidence  of  paraphyly  remains  between  geographically  proximate 
populations  of  these  taxa,  and  the  possible  lack  of  reproductive  isolation  between  certain 
distant  populations  is  not  a useful  criterion  for  species  recognition.  These  taxa,  if  marked 
by  a lack  of  discrete  evolutionary  potential,  seem  nevertheless  to  have  such  evolutionary 
potential  well  developed,  and  probably  have  had  it  so  developed  much  longer  than  have 
had  the  isolates  of  S.  tibialis.  Along  the  Rio  Conchos  and  Rio  Grande  in  Chihuahua,  clinal 
continuity  between  S.  pygmaeus  and  S.  scopaeus  is  suggested  in  several  statistical  character- 
istics (Fig.  243-246)  but  not  in  structures  of  the  male  genitalia  (Fig.  234).  I would  treat 
these  taxa  as  conspecific  only  if  unexpected  evidence  from  a geographically  intermediate 
sample  showed  my  interpretation  of  differences  in  the  male  genitalia  to  be  wrong. 

Subgenera  and  species  groups  of  Schizogenius 

In  this  section  I attempt  to  reconstruct  the  phylogeny  of  major  lineages  of  the  genus 
Schizogenius,  using  contrasting  phyletic  and  phenetic  methods.  For  purposes  of  this  dis- 
cussion, the  truquii,  lineolatus,  longipennis,  depressus,  brevisetosus,  pluripunctatus,  sallei, 
and  tristriatus  groups  are  treated  as  a monophyletic  group,  the  truquii  lineage.  The  capitalis, 
lindrothi,  quadripunctatus,  darlingtoni,  and  ocellatus  groups  form  another  monophyletic 
group,  the  capitalis  lineage.  All  other  species  groups  are  considered  as  separate  lineages. 

Relationships  of  the  genus  Schizogenius  to  other  genera  were  discussed  earlier,  and  in  a 
separate  paper  (1966b).  From  those  intergeneric  comparisons,  I judge  that  ancestral  charac- 
ter states  in  the  common  ancestor  of  living  Schizogenius  species  included  the  following: 
mandibles  more  or  less  arcuate  laterally;  lacinia  setose  along  outer  margin;  mentum  deeply 
emarginate  at  middle,  median  tooth  distinct;  gula  less  than  0.3  maximum  width  of  mentum; 
antennal  article  two  not  plurisetose;  eyes  not  bordered  by  dorsal  carinae;  frontal  carinae  not 
perfectly  regular,  parallel,  equidistant,  or  equally  raised;  occiput  punctate  dorsally,  extended 
laterad  along  posterior  margin  of  eyes;  hind  leg  with  tarsus  more  than  0.6  length  of  tibia; 
paramedian  carinae  of  sternum  three  strongly  developed;  and  pygidium  with  well  developed 
median  longitudinal  rows  of  files.  Additional  probable  plesiomorphic  character  states,  de- 
termined from  intergeneric  and  infrageneric  comparisons,  are  listed  in  Table  65.  Character 
states  so  listed  are  designated  or  coded  as  apomorphic  or  plesiomorphic,  based  on  what  I 
think  is  primitive  in  a group  or  lineage  (see  Ball  and  Erwin,  1969).  The  apomorphic  state 
of  several  characters  arose  in  more  than  one  lineage,  but  in  this  phylogeny  none  of  the 
characters  studied  reverted  to  the  plesiomorphic  state  except  the  elongate  paronychia  of  the 
arechavaletae-truquii-capitalis  and  strigicollis-elongatus-carinatus  lineages. 

The  genus  is  most  diverse  in  South  America,  which  I therefore  think  is  the  primary  center 
of  radiation.  Penetration  of  North  and  Middle  America  over  the  water  gap  that  separated 
this  area  from  South  America  during  Middle  Tertiary  is  considered  an  apochoric  zoogeo- 


Whitehead 


312 

graphic  characteristic.  Ancestors  of  only  the  cremilatas . ferrugineus,  and  truquii  lineages 
made  these  crossings.  These  zoogeographic  data  were  not  considered  in  reconstructing  the 
phylogeny,  but  are  not  in  conflict  with  it. 

Phytogeny  reconstructed  by  phyletic  techniques.  — The  reconstructed  phylogeny  of  ma- 
jor lineages  of  Schizogenius  is  shown  in  Fig.  255,  and  the  principal  data  used  are  listed  in 
Table  65:  the  time  scale  suggested  is  discussed  in  a later  section.  I here  summarize  principal 
features  of  this  phylogeny,  indicate  its  weaknesses  and  strengths,  and  note  apomorphic 
character  states  thought  to  have  arisen  in  more  than  one  lineage.  I am  confident  that  all 
branching  points  are  correctly  positioned,  and  have  indicated  apomorphic  character  states 
for  nearly  all  dichotomies.  I think  that  each  lineage  is  monophyletic.  and  that  this  phyloge- 
ny is  most  probably  a correct  representation  of  evolutionary  relationships:  this  reconstructed 
phylogeny  should  therefore  have  strong  heuristic  value  for  predicting  patristic  relationships. 

Ancestral  Schizogenius  differentiated  into  two  stocks,  the  subgenera  Genioschizus  and 
Schizogenius . Apomorphic  character  slates  in  ancestral  Genioschizus  included  flared  and 
pitted  lateral  channel  of  elytron,  and  reduced  numbers  of  discal  setae  on  elytron:  the  latter 
character  state  evolved  independently  in  several  lineages  of  Schizogenius.  In  ancestral  Schi- 
zogenius\ apomorphic  conditions  included  shortened  paronychia,  and  loss  of  paralateral  pro- 
notal  sulci:  the  latter  condition  evolved  independently  in  the  tenuis  group  of  Genioschizus. 

Genioschizus  differentiated  into  three  known  groups.  This  differentiation  took  place  com- 
paratively recently,  as  Genioschizus  is  far  less  diverse  than  is  Schizogenius.  Among  known 
forms,  the  crenulatus  group  is  the  most  distinctive.  Its  ancestor  entered  North  America 
across  a water  gap  between  North  and  South  America,  and  developed  the  apomorphic  con- 
dition of  truncated  mentum  lateral  lobes.  The  ancestor  of  its  sister  group  remained  in  South 
America,  developed  the  apomorphic  condition  of  elytral  intervals  carinate  at  apices,  and 
differentiated  into  the  quinquesulcatus  and  tenuis  groups.  An  apomorphic  condition  of 
ancestral  tenuis  group  was  loss  of  paralateral  pronotal  sulci,  an  apomorphic  condition  also 
of  ancestral  subgenus  Schizogenius.  A weakness  in  my  reconstructed  phylogeny  of  Ge- 
nioschizus is  that  I found  no  useful  synapomorphic  condition  to  characterize  the  quinque- 
sulcatus group. 

The  first  major  dichotomy  in  the  phylogeny  of  the  subgenus  Schizogenius  reflects  the 
differentiation  of  the  jacarensis-optirnus  lineage  from  the  remainder  of  the  subgenus.  The 
ancestor  of  its  sister  group  had  the  apomorphic  sharply  engraved  clypeal  suture:  this  condi- 
tion appeared  also  in  some  members  of  the  optimus  group.  Numerous  apomorphic  features 
distinguished  the  ancestor  of  the  jacarensis-optimus  lineage,  but  among  those  indicated  in 
Fig.  255  only  the  condition  of  elongate  tarsi  was  not  repeated  elsewhere  in  the  phylogeny 
of  the  genus.  The  jacarensis-optimus  lineage  subsequently  differentiated  into  the  optimus 
and  monotypic  jacarensis  groups.  Apomorphic  features  of  the  ancestor  of  the  optimus  group 
included  apex  of  male  median  lobe  abruptly  bent,  and  bristles  on  terminal  palpal  articles 
longitudinally  arranged. 

The  ancestor  of  the  ferrugineus  group  entered  Middle  and  North  America  where  it  lost 
the  female  paramedian  ambulatory  setae  from  sternum  seven,  but  this  loss  was  repeated 
later  in  the  phylogeny  of  its  sister  group.  Apomorphic  conditions  of  the  ancestor  of  that 
sister  group  were  antennae  filiform  and  female  pygidial  margin  crenulate. 

The  ancestor  of  the  South  American  basalis  group  differentiated  from  the  ancestor  of  its 
sister  group  when  the  latter  lost  the  female  paramedian  ambulatory  setae  from  sternum 
seven.  This  apomorphic  condition  also  characterized  ancestors  of  the  jacarensis-optimus  and 
ferrugineus  lineages.  A weakness  in  this  dichotomy  is  that  I found  no  synapomorphic  char- 
acteristic for  the  basalis  group.  In  most  members  of  the  sister  group  the  paronychia  are 
elongate  as  in  subgenus  Genioschizus. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


313 


Table  65.  Characters  and  character  states  used  in  phyletic  analysis  of  phylogeny  of  major 
lineages  of  Schizogenius. 


Character 

Charac 

Plesiomorphic 

ter  state 

Apomorphic 

Head 

( 1 ) Paramedian  clypeal  carinae: 

tuberculate 

not  tuberculate 

(2)  Clypeal  suture: 

shallow 

deep 

(3)  Occiput  punctation: 

uniform 

reduced  medially 

(4)  Antennae: 

moniliform 

filiform 

(5)  Arrangement  of  bristles  on 

terminal  palpal  articles: 

transverse 

longitudinal 

(6)  Mentum  lateral  lobes: 

produced 

truncate 

Pronotum 

(7)  Paralateral  sulci: 

present 

absent 

(8)  Paramedian  sulci: 

present 

obsolete 

(9)  Paralateral  carinae: 

absent 

present 

(10)  Hind  angles: 

rounded 

prominent 

Elytra 

(11)  Apex  of  lateral  channel: 

not  foveate 

foveate 

( 1 2)  Apices  of  intervals: 

not  carinate 

carinate 

(13)  Setae  of  interval  five: 

normal 

basal 

(14)  Setae  of  intervals  three, 
five,  and  seven: 

present 

absent  from 
interval  seven  (') 
interval  five  (") 
interval  three  ("') 

Legs 

(15)  Male  front  tarsi: 

narrow 

dilated 

(16)  Hind  tarsi: 

short 

long 

(17)  Paronychia: 

long 

(')  short 
(") long 

Abdomen 

(18)  Sternal  carinae: 

straight 

rounded  at  apices 

(19)  Patch  of  microsculpture  in 

coxal  area  of  sternum  three: 

present 

absent 

(20)  Female  anal  ambulatory  setae: 

present 

absent 

(21)  Female  pygidium  apex: 

entire 

crenulate 

(22)  Basal  collar  spines  of  internal 

sac  of  male  median  lobe: 

indistinct 

distinct 

314 


Whitehead 


The  strigicollis-elongatus-carinatus  lineage  developed  the  apomorphic  condition  of  strong- 
ly developed  paralateral  pronotal  carinae,  while  the  arechavaletae-truquii-capitalis  lineage 
had  as  apomorphic  the  curved  apices  of  paramedian  carinae  of  sternum  three.  Some  mem- 
bers of  the  truquii  lineage  have  weakly  developed  paralateral  pronotal  carinae,  obviously 
an  independent  apomorphy.  Members  of  the  strigicollis  lineage  have  accessory  setae  on 
submentum,  while  members  of  the  elongatus-carinatus  lineage  lack  discal  setae  on  elytron. 
Members  of  elongatus  lineage  have  shortened  paramedian  pronotal  sulci,  while  members  of 
carinatus  lineage  have  carinate  elytral  intervals.  Loss  of  discal  setae  and  reduced  pronotal 
sulci  are  also  characteristic  of  some  members  of  the  optimus  lineage,  and  in  the  quinque- 
sulcatus-tenuis  lineage  of  subgenus  Genioschizus  the  apices  of  elytral  intervals  are  carinate. 

The  ancestor  of  the  arechavaletae  lineage  lost  discal  setae  from  the  apical  half  of  interval 
seven,  and  in  the  ancestor  of  the  truquii-capitalis  lineage  the  paramedian  clypeal  carinae 
extended  to  median  clypeal  tooth.  The  ancestor  of  the  capitalis  lineage  lost  discal  setae 
from  interval  seven.  A weakness  of  this  dichotomy  is  that  I found  no  synapomorphic  mor- 
phological characteristic  of  the  truquii  lineage.  But  zoogeographic  evidence  supports  this 
dichotomy;  the  ancestor  of  the  truquii  lineage  entered  Middle  and  North  America,  while 
the  ancestor  of  the  capitalis  lineage  remained  in  South  America.  Loss  of  discal  setae  from 
interval  seven  occurred  in  ancestors  of  five  lineages:  Genioschizus,  jacarensis-optimus,  elon- 
gatus-carinatus, arechavaletae,  and  capitalis.  Discal  setae  were  lost  completely  in  ancestors 
of  the  jacarensis-optimus  and  elongatus-carinatus  lineages,  and  in  some  derived  members  of 
Genioschizus  and  the  capitalis  lineage.  In  the  ancestor  of  Genioschizus,  setae  of  interval  five 
were  restricted  to  the  base,  while  in  the  ancestor  of  the  arechavaletae  group  they  were  dis- 
tributed in  the  basal  half.  Some  members  of  the  strigicollis-elongatus-carinatus  lineage  have 
the  paramedian  clypeal  carinae  weakly  extended  to  median  tooth,  but  this  condition  is 
apparently  unstable;  in  the  truquii-capitalis  lineage  the  condition  is  synapomorphic,  stable, 
and  well  developed. 

Evident  weaknesses  in  this  reconstructed  phylogeny  concern  the  relative  positions  of  the 
Genioschizus  and  jacarensis-optimus  lineages,  and  the  ferrugineus  and  basalis  lineages.  Justi- 
fications for  my  interpretations  are  given  below.  Otherwise,  the  reconstructed  phylogeny 
appears  sound,  and  requires  no  further  discussion. 

On  first  examination,  it  is  difficult  to  decide  whether  the  crenulatus-quinquesulcatus- 
tenuis  ( Genioschizus)  lineage  or  the  jacarensis-optimus  lineage  was  the  first  to  differentiate 
from  the  rest  of  the  genus.  Members  of  the  former  are  plesiomorphic  in  most  characteristics, 
and  thus  appear  to  be  the  most  primitive  element  of  Schizogenius.  Members  of  the  jacaren- 
sis-optimus lineage  have  in  combination  numerous  apomorphic  characteristics,  and  thus 
appear  more  strongly  differentiated  than  do  members  of  the  Genioschizus  lineage.  My  inter- 
pretation is  based  on  the  following  observations.  In  the  Genioschizus  lineage  the  apomorphic 
characteristics  of  the  lateral  channel  form  a complex  not  found  in  other  carabid  beetles.  The 
one  apomorphic  characteristic  of  members  of  the  jacarensis-optimus  lineage  that  distin- 
guishes them  from  other  members  of  the  genus,  the  elongate  tibiae,  seems  a much  less  com- 
plex characteristic.  These  observations  alone  suggest  that  the  Genioschizus-Schizogenius 
dichotomy  is  the  most  likely.  An  even  more  compelling  reason,  if  the  simplest  system  of 
evolutionary  pathways  is  the  most  probable  one,  is  that  at  least  two  extra  evolutionary  steps 
are  needed  if  the  jacarensis-optimus  lineage  is  the  first  dichotomy.  These  are:  paronychia 
shortened  twice,  not  once;  and  paralateral  pronotal  sulci  lost  thrice,  not  twice.  No  evolution- 
ary steps  are  saved  if  the  jacarensis-optimus  lineage  is  regarded  as  the  sister  group  of  the  rest 
of  the  genus. 

An  extra  evolutionary  step  is  required  if  the  apomorphic  condition  of  lost  female  para- 
median ambulatory  setae  evolved  independently  in  the  ferrugineus  and  strigicollis-truquii 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


315 


lineages,  as  well  as  in  the  jacarensis-optimus  lineage.  But  the  evolutionary  significance  of  this 
character  condition  is  uncertain,  for  the  following  reasons.  Males  of  some  species  of  the 
optimus  group  lack  anal  ambulatory  setae.  Occasional  females  of  various  species  of  the 
truquii  and  capitalis  lineages  have  been  found  with  one  or  even  both  of  these  setae.  And 
females  of  S.  pluripunctatus  of  the  truquii  lineage  have  regained  both  setae  as  a clearly 
apomorphic  condition.  Future  genetic  studies  may  show  that  some  mechanism  such  as  a 
suppressor  gene  controls  the  presence  or  absence  of  female  ambulatory  setae.  If  so,  their  loss 
was  apomorphic  in  the  ancestor  of  the  subgenus,  and  they  were  regained  in  both  the  basalis 
group  and  in  S.  pluripunctatus.  Otherwise,  the  relative  positions  of  the  ferrugineus  and 
basalis  lineages  in  this  reconstructed  phylogeny  are  the  most  parsimonious.  The  following 
apomorphies  arose  once  rather  than  twice:  antennae  filiform;  female  pygidium  crenulate; 
basal  collar  spines  strongly  developed;  and  male  front  tarsi  markedly  dilated.  Of  these 
apomorphic  character  states,  the  most  stable  is  that  of  the  antennae;  all  members  of  the 
basalis-truquii  lineage  have  filiform  antennae,  though  of  variable  length.  The  other  three 
characters  do  not  appear  in  the  apomorphic  condition  in  all  members  of  the  basalis-truquii 
lineage,  but  most  probably  were  apomorphic  in  the  ancestor  of  that  lineage. 

Whatever  the  phylogeny  of  the  major  groups  of  Schizogenius  really  was,  many  character- 
istics of  external  morphology  are  convergent.  The  reconstructed  phylogeny  proposed  here 
is  the  most  parsimonious,  but  even  so  at  least  the  following  conditions  evolved  more  than 
once:  apices  of  elytral  intervals  carinate,  twice;  paralateral  pronotal  sulci  lost,  twice;  para- 
median ambulatory  setae  lost  from  female  sternum  seven,  three  times;  setae  lost  from  elytral 
interval  seven,  five  times;  setae  lost  from  apical  half  of  interval  five,  twice;  and  setae  entirely 
lost  from  interval  five,  twice.  The  apochoric  zoogeographic  characteristic,  penetration  of 
Middle  and  North  America,  happened  three  times.  The  paronychia  shortened  in  the  ancestor 
of  subgenus  Schizogenius , but  became  secondarily  elongated  in  some  of  the  more  derived 
lineages. 

Phylogeny  reconstructed  by  phenetic  techniques.  — Some  methods  of  numerical  cladistics 
were  proposed  by  Camin  and  Sokal  (1965)  for  deducing  phylogeny,  one  of  which  was  satis- 
factorily tested  for  a group  of  tiger  beetles  by  Willis  (1971).  Necessary  assumptions  are  that 
character  states  are  discrete  and  may  be  arranged  sequentially  from  primitive  to  derived, 
and  that  while  derived  character  states  may  have  arisen  repeatedly,  none  reverted  to  the 
ancestral  condition.  In  the  phylogeny  of  major  groups  of  Schizogenius  these  assumptions 
appear  valid,  except  that  the  paronychia  are  secondarily  elongate  in  some  members  of  the 
strigicollis-elongatus-carinatus-arechavaletae-truquii-capitalis  lineage.  I used  this  method  to 
test  the  logic  of  my  reconstructed  phylogeny  of  major  lineages;  as  a technique  for  deter- 
mining the  most  parsimonious  system  of  evolutionary  pathways,  it  should  produce  the  same 
result  as  my  phyletic  analysis.  In  the  phylogeny  of  species,  as  in  the  truquii  lineage,  character 
states  of  many  characters  have  evidently  reverted  to  the  ancestral  condition,  so  that  nume- 
rical cladistics,  at  least  in  their  simple  form  (Willis,  1971),  cannot  be  used. 

Coded  character  states  are  given  in  Table  66,  and  fitted  to  a data  matrix  in  Table  67  and 
a compatibility  matrix  in  Table  68.  The  character  state  of  any  given  character  is  the  one 
regarded  as  ancestral  in  the  operational  taxonomic  unit  (OTU)  concerned,  irrespective  of 
possible  specializations  in  various  derived  members  of  that  OTU.  The  secondarily  elongate 
paronychia  of  some  members  of  the  strigicollis-elongatus-carinatus-arechavaletae-truquii-ca- 
pitalis  lineage  are  coded  as  short.  Only  those  characteristics  represented  in  the  derived  char- 
acter state  by  two  or  more  OTU’s  were  used  in  this  analysis,  but  additional,  autapomor- 
phous,  characteristics  were  used  to  relate  terminal  elements  in  various  lineages.  If  two  or 
more  characteristics  produced  the  same  cladogram  pattern,  they  are  combined  and  coded 
accordingly. 


316 


Whitehead 


Table  66.  Characters  and  coded  character  states  used  in  phenetic  analysis  of  phylogeny  of 
major  lineages  of  Schizogenius. 


Character 


(1)  Elytral  lateral  channel: 

(2)  Apices  of  intervals: 

(3)  Elytral  intervals  with  discal  setae: 

(4)  Setae  on  interval  five: 

(5)  Antennae: 

(6)  Male  front  tarsi  + female  pygidium 
apex  + basal  collar  spines: 

(7)  Paralateral  pronotal  sulci: 

(8)  Anal  ambulatory  setae: 

(9)  Clypeal  suture: 

(10)  Hind  tarsi  + micro  sculpture  patch 
in  coxal  area  of  sternum  three: 

(11)  Paralateral  pronotal  carinae: 

( 1 2)  Paramedian  clypeal  carinae  + 
pronotal  hind  angles: 

(13)  Paronychia: 

(14)  Paramedian  sternal  carinae: 


Character  state 


0,  not  foveate;  1,  foveate 

0,  not  carinate;  1,  carinate 

0,  three,  five,  and  seven;  1,  three  and  five; 

2,  three;  3,  none 

0,  normal  or  absent;  1,  basal 

0,  moniliform;  1 , filiform;  2,  long 

0,  narrow  + not  crenulate  + not  developed; 

3,  dilated  + crenulate  + developed 
0,  present;  1,  absent 

0,  males  and  females;  1,  males  only 
0,  shallow;  1 deep 

0,  short  + present;  2,  elongate  + absent 
0,  absent;  1,  present 

0,  tuberculate  + rounded;  2,  not  tuberculate  + 
produced 
0,  long;  1,  short 
0,  straight;  1,  curved  at  apices 


As  seen  in  Table  68,  cladogram  patterns  one,  seven,  and  thirteen  are  the  most  parsimo- 
nious; each  requires  a minimum  of  four  extra  evolutionary  steps.  The  first  and  third  of  these 
link  all  three  species  groups  of  Genioschizus  as  a clade,  but  the  second  excludes  the  tenuis 
group.  Substudies  revealed  that  pattern  seven  is  less  parsimonious  than  patterns  one  and 
thirteen,  as  in  the  completed  phylogeny  extra  evolutionary  steps  are  required  to  account  for 
convergences  in  characteristics  one,  three,  and  four.  In  patterns  one  and  thirteen,  three 
OTU’s  cluster  on  one  branch,  and  ten  on  the  other.  Substudies  done  for  the  major  branch 
revealed  that  cladogram  pattern  ten  is  the  most  parsimonious,  with  two  OTU’s  on  one 
branch  and  eight  on  the  other.  In  the  final  cladogram  (Fig.  256),  33  evolutionary  steps  are 
needed,  12  more  than  the  21  minimum  steps  indicated  in  Table  67.  This  cladogram  is  the 
same  as  the  reconstructed  phylogeny  suggested  in  Fig.  255,  as  expected,  and  I arbitrarily 
fitted  it  to  the  same  time  scale  for  ready  comparison. 

The  truquii  lineage 

Relationships  among  species  and  species  groups  of  the  truquii  lineage  are  obscured  by  the 
lack  of  evident  and  strongly  developed  synapomorphous  characteristics.  My  reconstructed 
phylogeny  (Fig.  257)  is  provisional,  and  needs  to  be  tested  in  future  studies  by  analysis  of 
non-morphological  characteristics,  but  it  is  supported  by  zoogeographic  evidence.  Morpho- 
logical characteristics,  including  statistical  characteristics  obtained  from  Tables  8-64.  are 
listed  in  Table  69. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


317 


Table  67.  Data  matrix  for  phenetic  analysis  of  phylogeny  of  major  lineages 


OTU:  major  group  or  lineage 


Oh 

3 

O 

M 

00 

a 

»D 

a 

-C5 


(1) 

1 

1 

1 

0 

0 

0 

0 

0 

0 

0 

0 

0 

(2) 

0 

1 

1 

0 

0 

0 

0 

0 

1 

0 

0 

0 

(3) 

1 

1 

1 

2 

2 

0 

0 

0 

3 

3 

1 

0 

(4) 

1 

1 

1 

0 

0 

0 

0 

0 

0 

0 

1 

0 

(5) 

0 

0 

0 

0 

0 

0 

1 

2 

2 

1 

1 

2 

C/3 

(6) 

0 

0 

0 

0 

0 

0 

3 

3 

3 

3 

3 

3 

<L> 

o 

(7) 

0 

0 

1 

1 

1 

1 

1 

1 

1 

1 

1 

1 

cd 

(8) 

0 

0 

0 

1 

1 

1 

0 

1 

1 

1 

1 

1 

U 

(9) 

0 

0 

0 

0 

0 

1 

1 

1 

1 

1 

1 

1 

00) 

0 

0 

0 

2 

2 

0 

0 

0 

0 

0 

0 

0 

(11) 

0 

0 

0 

0 

0 

0 

0 

1 

1 

1 

0 

0 

(12) 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

2 

(13) 

0 

0 

0 

1 

1 

1 

1 

1 

1 

1 

1 

1 

(14) 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

1 

1 

of  Schizogenius. 


0 

0 

1 

0 

2 

3 

1 

1 

1 

0 

0 

2 

1 

1 


o 

C/D 

a> 

03 

C/D 

V- 

<L> 

CJ 

03 

*“H 

03 

hO 

u 


2 

2 

4 

2 

3 

4 
2 
2 
2 
3 
2 
3 
2 
2 


6 

C/D 

Oh 

a> 

C/D 

6 

3 

.g 

C 

2 


1 

1 

3 

1 

2 

3 

1 

1 

1 

2 

1 

2 

1 

1 

21 


The  truquii  lineage  includes  eight  species  groups.  Four  of  them,  the  truquii,  brevisetosus, 
sallei,  and  lineolatus  groups,  are  monobasic.  The  pluripunctatus,  tristriatus,  longipennis,  and 
depressus  groups  all  include  several  known  species  taxa.  In  the  following  discussion,  I treat 
first  the  suspected  relationships  between  groups,  and  then  the  suspected  relationships  with- 
in the  larger  groups. 

Relationships  of  species  groups.  — The  truquii  group,  the  only  group  of  the  lineage  not 
represented  north  of  the  Tropic  of  Cancer,  includes  one  living  species  from  central  Mexico. 
I suspect  the  ancestor  of  the  truquii  group  entered  Pacific  areas  of  southern  Mexico,  while 
the  ancestor  of  its  sister  group  penetrated  Atlantic  areas  in  northeastern  Mexico.  Specialized 
features  of  the  truquii  group  include  male  front  tarsus  narrowed,  female  pygidial  crenula- 
tions  lost,  and  paramedian  pronotal  sulci  extended  nearly  to  anterior  transverse  impression. 
Loss  of  female  pygidial  crenulations  was  probably  also  ancestral  in  the  brevisetosus  and 
pluripunctatus  groups,  and  in  some  species  of  the  depressus  group.  This  characteristic,  and 
the  narrowed  male  front  tarsi,  are  secondary  specializations  convergent  with  ancestral  condi- 
tions in  the  genus.  A weakness  of  this  dichotomy  is  the  lack  of  an  evident  synapomorphous 
characteristic  of  the  sister  group.  But  the  dichotomy  is  supported  by  distributional  data, 


318 


Whitehead 


Table  68.  Compatibility  matrix  for  phenetic  analysis  of  phylogeny  of  major  lineages  of 
Schizogenius. 


1 

2 

3 

4 

5 

6 

Patterns 
7 8 

9 

10 

11 

12 

13 

14 

Compatibilities 

Extra  steps 

(1) 

X 

1 

0 

0 

0 

0 

1 

0 

0 

0 

0 

0 

0 

0 

11 

2 

(2) 

1 

X 

1 

1 

1 

1 

1 

1 

1 

0 

1 

0 

1 

0 

3 

10 

(3) 

1 

3 

X 

1 

5 

2 

1 

1 

2 

2 

2 

1 

1 

1 

0 

23 

(4) 

1 

1 

0 

X 

1 

1 

1 

1 

1 

0 

0 

0 

1 

1 

4 

9 

(5) 

0 

2 

4 

1 

X 

1 

0 

1 

0 

0 

2 

9 

0 

2 

6 

14 

(6) 

0 

3 

6 

3 

3 

X 

0 

3 

0 

0 

3 

3 

0 

3 

5 

27 

(7) 

1 

1 

3 

1 

1 

1 

X 

1 

1 

1 

1 

1 

1 

1 

0 

15 

(8) 

0 

1 

3 

1 

1 

1 

0 

X 

1 

1 

1 

1 

0 

1 

3 

12 

(9) 

0 

1 

2 

1 

2 

1 

0 

0 

X 

0 

1 

1 

0 

1 

5 

10 

(10) 

0 

0 

0 

0 

0 

0 

0 

0 

0 

X 

0 

0 

0 

0 

13 

0 

(11) 

0 

1 

1 

0 

1 

0 

0 

0 

0 

0 

X 

0 

0 

0 

10 

3 

(12) 

0 

0 

2 

0 

0 

0 

0 

0 

0 

0 

0 

X 

0 

0 

12 

2 

(13) 

0 

1 

3 

1 

2 

1 

0 

1 

1 

1 

1 

1 

X 

1 

2 

14 

(14) 

0 

0 

1 

1 

1 

0 

0 

0 

0 

0 

0 

1 

0 

X 

9 

4 

Compati- 

bilities 

9 

3 

3 

4 

3 

6 

9 

6 

7 

9 

5 

5 

9 

5 

83 

— 

Extra 

steps 

4 

1 

15 

26 

11 

18 

8 

4 

9 

7 

5 

12 

11 

4 

11 

— 

145 

and  by  the  amount  of  phenetic  difference  between  members  of  the  truquii  group  and  mem- 
bers of  its  sister  group. 

Relationships  among  the  remaining  seven  species  groups  are  unclear  from  morphological 
evidence,  and  my  arrangement  is  suggested  partly  by  geographic  evidence  and  partly  by  the 
rule  of  parsimony.  Probable  sister  groups  are:  brevisetosus  and  pluripunctatus,  increased 
discal  setae;  sallei  and  tristriatus,  abdominal  microsculpture;  and  longipennis  and  depressus, 
reduced  hind  angles.  Members  of  the  brevisetosus , sallei,  and  tristriatus  groups  are  largely 
confined  to  limestone  regions,  and  their  pale  color  may  be  an  adaptation  to  that  habitat. 
Their  ancestor  was  a Texan  vicariant  of  the  Tamaulipan  ancestor  of  the  lineolatus,  longi- 
pennis, and  depressus  groups.  Ancestors  of  the  brevisetosus  and  pluripunctatus  groups  were 
Texan-Sonoran  vicariants,  and  ancestors  of  the  sallei  and  tristriatus  groups  were  Texan- 
Tamaulipan  vicariants.  If  the  ancestor  of  the  lineolatus  group  was  Texan,  the  ancestor  of  the 
longipennis  and  depressus  groups  was  Tamaulipan,  and  in  turn  the  ancestors  of  those  two 
groups  were  tropical-Tamaulipan  vicariants.  These  geographic  relationships  are  expressed  in 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


319 


Table  69.  Characters  and  character  states  in  truquii  lineage. 


Character 

Character  state 
Plesiomorphic 

Apomorphic 

(1) 

General 

Form  (mean  DP/LP): 

0.80-0.84 

(a)  0.75-0.79 

(2) 

Size  (mean  LE,  mm): 

2.50-2.95 

(b)  0.85-0.89 
(a')  2.21-2.49 

(3) 

Color: 

piceous 

(a")  under  2.21 
(b)  over  2.95 
(')  paler 

(4) 

Head 

Eye  size  (mean  WF/WH): 

0.60-0.63 

(")  piceous 
(')  0.64-0.67 

(5) 

Clypeal  field: 

narrow 

(")  0.68-0.71 
broad 

(6) 

Bases  of  frontal  carinae: 

free 

confused 

(7) 

Microsculpture  of  paramedian 
frontal  sulci: 

strong 

weak 

(8) 

Antennae: 

long 

short 

(9) 

Pro  thorax 

Form  (mean  LP/WP): 

0.89-0.96 

(a)  wider 

(10) 

Relative  size  (mean  LP/WE): 

0.75-0.80 

(b)  narrower 
(')  smaller 

(11) 

Length  of  paramedian  sulci 

0.67-0.74 

(”)  reverted 
(a)  shorter 

(12) 

(mean  PS/LP): 
Pronotal  hind  angles: 

prominent 

(b')  0.75-0.82 
(b") longer 
rounded 

(13) 

Paralateral  pronotal  carinae: 

absent 

evident 

(14) 

Accessory  marginal  setae: 

absent 

present 

(15) 

Pleura: 

impunctate 

punctate 

(16) 

Pleural  microsculpture: 

absent  or  weak 

strong 

(17) 

Elytral  setae 

Mean  number  per  elytron: 

23  or  fewer 

(')  24-34 

(18) 

Length: 

short 

(")  more 
long 

(19) 

Legs 

Tarsal  length  (mean  Ta/Ti): 

over  0.60 

under  0.60 

(20) 

Male  front  tarsi: 

dilated 

narrow 

(21) 

Front  tibia  near  base: 

narrow 

broad 

(22) 

Abdomen 

Microsculpture  of  coxal 
depressions  of  sternum  three: 

present 

absent 

(23) 

Median  microsculpture: 

absent 

present 

(24) 

Lateral  microsculpture: 

absent 

present 

(25) 

Female  pygidium  apex: 

crenulate 

not  crenulate 

320 


Whitehead 


Fig.  265-269,  and  further  discussed  under  ‘"Zoogeography”.  This  is  the  most  parsimonious 
arrangement  I can  develop,  is  consistent  with  the  premise  that  geographic  isolation  precedes 
speciation,  and  requires  the  shortest  sequence  of  branching  points.  Unfortunately,  I found 
no  synapomorphous  characteristics  to  justify  either  main  branch,  aside  from  the  pale  colora- 
tion of  members  of  the  brevisetosus,  sallei,  and  tristriatus  groups. 

Among  ancestors  of  species  groups  of  the  truquii  lineage,  my  reconstructed  phylogeny 
requires  at  least  the  following  character  convergences:  color  pale,  twice,  one  reversion;  pro- 
notum  small,  twice,  one  reversion;  paramedian  pronotal  sulci  long,  twice;  pronotal  hind 
angles  rounded,  twice;  and  female  pygidium  not  crenulate,  twice. 

Relationships  within  larger  species  groups.  — Relationships  among  species  and  subspecies 
taxa  of  the  pluripunctatus  group  are  quite  straightforward,  and  vicariant  relationships  are 
shown  in  Fig.  266.  If  the  ancestor  of  the  group  was  Sonoran,  the  ancestor  of  S.  seticollis 
and  the  ancestor  of  its  sister  group  were  Californian-Sonoran  vicariants  characterized  respec- 
tively by  accessory  pronotal  setae  and  relatively  short  antennae.  The  taxa  S.  seticollis 
seticollis  and  S.  seticollis  vandykei  diverged  from  one  another  as  north-south  vicariants.  A 
taxon  cycle  system  of  displacement  (Wilson,  1961)  would  account  for  the  general  distribu- 
tion of  members  of  the  pluripunctatus  subgroup  and  for  the  presence  of  accessory  pronotal 
setae  in  the  taxa  S.  seticollis  and  S.  plurisetosus.  It  would  not  account  for  various  similari- 
ties between  geographically  proximate  populations  of  S.  plurisetosus  and  S.  multisetosus. 
I suggest  that  the  ancestor  oi  the  taxa  S.  plurisetosus  and  S.  multisetosus  and  the  ancestor 
of  the  taxa  S.  pluripunctatus  and  S.  kulti  were  Sonoran-tropical  vicariants,  distinguished 
sharply  by  differences  in  form  of  male  median  lobe  (Fig.  131-136).  Subsequently,  S.  pluri- 
setosus and  S.  multisetosus  diverged  as  Tamaulipan-tropical  vicariants,  and  S.  pluripunctatus 
and  S.  kulti  diverged  as  Sonoran-tropical  vicariants.  The  apomorphic  condition  of  accessory 
pronotal  setae  is  convergent  in  the  taxa  S.  seticollis  and  S.  plurisetosus. 

The  tristriatus  group  is  divisible  into  four  clearly  distinguished  subgroups:  S.  tristriatus, 
tristriatus  subgroup;  S.  amphibius,  amphibius  subgroup;  S.  dilatus  and  S.  tibialis,  dilatus 
subgroup;  and  S.  planulatus,  S.  ozarkensis , and  S.  planuloides,  planulatus  subgroup.  Detailed 
relationships  among  species  taxa  of  the  planulatus  subgroup  were  discussed  above.  The^ra- 
phibius,  dilatus,  and  planulatus  subgroups  share  the  synapomorphic  characteristic  of  flat- 
tened body  form;  their  ancestor  and  the  ancestor  of  the  tristriatus  subgroup  were  Tamau- 
lipan-tropical vicariants.  Dark  coloration  is  a secondarily  acquired  characteristic  in  members 
of  the  tristriatus  subgroup,  and  relatively  enlarged  pronota  is  secondary  in  the  other  three 
subgroups.  Members  of  the  amphibius  and  planulatus  subgroups  differ  strikingly  in  size,  but 
agree  in  increased  numbers  of  elytral  setae  and  reduced  eye  size.  Their  ancestor  and  the 
ancestor  of  the  dilatus  subgroup  were  Texan-Tamaulipan  vicariants.  And,  as  judged  from  the 
distribution  of  living  species,  the  ancestors  of  the  amphibius  and  planulatus  subgroups  were 
temperate-Texan  vicariants.  Ancestors  of  the  taxa  S.  dilatus  and  S.  tibialis  were  clearly 
Tamaulipan-tropical  vicariants,  but  some  forms  of  S.  tibialis  reentered  the  Tamaulipan  area 
from  the  west,  and  even  entered  the  Texan  area  (Fig.  188).  These  vicariant  relationships  are 
shown  in  Fig.  267. 

Among  taxa  included  in  the  longipennis  group,  S.  longipennis  and  S.  neovalidus  are 
clearly  sister  species  linked  by  the  synapomorphous  condition  of  enlarged  virga  in  the  male 
endophallus,  and  S.  pacificus  with  elongate  tarsi  and  convex  body  is  the  most  divergent 
form.  My  views  on  vicariant  relationships  are  expressed  in  Fig.  268.  Three  of  the  four 
included  species  are  represented  in  Arizona,  two  apparently  as  relicts.  One  might  therefore 
suspect  that  ancestors  of  the  depressus  and  longipennis  groups  were  Tamaulipan-Sonoran 
vicariants,  rather  than  Tamaulipan-tropical.  But  no  members  of  the  group  are  known  either 
from  Texas  or  the  Mexican  Central  Plateau,  and,  further,  if  the  ancestor  of  the  group  was 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


321 


Sonoran  one  could  not  convincingly  explain  the  presence  of  the  Arizona  relicts.  I think  a 
taxon  cycle  system  of  displacement  (Wilson,  1961)  was  responsible  for  present  distributions. 
The  ancestor  of  the  group  spread  across  tropical  regions  of  southern  Mexico;  the  ancestor  of 
S.  pacificus  and  the  ancestor  of  its  sister  group  were  Pacific-Atlantic  vicariants.  When  the 
ancestor  of  that  sister  group  again  spread  to  the  Pacific  slopes,  S.  pacificus  was  well  differ- 
entiated, and  not  subject  to  displacement.  The  ancestor  of  S.  chiricahuanus  and  the  ancestor 
of  its  sister  group  were  Sonoran-tropical  vicariants,  but  the  range  of  S.  chiricahuanus  re- 
cessed when  its  sister  group  entered  the  Sonoran  region.  The  ancestors  of  S.  neovalidus  and 
S.  longipennis  were  again  Sonoran-tropical  vicariants,  and  the  range  of  S.  neovalidus  recessed 
when  S.  longipennis  entered  the  Sonoran  region. 

The  aggregate  range  of  members  of  the  depressus  group  is  nearly  as  great  as  that  of  the  en- 
tire genus.  Thus  evolutionary  relationships  are  difficult  to  interpret,  and  my  analysis  of  vicar- 
iant  relationships  (Fig.  269)  is  difficult  to  justify.  For  purposes  of  this  discussion,  I recognize 
three  subgroups:  S.  arimao,  S.  emdeni,  and  S.  “apicalis”,  arimao  subgroup;  S.  sulcifrons  and 
S.  litigiosus,  sulcifrons  subgroup;  and  S.  pygmaeus,  S.  scopaeus,  S.  falli,  S.  ochthocephalus, 
and  S.  depressus,  depressus  subgroup.  Members  of  the  arimao  subgroup  agree  in  the  synapo- 
morphous  characteristic  of  shortened  pronotal  sulci,  also  characteristic  of  members  of  the 
pluripunctatus  and  longipennis  groups.  I found  no  clear  synapomorphous  characteristic  of  its 
sister  group,  but  suspect  the  ancestor  of  that  group  had  slightly  increased  numbers  of  elytral 
setae.  Ancestors  of  the  arimao  subgroup  and  its  sister  group  were  probably  tropical-Tama- 
ulipan-Texan  vicariants.  I suspect  an  ancestral  condition  in  the  arimao  subgroup  was  the 
presence  of  weakly  developed  paralateral  pronotal  carinae,  subsequently  lost  in  S.  emdeni. 
The  ancestor  of  S.  arimao  may  have  entered  Cuba  via  a Middle  American  land  bridge,  and 
its  ancestor  and  the  ancestor  of  its  sister  group  thus  may  have  been  Cuban-Middle  American 
vicariants.  Whether  the  ancestors  of  S.  emdeni  and  S.  “ apicalis  ” were  early  Middle  Ameri- 
can-South American  vicariants,  or  whether  they  diverged  in  Middle  America,  is  uncertain. 

Chorological  data  suggest  that  the  ancestor  of  the  sulcifrons  and  depressus  subgroups  was 
Texan,  and  that  their  respective  ancestors  were  temperate-Texan  vicariants.  The  ancestor 
of  the  latter  subgroup  was  characterized  by  small  size  and  pale  color.  The  ancestor  of  the 
sulcifrons  subgroup  diverged  into  cool-adapted  western  and  eastern  vicariants  during  Pleisto- 
cene time.  The  common  ancestor  of  S.  falli,  S.  ochthocephalus,  and  S.  depressus  and  the 
common  ancestor  of  S.  pygmaeus  and  S.  scopaeus  were  Sonoran-Texan  vicariants,  as  sug- 
gested particularly  by  the  pattern  of  geographic  variation  in  S.  falli.  The  ancestor  of  S.  falli , 
S.  ochthocephalus,  and  S.  depressus  was  characterized  by  reduced  eye  size  and  flattened 
body,  while  the  ancestor  of  S.  scopaeus  and  S.  pygmaeus  was  characterized  by  reduced  body 
size.  The  ancestor  of  S.  depressus  and  the  ancestor  of  S.  falli  and  S.  ochthocephalus  diverged 
as  temperate-Sonoran  vicariants,  and  the  respective  ancestors  of  S.  falli  and  S.  ochthocepha- 
lus diverged  as  Sonoran-Californian  vicariants.  Simple  vicariance  does  not,  however,  explain 
the  evolutionary  history  of  the  taxa  S.  scopaeus  and  S.  pygmaeus.  Their  distribution  patterns 
are  best  explained  by  an  early  split  into  Texan-Tamaulipan+tropical  vicariants,  a later  split 
of  the  Texan  form  into  Texan-Sonoran  vicariants,  and  a still  later  reestablishment  of  repro- 
ductive continuity  between  the  tropical  and  Sonoran  vicariants. 

The  weakest  parts  of  this  reconstructed  phylogeny  (Fig.  257)  are  in  the  relationships  of 
the  lineolatus,  longipennis , and  depressus  groups,  and  in  relationships  within  the  depressus 
group.  In  particular,  present  distributions  of  all  members  of  both  the  arimao  and  sulcifrons 
subgroups  are  far  removed  from  both  the  Texan  and  Tamaulipan  regions,  so  that  my  sug- 
gested vicariant  relationships  are  not  well  justified.  In  general,  however,  the  suggested  vicar- 
iant  relationships  lend  support  to  the  reconstructed  phylogeny,  or  at  least  do  not  contradict 
it,  and  they  seem  to  satisfactorily  explain  known  distributions  of  modern  species  and  sub- 


322 


Whitehead 


species  taxa.  I think  they  suggest  the  most  probable  of  an  astronomical  number  of  possible 
evolutionary  pathways. 

Other  groups  and  lineages 

Hypothetical  relationships  among  species  and  subspecies  taxa  of  the  genus  Schizogenius, 
excluding  members  of  the  truquii  lineage,  are  shown  in  Fig.  258.  Most  of  these  taxa  are 
South  American,  and  as  the  South  American  fauna  is  poorly  known  this  reconstructed  phy- 
logeny  is  incomplete,  and  my  comments  on  it  are  brief. 

Among  the  three  included  taxa  of  the  quinquesulcatus  group,  S.  szekessyi  and  S.  janae 
share  maculate  coloration.  They  are  at  most  sister  species,  and  may  not  even  be  biologically 
distinct  entities. 

Within  the  tenuis  group,  S.  impressicollis  is  divergent  from  other  taxa  in  form  of  male 
genitalia,  but  plesiomorphic  in  having  parallel  rather  than  convergent  paramedian  frontal 
carinae.  Relationships  among  other  taxa  in  the  tenuis  group  are  unclear,  and  cannot  be  prop- 
erly interpreted  until  the  South  American  fauna  is  better  known.  I think  S.  sculptilis  is  least 
closely  related,  as  judged  by  amount  of  phenetic  difference;  in  particular,  I doubt  that  it 
shares  a direct  common  ancestry  with  S.  tenuis.  My  phylogeny  suggests  a possible  pre-Pleis- 
tocene  entry  into  Middle  America  for  the  ancestor  of  S.  sculptilis,  but  there  may  well  be  an 
as  yet  unknown  sister  species  in  South  America.  More  probably,  the  ancestor  of  S.  sculptilis 
entered  Middle  America  in  Early  Pleistocene,  and  S.  tenuis  followed  in  Late  Pleistocene. 

The  ancestor  of  the  optimus  group  gave  rise  to  one  line  with  shortened  pronotal  sulci,  and 
another  with  discal  setae  lost:  respectively,  S.  optimus,  S.  dyschirioides,  and  S.  clivinoides ; 
and  S.  bicolor  and  S.  grossus.  S.  optimus  and  S.  dyschirioides  are  evidently  sister  species, 
and  the  ancestor  of  S.  optimus  was  most  probably  an  Early  Pleistocene  entrant  into  Middle 
America. 

Among  the  four  included  taxa  of  the  basalis  group,  S.  basalis  and  S.  cearaensis  seem  most 
closely  related  to  one  another,  and  together  with  S.  negrei  agree  in  having  reduced  occiput 
punctation. 

Aside  from  the  truquii  lineage,  the  most  diverse  lineage  in  Schizogenius  is  the  capitalis 
lineage,  with  five  included  species  groups.  Similarities  among  some  members  of  the  darling- 
toni  and  ocellatus  group  in  size,  color,  and  ventral  microsculpture  suggest  possible  relation- 
ships, but  radical  specializations  of  members  of  the  ocellatus  group  imply  considerable 
antiquity  for  the  common  ancestor  of  these  two  groups.  Within  the  darlingtoni  group,  the 
taxa  S.  darlingtoni  and  S.  interstriatus  agree  in  having  extensive  abdominal  microsculpture. 
Of  the  remaining  three  species  groups,  the  quadripunctatus  group  is  the  most  divergent  and 
hence  probably  the  oldest.  Loss  of  discal  setae  from  all  but  elytral  interval  three  in  this 
and  the  ocellatus  group  is  probably  convergent.  The  ancestor  of  the  lindrothi  group  had 
reduced  pronotal  hind  angles  and  shortened  antennae,  as  in  some  groups  of  the  truquii 
lineage.  Ancestors  of  S.  lindrothi  and  S.  banningeri  were  probably  Early  Pleistocene  Middle 
American-South  American  vicariants. 

This  reconstructed  phylogeny  suggests  the  following  Middle  and  South  American  faunal 
exchanges.  Ancestors  of  the  crenulatus  group,  the  ferrugineus  group,  and  the  truquii  lineage 
entered  Middle  America  well  before  Pleistocene  time.  Early  Pleistocene  Middle  American 
immigrants  included  ancestors  of  S.  sculptilis,  S.  optimus,  and  S.  lindrothi;  the  ancestor  of 
S.  “apicalis”  was  a probable  Early  Pleistocene  immigrant.  During  Late  Pleistocene,  S.  tenuis 
spread  into  Middle  America,  and  S.  pygmaeus  spread  into  South  America.  Thus,  three  south- 
north  migrations  occurred  over  water,  three  south-north  migrations  and  one  north-south 
migration  occurred  soon  after  establishment  of  land  connections,  and  one  north-south  mi- 
gration and  one  south-north  migration  occurred  comparatively  recently. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


323 


crenulatus 

quinquesulcat  us 

ten  uis 

jacarensis 

optimus 

f errugineus 

basalis 

Strigicollis 

elongatus 

carinatus 

arechavaletae 

truquii 

capitalis 

255 


crenulatus 

quinquesulcatus 

tenuis 

jacarensis 

optimus 

f errugineus 

basalis 

strigicollis 

elongatus 

carinatus 

arechavaletae 

truquii 

capitalis 

256 


g.  255  256.  Phylogeny  of  major  groups  and  lineages  of  genus  Schizogenius;  crenulatus,  quinquesulcatus,  and  tenuis 
groups  are  m subgenus  Gemoschizus.  255.  Phylogeny  reconstructed  by  phyletic  techniques.  256.  Phylogeny  reconstructed 
by  phenetic  techniques. 


324 


Whitehead 


S.truquii 

S brevisetosus 

S s seticollis 
S s vandykei 
S plur ipunctatus 
S.kulti 

S plurisetosus 
S.multisetosus 

S sallei 

S tristriat us 
S dilatus 
S. tibialis 
S amphibius 
S planuloides 
S ozarkensis 
S planulatus 

S lineolatus 

S pacificus 
S chiricahuanus 
S longipennis 
S neovalidus 

S arimao 
S .emdeni 
S "apicalis" 

S sulcifrons 
S.litigiosus 
S pygmaeus 
S scopaeus 
S d«pressus 
S . f alii 

S ochthocephalus 

257 


Fig.  257.  Reconstructed  phylogeny  of  truquii  lineage  of  genus  Schizogenius. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


325 


I l l l 


i 


i 


S c crenulatus 
Sc  chiapatecus 

SquinqueSulcatus 
S szekessy i 
S janae 

S.impressicoilis 
S.  s c ulptilis 
S.suturalis 
S.maculatus 
S impuncticollis 
S. tenuis 

S jacarensis 

S optimus 
S.dyschirioides 
S.clivinoides 
S bicolor 
S grossus 

S f errugineus 
S auripennis 

S.multipunctatus 
S negrei 
S basalis 
S cearaensis 

S.strigicollis 

S.elonqatus 

S.costiceps 

S carinatus 
S.  cost  ipennis 

S.arechavaietae 
S reichardti 

S.capitalis 

S.putzeysi 

S.lindrothi 
S bonningeri 

S.quadr  ipunctat us 

S.darlingtoni 
S.  interstriatus 
S.riparius 

S.ocellatus 

S.truquii 


MILLIONS  OF  YEARS  BEFORE  PRESENT 


6 3 


258 


Fig.  258.  Reconstructed  phylogeny  of  subspecies  and  species  taxa  of  genus  Schizogenius,  excluding  components  of 
truquii  lineage. 


326 


Whitehead 


ZOOGEOGRAPHY 

Introduction  and  general  patterns  of  distribution 

Introduction 

Various  classes  of  clues  available  to  help  assess  past  movements  of  animals  include  number 
clues,  extent  of  area,  continuity  of  area,  degree  of  differentiation,  and  vicariance  (Darling- 
ton, 1957;  Erwin,  1970).  In  my  study  of  Schizogenius , I found  number  clues  and  vicariance 
clues  especially  useful. 

I base  much  of  my  argumentation  of  Maldonado-Koerdell’s  ( 1964)  summary  of  historical 
geography  of  Middle  America.  His  views  on  locations  and  times  of  opening  and  closing  of 
water  gaps,  and  of  land  connections  with  major  West  Indian  islands,  are  hotly  disputed; 
I here  attempt  to  reconcile  various  divergent  views. 

Biogeographers,  no  doubt  correctly,  dismiss  direct  land  connections  between  North  and 
South  America  before  Middle  to  Late  Tertiary  as  inconsistent  with  geologic  and  zoogeo- 
graphic facts.  Yet  the  possibility  of  indirect  connections  via  islandic  chains,  i.e.,  incomplete 
or  only  partially  effective  barriers  to  dispersal,  existing  through  Paleocene  should  not  be 
lightly  dismissed.  As  Patterson  and  Pascual  (1968)  state,  ancestral  forms  of  at  least  four 
mammalian  lineages  appeared  in  South  America  in  rapid  succession  about  the  Cretaceous- 
Tertiary  boundary,  while  subsequently,  during  the  first  half  of  Tertiary,  ancestors  of  perhaps 
only  three  more  mammalian  lineages  did  so.  Henceforth,  when  I speak  of  pre-Eocene  land 
bridges  or  land  connections,  I do  so  of  this  probable,  incomplete,  bridge.  The  geological 
history  of  not  only  Middle  America  but  of  the  whole  circum-Caribbean  land  mass  requires 
further  study. 

Patterson  and  Pascual  (1968)  doubt  that  land  bridges  from  Middle  America  to  Cuba  ever 
existed,  and  doubt  further  that  Middle  America  was  divided  into  smaller  land  masses  by 
water  portals  during  Middle  Tertiary.  Hershkovitz  (1969)  suspects  the  existence  of  both. 
For  riparian  species  of  Schizogenius,  water  gaps  such  as  the  supposed  Tehuantepec  portal 
are  not  required  for  barriers  to  exist.  Had  this  region  been  emergent  but  of  sufficiently  low 
relief  that  riparian  gravel  bars  did  not  exist,  it  would  have  been  perfectly  effective  as  a bar- 
rier. And  this  would  be  so  not  only  for  riparian  Schizogenius,  but  for  upland  terrestrial 
animals  of  many  kinds.  The  same  argument,  however,  could  well  apply  to  the  question  of 
Middle  American-Cuban  land  connections;  probably,  if  such  land  connections  did  exist, 
they  were  not  of  high  relief.  I accept  Maldonado-Koerdell’s  (1964)  postulations  of  these 
land  bridges  in  part  for  consistency  and  in  part  for  convenience,  but  without  specific  justi- 
fication. A land  bridge  to  Cuba  is  useful  to  explain  the  distribution  of  one,  and  only  one, 
species  of  Schizogenius’,  it  clearly  is  not  a requirement. 

Hershkovitz  (1969)  indicates  that  the  Panamanian  portal  closed  in  Late  Miocene-Early 
Pliocene  rather  than  Late  Pliocene-Early  Pleistocene,  and  surely  is  supported  in  this 
assertion  by  evidence  from  distributions  of  living  mammal  species.  Again,  however,  effect- 
ive land  connections  for  riparian  Schizogenius  species  require  sufficient  uplifting  for  form- 
ation of  riparian  habitats.  I am  satisfied  that  these  conditions  did  not  exist  before  Late 
Pliocene. 

General  patterns  of  distribution 

The  genus  Schizogenius  is  a sister  group  of  New  World  Halocoryza ; the  ancestor  of  these 
organisms  diverged  from  an  African  ancestor,  and  all  other  related  genera  are  of  Old  World 
origin.  The  genus  Schizogenius  is  best  developed  and  most  diverse  in  South  America, 
and  only  three  old  lineages  are  restricted  to  or  diversified  in  North  and  Middle  America. 
The  ancestor  of  the  genus  was  therefore  clearly  South  American;  and  it  must  have  differ- 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


327 


entiated  from  Halocoryza  stock  no  earlier  than  Eocene,  after  North  and  South  America 
became  well  isolated  by  truly  effective  Middle  American  water  barriers.  One  major  water 
gap  was  across  the  Isthmus  of  Tehuantepec,  and  another  across  Panama,  closing  respect- 
ively in  Late  Miocene  and  Late  Pliocene  (Maldonado-Koerdell,  1964).  These  gaps  approxi- 
mately defined  the  area  I recognize  as  Middle  America:  in  the  south  bounded  approxi- 
mately by  the  Panama-Costa  Rica  border,  and  in  the  north  including  Pacific  drainages 
of  southeastern  Chiapas  but  excluding  the  Usumacinta  and  Grijalva  Rivers  in  Guatemala. 
Known  southern  limits  of  two  Schizogenius  species,  S.  tenuis  and  S.  pygmaeus,  approxi- 
mate southern  limits  for  Middle  America  as  defined  by  Hershkovitz  (1969).  The  reality 
and  significance  of  this  geographic  boundary  as  it  pertains  to  members  of  Schizogenius 
cannot  be  further  assessed  at  present. 

Of  23  species  groups  in  Schizogenius , 10  evolved  in  North  America  and  13  in  South 
America.  North  American  species  groups  arose  from  three  early  migrations  from  South 
America.  One  early  immigrant  gave  rise  to  the  crenulatus  group,  with  one  subspecies  ranging 
into  northern  Middle  America.  A second  early  immigrant  was  the  ancestor  of  the  ferrugineus 
group,  one  species  of  which  extends  from  North  America  to  the  southern  limit  of  Middle 
America.  The  third  immigrant  gave  rise  to  eight  species  groups,  all  centered  in  North 
America.  One  species  of  the  pluripunctatus  group  enters  the  northern  part  of  Middle  Ameri- 
ca. One  species  of  the  tristriatus  group  extends  to  the  southern  limit  of  North  America.  One 
species  of  the  longipennis  group  extends  to  the  southern  limit  of  North  America,  and 
another  to  the  southern  limit  of  Middle  America.  And  in  the  depressus  group,  three  endemic 
Middle  American,  Cuban,  and  South  American  species  form  a monophyletic  subgroup  of 
old  Middle  American  ancestry,  while  another  extends  to  northern  South  America  from 
North  America.  I judge  that  the  endemic  South  American  form  arrived  there  about  Late 
Pliocene-Early  Pleistocene,  and  that  it  may  have  an  as  yet  unknown  vicariant  sister  species 
in  Middle  America. 

Six  of  the  13  South  American  groups  are  not  known  from  Panama,  Colombia,  or  Vene- 
zuela. Some  members  of  the  capitalis,  darlingtoni,  quinquesulcatus,  and  strigicollis  groups 
are  known  from  this  region  but  not  from  Middle  America.  The  lindrothi  and  optimus  groups 
include  vicariant  Middle  and  South  American  sister  species,  but  except  for  Floridian  popu- 
lations of  S.  lindrothi  are  otherwise  unrepresented  in  North  and  Middle  America.  Only  the 
tenuis  group,  with  one  species  endemic  in  southern  North  America  and  another  extending 
through  Middle  America  to  northern  Mexico,  is  represented  north  of  the  Isthmus  of  Tehuan- 
tepec. The  endemic  species,  S.  sculptilis,  probably  also  occurs  in  Middle  America;  its  ances- 
tor probably  arrived  there  soon  after  re-establishment  of  land  connections,  and  it  may  have 
an  as  yet  unknown  vicariant  sister  species  in  South  America. 

Middle  America  serves  as  a funnel  (Simpson,  1950)  between  North  and  South  America 
for  Schizogenius  species,  and  also  shows  a notable  filter  effect  at  both  ends.  Tropical  insect 
faunas  of  Mexico  and  Middle  America  derive  primarily  from  South  American  ancestors 
(Halffter,  1964),  but  for  Schizogenius  Halffter’s  notions  about  timing  of  immigrations  are 
inadmissible.  If  the  genus  originated  in  South  America  after  Eocene,  the  first  wave  of  migra- 
tions northward  was  not  earlier  than  mid-Tertiary.  After  closure  of  the  Panamanian  portal 
early  migrants  evolved  as  endemic  Middle  American  species.  Later  migrants,  S.  tenuis  and 
S.  pygmaeus,  were  more  probably  Late  Pleistocene  than  Late  Pliocene.  Despite  evidence  for 
two-way  mammalian  interchanges  between  Middle  and  South  America  since  Late  Miocene 
(Hershkovitz,  1966),  no  evidence  of  free  interchange  before  Late  Pliocene  exists  for  Schizo- 
genius species. 

The  Middle  American  Schizogenius  fauna  is  poorly  known,  but  probably  not  so  poorly 
as  suggested  by  the  limited  numbers  of  known  species.  Among  Mexican  species  not  yet 


328 


Whitehead 


recorded  from  Middle  America,  only  S.  sculptilis  is  really  likely  to  be  found  there.  Also 
expected  is  a sister  species  of  the  South  American  S.  “apicalis”,  and  perhaps  other  South 
American  species  enter  southern  parts  of  the  area.  Of  10  species  known  from  the  region, 
five  enter  from  North  America,  one  from  South  America,  and  four  are  endemic;  two 
endemics  have  South  American  sister  species,  while  the  others  differentiated  there  from  an 
earlier  penetration  from  North  America.  Through  much  of  Middle  Tertiary,  Middle  America 
was  an  isolated,  often  partitioned  land  mass,  and  despite  probable  continuity  with  Cuba 
during  much  of  this  period  was  relatively  small  in  area  (Maldonado-Koerdell,  1964).  Al- 
though the  Tehuantepec  portal  closed  in  Late  Miocene,  the  Isthmus  was  not  elevated  until 
Late  Pliocene;  the  Isthmus  may  have  been  a continuously  significant  barrier  since  Early 
Oligocene.  And,  since  Early  Miocene  and  particularly  since  Late  Pliocene,  Middle  America 
suffered  much  volcanic  activity.  These  observations  suggest  that  the  Schizogenius  fauna  of 
Middle  America  may  never  have  been  really  large.  Support  for  this  conclusion  is  evident 
from  known  distributions  of  species  in  the  area.  Aside  from  one  species  endemic  to  Cuba, 
all  species  except  S.  crenulatus  and  S.  kulti  are  known  to  be  widely  distributed  from  north 
to  south,  suggesting  lack  of  barriers  and  consequent  reduced  geographic  isolation  and  species 
diversification.  Thus,  while  mid-Tertiary  fracturing  of  Middle  America  may  have  contributed 
greatly  to  speciation  in  Trichoptera  (Ross,  1967),  it  had  no  apparent  effect  on  speciation 
in  Schizogenius . After  closure  of  the  Tehuantepec  portal  in  Late  Miocene,  some  North 
American  elements  penetrated  the  area,  and  may  have  replaced  some  endemic  forms. 
Further  elimination  of  endemic  forms  may  have  resulted  from  arrival  of  South  American 
immigrants  after  closure  of  the  Panamanian  portal  in  Late  Pliocene. 

Distribution  patterns  in  North  and  Middle  America 

Introduction 

The  genus  Schizogenius  is  represented  throughout  North  and  Middle  America,  south  of 
52°N  in  the  west  and  48°N  in  the  east,  except  in  the  West  Indies  where  known  only  from 
Cuba  and  Jamaica.  Since  this  fauna  originated  in  South  America,  most  included  species  and 
subspecies  are  tropical  or  subtropical,  and  relatively  few  have  acquired  truly  temperate  adap- 
tations. Their  distribution  patterns  were  regulated  by  the  development  of  Middle  and  Late 
Tertiary  climatic  and  physiographic  features,  and  were  modified  particularly  in  the  north  by 
Pleistocene  events.  I use  number  clues  to  evaluate  distribution  patterns,  generally  according 
to  techniques  developed  by  Ball  and  Freitag  (in  Freitag,  1969)  and  Erwin  (1970),  and  com- 
pare my  observations  with  some  of  their  observations  on  Evarthrus  and  Brachinus,  respec- 
tively. Then  I consider  vicariance  clues,  particularly  in  relation  to  the  truquii  lineage,  and 
attempt  to  reconstruct  the  historical  zoogeography  of  the  genus  in  North  and  Middle 
America. 

Methods  and  general  patterns 

Following  Erwin’s  (1970)  methods,  I show  on  a 5°  longitudinal  and  latitudinal  grid  map 
the  number  of  species  or  subspecies  known  to  occur  in  each  interval  (Fig.  261),  and  list 
“total  interval  values”  (TIV)  and  “average  landmass  interval  values”  (ALIV)  in  Table  70. 
The  number  of  species  is  maximum  in  southern  and  central  Mexico,  and  slightly  less  in 
Arizona.  Numbers  decrease  rapidly  in  all  directions  from  these  centers.  To  the  north,  num- 
bers decrease  least  rapidly  in  the  humid  forested  regions  of  the  Appalachians  and  the  Pacific 
coast.  To  the  south,  numbers  decrease  sharply  near  the  Panama-Costa  Rica  border.  And  the 
Schizogenius  fauna  of  Florida  and  the  West  Indies  is  notably  depauperate.  The  general 
pattern  is  similar  to  that  for  Brachinus , except  for  absence  of  an  evident  east-west  lateral 
asymmetry  and  reduced  numbers  in  extreme  southeastern  United  States. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


329 


These  general  patterns  are  readily  explained.  Few  species  are  cool-adapted,  and  none  are 
cold-adapted;  hence,  reduction  to  the  north,  particularly  in  central  North  America  and  in 
the  Rocky  Mountain  region.  Most  species  inhabit  riparian  gravel  bars;  hence  reduction  in 
extreme  southeastern  United  States,  and  lack  of  east-west  asymmetry  in  the  United  States. 
The  reduction  of  numbers  in  Middle  America  was  considered  previously. 

Maximum  linear  ranges  (Erwin,  1970)  are  given  in  Table  71,  and  compared  with  data  on 
linear  ranges  for  Brachinus  species  (Erwin,  1970)  and  Evarthrus  species  (Ball  and  Freitag,  in 
Freitag,  1969).  Maximum  ranges  of  Schizogenius  species  are  almost  exactly  intermediate 
between  those  of  Brachinus  and  Evarthrus ; average  maximum  distances  between  localities 
were  determined  as  about  700  miles  for  Evarthrus,  1000  miles  for  Schizogenius,  and  1300 
miles  for  Brachinus.  One  of  the  Schizogenius  species,  S.  pygmaeus,  has  a linear  distribution 
of  more  than  1000  miles  greater  than  any  species  of  Evarthrus  or  Brachinus. 

If  barriers  limit  distributions  of  Evarthrus  species,  and  broad  climatic  zones  limit  distri- 
butions of  Brachinus  species,  barriers  and  climatic  zones  together  may  account  for  distri- 
butions of  Schizogenius  species.  Barriers,  for  inhabitants  of  riparian  gravel  bars,  include  the 
following:  high  altitudes;  rivers  or  sections  of  rivers  without  gravel  bars;  and  land  or  water 
gaps  between  rivers.  Limiting  climatic  zones  correspond  roughly  to  tropical,  subtropical, 
warm  temperate,  and  cool  temperate.  Another  limiting  factor  in  distributions  of  some 
Schizogenius  species  and  subspecies  is  almost  certainly  one  of  ecological  displacement.  In 
North  and  Middle  America,  at  least,  no  more  than  seven  species  may  be  taken  in  gravel  bars 
at  a single  locality,  and  normally  no  more  than  five.  Ecological  displacement  is  particularly 
obvious  in  some  allopatric  sister  species,  which  inhabit  adjacent  river  systems  or  even 
separate  segments  of  the  same  river  system. 

An  example  of  a mountain  barrier  is  the  Sierra  Madre  Occidental.  Species  found  on  both 
sides  of  this  range,  such  as  S.  pluripunctatus  and  S.  pygmaeus,  evidently  lack  gene  flow 
across  the  mountains.  Others,  such  as  S.  depressus  to  the  east  and  S.  longipennis  to  the 
west,  have  ranges  limited  by  the  mountains.  Lack  of  suitable  habitat  along  portions  of  the 
Red  and  Mississippi  Rivers  probably  accounts  for  limits  of  ranges  in  S.  planuloides,  S.  ozark- 
ensis,  and  S.  planulatus.  And  the  apparent  isolation  of  S.  s.  seticollis  and  S.  s.  vandykei  is 
probably  due  to  a large  gap  between  suitable  habitats  in  northern  Baja  California. 

Climatic  zones  are  not  sharply  limited,  but  nonetheless  seem  at  least  partly  responsible 
for  range  limitations.  Of  13  species  groups  represented  in  North  and  Middle  America,  in- 
cluding representatives  of  three  South  American  groups,  all  are  represented  in  tropical  or 
subtropical  regions,  seven  in  warm  temperate  regions,  and  six  in  cool  temperate  regions. 
Numerous  species  or  subspecies  have  ranges  limited  near  the  Tropic  of  Cancer;  12  species  or 
subspecies  have  southern  or  northern  limits  within  31/2°S,  and  eight  within  31/2°N. 

Some  examples  of  allopatric  sister  taxa  found  in  adjacent  drainage  systems  are:  S.  kulti 
and  S.  pluripunctatus,  Rio  Grande  de  Santiago  and  Rio  Acaponeta;  S.  multisetosus'  and 
S.  plurisetosus , Rio  Panuco  and  Rio  Tamesi;  and  S.  c.  crenulatus  and  S.  c.  chiapatecus, 
Rio  Grande  de  Santiago  and  Rio  Ameca.  Adjacent  river  systems  may  also  limit  sharply 
differentiated  forms  within  a taxon,  as  in  S.  tibialis : large  eyed  forms  occur  from  the  Rio 
Panuco  southward,  moderate  eyed  forms  from  the  Rio  Panuco  to  the  Rio  Grande,  and  small 
eyed  forms  in  the  Nueces  River  basin.  An  example  of  displacement  on  a single  river  system 
is  the  occurrence  of  S.  pygmaeus  on  the  Rio  Conchos,  rather  than  S.  scopaeus  which  occurs 
elsewhere  along  the  Rio  Grande.  Somewhat  less  closely  related  taxa  may  also  be  subject  to 
displacement,  and  an  example  may  be  the  displacement  of  S.  dilatus  south  of  the  Rio 
Tamesi  by  S.  tristriatus  or  large  eyed  S.  tibialis  and  north  of  the  Rio  Sabinas  Hidalgo  by 
S.  sallei  or  S.  planuloides.  Faunal  compositions  along  river  systems  may  vary  from  down- 
stream to  upstream.  The  taxa  S.  multisetosus  and  S.  kulti  are  known  from  the  same  river 


330 


Whitehead 


Table  70.  Total  number  of  species  and  subspecies,  “average  landmass  interval  values” 
(ALIV),  and  “total  interval  values”  (TIV)  derived  from  Fig.  261. 


Interval 

No.  spp. 

ALIV 

TIV 

Interval 

No.  spp. 

ALIV 

TIV 

A 

r ~ 

— 

_ 

a 

1 

1.0 

1 

B 

5 

4.8 

12 

b 

4 

1.7 

10 

C 

5 

3.1 

14 

c 

10 

3.1 

33 

D 

11 

4.9 

22 

d 

14 

4.7 

45 

E 

16 

6.1 

32 

e 

20 

7.7 

48 

F 

20 

6.6 

43 

f 

19 

9.1 

32 

G 

22 

9.2 

46 

g 

20 

16.9 

38 

H 

17 

6.9 

26 

h 

17 

15.2 

38 

I 

13 

5.8 

23 

i 

6 

12.0 

12 

J 

13 

5.7 

23 

j 

7 

10.6 

8 

K 

7 

4.4 

13 

k 

1 

1.3 

1 

L 

6 

4.4 

10 

1 

- 

- 

- 

Table  71.  Frequency  distribution  of  maximum  linear  range  in  miles  of  species  of  Brachinus* , 
species  of  Evarthrus**,  and  species  and  subspecies  of  Schizogenius  in  North  and  Middle 
America. 


Class 

Brachinus 
No.  % 

Evarthrus 
No.  % 

Schizogenius 
No.  % 

3501-3750 

0 

0 

1 

3251-3500 

0 

0 

0 

0 

0 

3 

3001-3250 

0 

0 

0 

2751-3000 

0 

0 

1 

2501-2750 

5 

19 

0 

5 

2 

13 

2251-2500 

5 

0 

2 

2001-2250 

2 

2 

0 

1751-2000 

9 

1 

1 

1501-1750 

2 

39 

3 

16 

1 

24 

1251-1500 

8 

1 

1 

1001-1250 

5 

2 

6 

751-1000 

11 

8 

5 

501-  750 

4 

42 

5 

79 

5 

60 

251-  500 

3 

10 

6 

0-  250 

8 

12 

7 

* From  Erwin  (1970) 

**From  Ball  and  Freitag  (in  Freitag,  1969) 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


331 


systems  but  not  from  the  same  localities;  when  both  occur  on  a drainage  system,  & multi- 
setosus  is  found  upstream  from  S.  kulti.  Along  the  Rio  Grande,  seven  species  are  known 
from  the  Del  Rio  region,  three  of  them  from  the  Big  Bend  region,  and  one  of  them  plus 
another  from  northern  New  Mexico.  Part  of  these  faunal  changes  are  probably  due  to  lack 
of  suitable  habitat,  as  in  the  Las  Cruces-El  Paso  segment  of  the  Rio  Grande. 

Techniques  described  by  Erwin  (1970)  for  finding  centers  of  concentration  are  not  useful 
for  Central  America  and  southern  Mexico,  because  the  5°  grid  crosses  the  entire  continent 
and  obscures  possible  Atlantic  and  Pacific  centers.  Instead,  I plotted  centers  of  geographic 
distribution  of  all  species  and  subspecies  taxa  with  ranges  less  than  1250  miles,  and  circled 
these  centers  to  enclose  all  geographic  distributions  up  to  a 300  mile  radius  (Fig.  262).  Not 
surprisingly,  resulting  centers  of  concentration  correspond  to  presence  of  endemic  species. 
Centers  4 and  8 overlap,  but  contain  quite  distinct  faunas.  Based  on  these  centers  of  con- 
centration, areas  of  concentration  (Fig.  263)  were  determined  by  comparisons  of  geographic 
distributions  and  by  notions  about  barriers.  Area  5 is  judged  to  include  peninsular  Florida, 
because  of  the  presence  there  of  only  the  Middle  American  S.  lindrothi.  Area  6 is  of  doubt- 
ful reality,  as  it  includes  only  one  known  species  which,  though  endemic,  is  related  to  a 
species  in  area  5.  Area  7 is  also  of  doubtful  reality,  because  the  endemic  species  there,  S. 
sculptilis,  most  probably  ranges  into  Middle  America.  Distributions  of  species  and  subspecies 
are  compared  with  these  centers  and  areas  in  Table  72,  dissimilarity  values  (Erwin,  1970; 
Ball  and  Freitag,  1969)  are  listed  in  Table  73,  and  an  index  of  dissimilarity  is  given  in  Table 
74. 

Peripheral  areas  1,6,  11,  and  12  are  most  distinct,  as  were  Erwin’s  centers  1,  5,  and  8. 
Areas  5 and  7,  with  a dissimilarity  value  of  64,  are  least  distinct,  and  perhaps  not  distinct  at 
all.  In  general,  more  central  areas  are  least  distinct,  as  was  found  by  Erwin  for  Brachinus. 
All  contiguous  areas  share  one  or  more  taxa,  but  subtraction  patterns  are  evident.  Curiously, 
area  2 is  more  similar  in  faunal  composition  to  tropical  area  4 than  to  subtropical  or  temper- 
ate areas  1,  9,  10,  and  1 1.  Physical  barriers,  such  as  the  Sierra  Nevada,  Rocky  Mountains, 
Mexican  Plateau,  and  Colorado  Desert,  are  probably  responsible  for  this  pattern. 

In  general,  my  areas  of  concentration  for  Schizogenius  correspond  fairly  well  with  Erwin’s 
centers  of  concentration  for  Brachinus,  except  for  lack  of  a special  area  in  southeastern 
United  States  and  division  of  Mexico  and  Middle  America  into  several  separate  areas.  They 
are  not,  however,  as  well  correlated  with  general  climatic  patterns,  and  a more  detailed 
comparison  follows. 

My  area  1 differs  from  Erwin’s  center  1 mainly  by  including  arid  regions  south  of  the 
Tehachapi  crest.  In  Schizogenius,  the  Tehachapi  isolates  S.  falli  from  S.  ochthocephalus, 
and  also  isolates  distinctive  populations  of  S.  pygmaeus,  but  otherwise  is  not  evidently  an 
important  barrier.  Area  2 differs  from  Erwin’s  center  2 by  exclusion  of  western  Texas, 
southern  California  (area  1),  and  Baja  California  (area  3).  Although  the  endemic  fofm  in 
area  3 is  most  closely  related  to  one  in  area  1 , another  form  is  shared  with  area  2 but  not 
area  1,  and  populations  of  both  other  species  in  area  3 are  more  similar  to  populations  in 
area  2 than  area  1 . Areas  10  and  1 1 correspond  closely  with  Erwin’s  centers  4 and  3,  except 
that  western  Texas  is  included  in  area  10.  Area  12  includes  Erwin’s  center  6 plus  part  of 
center  7. 

Erwin’s  center  5 includes  approximately  my  areas  4,  5,  7,  8,  and  9.  Area  7,  which  may  be 
an  artifact,  and  area  5 comprise  Middle  America  plus  the  rest  of  the  area  east  of  the  Isthmus 
of  Tehuantepec.  This  region  is  not  sharply  differentiated  climatically  from  areas  4 and  8 in 
southern  Mexico,  but  probably  reflects  the  past  existence  of  a water  gap.  Areas  4 and  8 
form  the  rest  of  the  continental  tropical  region,  their  northern  limits  corresponding  closely 
to  northern  limits  of  rain-forest  formations.  When  the  distributions  of  the  various  taxa  of 


332 


Whitehead 


Table  72.  Distribution  of  North  and  Middle  American  species  and  subspecies  of  Schizo- 
genius  in  relation  to  centers  of  concentration. 


Centers 


Taxon 

1 

2 

3 

4 

5 

6 

7 

8 

9 

10 

11 

12 

amphibius 

arimao 

XX 

XX 

auripennis 

brevisetosus 

XX 

XX 

XX 

XX 

XX 

X 

XX 

chiapatecus 

chiricahuanus 

XX 

XX 

XX 

crenulatus 

XX 

X 

depressus 

XX 

XX 

X 

X 

X 

X 

dilatus 

XX 

emdeni 

falli 

X 

XX 

XX 

X 

XX 

XX 

XX 

XX 

X 

ferrugineus 

kulti 

XX 

XX 

X 

XX 

XX 

XX 

XX 

XX 

lindrothi 

XX 

lineolatus 

litigiosus 

longipennis 

XX 

XX 

XX 

XX 

XX 

XX 

XX 

XX 

XX 

XX 

multisetosus 

neovalidus 

ochthocephalus 

optimus 

ozarkensis 

XX 

XX 

XX 

XX 

XX 

XX 

pacificus 

planulatus 

planuloides 

X 

XX 

XX 

XX 

pluripunctatus 

plurisetosus 

XX 

XX 

pygmaeus 

sallei 

XX 

XX 

XX 

XX 

XX 

XX 

XX 

XX 

XX 

XX 

scopaeus 
sculp  tilis 
seticollis 
sulcifrons 

XX 

XX 

XX 

XX 

XX 

XX 

tenuis 

X 

XX 

XX 

XX 

XX 

tibialis 

XX 

XX 

XX 

XX 

tristriatus 

truquii 

vandykei 

X 

XX 

XX 

XX 

Totals,  x and  xx 

6 

13 

4 

13 

9 

1 

6 

8 

9 

9 

6 

6 

x-  in  area,  but  not  in  center 
xx-  in  center 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


333 


Table  73.  Dissimilarity  values  among  centers  of  concentration  of  the  genus  Schizogenius 
in  North  and  Middle  America. 


Center 


Center 

Statistics* 

2 

3 

4 

5 

6 

7 

8 

9 

10 

11 

12 

1 

t-c/t  X 

100 

84 

70 

84 

93 

100 

92 

86 

80 

87 

92 

92 

2 

t-c/t  X 

100 

— 

82 

65 

81 

100 

79 

81 

86 

86 

90 

90 

3 

t-c/t  X 

100 

— 

— 

82 

83 

100 

80 

92 

85 

92 

90 

90 

4 

t-c/t  X 

100 

— 

— 

— 

73 

100 

74 

76 

77 

86 

95 

95 

5 

t-c/t  X 

100 

— 

— 

— 

— 

100 

64 

75 

88 

100 

100 

100 

6 

t-c/t  X 

100 

— 

— 

— 

— 

100 

100 

100 

100 

100 

100 

7 

t-c/t  X 

100 

— 

— 

— 

— 

— 

— 

71 

87 

100 

100 

100 

8 

t-c/t  X 

100 

— 

— 

— 

— 

— 

— 

— 

76 

88 

100 

100 

9 

t-c/t  X 

100 

— 

— 

— 

— 

— 

— 

— 

72 

87 

93 

10 

t-c/t  X 

100 

— 

— 

— 

— 

— 

— 

— 

— 

— 

67 

80 

11 

t-c/t  X 

100 

— 

— 

— 

— 

— 

— 

— 

-— 

— 

— 

75 

* t = total  taxa,  c = taxa  in  common 


Table  74.  Index  of  dissimilarity  among  areas  of  concentration  of  the  genus  Schizogenius 
in  North  and  Middle  America,  determined  from  Table  73. 


1* 

Area  6 

1100 

2 

Area  12 

1015 

3 

Area  1 1 

996 

4 

Area  1 

980 

5 

Area  10 

968 

6 

Area  5 

957 

7 

Area  7 

947 

8 

Area  3 

946 

9 

Area  8 

945 

10 

Area  9 

931 

11 

Area  2 

924 

12 

Are^  4 

907 

* Ranked  in  order  from  most  to  least  dissimilar. 


334 


Whitehead 


area  2 are  better  known,  that  area  may  be  found  divisible  into  subunits  corresponding  at 
least  to  the  subtropical  area  9 and  the  warm  temperate  area  10. 

In  summary,  Schizogenius  species  are  generally  less  vagile  than  Brachinus  species  but 
more  vagile  than  Evarthrus  species,  and  their  distribution  patterns  are  accordingly  influ- 
enced by  a balance  of  factors  influencing  distributions  in  Brachinus  and  Evarthrus.  The 
riparian  gravel  bar  habitat  of  most  of  the  species  is  a further  restraining  variable,  but  also 
one  which  may  permit  penetration  of  otherwise  unfavorable  climatic  zones.  The  westward 
extension  of  area  10,  for  example,  is  one  which  simply  follows  the  Rio  Grande  drainage 
system;  its  Schizogenius  fauna  is  depauperate,  and  contains  some  elements  from  area  2,  but 
is  still  most  closely  related  to  the  fauny  of  the  rest  of  area  10. 

Historical  zoogeography 

Paleogeography . — According  to  Maldonado-Koerdell  (1964),  many  important  events 
occurred  in  Mexico  and  Middle  America  during  Tertiary  and  Quarternary  time.  For  pur- 
poses of  this  discussion,  I use  the  following  time  scale,  in  millions  of  years  before  present: 
Pleistocene  and  Recent,  0-3.0;  Pliocene,  3.1-13.0;  Miocene,  13.1-25.0;  Oligocene,  25.1-36.0; 
Eocene,  36.1-58.0;  and  Paleocene,  58.1-63.0.  North  and  South  America  were  separated  in 
Late  Paleocene  by  formation  of  the  Peruvian-Venezuelan  geosyncline.  During  Paleocene 
and  Eocene,  the  Balsas  portal  was  eliminated,  the  Sierra  Madre  Oriental  was  lifted,  the 
Mexican  Plateau  began  to  rise,  and  Middle  America  bulged  northward  to  include  Jamaica 
but  not  Cuba.  Near  Early  Oligocene,  Middle  America  became  separated  from  North  America 
by  the  Tehuantepec  portal,  and  extended  further  northward  to  include  Cuba.  Early  Miocene 
saw  the  rise  of  the  Sierra  Madre  Occidental,  continued  emergence  of  the  Mexican  Plateau, 
beginnings  of  volcanic  activity  in  Middle  America  and  along  the  Neo-Volcanic  Axis,  and 
approximate  delimitation  of  modern  climatic  zones.  In  Late  Miocene,  the  Tehuantepec 
portal  closed,  and  Cuba  and  Jamaica  became  isolated  from  Middle  America.  In  Middle 
Pliocene  volcanic  activity  was  generally  interrupted,  but  renewed  in  Late  Pliocene.  Also  in 
Late  Pliocene,  South  and  Middle  America  were  reconnected,  and  uplands  in  the  Isthmus  of 
Tehuantepec  were  formed. 

Pleistocene  events  are  more  difficult  to  interpret  and  collate.  Presumably,  the  first  two 
million  years  of  the  period  were  marked  by  a general  deterioration  in  climate,  followed  in 
the  last  million  years  by  a series  of  four  major  glaciations;  the  last  of  these  glaciations,  the 
Wisconsin,  commenced  about  100,000  years  ago.  In  Mexico  and  Middle  America,  inter- 
mittent but  intense  volcanic  activity  occurred  throughout  the  Pleistocene.  The  climate  was 
probably  warm  humid  in  Late  Pliocene,  cool  humid  in  Early  Pleistocene,  and  cool  dry 
during  the  first  glaciation. 

Subsequently  the  climate  varied  cyclically  through  warm  dry  to  warm  humid  during 
interglacials,  returning  through  cool  humid  to  cool  dry  during  glaciations.  Montane  glaciers 
in  central  Mexico  descended  to  as  low  as  2450  m,  or  about  half  the  lower  limits  of  present 
remnants  of  these  glaciers.  These  observations  suggest  that  climatic  conditions  at  low  eleva- 
tions even  in  the  south  fluctuated,  and  in  particular  tropical  conditions  probably  did  not 
extend  north  of  the  Neo-Volcanic  axis  during  glacial  maxima. 

Evolutionary  and  phylogenetic  rates.  — Evolutionary  and  zoogeographic  pathways  devel- 
oped interdependently,  and  phylogenetic  and  zoogeographic  analyses  should  therefore  be 
compatible  with  one  another.  One  method  for  testing  for  compatibility  is  to  correlate  the 
reconstructed  phylogeny  with  known  or  suspected  paleogeographical  events.  To  do  this,  I 
determined  an  average  time  between  branching  points  in  the  reconstructed  phylogeny,  based 
on  assumptions  discussed  in  the  section  on  phylogeny;  this  method  was  tested  and  found 
useful  for  comparing  phylogeny  and  zoogeography  in  Brachinus  (Erwin,  1970)  and  Evar- 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


335 


thrus  (Ball  and  Freitag,  in  Freitag,  1969).  The  average  time  between  branching  points  was 
determined  to  be  about  3,000,000  years,  from  the  following  lines  of  reasoning. 

For  Hawaiian  birds,  Bock  (1970)  suggests  that  time  required  for  speciation  may  range 
between  100,000  and  1,000,000  years.  For  recent  European  mammals,  Kurten  (1968) 
suggested  a time  range  of  between  30,000  and  3,000,000  years;  his  upper  limit  corresponds 
to  Early  Pleistocene,  while  the  lower  limit  may  at  least  in  part  reflect  inadequate  knowledge. 
According  to  Halffter  (1964),  La  Brea  (Pleistocene)  insects  are  no  more  than  subspecifically 
distinct  from  extant  species,  though  their  distributions  may  have  changed  greatly.  For  speci- 
ation in  Schizogenius  and  other  continental  North  American  carabid  beetle  genera  except 
those  with  highly  insular  distributions  such  as  Scaphinotus  (see  Ball,  1966),  Kurten’s  upper 
limit  may  be  close  to  an  average  figure. 

The  ancestor  of  Schizogenius  entered  South  America  after  formation  of  an  effective  bar- 
rier at  the  Panamanian  portal  in  Late  Paleocene,  no  more  and  probably  less  than  60,000,000 
years  ago.  According  to  my  reconstructed  phylogeny  (Fig.  257-258),  a minimum  of  16 
branching  points  are  required  to  reach  back  to  the  common  ancestor  of  American  Halo- 
coryza  and  Schizogenius.  Thus  the  maximum  possible  time  between  branching  points  is 

4.000. 000  years.  According  to  Halffter  (1964),  fossils  in  Upper  Oligocene-Lower  Miocene 
amber  from  Semojovel,  Chiapas,  about  25,000,000  years  in  age,  include  nine  species  of 
beetles,  all  extinct,  as  so  far  studied.  These  are  arrayed  in  seven  genera,  five  of  which  are 
extant;  none  are  carabid  beetles.  In  the  same  amber  deposit  is  a carabid  of  the  bembidiine 
genus  Tachys,  said  to  be  related  to  living  species  (T.  L.  Erwin,  1971).  The  extinct  genera 
may  represent  sideline  extinctions,  or  ancestral  forms,  and  relationships  of  the  extinct  spe- 
cies of  extant  genera  are  unclear.  But  the  Tachys  specimen,  at  least,  implies  that  ancestors 
of  existing  species  groups  of  carabid  beetles  may  already  have  evolved  at  that  time.  There- 
fore, ancestral  Schizogenius  had  probably  evolved  by  at  least  Late  Oligocene,  and  had 
probably  even  differentiated  into  at  least  the  major  lineages,  such  as  the  truquii  lineage,  by 
then.  If  the  ancestor  of  Schizogenius  had  diverged  from  Halocoryza  by  Late  Oligocene,  but 
not  yet  differentiated  into  subgenera,  the  average  time  between  dichotomies  would  be  about 
1,500,000  years.  If  the  truquii  lineage  had  differentiated  by  this  time  from  its  sister  group, 
but  not  yet  undergone  radiation,  the  time  required  between  dichotomies  would  be  close  to 

3.000. 000  years.  In  short,  clues  from  Semojovel  amber  suggest  by  analogy  that  at  least  the 
ancestor  of  Schizogenius  had  evolved  by  Late  Oligocene,  and  that  probably  some  differenti- 
ation had  taken  place. 

Matthews  (in  Hopkins,  et  al.,  1971)  discussed  a Late  Pliocene  insect  fauna  from  the 
Bering  Strait  region,  dated  at  approximately  5.7  million  years.  Among  carabid  beetles  re- 
ported in  this  paper,  some  apparently  represent  extant  species,  while  others  probably  repre- 
sent extinct  species  closely  related  to  extant  forms.  These  data  are  significant  in  their  impli- 
cations of  old  age  for  living  species  of  carabid  beetles. 

Faunal  exchanges  between  Middle  and  South  America  offer  a further  clue  to  the  age  of 
the  most  recent  dichotomies.  Only  three  lineages  are  likely  to  have  crossed  over  the  Pana- 
manian portal,  or,  perhaps,  over  emergent  land  of  low  relief  there.  I think  that  shortly  after 
this  barrier  disappeared,  one  invasion  of  South  America  and  three  of  Middle  America 
occurred,  and  resulted  in  differentiation  of  distinctive  species.  I judge  that  these  events  took 
place  between  three  and  five  million  years  ago,  more  probably  three  million  years  because 
there  is  no  reason  to  believe  immigration  proceeded  immediately.  The  Panamanian  area  is 
still  evidently  a significant  barrier  to  the  dispersal  of  Schizogenius  species,  as  only  two 
modem  species  are  known  from  both  Middle  and  South  America. 

Another  general  line  of  reasoning  concerns  Pleistocene  events  in  North  America.  In  S. 
falli , there  is  statistical  evidence  of  rapid  change  in  a single  characteristic  over  a short  period 


336 


Whitehead 


of  time  in  one  area:  50  years,  in  the  Dragoon  Mountains  of  southern  Arizona.  In  S.  depres- 
sus,  there  is  statistical  evidence  that  differentiation  of  eastern  and  western  groups  of  popula- 
tions in  the  southwestern  United  States  has  taken  place  within  the  last  10,000  years.  Perhaps 
in  the  north  further  differentiation  has  developed  much  more  recently,  as  a result  of  charac- 
ter displacement.  Also  in  the  north,  however,  is  evidence  that  genetic  differentiation  but  not 
speciation  is  associated  with  much  earlier  Pleistocene  events;  at  least  Wisconsin  (100,000 
years),  and  more  probably  Nebraskan  (1,000,000  years).  My  reasoning  here  presumes  that 
the  northern  forms  became  isolated  in  early  Pleistocene  because  of  deteriorating  climatic 
conditions,  were  subsequently  alternately  completely  and  partially  isolated,  and  that  by  the 
first  interglacial  were  partly  genetically  isolated.  The  common  ancestor  of  the  closely  related 
but  grossly  disjunct  S.  sulcifrons  and  S.  litigiosus  was  probably  transcontinental  in  Early 
Pleistocene,  but  with  the  deterioration  of  climate  formed  eastern  and  western  isolates  before 
the  Nebraskan  glaciation.  Similarly,  I suspect  that  geographic  isolates  of  the  common  ances- 
tor of  S.  planulatus  and  S.  ozarkensis  formed  in  Early  Pleistocene,  before  the  onset  of  glaci- 
ation. Alternatively,  one  might  date  the  initial  isolations  of  S.  sulcifrons  from  S.  litigiosus 
and  S.  planulatus  from  S.  ozarkensis  as  no  earlier  than  the  beginning  of  Wisconsin  glaciation. 
But  this  would  suggest  that  the  entire  evolution  of  the  genus  may  have  taken  place  within 
the  last  2,000,000  years.  If  these  isolations  occurred  in  Early  Pleistocene,  however,  they 
may  be  dated  as  about  3,000,000  years  and  thus  agree  with  datings  from  other  evidence. 

A particularly  difficult  problem  is  posed  by  those  species  complexes  in  which  included 
species  are  allopatric  but  proximate.  If  their  ranges  have  long  been  proximate,  and  fixed,  a 
period  of  3,000,000  years  seems  difficult  to  explain.  But  boundaries  unquestionably  were 
not  always  contiguous,  they  most  likely  are  not  fixed,  and  there  are  doubtless  interactions 
along  these  boundaries.  Competitive  exclusion  (Mayr,  1963)  is  the  probable  reason  for  main- 
tenance of  proximate  but  allopatric  distributions  in  Schizogenius  species.  For  instance,  S. 
pluripunctatus  and  S.  kulti  probably  were  isolated  by  Late  Pliocene  or  Early  Pleistocene 
vulcanism  along  the  Neo-Volcanic  Axis,  in  agreement  with  my  suggested  3,000,000  year 
time  limit.  Subsequently,  geographic  isolation  was  maintained  by  competitive  exclusion,  but 
boundaries  shifted  northward  with  the  northward  extension  of  tropical  climates  after  the 
end  of  the  Pleistocene.  If  two  species  which  are  closely  related  or  which  are  particularly 
similar  in  ecological  requirements  inhabit  adjacent  river  systems,  or  separate  segments  of  a 
single  river,  their  distributions  are  unlikely  to  overlap.  Undoubtedly,  there  is  contact  on  an 
individual  basis,  as  individuals  migrate  away  from  parental  habitat.  Evidence  that  this  does 
happen  is  found  in  allopatric  components  of  a species  complex  such  as  in  S.  falli  and  S. 
ochthocephalus  or  in  S.  kulti  and  S.  pluripunctatus,  or  even  in  eastern  and  western  forms 
of  S.  depressus  in  northwestern  United  States.  Where  ranges  approach  one  another,  proxi- 
mate populations  tend  to  be  the  most  strongly  differentiated,  which,  I think,  is  evidence  of 
character  displacement  (Brown  and  Wilson,  1956).  I have  used  this  observation,  of  inter- 
rupted gene  flow,  as  one  criterion  for  species  recognition.  But  I think  such  patterns  are 
best  explained  as  the  result  of  interspecific  interaction.  In  northeastern  Mexico,  where  S. 
scopaeus  and  S.  pygmaeus  are  parapatric  or  sympatric  in  distribution  they  are  also  strongly 
differentiated  morphologically  and,  probably,  ecologically;  elsewhere,  where  less  strongly 
differentiated,  they  are  allopatric.  Near  their  boundary  in  the  Big  Bend  Region  of  the  Rio 
Grande,  they  probably  are  not  strongly  differentiated  ecologically,  and  in  that  region  they 
show  evidence  of  character  displacement  in  form  of  male  median  lobe. 

In  summary,  the  separation  of  North  and  South  America  in  Late  Paleocene  suggests  an 
upper  limit  for  the  age  of  ancestral  Schizogenius  of  60,000,000  years,  and  paleontological 
data  from  the  Semojovel  fossils  suggests  a lower  limit  of  25,000,000  years.  In  turn  these 
datings  suggest  that  intervals  between  dichotomies  in  the  most  complete  lineage  of  the 


Classification,  phytogeny,  and  zoogeography  of  Schizogenius 


337 


reconstructed  phytogeny  of  the  genus  are  at  most  4,000,000  years  and  at  least  1,500,000 
years.  The  timing  of  overland  faunal  exchanges  between  South  and  Middle  America,  and 
Pleistocene  events  in  North  America,  suggest  that  this  interval  is  approximately  3,000,000 
years.  This  is  a crude  approximation,  and  is  an  average,  not  a constant;  I accept  this  interval 
as  a working  proposition. 

With  a 3,000,000  year  interval  between  dichotomies,  I judge  that  the  common  ancestor 
of  Schizogenius  evolved  some  50,000,000  years  ago,  about  Middle  Eocene.  Ancestors  of  the 
ferrugineus  group,  the  truquii  lineage,  and  the  crenulatus  group  entered  North  America  in 
Late  Eocene,  Middle  Oligocene,  and  Middle  Miocene,  respectively.  These  suggested  historical 
events  are  compared  with  paleogeological  events  in  the  following  section.  This  3,000,000 
year  interval  is  the  basis  of  the  time  scale  shown  in  my  reconstructed  phytogeny  (Fig.  255- 
258). 

Vicariance  and  historical  zoogeography.  — In  this  section,  I deal  briefly  with  matters  of 
vicariance  and  historical  zoogeography  pertaining  to  minor  groups  and  lineages  of  the  genus 
Schizogenius  that  have  penetrated  North  and  Middle  America,  and  at  some  length  as  they 
apply  to  the  truquii  lineage. 

The  origin  of  the  subgenus  Genioschizus  took  place  in  South  America  about  48,000,000 
years  ago.  This  group  is  now  poorly  represented,  with  three  closely  related  species  groups, 
and  extinctions  undoubtedly  account  for  its  present  lack  of  diversity.  The  ancestor  of  the 
crenulatus  group  entered  Middle  America  in  Early  to  Middle  Miocene.  The  ancestor  of  living 
taxa  entered  southwestern  North  America  after  closure  of  the  Tehuantepec  portal,  probably 
in  Middle  to  Late  Pliocene,  and  extended  northward  into  the  Sonoran  region.  Southern  and 
northern  isolates  formed  in  Early  Pleistocene  because  of  orogenic  activity  in  the  Neo- 
Volcanic  Axis,  but,  with  northward  extension  of  tropical  climates  in  more  recent  times, 
their  boundary  shifted  northward  to  its  present  location  near  the  Tropic  of  Cancer.  The  two 
living  subspecies  of  S.  crenulatus  are  north-south  vicariants.  Two  other  members  of  the  sub- 
genus Genioschizus  represented  in  North  and  Middle  America  are  distantly  related  members 
of  the  tenuis  group.  Ancestral  S.  sculptilis  entered  Middle  America  in  Early  Pleistocene,  and 
extended  westward  as  far  as  the  Usumacinta-Grijalva  area;  this  species  probably  remains  a 
component  of  the  Middle  American  fauna.  More  recently,  S.  tenuis  has  extended  northward 
from  Colombia  through  Middle  America,  as  far  as  the  northern  limits  of  the  tropics  in 
eastern  Mexico  and  the  northern  limits  of  the  subtropics  in  western  Mexico. 

As  do  members  of  the  subgenus  Genioschizus,  and  at  least  S.  lindrothi  of  the  lindrothi 
group,  members  of  the  ferrugineus  group  are  found  mainly  in  sandy  habitats.  I suggest  a 
particularly  early  entry  into  Middle  America  for  this  group,  perhaps  Early  Eocene,  before 
the  Tehuantepec  portal  opened.  The  group  is  characterized  by  lack  of  diversity,  no  close 
relationship  among  existing  species,  and  by  widespread  distributions  of  living  species.  The 
two  existing  species  are  largely  allopatric,  but  are  not  true  sister  species  and  therefore 
vicariant  relationships  are  doubtful.  The  group  is  not  represented  in  far  western  United 
States,  Florida,  or  eastern  Mexico.  These  gaps  in  distribution  are  difficult  to  explain.  S. 
ferrugineus,  or  a related  form,  may  until  comparatively  recently  have  been  represented  in 
subtropical  Florida,  but  displaced  by  the  immigrant  S.  lindrothi.  The  distribution  of  S. 
lindrothi,  Middle  America  and  Florida,  suggests  that  early  distributions  of  the  ferrugineus 
group  may  have  followed  a similar  pattern.  But  if  my  conclusions  about  timing  are  correct, 
Florida  did  not  then  exist  as  such,  and,  further,  there  was  no  barrier  to  dispersal  at  the 
Isthmus  of  Tehuantepec.  The  present  continuity  of  range  of  the  two  species  suggests  that 
Middle  America  and  southern  Mexico  were  depopulated  of  the  group  for  a tong  time,  with 
extinction  of  tropic-subtropic  elements  initiated  in  Late  Oligocene  or  Early  Miocene.  This 
may  have  been  caused  in  part  by  orogenic  activity  and  in  part  from  displacement  by  newer 


338 


Whitehead 


immigrants  of  the  truquii  lineage.  Surviving  forms  became  adapted  to  temperate  conditions 
in  northeastern  North  America.  A later  isolate  subsequently  became  established  in  the 
Sonoran  area,  from  which  it  successfully  penetrated  southward  along  the  Pacific  coast  of 
Mexico,  and  beyond  to  the  southern  limit  of  Middle  America. 

Other  minor  lineages  represented  in  North  and  Middle  America  are  the  lindrothi  and 
optimus  groups,  each  with  distinctive  vicariant  sister  species  in  Middle  and  South  America. 
Ancestors  of  Middle  American  species  arrived  there  in  Early  Pleistocene.  S.  lindrothi  subse- 
quently became  distributed  across  the  Gulf  of  Mexico  to  peninsular  Florida,  where  it  may 
have  displaced  S.  ferrugineus  from  sandy  habitats  in  subtropical  regions.  Although  Middle 
American  and  Floridian  forms  of  S.  lindrothi  are  quite  distinctive,  I suspect  the  species  may 
also  be  represented  in  the  West  Indies,  particularly  in  areas  of  Caribbean  pines  as  discussed 
by  Mirov  (1967). 

The  ancestor  of  the  truquii  lineage  passed  through  Middle  America  into  southern  Mexico 
in  Middle  Oligocene,  over  the  Tehuantepec  and  Panamanian  portals.  After  reaching  tropical 
Mexico,  it  gave  rise  to  two  lineages,  one  centered  in  southwestern  Mexico  and  the  other  in 
northeastern  Mexico.  One  lineage  is  today  represented  by  a single  species  and  species  group, 
S.  truquii.  The  other  lineage  diversified  into  seven  species  groups,  comprising  the  main  part 
of  the  genus  represented  at  present  in  North  and  Middle  America. 

The  major  part  of  the  evolutionary  history  of  the  truquii  lineage  unfolded  in  Mexico  and 
the  southern  part  of  the  United  States.  To  analyze  the  zoogeography  of  this  lineage,  I made 
detailed  comparisons  between  species  for  geographic  distributions,  geographic  variation, 
and  vicariance,  and  refined  limits  for  the  areas  of  concentration  in  Fig.  263  accordingly. 
Resulting  major  areas  of  vicariance  are  shown  in  Fig.  264.  Vicariance  areas  1 and  6 are  peri- 
pheral, corresponding  approximately  to  concentration  areas  1,  5,  6,  7,  11,  and  12.  High 
altitude  regions  of  northern  and  central  Mexico  form  a potent  barrier  to  the  dispersal  of 
Schizogenius  species,  as  suggested  by  greater  dissimilarity  between  eastern  and  western  areas 
of  concentration  than  between  northern  and  southern  areas.  Area  2 excludes  all  portions  of 
the  Colorado  River  except  the  Gila  system,  in  recognition  of  northern  limits  of  numerous 
species.  Area  5 was  recognized  as  separate  from  area  2 to  account  for  northern  limits  of 
several  southern  species,  to  conform  with  distributional  limits  of  S.  depressus,  and  to 
exclude  certain  species  endemic  to  area  2.  And  the  northwestern  limit  of  area  3 was  refined 
to  exclude  S.  depressus. 

Vicariance  areas  2 and  3,  Sonoran  and  Texan  regions,  have  similar  latitudinal  limits,  and 
climates  that  may  be  described  as  warm  temperate.  Vicariance  areas  4,  5,  and  6 may  be 
described  as  subtropical  regions,  respectively  the  Baja  Californian,  Sinaloan,  and  Tamaulipan 
regions.  All  these  areas  represent  broad  climatic  zones;  their  boundaries  are  inexact,  and 
surely  have  not  been  constant  in  the  past.  Probably,  for  example,  northern  boundaries  of 
tropical  areas  7 and  8 shifted  southward  to  the  Neo-Volcanic  Axis  during  Pleistocene  glacial 
maxima.  These  suggested  boundaries  are  intended  only  to  describe  general,  climatically 
limited,  centers  of  speciation,  not  specific  areas  of  distribution.  Thus,  exact  locations  of 
some  boundaries  such  as  those  between  areas  5 and  7 in  the  west  and  between  areas  3 and  6 
in  the  east  are  quite  arbitrary,  but  this  defect  is  more  apparent  than  real.  Other  boundaries, 
as  between  areas  6 and  8,  do  represent  abrupt  faunal  limits,  but  as  noted  above  have  not 
been  constant  over  long  periods  of  time. 

Fig.  265-269  are  vicariance  maps  showing  evolutionary  and  zoogeographic  pathways  for 
species  groups  of  the  truquii  lineage,  and  for  taxa  included  in  the  pluripunctatus,  tristriatus, 
longipennis,  and  depressus  groups.  These  vicariance  relationships  were  described  in  the 
section  on  the  phylogeny  of  the  truquii  lineage,  and  following  is  a correlation  of  evolu- 
tionary and  zoogeographic  pathways  with  paleogeological  events. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


339 


When  the  ancestor  of  the  truquii  lineage  arrived  in  North  America  in  Middle  Oligocene, 
orogenic  activity  in  Mexico  was  minimal.  After  divergence  into  eastern  and  western  lineages 
in  Late  Oligocene,  distributions  of  the  ancestors  of  these  lineages  were  maintained  initially 
through  competitive  exclusion.  In  Early  Miocene,  orogenic  and  volcanic  activity  commenced 
along  the  Neo-Volcanic  Axis  and  Sierra  Madre  Occidental,  rendering  vicariance  areas  2,  5, 
and  7 relatively  inhospitable  for  colonization.  In  areas  3 and  6,  however,  no  mountain  build- 
ing took  place  then  or  subsequently,  and  these  areas  were  thus  particularly  suitable  for 
speciation  of  Schizogenius  species. 

Nothing  certain  can  be  said  of  the  zoogeographic  history  of  the  truquii  group.  Doubtless 
some  species  diversification  did  take  place,  and  doubtless  vicariance  was  a factor,  but  all 
species  except  S.  truquii  subsequently  disappeared.  Limitations  of  habitat  caused  by  oro- 
genic disturbances  hindered  evolution  of  the  group,  and  when  elements  of  the  more  progres- 
sive sister  group  entered  regions  formerly  occupied  by  members  of  the  truquii  group,  the 
latter  were  simply  eliminated. 

Most  early  diversification  in  the  rest  of  the  truquii  lineage,  before  Middle  Pliocene  vol- 
canic quiescence,  took  place  in  Atlantic  areas  3,  6,  and  8,  and  resulted  from  isolations 
caused  by  shifts  in  climatic  tolerance.  The  ancestor  of  the  arimao  subgroup  of  the  depressus 
group  entered  Middle  America  over  the  Tehuantepec  portal  in  Late  Miocene,  and  spread 
to  Cuba  before  the  West  Indies  separated  from  Middle  America.  Also  in  Late  Miocene  or 
Early  Pliocene,  the  ancestor  of  the  pluripunctatus  group  entered  the  northern  part  of  area 
2,  north  of  the  intense  tectonic  activity  in  the  Sierra  Madre  Occidental  region.  In  Fig.  265  I 
show  the  split  of  the  ancestor  of  the  truquii  group  from  its  sister  group  in  southern  Mexico, 
latitudinal  vicariance  shifts  in  the  ancestry  of  other  groups  in  the  lineage,  and  the  immigra- 
tion to  the  Sonoran  region  of  the  ancestor  of  the  pluripunctatus  group. 

During  the  long  geological  quiescence  of  Middle  Pliocene,  ancestral  elements  of  the 
pluripunctatus  group  became  firmly  entrenched  in  the  Sonoran  region,  and  spread  west- 
ward and  southward  from  there.  One  form,  the  ancestor  of  S.  seticollis,  became  isolated 
in  California,  and  spread  into  southern  Baja  California;  in  Early  Pleistocene,  in  turn,  this 
southern  form  became  isolated  as  the  ancestor  of  the  living  subspecies  S.  s.  vandykei. 
Another  form,  the  ancestor  of  S.  plurisetosus  and  S.  multisetosus,  became  isolated  in 
southern  Mexico  in  Late  Pliocene,  perhaps  by  renewed  volcanic  activity  in  the  Neo-Volcanic 
Axis.  Vicariance  patterns  in  the  pluripunctatus  group  were  completed  in  Early  Pleistocene 
when  ancestral  forms  of  S.  pluripunctatus  and  S.  kulti,  and  of  S.  plurisetosus  and  S. 
multisetosus  were  divided  into  tropical  and  subtropical  isolates,  perhaps  also  by  vulcanism 
in  the  Neo-Volcanic  Axis.  Subsequently,  northern  limits  of  the  tropical  forms  shifted  north- 
ward to  their  present  locations,  and  the  more  progressive  S.  kulti  displaced  the  more 
conservative  S.  multisetosus  from  most  lowland  parts  of  its  range.  These  events  are  sum- 
marized in  Fig.  266. 

No  elements  of  the  tristriatus  group  evolved  in  Pacific  drainage  systems  (Fig.  267), 
although  one  species,  S.  tibialis  (Fig.  188),  has  successfully  penetrated  the  area.  The  ances- 
tor of  one  species,  S.  tristriatus , reacquired  tropical  adaptations  in  Late  Miocene  or  Early 
Pliocene,  but  probably  never  did  penetrate  south  of  the  then  active  Neo-Volcanic  Axis. 
The  ancestor  of  its  sister  group,  which  had  retained  subtropical  adaptations,  subsequently 
differentiated  into  warm  temperate  and  subtropical  lineages.  Of  the  subtropical  lineage, 
differentiation  into  subtropical  and  tropical  forms  led  to  the  speciation  of  S.  dilatus  and 
S.  tibialis.  Ancestral  S.  tibialis  spread  southward  to  the  Isthmus  of  Tehuantepec,  from 
there  into  the  Rio  Balsas  and  Rio  Grande  de  Santiago  drainages,  and  returned  northward 
across  the  highlands  of  central  Mexico  into  subtropical  area  6 and  even  into  southernmost 
warm  temperate  area  3.  Zoogeographic  history  of  the  warm  temperate  ancestor  of  the 


340 


Whitehead 


amphibius  and  planulatus  subgroups  was  one  of  a tendency  to  acquire  cool  temperate 
adaptations.  In  Middle  Pliocene,  ancestral  S.  amphibius  became  isolated  in  cool  temperate 
area  1 and  subsequently  spread  into  the  Appalachian  region,  to  which  it  was  restricted  in 
Pleistocene  time.  In  Late  Pliocene,  the  ancestor  of  S.  planulatus  and  S.  ozarkensis  was  iso- 
lated in  the  cool  temperate  area,  and  differentiated  into  those  species  in  Pleistocene. 

The  ancestor  of  the  longipennis  group  (Fig.  268)  was  the  first  element  of  the  subtropical 
and  warm  temperate  adapted  branch  of  the  truquii  lineage  to  return  to  tropical  areas  and 
extend  south  of  the  Neo-Volcanic  Axis.  In  Middle  Pliocene,  a western  vicariant,  ancestor  to 
S.  pacificus,  spread  through  western  areas  5 and  7.  The  eastern  vicariant  followed  later,  and 
in  Late  Pliocene  the  ancestor  of  S.  chiricahuanus  was  isolated  in  area  2 by  renewed  volcanic 
activity  in  the  Neo-Volcanic  Axis.  In  Early  Pleistocene,  the  ancestor  of  S.  neovalidus  and 
S.  longipennis  spread  into  area  2,  and  ancestral  S.  neovalidus  became  isolated  there  by 
Pleistocene  events.  More  recently,  the  progressive  S.  longipennis  also  extended  northward 
into  area  2,  and  displaced  the  more  conservative  S.  chiricahuanus  and  S.  neovalidus  from 
most  parts  of  their  ranges  except  relict  higher  elevation  strongholds. 

The  ancestor  of  the  arimao  subgroup  (Fig.  269)  acquired  tropical  adaptations  in  Late 
Miocene,  and  was  the  only  member  of  the  truquii  lineage  to  enter  Middle  America  over  the 
Tehuantepec  portal.  Indeed,  it  was  the  only  North  American  member  of  the  genus  to  enter 
Middle  America  before  Late  Pliocene  elevation  of  the  Isthmus  of  Tehuantepec.  This  ances- 
tral form  spread  into  Cuba  before  severance  of  Cuba  from  the  mainland,  but  since  Early 
Pliocene  S.  arimao  evolved  in  isolation.  The  ancestor  of  remaining  Middle  American  forms 
differentiated  into  the  ancestors  of  S.  emdeni  and  S.  “apicalis”.  In  Early  Pleistocene,  after 
closure  of  the  Panamanian  portal,  South  America  was  penetrated  and  widely  colonized  by 
the  ancestor  of  S.  ‘‘apicalis”. 

The  ancestor  of  the  rest  of  the  depressus  group  entered  warm  temperate  Texas,  and 
differentiated  into  one  line  that  continued  to  evolve  there  and  another  that  became  isolated 
in  cool  temperate  areas.  The  cool  temperate  form  spread  across  northern  parts  of  the  conti- 
nent, and  differentiated  into  modern  S.  litigiosus  and  S.  sulcifrons  in  Early  Pleistocene. 
In  Middle  Pliocene,  the  long  period  of  volcanic  quiescence  in  the  west,  one  line  of  the 
depressus  subgroup  became  isolated  in  the  Sonoran  Region.  The  ancestor  of  S.  depressus 
penetrated  northward,  acquired  cool  temperate  adaptations,  and  subsequently  extended 
southward  through  the  Sonoran  region  into  high  elevation  areas  of  central  Mexico.  In  Early 
Pleistocene,  another  cool  temperate  form,  ancestor  of  S.  ochthocephalus,  was  isolated  in 
northern  California.  Its  sister  species  spread  southward  through  area  5 into  northern  parts 
of  area  7,  northward  into  the  Colorado  River  area  of  western  part  of  area  1,  and  eastward 
into  area  3 from  which  it  spread  north  into  central  parts  of  area  1 and  south  into  area  6. 
Zoogeographic  origins  of  S.  pygmaeus  and  S.  scopaeus  are  less  well  shown  in  Fig.  269. 
Their  ancestor  was  widespread  in  eastern  areas  3,  6,  and  8 in  Middle  Pleistocene,  but  Late 
Pliocene  vulcanism  resulted  in  northern  and  southern  isolates.  In  Late  Pliocene  or  Early 
Pleistocene,  the  ancestor  of  the  northern  form  spread  westward  into  the  Sonoran  region, 
and  formed  an  isolate  there.  This  western  form  spread  northward  into  California,  and  south- 
ward into  southern  Mexico  where  it  reestablished  genetic  contact  with  the  southern  isolate. 
By  Late  Pleistocene,  however,  reproductive  isolation  had  developed  between  the  form  in 
Texas,  ancestral  to  S.  scopaeus,  and  the  southern  form  which,  reconnected  with  the  Sonoran 
form  constitutes  modern  S.  pygmaeus.  Also  in  Late  Pleistocene,  Californian  S.  pygmaeus 
evolved  into  statistically  differentiated  northern  and  southern  isolates. 

With  this  background,  more  can  be  said  about  distribution  of  the  ferrugineus  group. 
Probably  before  entry  of  the  truquii  lineage  into  the  area,  ancestral  forms  of  the  ferrugineus 
group  were  represented  throughout  Mexico  and  southern  United  States.  Paleogeological 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


341 


events  in  southern  and  western  Mexico  from  early  Miocene  contributed  to  elimination  of 
the  group  from  that  area.  Concurrently,  spread  of  progressive  and  actively  evolving  members 
of  the  truquii  lineage  into  eastern  areas  3,  6,  and  8 resulted  in  displacement  of  members  of 
the  ferrugineus  lineage  from  that  entire  region.  One  line  did  reach  cool  temperate  areas  of 
central  and  eastern  United  States,  and  there  perfected  adaptations  to  life  in  sandy  habitats, 
and,  consequently,  was  not  eliminated  when  gravel  bar  adapted  forms  of  the  truquii  lineage 
penetrated  cool  temperate  regions.  Western  United  States  meanwhile  may  have  been  gener- 
ally untenable  for  colonization  by  members  of  the  ferrugineus  lineage  because  of  tectonic 
and  orogenic  activity.  Alternatively,  members  of  the  group  living  there  lacked  special  adap- 
tations and  were  displaced  by  more  progressive  elements  of  the  truquii  lineage  from  Middle 
Pliocene  onward.  In  Middle  Pliocene,  however,  the  ancestor  of  S.  auripennis  succeeded  in 
penetrating  the  Sonoran  region,  and  subsequently  extended  southward  into  tropical  regions 
as  far  as  southern  Middle  America,  and  special  adaptations  prevented  competitive  displace- 
ment by  members  of  the  truquii  lineage. 

Before  Early  Pliocene,  members  of  relatives  of  the  truquii  group  may  well  have  occupied 
the  Sonoran  region  and  perhaps  even  California,  but  since  then  and  particularly  since  Middle 
Pliocene  were  in  competition  with  members  of  more  advanced  lineages,  and  gradually 
eliminated.  A study  of  ecological  specializations  of  S.  truquii  might  reveal  reasons  for  its 
apparent  success  in  continued  survival. 

Little  can  be  said  about  the  evolutionary  and  zoogeographic  histories  of  the  monotypic 
brevisetosus,  sallei,  and  lineolatus  groups,  all  with  probable  Texan  origins  (Fig.  265).  S. 
brevisetosus  seems  a relict  species,  and  other  elements  of  the  brevisetosus  group  may  have 
been  eliminated  through  competitive  displacement  by  other  forms  with  better  developed 
limestone  adaptations.  Curiously,  the  known  distribution  of  S.  brevisetosus  is  allopatric  in 
relation  to  distributions  of  members  of  the  pluripunctatus  group,  despite  loss  of  limestone 
adaptations  of  the  latter.  In  contrast,  distributions  of  S.  sallei  and  S.  lineolatus  are  not 
evidently  relict,  and  within  their  respective  ranges  both  tend  to  be  numerically  dominant 
over  all  others  in  the  genus.  Perhaps  the  relatively  great  vagility  of  these  species  is  a partial 
explanation  of  the  absence  of  close  relatives;  if  new  areas  are  colonized,  gene  flow  tends 
to  continue,  and  no  geographic  isolation  results.  The  only  clear  exception  to  this  pattern 
is  in  the  Rio  Grande  isolate  of  S.  lineolatus.  I suspect  this  form  is  a relict  from  cooler  Pleis- 
tocene times,  when  the  distribution  of  the  northern  form  of  the  species  extended  further 
south  than  it  presently  does. 

In  summary,  my  reconstructed  phylogeny  seems  completely  compatible  with  what  is 
known  about  paleogeological  events,  and  I think  I have  described  evolutionary  and  zoo- 
geographic pathways  that  have  a high  probability  of  correctness.  It  is  my  hope  that  future 
workers  will  test  these  hypotheses  in  comparative  studies  on  ecology,  physiology,  bio- 
chemistry, and  larval  taxonomy. 

ACKNOWLEDGEMENTS 

Many  individuals  contributed  much  toward  this  study,  and  I extend  my  sincerest  appre- 
ciation to  them  all. 

This  investigation  was  suggested  by,  and  constantly  encouraged  by,  G.  E.  Ball  of  the 
University  of  Alberta.  In  addition  to  acting  as  supervisor,  he  obtained  financial  support  for 
my  studies  (including  support  from  National  Science  Foundation  grant  GB-3312  and  Na- 
tional Research  Council  grant  A-1399). 

D.  A.  Craig,  B.  S.  Heming,  and  J.  G.  Packer  of  the  University  of  Alberta,  and  R.  L.  Wenzel 
of  the  Field  Museum  of  Natural  History,  read  and  criticized  this  manuscript  and  provided 


342 


Whitehead 


Fig.  259-260.  Evolutionary  zoogeography  of  the  milliped  genus  Cleptoria.  259.  As  suggested  by  R.  L.  Hoffman  (after 
Hoffman,  1967).  260.  As  reinterpreted.  Fig.  261.  Numbers  of  species  of  Schizogenius  in  5 intervals,  as  used  in  Table  70. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


343 


262 


264 


( 1000  mi ( 

Fig.  262.  Centers  of  concentration  of  species  of  the  genus  Schizogenius  in  North  and  Middle  America.  Fig.  263.  Areas 
of  concentration  of  species  of  the  genus  Schizogenius  in  North  and  Middle  America.  Fig.  264.  Areas  of  vicariance  for 
species  of  the  genus  Schizogenius  in  North  and  Middle  America;  shaded  area  represents  uplands. 


344 


Whitehead 


Fig.  265-269.  Evolutionary  zoogeography.  265.  Species  groups  of  truquii  lineage.  266.  Species  and  subspecies  of  pluri- 
punctatus  group.  267.  Species  of  tristriatus  group.  268.  Species  of  longipennis  group.  269.  Species  of  depressus  group. 


Classification,  phylogeny,  and  zoogeography  of  Schizogenius 


345 


numerous  useful  suggestions. 

Many  individuals  and  institutions  loaned  material.  I thank  them  all  heartily,  particularly 
those  who  made  special  efforts  to  secure  specimens  for  my  work.  I also  thank  D.  K.  Duncan 
and  F.  H.  Parker  of  Globe,  Arizona,  who  donated  specimens.  P.  J.  Darlington  and  P.  J. 
Spangler  were  courteous  and  helpful  hosts  when  I visited  the  Museum  of  Comparative  Zool- 
ogy and  the  United  States  National  Museum,  respectively. 

I am  particularly  indebted  to  numerous  colleagues  during  my  student  years  at  the  Univer- 
sity of  Alberta;  the  following  stand  out:  J.  Awram  and  A.  P.  Nimmo,  for  companionship  in 
the  field  as  well  as  continued  friendship;  B.  B.  Chiolino,  T.  L.  Erwin,  R.  Freitag,  H.  Goulet, 
D.  H.  Kavanaugh,  D.  J.  Larson,  and  R.  E.  Leech,  all  for  willingness  at  all  times  to  partake 
in  discussions. 


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Mirov,  N.  T.  1967.  The  genus  Pinus.  Ronald  Press,  New  York.  602  p. 

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fossil  mammal  fauna  of  South  America.  Quart.  Rev.  Biol.  43:409-45 1 . 

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Classification,  phytogeny,  and  zoogeography  of  Schizogenius 


349 


ANNOUNCEMENTS 


Biting  Fly  Control  and  Environmental  Quality  — Proceedings  of  a Symposium 
held  at  the  University  of  Alberta  in  Edmonton,  May  16,  17,  and  18,  1972. 

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Entomology  and  Education  — Proceedings  of  a Symposium  held 
at  the  University  of  Alberta  in  Edmonton,  May  19,  1972. 

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s. • o.  - Qg  7 V 


Quaestiones 


entomologicae 


MUS.  COMP.  ZOOL- 
LiBRARY 

NOV  3 1972 

harvard 

UNIVERSITY 


VOLUME  VIII 


A periodical  record  of  entomological  investigations, 
published  at  the  Department  of  Entomology, 
University  of  Alberta,  Edmonton,  Canada, 


NUMBER  4 OCTOBER  1972 


QUAESTIONES  ENTOMOLOGICAE 


A periodical  record  of  entomological  investigation  published  at  the  Department  of 
Entomology,  University  of  Alberta,  Edmonton,  Alberta. 

Volume  8 Number  4 1 October  1972 


CONTENTS 


Evans  — A temperature  controlled  capacitance-type  actograph  for 

cryptozoan  arthropods  351 

Doyen  — Familial  and  subfamilial  classification  of  the  Tenebrionoidea  (Coleoptera) 

and  a revised  generic  classification  of  the  Coniontini  (Tenebrionidae) 357 

Griffiths  — Studies  on  boreal  Agromyzidae  (Diptera).  II.  Phytomyza  miners 

on  Senecio,  Petasites  and  Tussilago  (Compositae,  Senecioneae) 377 

Announcement 406 


A TEMPERATURE  CONTROLLED  CAPACITANCE-TYPE 
ACTOGRAPH  FOR  CRYPTOZOAN  ARTHROPODS 

WILLIAM  G.  EVANS 
Department  of  Entomology 

University  of  Alberta  Quaestiones  entomologicae 

Edmonton,  Alberta  T6G  2E3  8 : 351-355  1972 

During  an  investigation  of  the  locomotor  activity  rhythms  of  intertidal  beetles  the  need 
arose  for  an  actograph  especially  suitable  for  these  animals  which  spend  a great  deal  of  then- 
time  in  crevices  or  under  stones.  The  actograph  also  needed  to  be  kept  at  a constant  temper- 
ature without  using  an  expensive  and  noisy  growth  chamber.  A review  of  the  literature  on 
the  many  kinds  of  actographs  used  with  insects  indicated  that  the  most  suitable  way  to 
detect  animal  movement  is  through  the  use  of  electronic  transducers.  In  the  capacitance- 
type  transducer  a movement  of  the  test  animal  causes  a change  in  capacitance  which  is 
detected  as  a frequency  change  and  amplified  into  a voltage  output  signal.  Therefore,  the 
electronic  circuit  of  the  sensing  apparatus  described  by  Schechter,  Dutky  and  Sullivan 
(1963)  and  later  modified  by  Grobbelaar  et  al.  (1967)  was,  with  minor  modification, 
adapted  for  the  actograph  described  here. 

The  sensors,  however,  differ  from  those  used  by  the  above  authors.  These  formed  a part 
of  two  parallel  5 mm  x 1 1.5  cm  x 1 1.5  cm  plexiglass  plates  between  which  the  insect  is 
placed  as  shown  in  Figure  1.  The  thin  wire  forming  the  electrode  on  the  bottom  of  the  top 
plate  is  positioned  to  form  a series  of  loops  starting  from  one  edge  of  a 10.5  cm  circle  and 
ending  at  the  other  edge  and  then  going  on  to  a ground  terminal.  In  contrast,  in  order  to 
obtain  a maximum  non-uniform  electric  field  between  the  two  plates,  the  other  electrode 
wire  consists  of  a 10.5  cm  spiral  starting  from  the  center  of  the  top  surface  of  the  bottom 
plate  and  eventually  leading  to  the  detector  circuit.  A change  in  capacitance  due  to  a move- 
ment of  the  test  insect  is  detected  at  the  junction  between  the  top  electrode  and  the  bottom 
one  so  that  the  insect  need  not  touch  either  of  the  wires.  The  closer  the  wire  spirals  or  the 
loops  or  the  distance  between  the  two  electrodes  the  more  sensitive  is  the  actograph  to  small 
movements.  The  3 1 gauge  electrode  wires  were  placed  into  grooves  scored  into  the  plates 
and  held  there  by  brushing  over  them  a thin  layer  of  plexiglass  dissolved  in  ethyl  acetate. 


352 


Evans 


The  thickness  of  the  spacer  is  determined  by  the  height  of  the  insect  so  that  ideally  the 
insect  is  able  to  move  freely  about  with  the  two  parallel  plates  as  close  as  possible  to  each 
other.  A series  of  plexiglass  spacers  were  made  ranging  in  thickness  from  0.25  to  3.5  mm 
and  any  of  these,  either  alone  or  in  combination,  can  be  used  for  insects  of  varying  heights. 
A test  insect  was  first  measured  for  height  before  placing  in  the  actograph  by  viewing  it  in 
the  bottom  of  a narrow  glass  vial  with  a micrometer  hand  lens. 

Sometimes  the  test  insect  is  kept  in  the  actograph  for  periods  of  10  to  14  days  and  in 
order  to  provide  the  high  humidity  and  water  necessary  for  hygrophilic  insects  two  methods 
were  employed,  both  involving  some  modification  of  the  bottom  plate.  Where  very  high 
humidity  was  necessary,  holes  were  drilled  between  the  wire  spirals  of  the  bottom  plate 
which  was  then  tightly  connected  with  screws  to  a plexiglass  container  of  water.  By  using  a 
graduated  glass  tube  connected  with  flexible  tubing  to  this  water  container  the  water  level 
could  be  adjusted  so  that  the  holes  in  the  bottom  plate  were  almost  filled  with  water.  Loss 
of  water  through  evaporation  was  measured  by  the  graduated  tube  and  water  was  added 
when  necessary  during  a test.  Where  high  humidity  is  not  critical  a cotton  wick  was  inserted 
into  a 2 cm  x 0.25  cm  inside  diameter  tube  which  was  inserted  into  the  center  of  the 
bottom  plate  and  suspended  over  a container  of  water.  The  top  of  the  wet  wick  just  barely 
protruded  above  the  top  of  the  bottom  plate.  A few  holes  can  be  drilled  between  the  wires 
in  the  top  plate  to  maintain  an  air  supply  and  a larger  hole  can  be  used  for  placing  the  insect 
into  the  space  between  the  plates.  This  larger  hole  is  then  corked  during  the  operation  of 
the  actograph. 

The  only  modification  of  the  electronic  circuit  described  by  Grobbelaar  et  al.  (1967) 
consisted  of  the  use  of  AC  power  with  two  regulated  power  supplies  instead  of  a battery 
source.  One  power  supply  (±  10  volts)  was  used  for  the  sensing  part  of  the  circuit  while  the 
other  (±15  volts)  powered  an  amplifier  and  a comparator  that  switched  a relay  at  selected 
output  voltage  peaks.  In  this  way  the  activity  record  could  consist  of  an  analog  tracing  or  a 
count  of  voltage  peaks  per  unit  time.  A potentiometer  allowed  a range  of  voltage  peaks  from 
0.1  volts  to  10  volts  to  switch  the  relay  thereby  providing  a means  of  sensitivity  adjustment. 
The  relay  signals  were  counted  and  printed  at  hourly  intervals  on  a Sodeco  counter-printer. 

Unless  walk-in  constant-temperature  chambers  are  available,  actographs  are  normally 
operated  in  constant-temperature  cabinets  such  as  plant  growth  chambers.  Unfortunately 
with  these  cabinets  the  cooling  compressor  is  often  exceedingly  noisy  and  the  very  notice- 
able on  and  off  cycle  of  the  compressor  could  influence  an  activity  rhythm  of  a test  animal. 
Figure  1 shows  a non-cyclical  constant-temperature  container  for  holding  the  actograph. 
The  container  consists  of  a fiberglass-encased  top  and  bottom  of  equal  dimensions  (34  cm  x 
34  cm  x 14  cm)  separated  by  two  13  mm  thick  foam  rubber  pads.  A length  of  6 mm  outside 
diameter  copper  tubing,  closely  coiled  in  circles  to  form  a 25  cm  inside  diameter  cylinder, 
was  placed  in  the  top  and  bottom  containers  with  inlet  and  outlet  connections  protruding 
from  the  sides.  The  spaces  between  the  tubing  and  the  walls  of  the  containers  were  filled 
with  poured-in  foam  insulation.  Provided  a styrofoam  cover  or  base  is  used  both  top  and 
bottom  containers  can  be  used  alone  or  they  can  be  used  together  as  shown  in  Figure  1. 
In  this  case  the  cables  from  the  temperature,  humidity  and  capacitance  sensors  as  well  as 
the  flexible  tubing  for  the  water  supply  fit  between  the  two  layers  of  foam  rubber.  The 
base  of  the  bottom  container  is  2.5  cm  thick  foam  insulation  encased  in  fiberglass  while 
the  top  of  the  top  container  which  can  be  used  for  viewing  the  actograph  consists  of  two 
sheets  of  6 mm  thick  plexiglass  separated  by  1.5  cm  of  air  space.  The  copper  tubing  is 
connected  to  a constant-temperature  circulating  bath  and  equilibrium  is  usually  obtained 
after  about  2 hours  depending  on  the  temperature  required  and  ambient  temperatures.  I 
have  been  using  two  complete  actographs  in  constant-temperature  containers  as  shown  in 


Actograph  for  cryptozoan  arthropods 


353 


Figure  1 connected  to  a Haake  circulating  water  bath  maintaining  a temperature  of  18.0° 
± 0.25°. 

An  example  of  the  kind  of  results  that  can  be  expected  with  this  type  of  actograph  is 
shown  in  Figure  2.  Specimens  of  Alphitobius  piceus  (Tenebrionidae)  were  kept  in  the  dark 
for  two  months  (group  A)  while  another  group  was  exposed  during  this  period  to  alternating 
12  hours  light  and  12  hours  darkness  (group  B).  After  this  time  five  specimens  of  each  group 
were  placed  in  the  actograph  under  dark  conditions  for  10  days.  As  is  evident  from  Figure  2 
the  group  kept  in  the  dark  for  two  months  were  arhythmic  while  the  other  group  main- 
tained a periodicity  of  23.75  hours  (periodogram  analysis;  see  Enright,  1965),  for  the  10 
days  of  activity  measurements. 

The  actograph  described  here  has  proved  to  be  very  suitable  for  carabid  beetles  (a  total 
of  18  species  representing  members  of  riparian,  forest  litter,  marsh,  intertidal  and  open- 
field  habitats  have  been  tested  so  far)  and  it  probably  can  be  used  successfully  with  other 
cryptozoan  animals  such  as  chilopods,  diplopods,  mites,  pseudoscorpions  as  well  as  beach- 
inhabiting  isopods  and  amphipods. 

ACKNOWLEDGEMENTS 

My  thanks  go  to  P.  Hardybala  and  W.  Diachuck  for  constructing  the  constant-temperature 
containers  and  the  actograph  respectively. 

REFERENCES 

Enright,  J.  T.  1965.  The  search  for  rhythmicity  in  biological  time  series.  J.  theor.  Biol.  8: 
426-468. 

Grobbelaar,  J.  H.,  G.  J.  Morris,  E.  E.  Baart  and  V.  C.  Moran.  1967.  A versatile,  highly  sensi- 
tive activity  recorder  for  insects.  J.  Insect  Physiol.  13:1843-1848. 

Schechter,  M.  S.,  S.  R.  Dutky  and  W.  N.  Sullivan.  1963.  Recording  circadian  rhythms  of  the 
cockroach  with  a capacity-sensing  device.  J.  econ.  Ent.,  56:76-79. 


354 


Evans 


double  clear 
plexiglass  top 


13mm  foam  rubbe 


top  plate 
(wire  loops  on  bottom) 


spacer 


6mm  copper  tubing 


to 


detector  circuit  ^ 


tube  for 
water  level 


13mm  foam  rubbe 


^^from 

circulating  bath 
to 


bottom  chamber 


to 

detector  circuit 
cork 


bottom  plate 
(wire  spiral  on  top) 


bottom  plate 
(wick  in  center) 


water  container 


from 


top  chamber 

^ to 

circulating  bath 


Figure  1.  An  exploded  view  of  the  sensors  of  the  capacitance-type  actograph  and  the  constant  temperature  containers. 


Actograph  for  cryptozoan  arthropods 


355 


RELATIVE 


ACTIVITY 


Figure  2.  A 10-day  activity  record  of  two  groups  of  five  specimens  of  Alphitobius  piceus  obtained  with  the  actograph 
in  darkness.  Group  A were  kept  in  the  dark  for  two  months  prior  to  the  test  while  group  B were  subjected  to  a DL 
12:12  at  the  same  time. 


TIME  IN  HOURS 


FAMILIAL  AND  SUBFAMILIAL  CLASSIFICATION  OF  THE 
TENEBRIONOIDEA  (COLEOPTERA)  AND  A REVISED 
GENERIC  CLASSIFICATION  OF  THE  CONIONTINI  (TENTYRIIDAE) 


JOHN  T.  DOYEN 
Division  of  Entomology 

University  of  California  Quaes tiones  entomologicae 

Berkeley,  California  94720  8 : 357-376  1972 


The  presence  of  visible  membranes  between  the  terminal  abdominal  stemites  of  Tenebri- 
onidae  and  related  families  is  correlated  with  the  presence  of  defensive  glands  which  empty 
posterad  of  sternite  7.  Species  which  lack  defensive  glands  consistently  have  the  aedeagus 
inverted  from  the  typical  position  in  most  Coleoptera.  These  complex,  correlated  characters 
indicate  a fundamental  division  among  the  Tenebrionoidea  and  several  changes  in  the  classi- 
fication of  these  beetles  are  proposed  accordingly. 

Morphological  comparisons  of  several  tribes,  principally  the  Coniontini,  Praocini  and 
Zophosini  are  summarized.  The  results  indicate  that  the  Zophosini  are  closely  related  to 
other  African  tribes  such  as  the  Adesmiini.  The  Praocini  show  close  affinities  to  the  South 
American  Nycteliini  and  Physogasterini.  The  North  American  Coniontini  and  Coelini  like- 
wise share  great  affinity.  While  the  taxa  from  these  geographic  regions  are  superficially 
similar,  they  differ  in  major  structural  features  and  are  not  closely  related. 

Finally,  taxonomic  changes  intended  to  simplify  the  classification  of  the  Coniontini  and 
Coelini  are  proposed,  with  a revised  generic  checklist  and  key. 

La  presence  de  membranes  visibles  entre  les  stemites  abdominales  terminates  des  Tene- 
brionidae  et  des  families  apparentees  est  en  rapport  avec  la  presence  de  glandes  defensives 
qui  s’ouvrent  derriere  le  sternite  7.  Les  especes  depourvues  regulierement  de  glandes  defen- 
sives ont  Taedeage  renverse  par  rapport  a la  position  normale  chez  Tensemble  des  coleop- 
teres.  La  correspondance  de  ces  carac teres  complexes  indique  une  division  fondamentale 
parmi  les  Tenebrionoidea  et  nous  proposons  en  consequence  plusieurs  modifications  dans  la 
classification  de  ces  coleop teres. 

Nous  resumons  les  comparaisons  morphologiques  entre  plusieurs  tribus  surtout  celles  des 
Coniontini,  des  Praocini  et  des  Zophosini.  Les  resultats  indiquent  que  les  Zophosini  sont  en 
rapport  etroit  avec  d’autres  tribus  africaines  telles  que  les  Adesmiini.  LI  y a des  affinites 
etroites  entre  les  Praocini  et  les  Nycteliini  et  les  Physogasterini  sud-americains.  Ces  rapports 
etroits  se  retrouvent  egalement  chez  les  Coniontini  et  les  Coelini  nord-americains.  Tandis 
que  les  groupes  de  ces  regions  geographiques  n accusent  que  des  ressemblances  superficielles, 
ils  different  par  leurs  carac  teristiques  structurales  fondamentales  et  ne  sont  pas  apparentes 
de  fagon  etroite. 

En  fin  de  compte,  nous  proposons  des  modifications  taxonomiques  qui  ont  pour  but  de 
simplifier  la  classification  des  Coniontini  et  des  Coelini  et  foumissons  a Vappui  une  liste 
generique  modifiee  ainsi  qu’une  clef. 

SUBFAMILY  CLASSIFICATION  OF  THE  TENEBRIONIDAE 

Watt  (1966)  reviewed  and  summarized  the  various  classifications  which  have  been  pro- 
posed for  the  Tenebrionidae.  Consequently,  only  the  most  important  works  will  be  men- 
tioned here.  Tenebrionidae  is  used  in  the  restricted  sense  of  Crowson  (1955)  and  Watt 


358 


Doyen 


(1966,  1967).  Based  on  the  structure  of  the  procoxae,  mesocoxae  and  aedeagus  they  ex- 
clude such  genera  as  Boros,  Dacoderus,  Tretothorax , Perimylops  and  Zopherus  et  al.  Watt 
(1967)  also  cites  evidence  indicating  a close  relationship  of  the  Alleculidae,  Lagriidae  and 
Nilionidae  to  the  Tenebrionidae.  These  families,  together  with  the  Zopheridae  and  Monom- 
midae,  are  referred  to  here  as  Tenebrionoidea.  In  his  analysis  of  the  taxonomic  position  of 
Petria  (Alleculidae:  Omophlinae)  Lawrence  (1971)  also  mentions  several  characters  which 
are  important  in  the  classification  of  the  Tenebrionoidea. 

The  most  recent  classification  of  the  Tenebrionidae  treating  the  world  fauna  is  that  of 
Lacordaire  (1859).  This  century-old  work  emphasizes  external  differences,  especially  in 
mesocoxal  structure  and  mouthparts.  In  a broad  sense,  Lacordaire’s  primary  divisions  into 
Sections  and  Cohortes  correspond  to  the  subfamilies  of  more  recent  classifications.  How- 
ever, the  Cohortes  of  his  Section  II  are  based  on  the  vestiture  of  the  tarsi.  As  noted  by  Watt 
(1966),  this  feature  is  probably  related  to  the  substrate  on  which  the  beetles  walk,  and  is 
not  a reliable  taxonomic  character.  Consequently,  Lacordaire’s  Section  II,  Cohorte  I con- 
tains an  assortment  of  unrelated  tribes  now  assigned  to  different  subfamilies.  It  should  be 
mentioned  that  nearly  all  Lacordaire’s  tribes  and  subtribes  are  recognized  as  tribes  today. 
Many  of  the  numerous  tribes  proposed  by  subsequent  workers,  especially  Casey  (1907, 
1908)  and  Reitter  (1917)  contain  very  few  genera.  These  small  tribes  have  served  mainly  to 
occlude  the  interrelationships  within  the  family. 

LeConte  (1862),  LeConte  and  Horn  (1883)  and  Horn  (1870),  considering  primarily  the 
North  American  fauna,  recognized  three  subfamilies.  Their  most  important  contribution  was 
an  appreciation  of  the  taxonomic  importance  of  the  intersegmental  membranes  between  the 
terminal  abdominal  sternites.  They  delimited  the  Tenebrioninae  by  the  presence  of  external 
intersegmental  membranes.  The  remainder  of  the  family,  with  internal  membranes,  they 
divided  between  the  Tentyriinae  (mesocoxal  cavities  enclosed  by  the  sterna;  mesotrochantin 
concealed)  and  the  Asidinae  (mesocoxal  cavities  open  laterally;  mesotrochantin  visible). 

Most  recent  classifications  (Gebien,  1910-11,  1937,  1938-44;  Arnett,  1960)  combine 
features  of  the  arrangements  of  LeConte  and  Lacordaire,  and  differ  from  one  another  in 
minor  ways.  An  exception  is  the  proposal  by  Koch  (1955)  to  divide  the  Tenebrionidae  into 
only  two  subfamilies,  depending  on  the  condition  of  the  membranes  between  the  terminal 
abdominal  sternites.  On  this  basis,  Koch  combined  the  Asidinae  with  the  Tentyriinae,  also 
pointing  out  that  the  mesocoxal  structure  of  the  African  Asidini  does  not  conform  to 
LeConte  and  Horn’s  criterion  of  the  presence  of  a distinct  trochantin.  Brown  (1971)  has 
shown  that  the  condition  of  the  trochantin  is  also  variable  in  the  North  American  Asidini. 
Koch’s  classification  is  supported  by  the  evidence  presented  below,  and  the  name  Tentyri- 
inae will  be  used  in  the  sense  he  advocated. 

MORPHOLOGICAL  AND  ECOLOGICAL  CHARACTERISTICS 
OF  THE  TENEBRIONOIDEA 

Adults 

Although  most  recent  workers  agree  that  the  presence  of  external  abdominal  membranes 
is  important  as  a diagnostic  character,  the  high  correlation  with  the  presence  of  abdominal 
glands  and  reservoirs  has  not  been  noticed.  The  reservoirs  are  paired,  cuticular  invaginations 
surrounded  by  diffuse  glandular  tissue.  They  empty  through  ducts  which  open  posterad  of 
sternite  7 (visible  sternite  5).  Morphological  studies  by  Blumberg  (1961),  Eisner,  McHenry, 
and  Salpeter  (1964)  and  Kendall  (1968)  show  that  the  glands  and  reservoirs  are  apparently 
homologous  throughout  the  Tenebrionidae,  Alleculidae  and  Lagriidae.  Dissections  made 
by  the  author  reveal  that  similar  glands  are  also  present  in  the  Nilionidae.  The  secretions 


Classification  of  Tenebrionoidea 


359 


produced  by  these  glands  are  probably  defensive  since  they  consist  largely  of  quinones 
and  other  irritants  (see  reviews  by  Eisner  and  Meinwald,  1966;  Schildknecht  et  al.,  1964). 
The  structural  details  of  the  glands  and  reservoirs  are  the  subject  of  a comprehensive  com- 
parative study  (Tschinkel,  in  progress ) which  may  clarify  the  relationships  among  those 
Tenebrionidae  which  possess  glands. 

A survey  by  the  author  of  over  45  tribes  of  Tenebrionidae,  as  well  as  members  of  the 
other  families  mentioned  above,  has  revealed  a nearly  perfect  correlation  between  the  pres- 
ence of  glands  and  external  membranes  between  abdominal  sternites  5,  6,  and  7.  The  single 
exception  is  the  Pimeliini,  where  membranes  are  visible,  as  noted  by  Watt  (1966),  but 
reservoirs  and  glands  are  absent. 

A further  correlation  exists  between  the  occurrence  of  glands  and  the  orientation  of  the 
aedeagus  in  the  retracted  position.  As  first  noted  by  Sharp  and  Muir  (1912)  and  elaborated 
by  Blaisdell  (1939),  the  aedeagus  is  rotated  180°  in  some  Tenebrionidae,  so  that  the  primi- 
tively ventral  surface  is  dorsal.  This  rotation,  which  also  occurs  in  the  Dacoderidae  (Watt, 
1967),  some  Salpingidae  (Spilman,  1952)  and  the  Monommidae  (Sharp  and  Muir,  1912)  is 
rather  inappropriately  termed  inversion  by  coleopterists.  With  one  known  exception,  Tene- 
brionidae with  the  aedeagus  inverted  lack  defensive  glands,  while  species  with  the  aedeagus 
in  the  normal  position  possess  them.  The  exception  is  the  Cossyphini,  a small  Palearctic- 
African  tribe  whose  members  are  specialized  for  living  beneath  bark.  In  these  beetles  the 
aedeagus  is  oriented  with  the  tegmen  dorsad,  but  defensive  glands  are  absent  and  the  termi- 
nal abdominal  membranes  are  internal.  This  tribe  is  highly  modified  morphologically,  parti- 
cularly in  thoracic  structure,  and  more  detailed  studies  will  be  required  to  clarify  its  rela- 
tionships. Possibly  the  defensive  glands  have  been  lost  secondarily. 

The  very  high  correlations  among  these  highly  complex  structures  is  a fundamental  differ- 
ence separating  the  Tenebrionoidea  into  two  distinct  groups.  The  tenebrionid  subfamily 
Tenebrioninae,  together  with  the  Alleculidae,  Lagriidae  and  Nilionidae  possess  defensive 
glands  and  external  abdominal  membranes  and  have  the  aedeagus  in  the  normal  position. 
The  subfamily  Tentyriinae  (sensu  Koch)  and  the  Monommidae  lack  defensive  glands,  have 
the  terminal  abdominal  membranes  internalized,  and  have  the  aedeagus  inverted.  In  addi- 
tion, these  two  groups  seem  to  have  evolved  in  different  ecological  situations.  The  Tenebri- 
oninae, etc.,  are  predominant  in  woodland  habitats  in  tropical  or  subtropical  climates,  and 
many  species  are  adapted  to  feeding  on  fungi  or  in  rotting  wood.  Most  of  the  species  of 
this  group  which  occur  in  temperate  regions  occupy  relatively  mesic,  woodland  habitats. 
Contrastingly,  the  Tentyriinae  occur  primarily  in  arid  or  subarid  habitats  in  temperate 
climates,  with  highly  distinct  faunas  in  the  deserts  of  southern  Africa,  Eurasia,  South  Ameri- 
ca, and  southwestern  North  America.  Nearly  all  of  the  species  in  this  group  are  soil  dwellers. 
There  are  exceptions  to  these  generalizations,  such  as  the  Eleodini  (Nearctic)  and  the  Blap- 
tini  (Palearctic),  which  contain  many  species  adapted  to  aridity,  but  clearly  belong  to  the 
Tenebrioninae.  Conversely,  the  Epitragini  (Tentyriinae)  are  widespread  in  tropical  regions, 
especially  in  the  western  hemisphere.  These  exceptions,  coupled  with  the  fact  that  the 
more  generalized  members  of  both  subfamilies  are  winged,  clearly  show  that  loss  of  defen- 
sive glands  and  flight  are  not  simply  associated  with  adaptation  to  arid  environments.  In- 
deed, the  diverse  array  of  components  in  the  defensive  secretions  (Tschinkel,  in  progress ), 
and  the  variability  of  the  associated  delivery  systems,  especially  in  those  Tenebrioninae 
inhabiting  arid  environments,  suggests  that  the  secretions  have  been  very  important  in  the 
evolution  of  these  beetles.  For  instance,  some  species  of  Blaps,  Eleodes  and  Centronopus 
are  capable  of  spraying  fine  jets  of  secretion  up  to  30  cm.  The  elytra  of  other  species,  such 
as  Cibdelis  blaschkei  Mannerheim  are  impressed  with  fine  canals  along  which  the  secretions 
flow,  rapidly  coating  the  posterior  portion  of  the  body. 


360 


Doyen 


Larvae 

The  great  majority  of  tenebrionid  larvae  which  have  been  associated  with  adults  are  in 
the  subfamily  Tenebrioninae  ( see  Korschevsky,  1943;  Van  Emden,  1947;  Hayashi,  1966, 
1968).  Recently,  larvae  of  several  tentyriine  tribes  have  been  adequately  characterized, 
including  Erodiini,  Akidini,  Epitragini,  Tentyriini,  and  Asidini,  chiefly  by  Russian  workers 
(Keleynokova,  1963,  1971;  Skopin,  1960,  1962,  1964).  Schulze  (1962,  1964)  and  Marcuzzi 
and  Rampazzo  (1960)  have  described  larvae  from  some  additional  tribes  (Lepidochorini, 
Adesmiini,  Coniontini,  Coelini).  Skopin  (1964)  and  Keleynokova  (1963)  have  attempted 
to  use  their  results  to  produce  larval  classifications.  Skopin’s  primary  division  splits  the 
Tenebrionoidea  into  two  groups,  based  on  the  structure  of  the  legs.  His  Pedobionta  in- 
cludes all  soil  inhabiting  larvae,  in  which  the  anterior  legs  are  enlarged  and  modified  for 
digging.  The  Pedobionta  correspond  to  the  Tentyriinae,  with  the  addition  of  the  Blap- 
tini,  Opatrini  and  Platyscelini  of  the  Tenebrioninae.  The  Eleodini  and  Scaurini  would  also 
belong  to  the  Pedobionta  on  the  basis  of  leg  structure.  Skopin’s  Heterobionta  incorpo- 
rates the  remainder  of  the  Tenebrioninae  as  well  as  the  families  Alleculidae,  Lagriidae  and 
Nilionidae. 

Clearly,  Skopin’s  primary  division  is  based  on  a highly  adaptive  feature.  Keleynokova 
(1963)  points  out  that  soil  inhabiting  larvae  are  characterized  by  the  absence  of  urogomphi 
as  well  as  enlarged  forelegs.  Urogomphi  are  commonly  present  in  the  Tenebrioninae,  and 
are  apparently  used  in  moving  backward  through  the  tunnels  these  larvae  excavate  in  lig- 
neous substrates.  In  Tenebrioninae  which  inhabit  soil  the  urogomphi  are  reduced  or  absent 
(e.g.,  Blaptini,  Eleodini,  Pedinini).  Tenebrio  (Tenebrionini)  is  exemplary  in  this  regard. 
Species  of  this  genus  infest  stored  grain  products,  which  are  probably  similar  to  soil  in 
physical  properties.  The  anterior  legs  of  Tenebrio  are  enlarged  and  the  urogomphi  are  much 
reduced.  For  these  reasons,  Skopin’s  placement  of  the  Blaptini,  Opatrini,  etc.,  in  the  Pedo- 
bionta is  judged  in  error.  The  same  conclusion  was  reached  by  Keleynokova  (1963),  who 
placed  these  tribes  in  a separate  subfamily  within  the  “tenebrioid  line.”  In  most  respects, 
however,  Keleynokova’s  classification  into  six  subfamilies  does  not  correspond  to  relation- 
ships indicated  by  adults.  As  noted  by  Watt  (1966),  her  subfamilies  are  not  clearly  defined, 
and  it  is  impossible  to  evaluate  them  at  present. 

One  larval  characteristic  which  is  not  stressed  by  Skopin  or  Keleynokova  is  the  configura- 
tion of  the  mandibles.  In  the  Tentyriinae  the  larval  mandibles  bear  a dorsolateral  promi- 
nence which  is  densely  set  with  coarse  setae.  Setae  on  the  mandibles  of  the  Tenebrioninae, 
Alleculidae,  etc.,  are  never  restricted  to  an  elevated,  dorsolateral  region.  The  distribution  of 
these  character  states  is  very  highly  correlated  with  the  presence  of  defensive  glands  and 
the  orientation  of  the  aedeagus,  and  further  supports  a primary  division  of  the  Tenebrionoi- 
dea into  two  taxa. 

Another  character  which  may  be  of  taxonomic  importance  is  egg  size.  In  the  few  tribes 
which  have  been  investigated,  the  eggs  are  relatively  large  in  the  Tentyriinae  (Asidini,  Coni- 
ontini, Nyctoporini,  Cryptoglossini)  and  small  in  the  Tenebrioninae  (Tenebrionini,  Coelo- 
metopini,  Ulomini)  (Doyen,  unpublished ). 

Skopin’s  (1964)  subdivisions  of  his  Heterobionta  suggest  several  interesting  relationships 
which  reinforce  the  conclusions  here  derived  from  adult  characteristics.  His  “Ulomimorpha” 
includes  the  Alleculidae  as  well  as  the  Ulomini,  and  he  flatly  states  that  these  should  be 
placed  in  the  same  family  on  the  basis  of  larval  features.  His  “Pycnocerimorpha”  includes 
the  Goniaderini,  Heterotarsini  and  Pycnocerini,  tribes  which  have  always  been  placed  in  the 
Tenebrioninae.  On  the  basis  of  larval  features,  however,  they  are  very  similar  to  the  Lagrii- 
dae. Hayashi  (1968)  also  reached  this  conclusion  from  his  studies  of  Japanese  members  of 
these  taxa. 


Classification  of  Tenebrionoidea 


361 


Evidence  concerning  the  Nilionidae  is  not  clear-cut.  The  adults  greatly  resemble  members 
of  the  Leiochrini  (Tenebrioninae),  and  Boving  and  Craighead’s  larval  characterization  is 
based  on  Leiochrodes  sp.  The  characters  listed  by  Skopin  (1964)  for  the  Nilionidae  fit 
known  larvae  of  the  Leiochrini.  However,  he  does  not  specify  the  taxa  on  which  his  con- 
cept is  based,  and  the  larva  of  Nilio  appears  to  be  unknown. 

RECLASSIFICATION  OF  THE  TENEBRIONOIDEA 

The  major  morphological  and  ecological  differences  described  above,  and  the  extreme 
diversity  in  body  form  in  both  subfamilies,  suggest  that  they  have  been  evolving  separately 
for  a long  time,  and  that  the  Tenebrionidae  as  now  constituted  are  probably  polyphyletic. 
Biogeographic  evidence  also  lends  tentative  support  to  this  conclusion.  The  Tenebrioninae, 
Alleculidae  and  Lagriidae  enjoy  a world-wide  distribution,  especially  in  the  tropics.  Con- 
trastingly, the  Tentyriinae,  although  widely  distributed  in  temperate  regions,  are  almost 
entirely  absent  from  Australia  and  New  Zealand,  where  a few  tribes  of  the  Tenebrioninae 
have  radiated  extensively  into  arid  habitats.  The  few  Australian  genera  previously  included 
in  the  Tentyriinae  have  mostly  been  removed  to  other  families  (e.g.,  Tretothorax,  Zophero- 
sis : see  Boving  and  Craighead,  1931;  Crowson,  1955;  Watt,  1966,  1967).  These  distri- 
butions suggest  that  the  evolutionary  line  producing  the  Tenebrioninae  arose  before  the 
separation  of  the  Australian  land  mass.  The  Tentyriinae  apparently  differentiated  after 
the  separation  of  Australia,  probably  in  southern  Africa,  which  unquestionably  supports 
the  most  diverse  and  distinct  fauna  now  known  (Koch,  1955). 

If  the  correlated  differences  described  above  are  to  be  reflected  in  the  classification  of 
the  Tenebrionoidea,  the  Tenebrioninae,  Alleculidae,  Lagriidae  and  Nilionidae  should  be 
treated  as  a single  taxon,  coordinate  with  the  Tentyriinae.  I feel  that  these  relationships 
are  best  reflected  by  recognizing  a family  Tentyriidae  and  placing  the  Alleculidae,  Lagriidae, 
Nilionidae  and  Tenebrioninae  as  subfamilies  of  the  Tenebrionidae.  The  Monommidae,  which 
share  most  of  the  characters  of  the  Tentyriidae  are  differentiated  by  having  all  the  abdomi- 
nal sternites  flexibly  connected  by  internal  membranes  and  the  front  coxal  cavities  open, 
and  should  clearly  be  recognized  at  the  family  level.  The  proposed  arrangement  is  compared 
with  previous  classifications  in  Table  1. 

In  addition,  a number  of  tribes  are  incorrectly  placed  in  the  Tenebrioninae  in  recent 
classifications  (Gebien,  1938-44;  Arnett,  1960).  According  to  the  criteria  described  above, 
the  Coniontini,  Coelini,  Branchini,  Physogasterini,  Praocini  and  Pimeliini  are  members  of 
the  Tentyriidae.  Interestingly,  all  the  early  American  workers  agreed  that  these  tribes  be- 
longed to  the  Tentyriinae  (Blaisdell,  1939;  Casey,  1908;  Horn,  1870;  LeConte  and  Horn, 
1883).  In  addition  the  genera  Eupsophulus  Cockerell  and  Alaephus  Horn,  currently  placed 
in  the  Tenebrioninae  (Tenebrionini),  clearly  belong  to  the  Tentyriidae,  although  their  exact 
affinities  are  uncertain.  Horn  (1870)  realized  that  these  genera  exhibited  characteristics  of 
both  subfamilies  and  judged  their  classification  as  tentative. 

It  must  be  emphasized  that  the  infrafamilial  classification  of  these  beetles  remains  in  a 
confused  state.  The  numerous  tribes  of  the  Tentyriidae  are  frequently  very  distinct  and 
without  intermediates  (e.g.,  Stenosini,  Triorophini,  Nyctoporini).  Conversely,  while  some 
of  the  tribes  assigned  to  the  Tenebrioninae  are  disjunct  (e.g.,  Diaperini,  Scaurini,  Cossy- 
phini),  many  are  founded  on  superficial  characters  which  have  arisen  independently  many 
times  (e.g.,  Coelometopini,  which  are  distinguished  from  the  Tenebrionini  by  being  apter- 
ous). 


362 


Doyen 


Table  1.  Comparison  of  classifications  of  the  Tenebrionoidea.* 


Lacordaire,  1859  LeConte  and  Gebien  Koch,  1955 

Horn,  1883  1937-44 


Tentyriidae 

Tenebrionides 
(Section  1; 
Section  II, 
Cohorte  I, 
in  part) 

Tentyriinae, 

Asidinae 

Tentyriinae, 
Asidinae  & 
Coniontini, 
Praocini, 
Pimeliini,  etc. 

Tentyriinae 

Tenebrionidae 

Tenebrioninae 

Tenebrionides 
(Section  II, 
Cohorte  I, 
in  part; 
Cohorte  II) 

Tenebrioninae 

Tenebrioninae 

Tenebrioninae 

Alleculinae 

Cistelides 

(=Alleculidae) 

Cistelidae 

(=Alleculidae) 

[Alleculidae] 

[Alleculidae] 

Lagriinae 

Lagriides 

Lagriidae 

[Lagriidae] 

[Lagriidae] 

Nilioninae 

Nilionides 

[Nilionidae] 

Leiochrini  & 
[Nilionidae] 

[Leiochrini  & 
Nilionidae] 

* The  arrangement  proposed  here  is  listed  on  the  left.  The  characters  on  which  this  classi- 
fication is  based  are  described  fully  in  the  text.  The  Monommidae,  considered  a distinct 
family  by  all  workers,  is  not  included.  Names  in  brackets  are  not  specifically  mentioned  in 
the  works  cited  because  these  taxa  were  outside  the  geographic  or  taxonomic  scope  of  the 
respective  classifications. 


AFFINITIES  OF  THE  CONIONTINI,  COELINI  AND  BRANCHINI 

Say  (1824)  in  his  original  description  of  Eusattus  reticulatus,  assigned  it  to  Zophosis 
(Zophosini),  an  old  world  genus  that  is  superficially  similar.  Subsequently,  Casey  (1908) 
speculated  that  the  Branching  Coniontini,  Zophosini,  Praocini  and  Nycteliinae  comprise 
a closely  related  group  which  he  designated  the  Coniontinae.  LeConte  and  Horn  (1883) 
and  Champion  (1884)  also  noticed  the  superficial  similarity  of  the  Branchini  to  Praocis 
and  Nyctelia,  and  suggested  a relationship  to  these  South  American  tribes.  In  the  present 
study,  mouthparts,  male  and  female  genitalia,  and  internal  thoracic  structures  were  com- 
pared among  the  following  taxa  in  order  to  reassess  their  interrelationships. 

Coniontini 

Coniontides  latus  LeConte 
Coniontis  viatica  Eschscholtz 


Classification  of  Tenebrionoidea 


363 


Coniontellus  inflatus  Casey 

Coelo taxis  punctulata  Horn 

Coelosattus  fortineri  Blaisdell 

Eusattus  robustus  LeConte 
erosus  Horn 
dubius  LeConte 
reticulatus  (Say) 
muricatus  LeConte 

Coelini 

Coelus  globosus  LeConte 
ciliatus  Eschscholtz 
remotus  Casey 

Coelomorpha  maritima  Casey 
Zophosini 

Zophosis  plana  Fabricius 
Praocini 

Praocis  chiliensis  (Gray)  (Det.  L.  E.  Pena) 
penai  Kulzer  (Det.  L.  E.  Pena) 
pilula  Laporte  (Det.  L.  E.  Pena) 

Nycteliini 

Nyctelia  varipes  Fairmaire  (Det.  L.  E.  Pena) 

Gyriosomus  modestus  Kulzer  (Det.  L.  E.  Pena) 

Physogasterini 

Entomochilus  varius  laevis  Kulzer  (Det.  L.  E.  Pena) 

Branchini 

Branchus  floridanus  LeConte 

Branchus  woodii  LeConte 

Oxinthas  praocioides  Champion 

MOUTHPARTS 

The  mentum  is  typically  large  in  most  Tentyriidae,  concealing  the  maxillae  and  ligula. 
which  is  usually  membranous.  In  the  Coniontini  and  Coelini  the  mentum  is  relatively  small 
exposing  the  maxillae  and  ligula,  which  is  always  ventrally  sclerotized  and  articulated  with 
the  mentum  by  a narrow  membrane  (Fig.  1).  The  labial  palp  hinges  with  a sclerotized 
palpifer  which  is  embedded  dorsally  in  the  membrane  above  the  ligular  articulation.  The 
only  appreciable  variation  in  these  structures  involves  the  size  and  pattern  of  the  setae  on 
the  dorsal  surface  of  the  ligula.  The  coarse  bristles  shown  in  Fig.  1 are  characteristic  of 
several  fossorial  species  of  Eusattus  as  well  as  Coelus.  In  more  generalized  species  of  Eusat- 
tus and  in  Coniontis  the  ligular  setae  are  much  finer,  more  numerous  and  brush-like,  as  in 
Branchus  (Fig.  2).  In  general  shape  and  morphology  the  labial  structure  of  Branchus  shows 
no  important  differences  from  the  Coniontini. 

The  mentum  of  the  Physogasterini  and  Praocini  is  relatively  smaller,  compared  to  the 
ligula,  which  is  articulated  by  a broad  membranous  band  (Fig.  3).  Portions  of  the  ventral 
surface  of  the  ligula  are  usually  membranous,  especially  in  the  Praocini.  The  greatest  diver- 
gence from  the  Coniontini  involves  the  insertion  of  the  labial  palps.  Whereas  these  are 
attached  dorsally  to  sclerotized  palpifers  in  the  Coniontini  they  are  articulated  with  the 
ventral  (external)  surface  of  the  ligula  in  the  Praocini  and  Physogasterini,  and  the  palpifer 
is  absent.  The  labial  structure  is  very  similar  in  the  Nycteliini,  but  in  Nyctelia  the  ligula 


364 


Doyen 


is  relatively  small  and  retracted  beneath  the  mentum.  In  Gyriosomus  the  ligula  is  large  and 
protuberant,  as  in  the  Praocini. 

The  mouthparts  of  Zophosis  (Fig.  4)  are  distinctly  tentyrioid  in  structure.  The  mentum 
is  large,  concealing  the  ligula  and  the  maxillae.  The  membranous  ligula  is  relatively  small, 
with  the  labial  palps  articulated  dorsally.  The  palpifer  is  absent.  A similar  structural  arrange- 
ment occurs  in  many  other  tribes  of  Tentyriidae,  including  the  Tentyriini,  Adesmiini,  and 
Triorophini.  Zophosis  also  differs  from  the  other  taxa  considered  here  in  the  structure 
of  the  maxilla.  In  these  other  tribes,  the  lacinia  is  bidentate,  and  the  galea  brush-like.  In 
Zophosis,  both  lacinia  and  galea  are  densely  setate  and  brush-like. 

GENITALIA 

The  aedeagus  and  penis  of  most  Tentyriidae  are  simple  fusiform  tubes  (Koch,  1955). 
This  structure  is  exemplified  by  the  Coniontini  and  Coelini.  In  these  tribes  the  tegmen 
consists  of  lateral  struts,  connected  proximally.  The  sclerotized  parts  of  the  penis  are 
narrow,  lateral  rods  (Fig.  5,  6).  In  Branchus  floridanus  (Fig.  7,  8)  the  tegmen  is  a sclerotic 
tube,  with  only  a small  ventral  membrane  (the  homologous  membrane  is  dorsal  in  the 
Tenebrionidae).  The  paramere  is  strongly  ridged  proximally,  and  is  apically  truncate,  with 
sharp  lateral  spurs.  The  penis  is  exceptionally  elongate  and  distally  curved  and  enlarged. 
The  functional  significance  of  these  remarkably  modified  structures  is  unknown.  In  Bran- 
chus woodii  and  Oxinthas  the  male  genitalia  are  essentially  similar  to  those  of  the  Coni- 
ontini. The  Praocini,  Nycteliini  and  Physogasterini  are  very  similar  to  the  Coniontini  in 
respect  to  male  genitalia,  with  minor  differences  in  the  degree  of  sclerotization  and  shape 
and  proportions  of  the  aedeagus  and  penis.  It  may  be  significant  that  the  aedeagus  bears 
lateral,  subterminal  patches  of  setae  in  all  four  tribes.  The  aedeagus  and  penis  of  Zo- 
phosis (not  illustrated)  are  relatively  much  shorter  and  thicker  and  lack  the  subterminal 
setae. 

Female  genitalia  in  these  beetles  consist  of  an  elongate  1st  valvifer  and  short,  distally 
spatulate  and  strongly  sclerotized  2nd  valvifer  (Fig.  9-11).  The  2nd  valvifer  is  distally  modi- 
fied as  a sclerotized  process  in  many  Tentyriidae  (e.g.,  Adesmiini,  Asidini,  Cryptoglossini), 
probably  for  penetrating  the  oviposition  substrate,  and  frequently  the  ovipositor  tube  is 
very  elongate  as  compared  to  that  of  the  Tenebrionidae.  In  the  Coniontini,  Coelini  and 
Branchini  the  1st  valvifer  is  a weakly  sclerotized  plate  with  a marginal  baculus  (Fig.  9). 
The  second  valvifer  is  continuously  sclerotized,  with  the  distal  process  oriented  horizon- 
tally. A subterminal  membranous  area  marks  the  position  of  the  gonostylus,  which  is 
recognizable  as  a group  of  elongate  setae.  This  configuration  is  nearly  constant  throughout 
these  tribes,  the  only  significant  variation  involving  size  and  slight  differences  in  shape. 
The  ovipositor  is  similar  in  the  Praocini,  Physogasterini  and  Nyctelia  (Fig.  10)  with  the 
following  differences.  The  ovipositor  tube  is  usually  more  elongate  and  the  baculus  of 
the  1st  valvifer  is  submarginal.  The  2nd  valvifer  is  oriented  obliquely  or  nearly  vertically 
and  bears  a sulcus  about  two-thirds  of  the  distance  to  the  base.  Two  features  shared  with 
the  Coniontini  are  the  setal  clothing  of  the  2nd  valvifer  and  the  median,  ventral  sclerite 
situated  in  the  membrane  between  the  2nd  valvifers.  In  Gyriosomus  (Nycteliini)  the  entire 
ovipositor  is  densely  setate,  the  sulcus  on  the  2nd  valvifer  is  very  strong,  and  the  median 
ventral  sclerite  is  absent. 

The  ovipositor  of  Zophosis  (Fig.  1 1 ) is  relatively  short  and  thick.  Both  valvifers  are 
densely  setate  and  the  2nd  valvifer  consists  of  a basal  sclerotized  plate  and  baculus  with 
a narrow,  lateral  sclerotization  articulating  with  the  strongly  sclerotized,  terminal  process. 
There  are  no  suggestions  of  the  gonostylus  or  the  median,  dorsal  sclerite. 


Classification  of  Tenebrionoidea 


365 


Fig.  1-4.  Labial  structures.  The  left  side  of  each  figure  represents  the  ventral  (external)  surface.  The  right  side  repre- 
sents the  dorsal  (internal)  surface.  1,  Eusattus  muricatus;  2,  Branchus  floridanus-,  3,  Praocis  penar,  4,  Zophosis  plana. 


366 


Doyen 


Fig.  5-8.  Male  genitalia.  5,  Eusattus  reticulatus,  ventral  aspect  of  aedeagus;  6,  Eusattus  reticulatus,  lateral  aspect  of 
aedeagus  and  penis;  7,  Bronchus  floridanus,  ventral  aspect  of  aedeagus;  8,  Bronchus  floridanus,  lateral  aspect  of  aedeagus 
and  penis.  Fig.  9-11.  Ventral  aspect  of  ovipositors.  9,  Eusattus  dubius\  10,  Praocis  penai\  11,  Zophosis  plana. 


Classification  of  Tenebrionoidea 


367 


INTERNAL  THORACIC  STRUCTURE 

Flightless  tenebrionids  and  tentyriids  illustrate  extreme  modifications  of  thoracic  struc- 
ture. Smith  (1964)  described  reductions  of  flight  musculature  and  the  accompanying  de- 
sclerotization  and  reduction  in  size  of  the  metathoracic  terga  in  a number  of  micropterous 
and  apterous  beetles.  However,  most  of  his  examples  appear  to  represent  relatively  early 
stages  in  the  specialization  for  ambulatory  life.  Many  groups  of  Tentyriidae  have  apparently 
been  apterous  for  a very  long  time.  In  these  the  metanotum  is  completely  membranous 
and  the  mesonotum  is  reduced  to  a narrow,  leathery,  transverse  sclerite  which  extrudes 
externally  as  the  scutellum.  In  highly  modified  forms  (e.g.,  Edrotes,  Epiphysa;  Doyen, 
1968)  the  mesonotum  is  further  reduced,  with  no  external  indication  of  the  scutellum,  and 
the  mesosterna  and  prosterna  are  fused  by  cuticular  extensions  of  the  sternal  apophyses. 
These  specializations  are  usually  accompanied  by  elongation  and  thickening  of  the  metendo- 
stemite,  which  frequently  becomes  fused  with  the  mesocoxal  inflections  and  the  meso- 
pleura,  especially  in  fossorial  species. 

Most  of  the  skeletal  adaptations  described  above  are  represented  in  the  taxa  discussed 
here.  In  the  Coniontini,  Coelini  and  Branchini  the  arms  of  the  metendostemite  are  extreme- 
ly elongate  and  approximated  to  the  mesocoxal  inflections,  or  fused  to  the  inflections  in 
more  highly  modified  species  (Fig.  12,  Table  2).  The  arms  extend  to  the  vicinity  of  the 
mesepisterna,  terminating  in  muscle  disks  which  are  fused  with  the  episterna  in  the  most 
specialized  species.  The  mesapophyseal  arms  are  relatively  short,  and  may  be  expanded  as 
vertically  oriented  flanges  in  fossorial  species  (Fig.  12).  Structural  details  of  the  pterothorax 
of  various  species  of  these  tribes  are  summarized  in  Table  2. 

The  Praocini,  Nycteliini  and  Physogasterini  share  a distinct  thoracic  structure.  The  arms 
of  the  metendostemite  are  broadly  fused  with  the  mesocoxal  inflections,  but  terminate  as 
short  prongs  without  terminal  muscle  disks,  and  never  approach  the  mesopleura  (Fig.  13). 
The  elongate,  slender  arms  of  the  mesendostemite  extend  dorsolaterally  almost  to  the 
dorsal  margin  of  the  mesepisternum.  In  Nyctelia  and  Gyriosomus  the  mesothorax  and 
prothorax  are  rigidly  joined  by  strong,  ligamentous  thickenings  of  the  intersegmental  mem- 
brane, while  in  Praocis  and  Entomochilus  the  prothorax  is  relatively  mobile  as  in  the 
Coniontini. 

The  thoracic  modifications  of  Zophosis  (Fig.  14)  are  strikingly  similar  to  those  of  Ades- 
mia  and  Epiphysa  ( see  Doyen,  1968).  The  metendostemite  is  fused  with  the  mesocoxal 
inflections  and  the  arms  extend  anterodorsally  to  the  mesopleura,  terminating  in  large 
muscle  disks,  but  are  not  fused  with  the  mesopleura.  The  mesendostemite  consists  of  short, 
horizontal  arms  with  large  terminal  muscle  disks  which  are  opposed  to  similar  disks  formed 
by  the  proapophyseal  arms.  The  prothorax  is  attached  to  the  pterothorax  by  a stout, 
ligamentous  membrane,  permitting  little  flexibility. 

Zophosis  has  some  other  noteworthy  structural  modifications.  The  anterior  three  ab- 
dominal sternites,  which  are  connate  in  all  tenebrionids,  are  rigidly  fused  with  the  meta- 
sternum  by  continuous  cuticular  bands  laterad  of  the  metacoxal  cavities.  The  pterothorax 
and  abdomen  thus  become  a single,  rigid  unit.  In  all  other  Tentyriidae,  Tenebrionidae 
and  other  Coleoptera  which  have  been  examined,  the  mesothoracic-abdominal  articula- 
tion is  flexible,  although  in  most  apterous  tenebrionids  movement  is  prevented  by  inter- 
locking joints  between  the  elytra  and  the  abdominal  sternites  and  thoracic  pleurites.  In 
Zophosis  there  is  a pair  of  sclerotized,  dorsolateral  projections  from  the  region  of  the 
lateral  metacoxal  articulations.  The  function  of  these  projections,  which  are  unique  among 
known  Coleoptera,  is  uncertain,  but  they  may  help  secure  the  elytra  against  the  abdominal 
sternites. 


368 


Doyen 


mesapophyseal 

flange 

mesosternum 


/metafurcal  muscle  disk 
— - mesepimeron  12 
— metapleuron 


mesapophyseal 
muscle  disk 


metasternum 

mesepisternurm 
mesosternum  -- 

metasternum' 

mesapophyseal 

arm 

mesepisternum 

' \ 

\ 

\ 

mesosternum 


metafurcal 
^'muscle  disk 


^ mesepimeron 


mesocoxal 

inflection 


metacoxal 

inflection 

14 


mesepimeron 

'metafurcal  arm 
metapleuron 

mesocoxal 

inflection 


metasternum  metafurcal 

stalk 


Fig.  12-14.  Internal  thoracic  structures.  The  figures  are  from  an  oblique  posterodorsal  aspect,  with  the  thoracic  nota 
and  dorsal  abdominal  membranes  removed.  12,  Coelosattus  fortineri ; 13,  Praocis  penai',  14,  Zophosis  plana. 


Classification  of  Tenebrionoidea 


369 


Table  2.  Comparison  of  certain  thoracic  features  among  selected  species  of  Coniontini.* 


mentendosternite 

mesapophyseal  arms 

(a) 

mesocoxal  fusion 

(b) 

muscle  disks 

(c) 

mesopleural  fusion 

(d) 

length 

(e) 

basal  flange 

Coelus  ciliaris 

_ 

_ 

_ 

- 

- 

Coelomorpha  maritima 

- 

- 

- 

- 

- 

Coniontellus  obesa 

- 

- 

- 

- 

- 

Coelo taxis  punctulata 

- 

+ 

- 

- 

- 

Coniontides  latus 

- 

+ 

- 

- 

- 

Coniontis  viatica 

- 

+ 

- 

- 

- 

Branchus  floridanus 

- 

+ 

- 

+ 

- 

Oxinthas  praocioides 

- 

+ 

- 

+ 

+ 

Eusattus  dubius 

- 

+ 

+ 

- 

- 

robustus 

- 

+ 

+ 

+ 

- 

erosus 

- 

+ 

+ 

+ 

- 

reticulatus 

- 

+ 

+ 

+ 

+ 

muricatus 

+ 

+ 

+ 

+ 

+ 

Coelo  sat  tus  fortineri 

+ 

+ 

+ 

+ 

+ 

*(a)  Arms  of  metendostemite  fused  with  mesocoxal  inflection  (+)  or  free  (-);  (b)  arms 
terminated  as  enlarged  muscle  discs  (+)  or  unmodified  (-);  (c)  muscle  disks  fused  with 
mesopleuron  (+)  or  free  (-);  (d)  mesapophyseal  arms  short,  thick,  extending  no  more  than 
1/2  the  distance  to  the  mesepisternal  process  (+)  or  more  slender  and  extending  at  least 
1/3  the  distance  to  the  mesepisternal  process  (-);  (e)  mesapophyseal  arms  expanded  as 
flattened  flanges  basally  (+)  or  unmodified  (-).  Taxa  are  arranged  in  order  of  increasing 
specialization,  which  has  occurred  in  two  ways.  In  Branchus  and  Oxinthas,  the  mesa- 
pophyseal arms  are  shortened  and  flanged,  while  the  metafurcal  arms  are  unmodified. 
Conversely,  in  all  Eusattus  the  metafurcal  arms  are  subject  to  fusions  with  other  thoracic 
structures,  while  the  mesapophyseal  arms  are  modified  only  in  fossorial  species.  Speciali- 
zation in  internal  thoracic  structures  is  not  always  concordant  with  trends  in  other  charac- 
teristics. For  example,  the  protarsi  and  antennae  of  Coelus  and  Coelomorpha  are  highly 
specialized  for  burrowing  (Fig.  19-20). 


Externally,  Zophosis  shows  some  other  puzzling  structural  features.  The  metepistema, 
which  are  separated  from  the  metasterna  by  membranous  clefts  in  winged  Coleoptera, 
have  apparently  coalesced  with  the  sterna.  A pair  of  oblique  grooves  arising  near  the 
lateral  metacoxal  articulations  and  terminating  near  the  mesocoxal  inflections  may  repre- 
sent the  metepistemal  sutures,  but  more  likely  are  secondary  grooves  which  strengthen 


370 


Doyen 


the  metastemum.  Zophosis  also  has  a pair  of  “oblique  sutures”  running  anterolaterad  from 
the  intercoxal  process.  These  probably  strengthen  the  metastemum.  None  of  the  beetles 
discussed  here  possess  antecoxal  grooves,  which  are  almost  universally  present  in  winged 
forms.  The  antecoxal  grooves  run  from  the  medial  metacoxal  articulation  to  the  lateral 
metacoxal  articulation,  and  probably  reinforce  the  posterior  sternal  region. 

The  evidence  described  above  indicates  close  affinities  between  geographically  related 
taxa,  but  does  not  clarify  the  relationships  among  taxa  in  different  zoogeographic  regions. 
Zophosis  shows  clear  relationships  to  Adesmia  and  Epiphysa,  all  inhabiting  Africa  and 
southern  Eurasia.  Nyctelia , Gyriosomns,  Praocis  and  Entomochilus,  all  endemic  to  South 
America,  share  a strong  structural  consistency,  particularly  in  internal  thoracic  features. 
Likewise  the  Coniontini,  Coelini  and  Branchini  (North  America)  are  very  similar  morpho- 
logically. Intermediate  forms  between  the  African,  South  American  and  North  American 
taxa  are  unknown,  and  the  adaptations  of  each  of  these  groups  are  so  different  that  it  is 
premature  to  speculate  upon  their  affinities. 

CLASSIFICATION  OF  THE  CONIONTINI,  COELINI  AND  BRANCHINI 

Current  tribal  and  generic  classifications  of  these  beetles  largely  follow  the  arrangement 
of  Casey  (1908,  1924)  which  emphasizes  external  differences  in  leg  and  antennal  structure 
and  size  and  shape  of  the  epipleura.  These  features  are  variously  modified  to  facilitate 
burrowing,  and  intermediate  forms  sometimes  relate  the  specialized,  fossorial  species  to 
the  more  generalized,  ambulatory  ones.  This  is  especially  evident  in  Casey’s  group  Eusatti, 
as  recognized  by  Triplehom  (1968),  who  synonymized  all  Casey’s  genera  under  Eusattus, 
greatly  simplifying  the  previously  arbitrary  and  unworkable  classification.  Several  other 
generic  and  tribal  changes  are  proposed  here,  based  on  adult  characteristics.  Larval  features 
are  largely  concordant,  and  will  be  described  elsewhere. 

As  indicated  above,  the  morphological  features  of  coniontine  tentyriids  strongly  reflect 
their  mode  of  life.  Most  species  of  Coniontis  dwell  on  substrate  surfaces,  and  the  body 
is  oval  and  relatively  elongate.  The  arms  of  the  metendostemite  are  elongate,  but  not 
fused  with  the  mesocoxal  inflections  (Table  2).  Coelotaxis,  distinguished  by  an  elongate 
basal  protarsomere  and  “minute”  scutellum,  is  extremely  similar  to  Coniontis  in  all  other 
external  and  internal  characters.  There  is  considerable  variation  in  the  tarsal  character, 
and  some  individuals  are  scarcely  distinguishable  from  Coniontis  (Fig.  15-18).  Further- 
more, the  scutellum  is  relatively  large,  but  is  frequently  hidden  by  the  pronotum  in  pinned 
specimens.  Therefore,  Coelotaxis  is  placed  as  a synonym  of  Coniontis.  Coniontellus  is  dif- 
ferentiated from  Coniontis  by  having  the  eyes  completely  divided  by  a median  canthus. 
However,  there  is  considerable  interspecific  variation  in  the  degree  of  constriction  of  the 
eyes  in  Coniontis,  and  in  some  specimens  of  Coniontellus  the  eyes  are  not  completely 
divided.  Coniontides  and  Conisattus  differ  from  Coniontis  only  in  minor  body  propor- 
tions. These  three  genera  are  also  placed  as  synonyms  of  Coniontis.  Detailed  revisionary 
studies  of  Coniontis  may  show  that  some  of  these  names  should  be  recognized  as  sub- 
genera,  but  even  in  the  expanded  sense  proposed  here  Coniontis  is  much  more  mono- 
morphic  than  Eusattus  or  Praocis. 

The  species  of  Eusattus  are  usually  stouter  bodied  than  Coniontis,  and  several  are  highly 
modified  for  burrowing  in  aeolian  sand  ( E . muricatus,  E.  ciliatus  Horn,  E.  puberulus  LeC.). 
Within  Eusattus  ( sensu  Triplehorn)  the  degree  of  modification  of  internal  thoracic  structure 
ranges  from  the  generalized  condition  in  E.  dubius  to  that  in  E.  muricatus  (Table  2).  Coelo- 
sattus,  which  was  placed  in  the  Coelini  by  Blaisdell  (1927),  differs  from  E.  muricatus  chiefly 
in  having  broadly  expanded  protibiae  (Fig.  22-23)  and  strongly  arcuate  middle  and  hind 


Classification  of  Tenebrionoidea 


371 


tibiae.  It  lacks  the  specialized  tarsal  and  antennal  characters  of  the  Coelini,  while  the  spe- 
cialized internal  thoracic  structure  (Fig.  12)  is  nearly  identical  to  that  of  E.  muricatus.  The 
thoracic  structure  of  Coelus  and  Coelomorpha  is  similar  to  that  of  Coniontis.  For  these 
reasons,  Coelosattus  is  placed  as  a synonym  under  Eusattus. 

The  Coelini  comprise  a small,  monomorphic  group  of  fossorial  species  restricted  to  mari- 
time sand  dunes  along  the  Pacific  coast  of  North  America.  Superficially  they  greatly  resem- 
ble certain  species  of  Praocis  (e.g.,  P.  pilula ),  but  show  distinct  differences  in  mouthparts 
and  internal  thoracic  structures,  as  described  earlier.  The  most  important  characters  differ- 
entiating the  Coelini  from  the  Coniontini  are  the  enlarged,  spatulate  basal  protarsomeres 
(Fig.  19,  20)  and  the  extremely  short  antennae.  In  all  other  characters  they  greatly  re- 
semble the  Coniontini,  particularly  the  fossorial  species  of  Eusattus.  That  they  apparently 
evolved  independently,  perhaps  from  Coniontis , is  indicated  by  the  generalized  nature  of 
the  internal  thoracic  structures  (Table  2).  However,  the  general  similarity  to  the  Coniontini 
suggests  that  the  Coelini  should  not  be  recognized  as  a separate  tribe.  This  classification  was 
also  favored  by  LeConte  (1866),  LeConte  and  Horn  (1883)  and  Horn  (1870).  One  further 
point  concerns  the  genera  Coelus  and  Coelomorpha,  which  show  very  similar  modifications 
in  protarsal  and  antennal  structure,  but  differ  in  the  number  of  antennal  segments  (10  in 
Coelomorpha',  11  in  Coelus ).  Because  of  their  overall  similarity,  I propose  that  Coelomorpha 
be  placed  in  synonymy  under  Coelus. 

The  Branchini  were  originally  differentiated  from  the  Coniontini  by  LeConte  (1862).  He 
felt  that  the  anteriorly  confluent  gular  sutures  and  abrupt  basal  expansion  of  the  epipleura 
indicated  an  affinity  to  the  Nycteliini  and  Praocini,  respectively.  Later,  LeConte  (1866: 1 13) 
realized  that  most  of  the  character  states  he  used  to  separate  these  tribes  were  represented  in 
the  single  genus  Eusattus,  and  he  suggested  that  the  Nycteliini,  Praocini  and  Branchini 
should  possibly  be  placed  in  synonymy  under  the  Coniontini.  LeConte  was  unaware  of  the 
internal  thoracic  differences  of  the  first  two  tribes,  but  his  judgement  concerning  the  Bran- 
chini was  undoubtedly  correct.  The  only  major  structural  feature  differentiating  the  Bran- 
chini is  the  absence  of  the  submental  sclerite  (Fig.  24).  However,  the  submentum  is  very 
small  in  some  Eusattus  (Fig.  25).  For  these  reasons  the  Branchini  are  placed  in  synonymy 
under  the  Coniontini. 

The  taxonomic  changes  proposed  here  are  intended  to  consolidate  the  classification  of 
the  Coniontini  so  that  the  degree  of  variation  encompassed  is  similar  to  that  of  other  differ- 
entiated tribes,  such  as  the  Adesmiini,  Zophosini,  Praocini  and  Eleodini.  Extensive  elucida- 
tion of  the  generic  and  tribal  relationships  will  be  necessary  before  the  patterns  of  variation 
and  affinity  in  the  Tenebrionidae  and  Tentyriidae  can  be  used  to  reach  more  general  evolu- 
tionary and  biogeographic  conclusions. 

The  proposed  taxonomic  changes  are  summarized  in  the  following  checklist. 

Tribe  Coniontini 

Coniontini  Lacordaire,  1859:218;  Horn,  1870:291;  LeConte  and  Horn,  1883:371;  Casey, 

1908:55. 

Coelini  Casey,  1908:150. 

Branchini  LeConte,  1862:222. 

Body  stout,  oval  to  subglobose,  apterous.  Mentum  small,  trapezoidal,  weakly  emarginate 
anteriorly;  ligula  large,  sclerotized,  projecting  anteriorly  beyond  mentum;  labial  palps  in- 
serted dorsally  on  distinct  palpifers;  maxillae  exposed  laterad  of  labium,  lacinia  dentiform. 
Metendosternite  with  arms  elongate,  extending  to  region  of  metapleural  wing  process.  Ovi- 
positor with  1st  valvifers  elongate,  weakly  sclerotized;  2nd  valvifers  prolonged  posteriorly 


Doyen 


372 


•25  mm 


22 


Fig.  15-20.  Basal  tarsomeres  of  forelegs,  posteroventral  aspect;  15,  Coniontis  hoppingi  Blaisdell;  16,  Coniontides  latus\ 
17,  18,  Coelotaxis  punctulata\  19,  Coelomorpha  maritima;  20,  Coelus  globosus.  Fig.  21-23.  Posterior  aspect  of  forelegs; 
21,  Coniontides  latus\  22,  Eusattus  reticulatus ; 23,  Coelosattus  fortineri.  Fig.  24-25.  Ventral  aspect  of  crania;  maxillae, 
mandibles,  ligula  and  clypeus  excised;  24,  Oxinthas  praocioides;  25,  Eusattus  muricatus. 


Classification  of  Tenebrionoidea 


373 


as  strongly  sclerotized,  spatulate  prongs;  gonostyli  represented  by  several  elongate  setae  in- 
serted in  membranous  foramen  situated  medially  on  2nd  valvifers.  Aedeagus  inverted;  teg- 
men  with  variable  desclerotized  area  posteroventrally;  paramere  longer  than  tegmen,  bearing 
several  lateral  setae  at  apex. 

Coniontis  Eschscholtz,  1829.  Type  species:  Coniontis  viatica  Eschscholtz,  Casey  designation, 
1908:57. 

syn.  Coniontellus  Casey,  1890.  Type  species:  Coniontis  obesa  LeConte,  1851,  Casey 
designation,  1908:57. 

syn.  Coniontides  Casey,  1908.  Type  species:  Coniontis  lata  LeConte,  1866,  by  original 
designation,  p.  57. 

syn.  Conisattus  Casey,  1908.  Type  species:  Conisattus  rectus  Casey,  1908:57,  mono- 
basic. 

syn.  Coelo taxis  Horn,  1876.  Type  species:  Coelo taxis  punctulata  Horn,  1876,  Casey 
designation,  1908:57. 

Eusattus  LeConte,  185 1.1  Type  species:  Eusattus  difficilis  LeConte,  1852,  Casey  designa- 
tion, 1908:56. 

syn.  Eusattodes  Casey,  1908.  Type  species:  Eusattus  laevis  LeConte,  1866,  Casey 
designation,  1908:56. 

syn.  Megasattus  Casey,  1908.  Type  species:  Eusattus  erosus  Horn,  1870,  by  original 
designation,  p.  56. 

syn.  Nesostes  Casey,  1908.  Type  species:  Eusattus  robustus  LeConte,  1866,  by  original 
designation,  p.  56. 

syn.  Sphaeriontis  Casey,  1908.  Type  species:  Eusattus  muricatus  LeConte,  1851,  by 
original  designation,  p.  56. 

syn.  Coelosattus  Blaisdell,  1927.  Type  species:  Coelosattus  fortineri  Blaisdell,  1927, 
monobasic. 

syn.  Discodemus  LeConte,  1862.  Type  species:  Zophosis  reticulata  Say,  1824,  mono- 
basic. 

syn.  Conipinus  LeConte,  1862.  Type  species:  Eusattus  dubius  LeConte,  1851,Gebien 
designation,  1938:284. 

Coelus  Eschscholtz,  1829.  Type  species:  Coelus  ciliatus  Eschscholtz,  1829,  monobasic. 

syn.  Coelomorpha  Casey,  1890.  Type  species:  Coelomorpha  maritima  Casey,  1890, 
Casey  designation,  1908:151. 

Branchus  LeConte,  1862.  Type  species:  Bronchus  floridanus  LeConte,  monobasic. 
Oxinthas  Champion,  1892.  Type  species:  Oxinthas  praocioides  Champion,  monobasic. 
Anectus  Horn,  1866.  Type  species:  Anectus  vestitus  Horn,  monobasic. 

Detailed  keys  to  species  will  be  presented  in  future  revisionary  work  on  each  genus.  The 
genera  are  keyed  below. 


Key  to  the  genera  of  Coniontini 

1.  Submentum  clearly  defined,  though  sometimes  small,  transverse  (Fig.  25)  2 

— Submentum  extremely  small,  invisible  externally  (Fig.  24)  4 


2.  Basal  protarsomere  truncate  or  with  process  shorter  than  second  tarsomere  (Fig.  15-18) 
3 


1 Generic  synonymy  for  Eusattus  is  adapted  from  Triplehom  (1968)  with  the  addition  of  Coelosattus  Blaisdell. 


374 


Doyen 


Basal  protarsomere  extending  beyond  second  tarsomere  as  spatulate  process  (Fig.  19- 


20)  Coelus 

3.  Protibia  abruptly  expanded  apically  as  an  acute  process  (Fig.  22-23)  Eusattus 

— Protibia  gradually  enlarged  apically  (Fig.  21 ) Coniontis 

4.  Intercoxal  process  of  abdomen  broadly  rounded 5 

— Intercoxal  process  of  abdomen  rectangularly  truncate2 Anectus 

5.  Protibia  abruptly  expanded  apically  as  a short,  acute  process Branchus 

— Protibia  gradually  enlarged  apically Oxinthas 


ACKNOWLEDGEMENTS 

The  ideas  contained  in  this  paper  have  been  discussed  extensively  with  several  other 
workers,  especially  W.  Tschinkel,  Florida  State  University,  J.  F.  Lawrence  and  T.  Hlavac, 
Museum  of  Comparative  Zoology,  Harvard  University  and  J.  A.  Chemsak,  University  of 
California.  Their  comments  and  criticisms  have  greatly  clarified  many  of  the  problems 
encountered.  Various  drafts  of  the  manuscript  were  read  by  J.  A.  Chemsak,  H.  V.  Daly 
and  J.  F.  Lawrence,  who  offered  helpful  suggestions  regarding  writing  and  organization. 
I thank  all  of  these  persons  for  their  help. 

Specimens  of  Branchus  and  Oxinthas  were  loaned  by  T.  J.  Spilman,  United  States  Na- 
tional Museum,  and  J.  F.  Lawrence,  to  whom  I am  thankful.  Without  the  specimens  the 
second  portion  of  the  paper  would  have  been  incomplete.  H.  B.  Leech,  California  Academy 
of  Sciences,  generously  allowed  dissections  to  be  made  of  specimens  of  several  species  which 
would  have  otherwise  been  unavailable.  Important  material  for  comparison  was  also  ob- 
tained from  the  collections  under  the  care  of  the  following  individuals,  to  whom  I am 
grateful:  W.  Barr,  University  of  Idaho,  Moscow;  D.  E.  Bright,  Canadian  National  Collection, 
Ottawa;  J.  P.  Donahue  and  R.  R.  Snelling,  Los  Angeles  County  Museum;  H.  Dybas,  Field 
Museum  of  Natural  History,  Chicago;  L.  H.  Herman,  American  Museum  of  Natural  History, 
New  York;  J.  F.  Lawrence,  Museum  of  Comparative  Zoology,  Cambridge;  H.  B.  Leech, 
California  Academy  of  Sciences,  San  Francisco;  W.  W.  Moss,  Academy  of  Natural  Sciences, 
Philadelphia;  L.  L.  Pechuman,  Cornell  University  Collection,  Ithaca;  P.  O.  Ritcher,  Oregon 
State  University,  Corvallis;  T.  J.  Spilman,  United  States  National  Museum,  Washington, 
D.  C.;  C.  A.  Triplehorn,  Ohio  State  University,  Columbus;  W.  Turner,  Washington  State 
University,  Pullman. 


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STUDIES  ON  BOREAL  AGROMYZIDAE  (DIPTERA).  II. 
PHYTOMYZA  MINERS  ON  SENECIO,  PETASITES  AND  TUSSILAGO 
(COMPOSITAE,  SENECIONEAE) 


GRAHAM  C.  D.  GRIFFITHS 
Department  of  Entomology 

University  of  Alberta  Quaes  tiones  entomologicae 

Edmonton,  Alberta  T6G  2E3  8 : 377-405  1972 


Twelve  species  of  Phytomyza  are  known  as  miners  of  Senecio,  Petasites  and  Tussilago 
in  boreal  areas.  These  belong  to  three  species-groups,  the  albiceps  group  (4  species),  the 
syngenesiae  group  ( 3 species)  and  the  robustella  group  ( 5 species).  Three  new  species  of 
the  robustella  group  are  described,  as  follows:  Phytomyza  hyperborea  n.  sp.  ( type-locality 
Walker  Fork,  Alaska),  P.  hypophylla  n.  sp.  (type-locality  Eagle  Summit,  Alaska)  and  P.  lu- 
gentis  n.  sp.  ( type-locality  Summit  Lake  Pass,  British  Columbia).  In  the  albiceps  group  one 
new  subspecies  is  described,  Phytomyza  tussilaginis  kevani  n.  ssp.  ( type-locality  Richards 
Island,  Northwest  Territories);  the  North  American  P.  petasiti  Spencer  is  considered  a 
geographical  subspecies  of  P.  tussilaginis  Hendel,  described  from  Europe;  and  P.  alpina 
Groschke,  previously  known  from  Scotland  and  the  Alps,  is  recorded  for  British  Columbia, 
Yukon  and  Alaska. 


Douze  especes  de  Phytomyza  sont  connues  dans  les  regions  boreales  comme  mineuses 
du  Senecio,  du  Petasites  et  de  la  Tussilago.  Ces  especes  appartiennent  a trois  groupes 
d’ especes,  le  groupe  albiceps  (4  especes),  le  groupe  syngenesiae  (3  especes)  et  le  groupe 
robustella  (5  especes).  Trois  nouvelles  especes  sont  decrites  dans  le  groupe  robustella,  tel 
que:  Phytomyza  hyperborea  n.  sp.  (localite-type  Walker  Fork,  Alaska),  P.  hypophylla  n. 
sp.  (localite-type  Eagle  Summit,  Alaska)  et  P.  lugentis  n.  sp.  (localite-type  Summit  Lake 
Pass,  Colombie  britannique ).  Dans  le  groupe  albiceps  une  nouvelle  sous-espece  est  de- 
crite,  Phytomyza  tussilaginis  kevani  n.  ssp.  (localite-type  Richards  Island,  Territoires  du 
nord-ouest);  P.  petasiti  Spencer  d’Amerique  du  nord  est  consideree  comme  sous-espece 
geographique  de  P.  tussilaginis  Hendel,  d’Europe;  et  P.  alpina  Groschke,  connue  dans  le 
passe  d’Ecosse  et  des  Alpes,  est  maintenant  notee  pour  la  Colombie  britannique,  pour  le 
Yukon  et  pour  V Alaska. 


Zwolf  Phytomyza -Arten  werden  als  Minierer  von  Senecio,  Petasites  und  Tussilago  in 
borealischen  Gebieten  besprochen.  Diese  gehoren  zu  drei  Arten-Gruppen,  der  albiceps- 
Gruppe  (4  Arten),  der  syngenesia e-Gruppe  (3  Arten)  und  der  robustella -Gruppe  (5  Arten). 
Drei  Arten  der  robustella -Gruppe  sind  neu  beschrieben,  wie  folgt:  Phytomyza  hyperborea 
n.  sp.  (Fundort  vom  Typus  Walker  Fork,  Alaska),  P.  hypophylla  n.  sp.  (Fundort  vom 
Typus  Eagle  Summit,  Alaska)  und  P.  lugentis  n.  sp.  (Fundort  vom  Typus  Summit  Lake 
Pass,  British  Columbia).  Bei  der  albiceps  -Gruppe  wird  eine  geographische  Unterart  neue 
beschrieben,  Phytomyza  tussilaginis  kevani  n.  ssp.  (Fundort  vom  Typus  Richards  Island, 
Northwest  Territories);  die  nordamerikanische  P.  petasiti  Spencer  wird  als  Unterart  von 
P.  tussilaginis  Hendel  (aus  Europa  beschrieben)  vermutet;  P.  alpina  Groschke,  vorher  nur 
aus  Schottland  und  aus  den  Alpen  bekannt,  wird  fur  British  Columbia,  fiir  Yukon  und  fur 
Alaska  besprochen. 


378 


Griffiths 


The  present  paper  deals  with  the  Phytomyza  miners  of  part  of  the  Senecioneae.  All 
known  Phytomyza  miners  of  Tussilago  and  Petasites  are  treated;  but  I leave  out  of  consider- 
ation (as  outside  my  geographical  area  of  interest)  the  following  miners  of  Senecio:  Phyto- 
myza burchardi  Hering  (Canary  Isles)  and  P.  seneciovora  Spencer  (Africa).  K.  A.  Spencer  in- 
forms me  (in  correspondence)  that  he  has  recently  obtained  an  undescribed  species  close  to 
seneciovora  from  a tree  Senecio  in  Kenya.  He  will  discuss  these  species  in  a later  work.  In 
his  opinion  both  belong  to  the  syngenesiae  group. 

The  species  of  Phytomyza  here  treated  are  referred  to  three  groups,  the  albiceps  group, 
the  syngenesiae  group  and  the  robust ella  group. 

The  flies  from  Holland  described  by  de  Meijere  (1924)  as  Phytomyza  jacobaeae  were 
almost  certainly  not  bred  from  Senecio.  No  one  has  since  succeeded  in  obtaining  from 
Senecio  flies  which  agree  with  his  description.  I have  seen  two  female  syntypes  of  this  spe- 
cies, and  in  my  opinion  they  belong  to  Phytomyza  milii  Kaltenbach,  a well-known  grass- 
feeder.  The  puparia  from  Senecio  jacobaea  L.  associated  with  these  flies  by  de  Meijere 
doubtless  belonged  to  a species  of  the  Phytomyza  syngenesiae  group;  for  de  Meijere  (1926: 
267)  stated  that  he  could  find  no  morphological  difference  between  these  puparia  and 
those  of  that  group  (as  “ atricornis ”).  I suggest  that  the  description  of  “ jacobaeae ” was  the 
result  of  an  error  in  associating  the  wrong  flies  with  these  puparia. 

The  presentation  followed  in  this  paper  is  similar  to  that  of  the  first  paper  of  this  series 
(Griffiths,  1972),  which  should  be  consulted  for  explanation  of  the  terms  and  abbreviations 
used  in  my  descriptions.  My  use  of  names  of  North  American  plants  again  follows  Hulten 
(1968).  The  holotypes  of  the  new  species  and  subspecies  described  in  this  paper  will  be 
deposited  in  the  Canadian  National  Collection  (Ottawa). 

DIAGNOSIS 

The  species  treated  in  this  paper  all  belong  to  groups  in  which  identification  is  based 
largely  on  the  form  of  the  male  genitalia,  particularly  the  aedeagus.  The  new  species  whose 
male  is  unknown  ( Phytomyza  hyperborea  n.  sp.)  may  be  distinguished  from  its  close  rela- 
tives by  its  long  antennal  pubescence.  The  difference  in  the  form  of  the  aedeagus  between 
the  species  of  the  albiceps  and  robustella  groups  treated  here  is  rather  striking,  as  indicated 
in  my  figures.  No  difficulty  should  be  experienced  in  identifying  males  of  these  species, 
if  a suitable  technique  of  dissection  is  used.  For  further  information  on  the  more  difficult 
P.  syngenesiae  group,  only  briefly  discussed  here,  see  my  earlier  revision  (Griffiths,  1967). 

Keys  with  worldwide  coverage  to  the  mines  of  Phytomyza  larvae  on  Senecio,  Petasites 
and  Tussilago  are  given  below.  Other  genera  of  Agromyzidae  whose  larvae  are  known  to 
attack  these  host  genera  are  Liriomyza,  Calycomyza,  Ophiomyia  and  Melanagromyza.  Some 
species  of  the  albiceps  and  syngenesiae  groups  cannot  be  separated  on  the  basis  of  their 
mines  and  larvae,  as  indicated  in  the  keys. 

The  new  species  of  the  robustella  group  described  in  this  paper  may  be  included  in 
Spencer’s  (1969b)  key  to  the  Phytomyza  species  of  Canada  and  Alaska  by  the  extensions 
given  below.  I am  not  attempting  to  revise  the  couplets  to  which  the  species  of  the  albiceps 
group  would  be  referred  (couplets  12-16  and  18-25),  as  I think  that  this  part  of  the  key 
will  need  to  be  substantially  rewritten.  The  distinctions  in  the  range  of  the  costal  ratio 
drawn  in  Spencer’s  couplets  12  and  18  are  a particular  source  of  difficulty,  as  the  range 
in  some  species  transgresses  the  boundaries  indicated. 


26.  Third  antennal  segment  with  conspicuously  long  pubescence 26a 

Third  antennal  pubescence  normal  27 


Boreal  Agromyzidae 


379 


26a.  Third  antennal  segment  enlarged  in  female  (Spencer  1969b,  Fig.  450).  Aedeagus  as 

in  Spencer’s  Fig.  451 lactuca  Frost 

Third  antennal  segment  not  enlarged  in  female  (Fig.  31).  (<J  unknown)  

hyperborea  n.  sp. 

66.  Distal  section  of  aedeagus  with  large  distiphallus  containing  bifid  terminal  portion  of 
ejaculatory  duct  66a 

— Aedeagus  not  of  above  type;  “supporting  sclerites”  arising  from  base  of  distal  section 
66b 

66a.  Orbits  distinctly  projecting  above  eye;  aedeagus  as  in  Figs.  23-24  

hypophylla  n.  sp. 

Orbits  not  projecting  above  eye;  aedeagus  as  in  Figs.  17-18 lugentis  n.  sp. 

66b.  (as  Spencer’s  couplet  66) 

Key  to  Phytomyza  mines  on  Senecio 

1.  Larva  normally  leaving  leaf  before  formation  of  strongly  arched  brown  or  black 
puparium  (Fig.  26).  Posterior  spiracles  of  puparium  and  third  instar  larva  with 

19-36  bulbs  (Fig.  28) ( albiceps  group) 2 

Puparium  formed  inside  leaf,  with  anterior  spiracles  turned  downwards,  projecting 
through  epidermis  (Fig.  27).  Posterior  spiracles  of  puparium  and  third  instar  larva 
with  fewer  bulbs 3 

2.  Canary  Isles.  Linear  mines  on  S.  papyraceus  DC P.  burchardi  Hering 

— Europe.  Mines  linear  throughout,  not  more  than  1.5  mm  wide  terminally  (Fig.  37), 
on  S.  nemorensis  L.,  S.  fuchsii  Gmel.,  S.  subalpinus  Koch  and  S.  fluviatilis  Wallr. 

P.  senecionis  Kaltenbach 

Europe.  Mines  initially  linear,  but  blotchy  terminally  (Fig.  36).  On  S.  alpinus  (L.) 
and  S.  jacobaea  L P.  alpina  Groschke 

— North  America.  Linear  mines  on  S.  lugens  Richards  and  S.  pauperculus  Michx. 

(Fig.  35)  P.  alpina  Groschke 

Japan.  Linear  mines  on  S.  palmatus  Pall P.  ravasternopleuralis  Sasakawa 

3.  Puparium  bright  green;  hind  spiracles  distinctly  horned,  with  about  20  bulbs.  Africa. 

P.  seneciovora  Spencer 

Puparium  white,  brown  or  black;  hind  spiracles  knob-shaped,  with  not  more  than 
12  bulbs  4 

4.  Mine  with  narrow  linear  channel,  not  more  than  2 mm  wide  (if  with  blotchy  areas, 

these  formed  by  convolutions  of  mine  channel  or  by  coalescence  of  mines  of  differ- 
ent larvae) P.  syngenesiae  group 

Three  species  of  this  group,  P.  syngenesiae  (Hardy),  P.  horticola  Goureau  and  P.  senecionella  Sehgal,  are 
reported  from  Senecio. 

Mine  channel  broader,  often  with  blotchy  areas.  North  America.  On  S.  lugens  Rich- 
ards and  S.  sheldonensis  Pors P.  lugentis  n.  sp. 

Key  to  Phytomyza  mines  on  Petasites  and  Tussilago 

1.  Larva  normally  leaving  leaf  before  formation  of  strongly  arched  brown  or  black 
puparium  (Fig.  26).  Mines  linear,  on  upper  surface  of  leaf.  Posterior  spiracles  of 

puparium  and  third  instar  larva  with  19-3 1 bulbs  (Fig.  28) 

(< albiceps  group) 2 

Puparium  formed  inside  leaf  or  petiole,  with  anterior  spiracles  turned  downwards, 


380 


Griffiths 


projecting  through  epidermis  (Fig.  27)  3 

2.  Europe.  On  Tussilago  and  Petasites P.  tussilaginis  tussilaginis  Hendel 

— Japan.  On  Petasites P.  ravasternopleuralis  Sasakawa 

— North  America.  On  Petasites P.  tussilaginis  petasiti  Spencer 

or  P.  tussilaginis  kevani  n.  ssp. 

or  P.  alpina  Groschke 

3.  Larvae  feeding  mainly  in  petiole  of  leaf,  in  some  cases  also  in  veins.  Posterior  spir- 

acles of  puparium  and  third  instar  larva  with  18-22  bulbs.  Europe.  On  Petasites  and 
Tussilago P.  buhriella  Spencer 

— Larvae  feeding  on  parenchyma  of  leaf,  not  in  veins  or  petioles  (although  the  mine 

channel  may  run  besides  some  of  the  veins).  Posterior  spiracles  of  puparium  and 
third  instar  larva  with  fewer  bulbs 4 

4.  Posterior  spiracles  of  puparium  and  third  instar  larva  with  about  1 5 bulbs.  Puparium 

reddish  yellow.  North  America.  On  Petasites P.  hyperborea  n.  sp. 

— Posterior  spiracles  of  puparium  and  third  instar  larva  with  7-12  bulbs  (Fig.  29). 

Puparia  mostly  white  (but  some  overwintering  puparia  of  syngenesiae  group  brown 
or  black) 5 

5.  Mines  mainly  on  lower  surface  of  leaf,  normally  convolute  within  restricted  area 

(Fig.  34).  North  America.  On  Petasites P.  hypophylla  n.  sp. 

Mines  on  upper  surface  of  leaf,  narrowly  linear  (about  1 mm  wide  terminally),  24- 

28  cm  long  (Fig.  33A).  Europe.  On  Tussilago  and  Petasites 

P.  farfarae  Hendel 

— Mines  linear,  on  upper  or  lower  surface,  much  shorter  than  those  of  farfarae  (less 

than  1 0 cm  long),  in  most  cases  over  1 mm  wide  terminally 

P.  syngenesiae  group 

Two  species  of  this  group,  P.  horticola  Goureau  and  P.  senecionella  Sehgal,  are  reported  from  Petasites. 

TREATMENT  OF  SPECIES 
(a)  the  Phytomyza  albiceps  group 

Nowakowski  (1962b)  has  already  discussed  the  possibility  of  defining  a “natural  group  or 
subgenus”  in  this  sense.  In  the  albiceps  group  the  puparia  are  strongly  arched  and  dark  in 
colour  (brown  or  black),  formed  outside  the  mine  (Fig.  26);  the  aedeagus  is  characterized  by 
narrow,  strongly  sclerotized  basal  sclerites  (“arms  of  basiphallus”)  and  in  many  species  also 
by  the  presence  of  spine-like  cuticular  processes;  and  the  posterior  ors  is  variable  in  length 
(in  most  individuals  shorter  than  the  anterior  ors  or  completely  absent).  The  limits  of  this 
group  have  not  yet  been  clarified,  but  it  is  evident  that  numerous  species  whose  larvae  mine 
Compositae  and  Umbelliferae  belong  here.  The  black-frons  species  which  have  been  referred 
to  as  the  “ obscurella  group”  should  in  my  opinion  also  be  included  in  the  albiceps  group. 

Identification  of  many  species  of  the  albiceps  group  is  only  possible  through  study  of  the 
male  genitalia.  Nowakowski  (1962b)  has  rightly  remarked  that  the  reliance  on  colour  differ- 
ences in  Hendel’s  (1935-6)  key  has  led  to  artificial  separation  of  closely  related  species. 
My  present  study  indicates  that  gross  differences  in  colour  can  be  shown  not  only  by  closely 
related  species,  but  even  by  races  of  the  same  species.  Another  character  whose  high 
variability  has  not  been  appreciated  is  the  length  of  the  posterior  orbital  bristle  (ors).  For 
instance,  in  Phytomyza  tussilaginis  Hendel  the  range  of  variation  in  the  development  of 
this  bristle  varies  from  equal  length  to  the  anterior  ors  to  complete  absence  (the  extremes 
can  even  be  shown  on  either  side  of  the  same  individual!).  Therefore  all  claims  that  the 


Boreal  Agromyzidae 


381 


length  of  the  posterior  ors  can  be  used  for  differentiating  species  of  the  albiceps  group 
should  be  regarded  as  doubtful. 

K.  A.  Spencer  has  in  press  a note  on  the  type  specimens  of  Phytomyza  albiceps  Meigen.  The  customary  use  of  this 
name  for  a species  mining  Artemisia  in  Europe  has  proved  to  be  incorrect.  Spencer  thinks  that  the  female  lectotype 
probably  belongs  to  the  species  since  described  as  Phytomyza  rydeniana  Hering.  Fortunately  the  latter  species  is  refer- 
able to  the  albiceps  group  in  the  wide  sense  here  followed,  so  there  is  no  need  to  propose  a different  group  name. 

Phytomyza  tussilaginis  Hendel  1925 
(synonymy  below  under  subspecies) 

Adult.  — Head  with  orbits  not  or  only  very  narrowly  projecting  above  eye  in  lateral 
view;  genae  in  middle  1/4  to  1/3  of  eye  height;  eyes  with  only  sparse  fine  pubescence  or 
apparently  bare.  Frons  at  level  of  front  ocellus  about  twice  width  of  eye.  Ors  directed 
posteriorly,  ori  directed  inwardly;  posterior  ors  variably  developed,  in  most  individuals 
1/2  to  2/3  as  long  as  anterior  ors,  but  ranging  from  fully  as  long  (as  in  the  holotype  of 
subspecies  petasiti ) to  completely  absent;  anterior  ori  usually  1/2  to  2/3  as  long  as  posterior 
ori  (but  in  some  individuals  absent  on  one  side  according  to  Hendel,  1935);  orbital  setulae 
one-rowed.  Peristomal  margin  with  vibrissa  and  4-6  upcurved  peristomal  setulae.  Third 
antennal  article  rounded  distally,  with  short  pubescence. 

3 + 1 dc;  acr  numerous,  in  4-6  rows  anteriorly;  presutural  ia  numerous;  7-12  postsutural 
ia;  inner  pa  1/3  to  1/2  as  long  as  outer  pa. 

Second  cross-vein  (m-m)  absent.  Costal  ratio  mg2/mg4  2. 6-4.0.  Wing  length  2. 2-2. 8 mm 
(see  below  under  subspecies). 

Colour  geographically  variable  (see  below  under  subspecies). 

Male  postabdomen  with  8th  sternum  fused  with  6th  tergum.  Telomeres  not  clearly  de- 
limited from  periandrium,  bearing  only  fine  setulae.  Pregonites  extending  ventrally,  shielding 
base  of  aedeagus  at  rest.  Aedeagus  as  Fig.  6-7;  basal  sclerites  narrow,  strongly  divergent 
distally,  with  row  of  spinules  distally  along  their  dorsal  margins;  sclerites  of  medial  lobe 
widely  separated  anteriorly,  convergent  and  bent  upwards  posteriorly;  distal  section  with 
small  distiphallus  consisting  of  pair  of  tubules  widely  separated  from  basal  section  by  clear 
membranous  area  (without  or  at  most  with  fine  linear  traces  of  pigmentation).  Ejaculatory 
apodeme  small  (Fig.  8). 

A figure  of  the  aedeagus  has  also  been  published  by  Spencer  (1969b). 

Puparium  and  third  instar  larva.  - Mandibles  with  two  alternating  teeth;  right  mandible 
longer  than  left.  Anterior  spiracles  with  two  short  horns,  with  12-15  bulbs  in  irregular 
ellipse.  Posterior  spiracles  borne  on  short  conical  processes,  with  25-31  bulbs  in  irregular 
ellipse.  Puparium  dark  brown  or  black,  2. 0-2. 3 mm  long  (Fig.  26). 

Mine.  — Larvae  leaf-miners  on  Tussilago  and  Petasites.  Mine  (Fig.  33 B)  entirely  linear, 
up  to  25  cm  long,  2-4  mm  wide  terminally;  faeces  deposited  as  discrete  particles  (separated 
by  over  1 mm  in  mines  on  Tussilago,  but  closer  together  in  mines  on  Petasites)',  mine 
entirely  on  upper  surface  of  leaf,  appearing  white  (on  Petasites ) or  whitish  green  (on 
Tussilago ) in  reflected  light;  larvae  leaving  leaf  through  semicircular  slit  on  upper  surface 
before  puparium  formation. 

A figure  has  also  been  given  by  Beiger  (1960)  of  a mine  on  Petasites  albus  (L.). 

Remarks.  — This  is  the  first  species  of  the  albiceps  group  in  which  geographical  colour 
variation  has  been  discovered.  Since  the  colour  forms  do  not  differ  in  morphology  or  in 
life-history,  I conclude  that  they  are  probably  geographical  races  of  the  same  species.  Thus 
I interpret  tussilaginis  as  a Rassenkreis  or  polytypic  species.  All  specimens  obtained  from 
forested  areas  in  North  America  are  referable  to  the  yellow  subspecies  {petasiti  Spencer), 
while  the  new  dark  subspecies  described  below  was  obtained  from  arctic  tundra  (see  Fig. 


382 


Griffiths 


38).  All  European  specimens  so  far  described  are  referable  to  the  nominate  subspecies 
( tussilaginis  Hendel).  However  all  these  specimens  are  from  moderate  latitudes,  so  that  the 
possibility  of  other  races  occurring  in  more  northern  areas  of  Europe  remains  open. 

Spencer  (1969b)  has  suggested  that  the  leaf  mine  of  subspecies  petasiti  is  shorter  and 
broader  than  that  of  subspecies  tussilaginis.  But  I can  find  no  such  difference  in  the  addi- 
tional material  now  available  to  me.  The  mine  figured  by  Spencer  seems  to  me  untypically 
short  and  convolute. 

This  species  has  only  been  bred  with  certainty  from  Tussilago  and  Petasites.  I am  doubt- 
ful whether  Hendel’s  (1935)  record  of  Adenostyles  as  a host-plant  is  correct,  for  that  genus 
does  not  belong  to  the  Senecioneae.  The  fly  in  Hendel’s  collection  bred  from  Adenostyles  is 
a female,  whose  specific  identity  will  remain  uncertain  until  males  can  be  associated  with  it. 

The  type  series  of  tussilaginis  was  bred  by  Hendel  from  Tussilago  farfara  L.  in  Austria. 
Since  only  a female  from  this  series  has  been  traced  (listed  below  under  subspecies  tussila- 
ginis),  the  application  of  Hendel’s  name  is  open  to  doubt.  I here  follow  the  accepted 
opinion  that  flies  on  Petasites  and  Tussilago  in  Europe  belong  to  the  same  species.  But  no 
male  has  yet  been  bred  from  Tussilago  to  confirm  this. 

Phytomyza  tussilaginis  tussilaginis  Hendel  1925 

Phytomyza  tussilaginis  Hendel.  Hendel,  1925:308.  —1935:493.  Hering,  1927:1 14.  Holotype 

lost;  type-locality,  Vienna  (Austria). 

Adult.  — Frons  largely  clear  yellow,  but  with  dark  ocellar  plate  and  dark  vertex  (both  vt 
on  dark  ground);  in  some  specimens  upper  part  of  orbits  also  slightly  infuscated  along  eye 
margins.  Face  partly  yellow,  but  becoming  brown  in  antennal  pits  and  in  some  specimens 
also  around  antennal  bases.  Genae  yellow.  Occiput  dark.  Antennae  with  first  article  yellow 
or  yellow-brown,  second  article  brown  or  black,  third  article  black.  Palpi  brown  or  black; 
labella  yellow. 

Mesonotum  weakly  shining,  finely  grey-dusted,  black  centrally,  brown  on  sides  with 
traces  of  brighter  coloration  (yellow-brown  or  whitish  yellow)  around  margins  of  humeral 
callus  and  on  upper  part  of  sutural  triangle.  Scutellum  black.  Pleura  black  except  narrow 
white  band  along  dorsal  margin  of  mesopleuron  and  white  seam  of  mesopleural  suture. 
Wing  base  and  squamae  contrastingly  white,  but  latter  with  dark  fringe.  Legs  largely  dark, 
with  tips  of  femora  contrastingly  yellow;  tibiae  and  tarsi  brown. 

Abdomen  largely  brown,  becoming  yellow-brown  on  sides  at  base.  Basal  cone  of  oviposi- 
tor (9)  grey-dusted  on  about  basal  third  to  half. 

Wing  length  2.4-2. 8 mm.  Costal  ratio  mg2/mg4  2. 6-4.0. 

Material  examined.  — 1 9 paratype  from  larva  17.x. 23  on  Tussilago  farfara  L.,  Vienna 
University,  Austria,  emerged  18.iii.24  (in  Zoological  Museum  of  Humboldt  University,  Ber- 
lin). 1 6,  3 99  from  larvae  29.viii.53  on  Petasites  hybridus  (L.),  Boxhill,  Surrey,  England, 
emerged  16-18. ix. 53,  leg.  G.  C.  D.  Griffiths;  1 <3,  1 9 from  larvae  12.ix.54,  same  plant  and 
locality,  emerged  4.x. 54  and  29.iii.55,  leg.  G.  C.  D.  Griffiths.  1 6 from  larva  21.viii.53  on 
Petasites  hybridus  (L.),  Millers  Dale,  Derby,  England,  emerged  10.ix.53,  leg.  K.  A.  Spencer. 

Other  records.  — The  following  records  are  referred  to  this  subspecies,  on  the  assumption 
that  it  is  the  only  member  of  the  albiceps  group  whose  larvae  mine  Petasites  and  Tussilago 
in  Europe. 

Ireland  — Tipperary,  29.viii.69,  mines  on  Tussilago  farfara  L.  (K.  A.  Spencer). 

Austria  — Volderbad  and  Haller  Strasse  (Tirol)  on  Tussilago  farfara  L.  (Hendel,  1925); 

Mosern  (Tirol)  on  Tussilago  farfara  L.  (Griffiths,  1966:807);  also  sheets  in 
Hering’s  mine  herbarium  for  Carinthia  (Plockenpass  on  Tussilago  farfara  L., 


Boreal  Agromyzidae 


383 


and  Mauthen  on  Petasites  albus  [ L.  ] ). 

Germany  — Neubrandenburg  (Mecklenburg)  on  Petasites  hybridus  (L.)  (Buhr,  1941a); 

Gottesberg,  Bad  Elster  and  Oberwiesenthal  (Saxony)  on  Tussilago  farfara  L., 
Petasites  hybridus  (L.)  and  P.  albus  (L.)  (Buhr,  1964);  also  sheets  in  Hering’s 
mine  herbarium  for  Muhlhausen  (Thuringia)  on  Tussilago  farfara  L.,  Soritz 
(Bautzen)  on  Tussilago  farfara  L.,  Falkenstein  (Bavaria)  on  Petasites  albus 
(L.),  Tolz  (Bavaria)  on  Petasites  hybridus  (L.),  Heimkehle  (Alter  Stolberg) 
on  Petasites  albus  (L.),  and  Berlin  Botanical  Gardens  on  Petasites  hybridus 
(L.). 

Italy  — Rionero  in  Vulture,  on  Tussilago  farfara  L.,  leg.  Ricchello  (sheet  in  Hering’s 
mine  herbarium). 

Czechoslovakia  — Tisova,  Orlik  and  Jesenik  (Stary,  1930),  mines  on  Tussilago  farfara  L. 

Roumania  — Herculesbad  (Banat),  mines  on  Petasites  sp.  and  Tussilago  farfara  L.  (Hering, 
1924,  nos.  60  and  104);  Sinaia,  on  Tussilago  farfara  L.,  leg.  Sienkiewicz 
(sheet  in  Hering’s  mine  herbarium). 

Poland  — Tatry  Mountains,  on  Tussilago  farfara  L.,  leg.  Nowakowski  (Griffiths,  1966: 
807);  Ojkow  National  Park,  on  Petasites  albus  (L.)  (Beiger,  1960). 

Denmark  — H0rsholm,  on  Petasites  hybridus  (L.)  (sheet  in  Hering’s  mine  herbarium). 

Finland  — mines  on  Petasites  frigidus  (L.)  at  Kemi  (Ostrobothnia  borealis)  (Linnaniemi, 
1913)  and  Viborg  (Karelia  australis)  (Frey,  1937). 

Russia  — Moscow  region,  mines  on  Tussilago  farfara  L.  (Rohdendorf,  1960). 

Hendel  (1935)  also  lists  Petasites  paradoxus  (Retz.)  as  a host-plant,  in  addition  to  host 
species  recorded  above. 

Phytomyza  tussilaginis  petasiti  Spencer  1969,  new  status 

Phytomyza  petasiti  Spencer.  Spencer,  1969b: 266.  Holotype  6,  Alberta  (Canada),  in  K.  A. 

Spencer’s  collection. 

Adult.  — Colour  of  head  as  described  for  subspecies  tussilaginis,  but  with  dark  coloration 
of  vertex  less  extensive,  not  enclosing  bases  of  vt  in  specimens  from  Alberta  and  British 
Columbia  (however  vte  on  dark  ground  in  specimens  from  Yukon  and  Alaska);  face  com- 
pletely yellow,  or  at  most  with  weak  traces  of  brown  in  antennal  pits. 

Mesonotum  dark  centrally  (weakly  shining,  finely  grey-dusted),  but  with  strongly  con- 
trasting broad  whitish  side  bands  which  anteriorly  extend  inwards  along  its  anterior  margin 
to  level  of  either  row  of  dc,  and  posteriorly  to  scutellar  suture;  outer  pa  on  yellow  ground  or 
on  boundary  between  yellow  and  dark  ground;  humeral  calli  with  traces  of  infuscation  (a 
distinct  dark  spot  in  some  specimens).  Scutellum  largely  dark,  but  with  traces  of  pale 
coloration  at  its  basal  corners.  Pleura  extensively  whitish,  but  with  dark  anteroventral  area 
of  variable  size  on  mesopleuron  and  in  some  specimens  with  parts  of  pteropleuron  infus- 
cated;  sternopleuron  dark  ventrally,  with  pale  dorsal  band;  hypopleuron  largely  dark.  Wing 
base  and  squamae  white,  latter  with  white  or  ochreous  fringe.  Coxae  pale  apically,  dark  at 
base;  femora  largely  dark  with  contrasting  yellow  tips;  tibiae  and  tarsi  entirely  yellow  or 
yellow-brown. 

Abdomen  extensively  yellowish  (especially  towards  sides),  in  female  with  contrasting 
black  basal  cone  of  ovipositor  (grey-dusted  on  basal  third  to  half). 

Wing  length  2. 4-2. 8 mm.  Costal  ratio  mg2/mg4  2. 6-4.0. 

Material  examined.  — 1 <3,  1 9 from  larvae  4.vii.71  on  Petasites  palmatus  (Ait.),  Elk  Island 
National  Park,  Alberta,  emerged  27.vii.71  and  28.V.72,  leg.  G.  C.  D.  Griffiths;  1 6,  1 9 from 
larvae  25.vii.71  on  Petasites  (?  palmatus  X frigidus),  same  locality,  emerged  1 3 .viii.7 1 and 


384 


Griffiths 


15.V.72,  leg.  G.  C.  D.  Griffiths;  1 9 from  larva  26.ix.71  on  Petasites  sagittatus  (Banks),  same 
locality,  emerged  3.vi.72,  leg.  G.  C.  D.  Griffiths.  1 9 from  larva  6.viii.70  on  Petasites 
palmatus  (Ait.),  Summit  Lake  Pass  (4200  feet  elevation;  Alaska  Highway  mile  392),  British 
Columbia,  emerged  19.V.71,  leg.  G.  C.  D.  Griffiths.  1 6,  1 9 from  larvae  31.viii.69  on  Peta- 
sites sagittatus  (Banks),  East  shore  of  Lake  Teslin,  Yukon  Territory,  emerged  16  & 22.V.70, 
leg.  G.  C.  D.  Griffiths.  1 6 from  larvae  2-3.viii.68  on  Petasites  frigidus  (L.),  Walker  Fork, 
Taylor  Highway,  Alaska,  emerged  23.x. 68  (forced),  leg.  G.  C.  D.  Griffiths. 

Other  records.  — The  holotype  was  bred  from  leaves  of  Petasites  frigidus  (L.)  (=  vitifolius ) 
collected  at  Blairmore,  Alberta  (Spencer,  1969b).  Sehgal  (1971)  records  specimens  bred 
from  Petasites  sagittatus  (Banks)  at  Edmonton  and  Elk  Island  Park  (Alberta). 

Phytomyza  tussilaginis  kevani  new  subspecies 

Adult.  — Frons  yellow  centrally,  with  ocellar  plate  and  vertex  contrastingly  shining 
black  (both  vt  on  dark  ground);  orbits  somewhat  infuscated,  especially  along  eye  margins 
and  around  bases  of  orbital  setae.  Face  clear  yellow  only  on  margins,  extensively  infuscated 
in  antennal  pits.  Genae  yellow.  Occiput  black,  somewhat  shining.  Antennae  with  first  article 
brown,  second  and  third  articles  black.  Palpi  black;  labella  yellow. 

Thorax  weakly  shining,  finely  grey-dusted,  almost  entirely  black,  with  traces  of  pale 
coloration  only  at  margins  of  humeral  calli  (especially  around  anterior  spiracles);  seams  of 
notopleural  and  mesopleural  sutures  white;  wing  base  and  squamae  contrastingly  white, 
latter  with  dark  fringe.  Legs  largely  dark  with  tips  of  femora  yellow  (but  only  those  of  front 
legs  distinctly  so  in  holotype);  tibiae  and  tarsi  brown  or  black.  Abdomen  entirely  dark. 

Wing  length  2. 2-2. 3 mm.  Costal  ratio  mg2/mg4  2. 6-2. 7 (lower  end  of  range  of  values  for 
other  subspecies). 

Types.  — Holotype  6 from  larva  31.vii.70  on  Petasites  frigidus  (L.),  South  shore  of  Yaya 
Lake,  Richards  Island,  Northwest  Territories  (Canada),  emerged  6.V.71,  leg.  P.  G.  Kevan. 
1 6 paratype  from  larva  18.viii.70  on  Petasites  frigidus  (L.),  Triple  Summit  (132°  54'  W,  69° 
32'  N),  Northwest  Territories,  emerged  4.V.71,  leg.  P.  G.  Kevan. 

I am  pleased  to  name  this  subspecies  after  Dr.  Peter  G.  Kevan,  who  collected  material  for 
me  while  working  in  the  Arctic. 

Phytomyza  alpina  Groschke  1957 

Phytomyza  alpina  Groschke.  Groschke  and  Hering,  1957:122.  Holotype  6,  Bavaria  (Ger- 
many), in  Staatliches  Museum  fur  Naturkunde,  Ludwigsburg. 

Adult.  — Head  (Fig.  32)  with  orbits  not  or  only  very  narrowly  projecting  above  eye  in 
lateral  view;  genae  in  middle  1/4  to  1/3  of  eye  height;  eyes  with  only  sparse  fine  pubescence. 
Frons  at  level  of  front  ocellus  about  twice  width  of  eye.  Ors  directed  posteriorly,  ori 
directed  inwardly;  posterior  ors  variably  developed,  in  most  specimens  about  2/3  as  long  as 
anterior  ors,  but  ranging  from  fully  as  long  to  completely  absent;  anterior  ori  in  most  speci- 
mens 1/3  to  1/2  as  long  as  posterior  ori,  but  in  some  absent  or  represented  only  by  very 
small  setulae;  orbital  setulae  more  or  less  one-rowed.  Peristomal  margin  with  vibrissa  and  4-7 
upcurved  peristomal  setulae.  Third  antennal  article  rounded  distally,  with  short  pubescence. 

3 + 1 dc;  acr  in  4-5  rows;  5-10  presutural  ia;  4-10  postsutural  ia;  inner  pa  over  half  as  long 
as  outer  pa. 

Second  cross-vein  (m-m)  absent.  Costal  ratio  mg2/mg4  2.8-3 .4.  Wing  length  2. 2-3. 2 mm. 

Frons  clear  yellow  centrally,  with  ocellar  plate  and  vertex  contrastingly  black  (vte  on 
dark  ground;  vti  on  boundary  between  dark  and  pale  ground);  orbits  largely  yellow,  but  with 


Boreal  Agromyzidae 


385 


traces  of  infuscation  along  eye  margins  and  around  bases  of  orbital  setae.  Face  partly  yellow, 
but  infuscated  in  antennal  pits.  Genae  yellow.  Occiput  black.  Antennae  with  first  article 
yellow-brown,  second  and  third  articles  black.  Palpi  black;  labella  yellow.  Thorax  largely 
dark,  strongly  grey-dusted,  scarcely  shining;  sides  of  mesonotum  with  limited  area  of  pale 
coloration  around  margins  of  humeral  calli  and  on  upper  part  of  sutural  triangle;  scutellum 
dark;  mesopleuron  with  narrow  whitish  dorsal  band  along  notopleural  suture;  seam  of  meso- 
pleural  suture  whitish;  wing  base  and  squamae  yellowish  white,  latter  with  dark  fringe.  Legs 
largely  dark,  with  tips  of  femora  contrastingly  yellow.  Abdomen  largely  black  or  brown. 
Basal  cone  of  ovipositor  (9)  grey-dusted  on  about  basal  half. 

Male  postabdomen  with  8th  sternum  fused  with  6th  tergum.  Telomeres  not  clearly  delim- 
ited from  periandrium,  bearing  only  fine  setulae.  Pregonites  extending  ventrally,  shielding 
base  of  aedeagus  at  rest.  Aedeagal  hood  with  two  pairs  of  lateral  sclerites.  Aedeagus  as  Fig. 
1,  2 and  5;  basal  sclerites  narrow,  slightly  convergent  distally;  dense  group  of  spinules  on 
left  side  near  dorsal  margin  of  left  basal  sclerite;  on  right  side  less  dense  group  of  more  dor- 
sally  situated  spinules  nearer  centre-line;  sclerotization  of  medial  lobe  forming  loop,  con- 
fluent anteriorly  with  basal  sclerites;  distal  section  of  aedeagus  with  pair  of  slender  para- 
mesophalli  and  distiphallus  consisting  of  more  or  less  parallel,  paired  tubules.  Ejaculatory 
apodeme  small  (Fig.  3-4). 

In  most  European  specimens  the  tubules  of  the  distiphallus  appear  rounded  dorsally  in 
lateral  view  (as  Fig.  5).  But  they  appear  more  or  less  angulate  in  some  British  specimens,  as 
in  all  the  specimens  from  North  America  (Fig.  1).  No  clear-cut  morphological  distinction 
can  be  made  between  populations  from  the  two  areas. 

Puparium  and  third  instar  larva.  — Similar  to  those  of  tussilaginis . In  my  British  series 
there  is  an  unusually  wide  range  of  variation  between  individuals  in  the  number  of  spiracular 
bulbs  (anterior  spiracles  with  14-18  bulbs;  posterior  spiracles  with  22-36  bulbs).  The  varia- 
tion is  less  in  North  American  material  (anterior  spiracles  with  12-15  bulbs;  posterior 
spiracles  with  19-25  bulbs)  (Fig.  28).  Puparium  2.0-2. 5 mm  long. 

Mine.  - Larvae  leaf-miners  on  Senecio  and  Petasites,  leaving  leaf  through  semicircular 
slit,  in  most  cases  on  upper  surface,  before  puparium  formation.  Mines  on  upper  surface  of 
leaf,  appearing  white  or  greenish  white  in  reflected  light,  geographically  variable  in  shape 
(Fig.  35-36). 

In  Europe  mines  have  been  reported  only  on  Senecio  alpinus  (L.)  and  S.  jacobaea  L. 
Mine  (Fig.  36)  initially  linear  but  becoming  progressively  broader  and  more  or  less  blotchy 
terminally;  faeces  deposited  as  discrete  particles,  well  separated  (mostly  by  over  1 mm)  in 
terminal  part  of  mine. 

In  North  America  I have  bred  this  species  from  mines  on  Petasites,  as  well  as  on  Senecio 
(see  records  below).  Mine  (Fig.  35)  retaining  its  linear  appearance  throughout,  up  to  15  cm 
long,  2-3  mm  wide  terminally;  faecal  particles  separated  by  about  2 mm  in  terminal  part  of 
mines  on  Senecio  lugens  Richards,  but  more  numerous  and  separated  by  less  than  1 mm  in 
mines  on  Petasites. 

Material  examined.  — 1 6 paratype  from  larvae  on  Senecio  alpinus  (L.),  Partnachklamm, 
Bavaria,  Germany,  emerged  27.viii.51,  leg.  F.  Groschke.  5 66, 7 99  from  larvae  30.vii.62  on 
Senecio  jacobaea  L.,  Ingleborough,  Yorks.,  England,  emerged  26-3 1 .viii.62,  leg.  G.  C.  D. 
Griffiths.  8 66,  5 99  from  larvae  6.ix.64  on  Senecio  jacobaea  L.,  Gorsdale  Scar,  Yorks., 
England,  emerged  5- 12.x. 64'  and  22.iv.65  (1  d),  leg.  G.  C.  D.  Griffiths.  6 66,  1 9 from  larvae 
10.vi.65  on  Senecio  jacobaea  L.,  Mullagh  More,  Clare,  Ireland,  emerged  2-8.vii.65,  leg. 
G.  C.  D.  Griffiths.  2 66  from  larvae  31.viii.66  on  Senecio  jacobaea  L.,  Derreen,  Clare,  Ire- 
land, emerged  6-27.iii.67,  leg.  G.  C.  D.  Griffiths. 

1 6,  2 99  from  larvae  3 1 .viii.69  on  Petasites  sagittatus  (Banks),  on  East  shore  of  Lake 


386 


Griffiths 


Teslin,  Yukon  Territory,  emerged  17-26.V.70,  leg.  G.  C.  D.  Griffiths.  1 d,  1 9 from  larvae 
30.viii.69  on  Senecio  pauperculus  Michx.,  Lake  Laberge,  Yukon  Territory,  emerged  17-19.V. 
70,  leg.  G.  C.  D.  Griffiths.  1 9 from  larvae  19-26.vii.68  on  Petasites  frigidus  (L.),  Eagle  Sum- 
mit (3900  feet  elevation),  Steese  Highway,  Alaska,  emerged  8.X.68  (forced),  leg.  G.  C.  D. 
Griffiths.  3 dd,  2 99  from  larvae  3-ll.viii.70  on  Senecio  lugens  Richards,  Summit  Lake 
Pass  (4200  feet  elevation;  Alaska  Highway  mile  392),  British  Columbia,  emerged  12-13.V.71, 
leg.  G.  C.  D.  Griffiths;  1 9 from  larva  9.ix.71,  same  plant  and  locality,  emerged  ll.v.72, 
leg.  G.  C.  D.  Griffiths. 

Other  records.  — Groschke’s  original  material  was  bred  from  Senecio  alpinus  (L.)  in  the 
Bavarian  Alps  (Partnachklamm  and  Lenggries).  Other  localities  where  mines  on  Senecio 
alpinus  (L.)  have  been  recorded  are  Tolz  (Bavaria),  Kleiner  Walsertal  and  Eisenerzer  Reichen- 
stein  (Austria)  (in  Groschke  and  Hering,  1957),  and  Maloja,  Switzerland  (in  Griffiths,  1966). 
The  first  British  record  was  from  Kinlochewe,  Ross  (Scotland),  mines  on  Senecio  jacobaea 
L.  collected  by  O.  W.  Richards  on  10.vii.53  (Spencer,  1956).  Other  Irish  localities  (all  in 
the  Burren  area  of  County  Clare)  are  given  by  Griffiths  (1968). 

Remarks.  — The  known  distribution  of  this  species  is  indicated  on  Fig.  39. 

I am  not  able  to  distinguish  the  leaf  mines  of  this  species  on  Petasites  in  North  America 
from  those  of  tussilaginis.  Although  the  maximum  length  of  mines  of  tussilaginis  in  the  avail- 
able samples  is  longer,  there  is  overlap  between  the  species  in  respect  of  this  measurement. 

The  species  described  by  Nowakowski  (1962b)  as  Phytomyza  aronici  is  the  sister-species 
of  alpina.  The  aedeagus  of  aronici  is  very  similar  to  that  of  alpina,  particularly  in  respect  of 
the  asymmetrical  development  of  the  groups  of  spinules  (an  undoubtedly  apomorphous 
character).  The  only  clear  differences  which  I have  noted  involve  the  shape  of  the  parameso- 
phalli  and  the  situation  of  the  left  group  of  spinules  closer  to  the  left  basal  sclerite  in 
aronici.  The  type  series  of  aronici  was  bred  from  mines  on  Doronicum  clusii  (All.),  a mem- 
ber of  the  Senecioneae,  in  the  Tatry  Mountains  of  Poland  (1600-2400  metres  elevation). 

Phytomyza  senecionis  Kaltenbach  1 869 

Phytomyza  senecionis  Kaltenbach.  Kaltenbach,  1869:176.  —1874:364.  Hendel,  1935:478. 

Types  lost;  type-locality,  Germany. 

Adult.  — Head  with  orbits  not  or  only  narrowly  projecting  above  eye  in  lateral  view; 
genae  in  middle  about  1/4  of  eye  height;  eyes  with  only  sparse  fine  pubescence.  Frons  at 
level  of  front  ocellus  about  twice  width  of  eye.  Ors  directed  posteriorly,  ori  directed  in- 
wardly; posterior  ors  variably  developed,  ranging  from  two-thirds  as  long  as  anterior  ors  to 
completely  absent;  anterior  ori  short,  not  more  than  half  as  long  as  posterior  ori;  orbital 
setulae  irregularly  one-rowed.  Peristomal  margin  with  vibrissa  and  4-5  upcurved  peristomal 
setulae.  Third  antennal  article  rounded  distally,  with  short  pubescence. 

3 + 1 dc;  acr  in  4-5  rows;  7-10  presutural  ia;  8-1 1 postsutural  ia;  inner  pa  about  half  as 
long  as  outer  pa. 

Second  cross-vein  (m-m)  absent.  Costal  ratio  mg2/mg4  3.3-4. 1.  Wing  length  2. 1-2.6  mm. 

Frons  clear  yellow  except  dark  ocellar  plate  (both  vt  on  yellow  ground).  Face  largely  or 
completely  yellow,  at  most  infuscated  in  antennal  pits.  Genae  yellow.  Occiput  dark.  Anten- 
nae with  first  article  yellow-brown,  second  and  third  articles  brown  to  black.  Palpi  brown  or 
black;  labella  yellow.  Mesonotum  dark  centrally  (weakly  shining,  finely  grey-dusted),  but 
with  strongly  contrasting  broad  whitish  side  bands  which  enclose  the  humeral  calli  (indi- 
cated by  small  dark  spot)  and  extend  posteriorly  to  the  scutellar  suture;  outer  pa  on 
boundary  between  yellow  and  dark  ground;  scutellum  dark;  mesopleuron  whitish  on  upper 
third  to  half,  dark  ventrally;  other  pleura  largely  dark,  but  with  some  pale  coloration  along 


Boreal  Agromyzidae 


387 


sutures.  Wing  base  and  squamae  white,  latter  with  contrastingly  dark  fringe.  Coxae  dark; 
femora  largely  dark  with  contrasting  yellow  tips;  tibiae  and  tarsi  deep  yellow  or  yellow- 
brown.  Abdomen  largely  brown,  but  yellow  on  sides  at  base.  Basal  cone  of  ovipositor  (9) 
grey-dusted  on  basal  half  to  two-thirds. 

Male  postabdomen  with  8th  sternum  fused  with  6th  tergum.  Telomeres  not  clearly  de- 
limited from  periandrium,  bearing  only  fine  setulae.  Ventral  extensions  of  pregonites  incon- 
spicuous, more  or  less  membranous.  Aedeagal  hood  with  two  pairs  of  lateral  sclerites. 
Aedeagus  as  Fig.  11;  basal  sclerites  with  angular  notches  (notch  on  left  sclerite  lower  than 
that  on  right)  near  each  of  which  lies  a group  of  spinules;  sclerotization  of  medial  lobe  form- 
ing loop;  distal  section  small,  with  conspicuous  spine  on  left  side,  with  poorly  differentiated 
distiphallus  (pigmented  only  around  its  margins).  Ejaculatory  apodeme  rather  large  (Fig.  12). 

Puparium  and  third  instar  larva.  — Similar  to  those  of  tussilaginis.  Anterior  spiracles  with 
9-11  bulbs;  posterior  spiracles  with  21-28  bulbs.  Puparium  1. 7-2.1  mm  long. 

Mine.  — Larvae  leaf-miners  on  Senecio.  Mine  (Fig.  37)  entirely  linear,  up  to  23  cm  long, 
in  many  cases  following  midrib  for  long  distance,  remaining  narrow  terminally  (not  more 
than  1.5  mm  wide);  faeces  deposited  as  fine  particles,  in  some  cases  forming  short  strips; 
most  mines  confined  to  upper  surface  of  leaf  (but  some  beginning  on  lower  side  accord- 
ing to  Hering,  1957b),  appearing  contrastingly  white  in  reflected  light;  larvae  leaving  leaf 
through  semicircular  slit  on  upper  surface  before  puparium  formation. 

A figure  of  the  mine  has  also  been  given  by  Hering  (1957b). 

Material  examined.  - 1 6 from  larva  10.vi.54  on  Senecio  fuchsii  Gmel.,  Kunnersdorf  (near 
Gorlitz),  Germany,  emerged  28.vi.54,  leg.  E.  M.  Hering  (no.  6040).  2 99  from  larvae  23.viii.63 
on  Senecio  nemorensis  L.,  North  of  Como,  Italy,  emerged  17-22.ix.63,  leg.  G.  C.  D.  Griffiths. 

Other  records.  — According  to  Hering  (1957b)  the  larvae  of  this  species  occur  commonly 
on  Senecio  nemorensis  L.  and  S.  fuchsii  Gmel.  in  Central  Europe.  Records  are  as  follows. 

Holland  — Berg  en  Dal  and  Valkenburg  on  Senecio  fuchsii  Gmel.  (de  Meijere,  1926,  as 
“ Phytomyza  lappae  Gour.”). 

Germany  — Kaltenbach’s  original  material  was  bred  from  Senecio  nemorensis  L.  (locality 
not  stated).  Buhr  (1964)  lists  localities  in  Saxony,  where  he  reports  this  spe- 
cies as  common  on  Senecio  fuchsii  Gmel.  Voigt  (1929)  records  mines  at  Gei- 
senheim  and  Laacher  See  (Rheingau);  Zoerner  (1969)  at  Saarensee  (Middle 
Elbe  region).  There  are  also  sheets  of  Senecio  fuchsii  Gmel.  in  Hering’s  mine 
herbarium  for  Frankenhausen  (Thuringia),  Alter  Stolberg  (Siidharz),  Siegen 
(Westphalia),  Schlosspark/Torga  (Lausitz),  Lowenburg/Rhondorf  (Rheinland) 
and  the  Mosel  Valley  (Rheinland). 

Austria  — Sheets  of  Senecio  fuchsii  Gmel.  in  Hering’s  mine  herbarium  for  Linz  (Donau) 
and  Tal  der  Grossen  Miihlviertel. 

Czechoslovakia  — Stary  (1930)  lists  localities  where  larvae  were  collected  on  Senecio 
fuchsii  Gmel.  He  also  reports  this  species  on  Senecio  jacobaea  L.  at 
Tisova,  which  record  needs  checking  as  this  plant  is  a host  of  Phyto- 
myza alpina  Groschke  (not  yet  described  when  Stary  wrote). 

Bulgaria  — Rila  Mountains,  on  Senecio  nemorensis  L.  (Buhr,  1941b). 

Poland  — Collected  by  Nunberg  (1948)  and  Nowakowski  (1962a:  152)  on  Senecio  nemo- 
rensis L.,  S.  fuchsii  Gmel.  and  S.  subalpinus  Koch  at  various  localities  in  the 
Tatry  Mountains  (see  also  Griffiths,  1966:797,  807,  and  809);  also  found  by 
Nowakowski  on  Senecio  fluviatilis  Wallr.,  near  Sztum  and  Warsaw  (Nowakow- 
ski, 1962a:  152);  Beiger  (1959,  1965)  gives  records  for  the  district  of  Wieliczka 
(on  Senecio  fuchsii  Gmel.)  and  for  the  Krakow-Wieluri  Jura  (on  Senecio  nemo- 
rensis L.). 


388 


Griffiths 


Denmark  - Maribo  (S^nderup,  1949). 

Whether  this  species  occurs  in  Scandinavia  has  not  been  established.  Ryden’s  (1952) 
record  for  Sweden  is  doubtful,  since  it  is  based  on  a caught  female. 

Phytomyza  ravasternopleuralis  Sasakawa  1955,  new  status 

Phytomyza  senecionis  ravasternopleuralis  Sasakawa.  Sasakawa,  1955:19.  — 1 96 1 a: 468.  Holo- 

type  <3,  Aomori  prefecture  (Japan),  in  Entomological  Laboratory,  Saikyo  University. 

Adult.  — As  described  for  senecionis , except  as  follows. 

Stemopleuron  entirely  dark,  without  pale  dorsal  band  along  suture. 

Distal  section  of  aedeagus  (Fig.  9)  longer,  with  weakly  differentiated  paramesophalli  and 
slender  distiphallus  whose  tubules  are  almost  parallel  apically.  Ejaculatory  apodeme  small 
(Fig.  10). 

For  further  description  and  figures  see  Sasakawa  (1955,  1961a). 

Puparium  and  third  instar  larva.  - Similar  to  those  of  tussilaginis  and  senecionis.  Anterior 
spiracles  with  11-14  bulbs;  posterior  spiracles  with  22-28  bulbs.  Puparium  about  2 mm  long. 
For  further  description  and  figures  see  Sasakawa  (1961a). 

Mine.  — Larvae  leaf-miners  on  Senecio  and  Petasites.  Sasakawa  (1955)  describes  the  mines 
as  linear  throughout  (“ophionome”),  whitish,  yellowish  or  yellowish  green  in  colour,  7-13.5 
cm  long,  on  upper  surface  of  leaf;  faeces  deposited  in  short  strips  or  as  fine  particles;  larvae 
leaving  leaf  through  semicircular  slit  on  upper  surface  before  puparium  formation.  Figures 
of  the  mine  are  given  by  Sasakawa  (1955,  1961a). 

Material  examined.  — 1 6 from  larvae  on  Petasites  japonicus  Miq.,  Mominoki,  Mount 
Sara,  Ehime  prefecture  (Shikoku),  Japan,  leg.  T.  Yano  (May  1954). 

Other  records.  — The  type  series  was  bred  from  Senecio  palmatus  Pall,  at  Towada  Park, 
Aomori  prefecture,  Japan  (Sasakawa,  1955). 

Remarks.  — I propose  to  consider  ravasternopleuralis  a full  species,  rather  than  a sub- 
species of  senecionis  (as  it  was  described  by  Sasakawa),  because  the  form  of  the  disti- 
phallus is  substantially  different  from  that  of  the  only  male  of  senecionis  available  to  me. 
But  this  judgement  is  only  tentative,  in  view  of  the  limited  material  available. 

Sasakawa  (1961a)  referred  specimens  bred  from  Petasites  japonicus  Miq.  to  “ Phytomyza 
lappae  Robineau-Desvoidy”.  However  the  aedeagus  of  the  male  from  this  plant  lent  me  by 
Sasakawa  (Fig.  9)  does  not  agree  with  his  figure  of  “ lappae ”,  but  rather  with  his  figure  of 
ravasternopleuralis . I conclude  that  the  Japanese  flies  from  Petasites  should  be  referred  to 
the  latter  taxon. 


(b)  the  Phytomyza  syngenesiae  group 

I have  discussed  the  definition  of  the  Phytomyza  syngenesiae  group  and  given  descrip- 
tions of  species  in  my  1967  revision.  One  additional  species  ( senecionella ) has  since  been 
described  by  Sehgal  (1971).  In  this  group  the  puparia  remain  inside  the  leaf,  as  in  the 
robustella  group.  But  the  form  of  the  aedeagus  is  very  different,  characterized  ( inter  alia) 
by  inclusion  of  the  terminal  part  of  the  ejaculatory  duct  in  an  unpaired  distal  tubule. 

The  two  polyphagous  species  of  this  group,  Phytomyza  syngenesiae  (Hardy)  and  P. 
horticola  Goureau,  probably  occur  commonly  on  Senecio  (see  records  below).  Whether 
they  also  sometimes  attack  Tussilago  and  Petasites  requires  confirmation.  Records  of  Phy- 
tomyza atricornis  Meigen”  (a  name  formerly  used  for  species  of  this  group)  on  Petasites 
and  Tussilago  in  Europe  were  published,  for  instance,  by  Hering  (1924,  1927),  de  Meijere 
(1926)  and  Stary  (1930).  But  these  records  antedate  the  description  of  P.  farfarae  Hendel, 


Boreal  Agromyzidae 


389 


and  confusion  with  that  species  must  therefore  be  suspected.  In  1967  I dissected  three 
males  identified  as  “ atricornis ” in  Hering’s  collection  and  they  all  proved  to  belong  to 
farfarae.  The  only  European  record  since  the  description  of  farfarae  is  S^nderup’s  (1949) 
report  of  “ atricornis ” on  both  Tussilago  and  Petasites  in  Denmark.  This  record  cannot 
be  checked,  as  no  adult  flies  are  known  to  have  been  obtained  from  his  samples. 

Phytomyza  syngenesiae  (Hardy) 

I have  given  firm  records  for  Senecio  jacobaea  L.,  S.  cruentus  DC.  and  S.  squalidus 
L.  in  Europe  (Griffiths,  1967).  There  are  many  other  published  records  of  “ Phytomy- 
za atricornis  Meigen”  on  species  of  Senecio  in  Europe,  but  I cannot  determine  whether 
they  refer  to  syngenesiae  or  horticola.  In  North  America  Senecio  cruentus  DC.,  S.  mikani- 
oides  Otto  and  Petasites  sp.  are  recorded  as  hosts  (Frick,  1959;  Griffiths,  1967).  The 
last  record  requires  confirmation  in  view  of  possible  confusion  with  species  of  the  robus- 
tella  group. 

My  previous  opinion  (Griffiths,  1967)  that  this  species  dispersed  across  the  Bering  land 
bridge  between  Siberia  and  Alaska  is  now  withdrawn.  The  most  northern  locality  for  syn- 
genesiae in  North  America  is  the  City  of  Edmonton  (Alberta),  where  it  has  been  collected 
only  once  and  does  not  seem  established.  The  species  does  not  occur  in  the  boreal  forest 
nor  in  coastal  regions  of  Alaska,  where  trans-Beringian  migrants  are  expected  to  occur. 
The  only  species  of  the  syngenesiae  group  so  far  found  in  such  northern  areas  is  senecio- 
nella.  I now  think  it  more  likely  that  syngenesiae  was  introduced  into  North  America  with 
horticultural  plants. 

Phytomyza  horticola  Goureau 

I have  given  firm  records  for  Senecio  vulgaris  L.,  S.  vernalis  W.  et  K.  and  S.  doria  L.  in 
Europe  (Griffiths,  1967).  For  Japan  Sasakawa  (1961b)  lists  Senecio  vulgaris  L .,S.  cruentus 
DC.  and  Petasites  japonicus  Miq.  as  host  plants. 

Phytomyza  senecionella  Sehgal 

Sehgal  (1971)  has  described  this  species  on  the  basis  of  material  bred  from  Senecio  con- 
gestus  (R.  Br.)  var.  palustris  (L.)  at  Elk  Island  National  Park  (shores  of  Tawayik  Lake), 
Alberta.  His  figure  123  indicates  that  the  distal  tubule  of  the  aedeagus  has  a characteristic 
sinuate  shape,  by  which  the  species  may  be  distinguished  from  syngenesiae.  I have  figured 
(Fig.  13)  the  aedeagus  of  a male  bred  from  Petasites  frigidus  (L.)  at  Eagle  Summit  (3900 
feet  elevation),  Steese  Highway,  Alaska  (emerged  26.vii.68  from  puparium  collected  19.vii. 
68,  leg.  G.  C.  D.  Griffiths).  Probably  Petasites  is  only  an  occasional  host  of  this  species,  for 
only  a single  mine  was  found.  The  main  host  at  Eagle  Summit  was  Senecio  atropurpureus 
(Ledeb.)  subsp.  tomentosus  (Kjellm.)  (1  6,  2 99  emerged  24-26.vii.68  from  puparia  collected 
17.vii.68).  Mines  were  also  found  the  same  day  on  Senecio  yukonensis  Pors.  While  the  male 
genitalia  of  these  Alaskan  specimens  agree  substantially  with  Sehgal’s  figure,  the  coloration 
of  the  head  is  darker  than  in  the  original  series,  with  the  frons  largely  orange-brown  with 
grey-dusted  orbits. 

Leaf  mines  collected  at  Eagle  Summit  are  6-7  cm  long,  1. 5-2.0  mm  wide  terminally; 
faecal  particles  discrete,  mostly  separated  by  0.75-1.00  mm  in  terminal  part  of  mine;  mines 
formed  mainly  on  upper  surface  of  leaf,  with  puparium  formation  following  in  most  cases 
on  lower  surface. 


390 


Griffiths 


(c)  the  Phytomyza  robustella  group 

The  term  “ Phytomyza  robustella  group”  has  been  used  to  include  certain  species  whose 
larvae  produce  gall-like  swellings  in  the  midrib  of  the  leaves  of  Compositae.  However  no 
clear  morphological  distinction  can  be  drawn  between  these  species  and  certain  other  species 
whose  larvae  are  leaf-  or  stem-miners  on  Compositae,  not  forming  swellings.  Spencer  (1971) 
has  already  referred  to  this  group  one  such  species,  P.  buhriella  Spencer  (as  notabilis ).  In  the 
present  work  I treat  that  species  and  some  additional  leaf-mining  species  whose  male  geni- 
talia are  of  similar  type.  The  puparia  of  all  species  referred  to  this  group  remain  in  the  plant 
tissue,  with  the  anterior  spiracles  turned  downwards  so  that  they  project  ventrally  through 
the  epidermis.  A similar  apomorphous  type  of  puparium  is  shown  by  the  Phytomyza  syn- 
genesiae  group  and  related  groups,  as  discussed  in  the  first  paper  of  this  series  (Griffiths, 
1972).  Species  of  the  Phytomyza  robustella  group  differ  from  those  groups  most  obviously 
in  respect  of  the  structure  of  the  aedeagus,  retaining  a forked  distiphallus  (containing  the 
bifid  terminal  portion  of  the  ejaculatory  duct)  and  lacking  “supporting  sclerites”.  However 
since  these  aedeagal  characters  are  probably  plesiomorphous  for  Phytomyza , they  do  not 
provide  a satisfactory  indication  of  the  limits  of  the  group.  My  delimitation  of  the  “ Phyto- 
myza robustella  group”  is  therefore  only  tentative. 

Phytomyza  buhriella  Spencer  1969 

Phytomyza  spec.  Hering,  1957a:93.  —1957b: 746  (no.  3604a). 

Phytomyza  buhriella  Spencer.  Spencer,  1969a:21.  Holotype  6,  Miihlhausen  (Germany),  in 

K.  A.  Spencer’s  collection. 

Phytomyza  notabilis  Spencer.  Spencer,  1971:182.  Holotype  6,  Edinburgh  (Scotland),  in 

University  Museum,  Oxford.  New  synonymy. 

The  synonymy  of  notabilis  has  been  pointed  out  in  correspondence  by  M.  von  Tschim- 
haus. 

Adult.  — External  form  and  colour  as  described  by  Spencer  (1969a). 

Male  postabdomen  with  8th  sternum  fused  with  6th  tergum.  Telomeres  not  clearly  de- 
limited from  periandrium  (without  suture  on  outer  side),  bearing  numerous  fine  setulae. 
Pregonites  not  extending  ventrally.  Aedeagal  hood  with  one  pair  of  lateral  sclerites.  Aedea- 
gus as  in  Fig.  15;  basal  sclerites  rather  broad,  more  or  less  parallel  distally;  medial  lobe  large, 
with  distinct  sclerites;  distal  section  with  broad  paramesophalli  whose  sclerotization  is  con- 
fluent with  V-shaped  distiphallus.  Ejaculatory  apodeme  as  Fig.  16. 

Puparium  and  third  instar  larva.  — See  the  detailed  larval  description  given  by  Hering 
(1957a: 93)  (as  Phytomyza  spec.).  The  larvae  can  be  distinguished  from  those  of  all  other 
species  treated  in  this  paper  by  the  presence  of  a pair  of  sclerotized  processes  of  the  mandi- 
bular adductor  apodeme  on  either  side  of  the  labial  sclerite  (as  also  in  gall-forming  species  of 
the  robustella  group),  and  the  large  size  and  annulate  appearance  of  the  frontal  process. 
Spiracles  (both  anterior  and  posterior)  with  18-22  bulbs.  Puparia  white. 

Mine.  - Larvae  miners  on  Petasites  and  Tussilago.  Mines  linear,  formed  mainly  in  petioles 
of  basal  leaves  (but  in  some  cases  beginning  in  leaf  vein);  faecal  particles  inconspicuous 
(Hering,  195 7b:  746).  Puparium  with  its  ventral  surface  adjacent  to  surface  of  petiole,  with 
its  anterior  spiracles  projecting  ventrally  through  epidermis. 

Material  examined.  — 5 66  swept  on  Tussilago  farfara  L.,  28.V.67,  Ihlkate,  SW  Kiel, 
Germany,  leg.  M.  von  Tschirnhaus. 

Other  records.  — The  type  series  of  buhriella  was  bred  by  H.  Buhr  from  Petasites 
albus  (L.)  at  Miihlhausen,  Thuringia,  Germany.  Buhr  (1964)  has  also  reported  mines  from 


Boreal  Agromyzidae 


391 


Oberwiesenthal  (Erzgebirge).  Von  Tschirnhaus  has  obtained  further  material  (both  swept 
and  bred)  from  Tussilago  farfara  L.  in  Schleswig-Holstein.  In  Britain  this  species  is  known 
from  a male  taken  at  Edinburgh,  Scotland,  2.vi.  1905  (holotype  of  notabilis );  and  I found 
puparia  in  petioles  of  Tussilago  farfara  L.  at  Leeds,  Yorks.,  in  October  1964. 

Phytomyza  farfarae  Hendel  1935 

Phytomyza  farfarae  Hendel.  Hendel,  1935:400.  De  Meijere,  1938:90.  Holotype  9,  without 

locality  label  (presumably  Austria),  in  the  Naturhistorisches  Museum,  Vienna. 

Adult.  — Head  with  orbits  only  very  narrowly  projecting  above  eye  in  lateral  view;  genae 
in  middle  about  1/3  of  eye  height;  eyes  with  sparse  fine  pubescence  or  virtually  bare. 
Frons  at  level  of  front  ocellus  about  twice  width  of  eye.  Two  ors,  of  equal  length,  pos- 
teriorly directed;  only  one  strong  ori,  inwardly  directed  (anterior  ori  absent  or  represented 
by  short  setula);  orbital  setulae  one-rowed.  Peristomal  margin  with  vibrissa  and  3-4  upcurved 
peristomal  setulae.  Third  antennal  article  rounded  distally,  with  short  pubescence. 

3 + 1 dc;  acr  few,  in  two  rows;  ia  few  (4-5  presutural;  1-3  postsutural);  inner  pa  1/3  to 
1/2  as  long  as  outer  pa. 

Second  cross-vein  (m-m)  absent.  Costal  ratio  mg2/mg4  2. 1-2.7.  Wing  length  2. 1-2.7  mm. 

Frons  deep  yellow  or  brown  centrally,  with  ocellar  plate  and  vertex  contrastingly  dark 
(both  vt  on  dark  ground,  or  vti  on  boundary  between  dark  and  pale  ground);  orbits  grey 
along  eye  margins.  Face  largely  dark  brown,  in  some  specimens  becoming  yellow  towards 
sides.  Genae  deep  yellow  or  brown.  Occiput  black.  Antennae  with  first  article  yellow-brown 
or  brown,  second  and  third  articles  dark  brown  or  black.  Palpi  black;  labella  yellow.  Thorax 
dark,  strongly  grey-dusted,  scarcely  shining,  with  pale  coloration  only  along  notopleural  and 
mesopleural  sutures;  wing  base  yellow  or  ochreous;  squamae  with  dark  margin  and  fringe. 
Legs  largely  dark,  with  tips  of  femora  contrastingly  yellow.  Abdomen  largely  brown.  Basal 
cone  of  ovipositor  (9)  grey-dusted  on  about  basal  two-thirds. 

Male  postabdomen  with  8th  sternum  fused  with  6th  tergum.  Telomeres  delimited  from 
periandrium  by  distinct  suture  on  outer  side,  bearing  numerous  fine  setulae.  Pregonites  not 
extending  ventrally.  Aedeagal  hood  with  one  pair  of  lateral  sclerites.  Aedeagus  as  in  Fig. 
20-21 ; basal  sclerites  rather  broad,  divergent  distally;  medial  lobe  without  or  with  only  weak 
traces  of  sclerotization;  distal  section  with  pair  of  small  paramesophalli  and  large  Y-shaped 
distiphallus.  Ejaculatory  apodeme  as  Fig.  22. 

Puparium  and  third  instar  larva.  - Mandibles  with  two  alternating  teeth;  right  mandible 
longer  than  left.  Spiracles  as  described  and  figured  by  de  Meijere  (1938:90);  anterior  spira- 
cles knob-shaped,  with  about  9 bulbs;  posterior  spiracles  also  knob-shaped,  about  same  size 
as  anterior  spiracles,  with  7-11  bulbs.  Puparium  white,  2. 2-2. 7 mm  long. 

Mine.  — Larvae  leaf-miners  on  Tussilago  and  Petasites.  Mine  (Fig.  33A)  interparenchymal, 
pale  green  in  reflected  light,  entirely  linear,  24-28  cm  long,  about  1 mm  wide  terminally, 
in  many  cases  with  long  straight  stretches  besides  some  of  main  veins;  faeces  deposited 
in  very  fine  particles,  often  forming  long  beaded  strips  (in  Perlschniiren  zusammenhan- 
gend)  on  one  side  of  mine;  main  part  of  mine  formed  on  upper  surface  of  leaf,  but 
with  puparium  formed  at  end  of  short  channel  on  lower  surface.  Puparium  with  its  ventral 
surface  adjacent  to  surface  of  leaf,  with  its  anterior  spiracles  projecting  ventrally  through 
epidermis. 

Material  examined.  — 3 66,  4 99  from  puparia  19.viii.64  on  Tussilago  farfara  L.,  Mosem 
(1250  metres  elevation),  Tirol,  Austria,  emerged  28.viii-7.ix.64,  leg.  G.  C.  D.  Griffiths.  1 6 
from  puparium  4.ix.55  on  Petasites  sp.,  Garmisch,  Bavaria,  Germany,  emerged  21.ix.55, 
leg.  K.  A.  Spencer. 


392 


Griffiths 


Other  records.  — This  species  can  be  definitely  accepted  as  occurring  only  in  central 
Europe  and  the  Balkans.  Additional  firm  records  are  as  follows. 

Austria  — found  “everywhere”  according  to  Hendel  (1935);  Stanzach  im  Lechtal,  on 
Tussilago  farfara  L.  (de  Meijere,  1938);  also  sheets  in  Hering’smine  herbarium 
for  Mauthen  (Carinthia)  on  Petasites  albus  (L.),  Tal  der  Grossen  Miihlviertel 
on  Petasites  paradoxus  (Retz.),  Heiligenbliit  (Tauern)  on  Tussilago  farfara  L., 
and  Warscheneck-Gebirge  (Linzershaus,  1400  metres  elevation)  on  Tussilago 
farfara  L. 

Germany  - Miihlhausen,  Thuringia,  on  Petasites  albus  (L.)  (Buhr,  1960);  Lenggries,  Ba- 
varia, on  Tussilago  farfara  L.,  leg.  Groschke  (Griffiths,  1966). 

Poland  — common  in  the  Tatry  Mountains  (Nowakowski,  in  correspondence). 

Roumania  — Sinaia,  ix.57  (sheet  in  Hering’s  mine  herbarium). 

Bulgaria  — Rila  Mountains,  on  Tussilago  farfara  L.  and  Petasites  albus  (L.)  (Buhr,  1941b). 

Records  for  Denmark  (Ryden,  Lyneborg  and  Nielsen,  1963)  and  Angermanland,  Sweden 
(Ryden,  1956)  cannot  be  accepted,  as  they  are  based  on  caught  specimens  which  have  not 
been  dissected.  Ry den’s  (1947)  record  for  Jamtland,  Sweden,  is  almost  certainly  erroneous, 
since  the  fly  was  bred  from  Solidago.  Hendel  (1935)  also  referred  to  this  species  a series  of 
caught  specimens  from  Jakutsk  (Siberia)  in  the  Leningrad  Museum.  They  were  misidenti- 
fied;  on  dissection  I found  that  they  belong  to  the  grass-feeding  Phytomyza  fuscula  Zetter- 
stedt,  in  the  sense  clarified  by  Spencer  (1969b). 

Phytomyza  hyperborea  new  species  (9) 

Adult.  — As  described  for  farfarae,  except  as  follows. 

Third  antennal  article  with  long  upcurved  pubescence  distally  (Fig.  31). 

Costal  ratio  mg2/mg4  2.6.  Wing  length  2.4  mm. 

Frons  largely  deep  yellow,  slightly  grey-dusted  along  eye-margins,  with  ocellar  plate  con- 
trastingly dark;  dark  coloration  of  vertex  less  extensive  (vti  on  yellow  ground,  vte  on  boun- 
dary between  dark  and  yellow  ground).  Face  deep  yellow,  only  slightly  infuscated  in  anten- 
nal pits.  Genae  deep  yellow.  Wing  base  contrastingly  yellow;  squamae  pale  with  ochreous 
fringe.  Abdomen  largely  brown,  but  yellow  on  sides  at  base  and  with  yellow  bands  on  hind 
margins  of  all  terga. 

Puparium  and  third  instar  larva.  — Mandibles  with  two  alternating  teeth;  right  mandible 
longer  than  left.  Spiracles  knob-shaped,  anterior  with  about  20  bulbs,  posterior  with  15 
bulbs.  Puparium  reddish  yellow,  2.6  mm  long. 

Mine.  - The  single  specimen  was  bred  from  a puparium  found  at  the  end  of  a linear  mine 
with  widely  spaced  faecal  particles  on  the  upper  surface  of  a leaf  of  Petasites  frigidus  (L.). 
Since  much  of  the  mine  had  been  destroyed  through  the  feeding  of  a large  tephritid  larva  in 
the  same  leaf,  a full  description  is  not  possible.  Puparium  with  its  ventral  surface  adjacent 
to  upper  surface  of  leaf,  with  its  anterior  spiracles  projecting  ventrally  through  epidermis. 

Type.  - Holotype  9 from  puparium  2.viii.68  on  Petasites  frigidus  (L.),  Walker  Fork, 
Taylor  Highway,  Alaska,  emerged  2 1.x. 68,  leg.  G.  C.  D.  Griffiths. 

Remarks.  — Long  pubescence  on  the  third  antennal  article  is  also  shown  by  Phytomyza 
ciliata  Hendel,  a European  species  of  the  robustella  group  whose  larvae  mine  the  leaves  of 
Chrysanthemum  leucanthemum  L.  Phytomyza  hyperborea  differs  from  that  species  as  fol- 
lows: (1)  orbits  not  projecting  above  eye  in  lateral  view;  (2)  higher  costal  ratio  mg2/mg4 
(less  than  2.0  in  ciliata );  and  (3)  more  extensive  yellow  coloration  (vti  on  yellow  ground, 
face  largely  yellow,  abdomen  yellow  on  sides  at  base  and  with  yellow  bands  on  hind  margins 
of  all  terga). 


Boreal  Agromyzidae 


393 


Under  natural  conditions  the  holotype  would  not  have  emerged  until  the  following 
spring.  The  late  autumn  emergence  was  due  to  delay  in  my  obtaining  outdoor  storage 
facilities. 


Phytomyza  hypophylla  new  species 

Adult.  — As  described  for  farfarae,  except  as  follows. 

Head  (Fig.  30)  with  orbits  more  distinctly  projecting  above  eye  in  lateral  view;  genae  in 
middle  1/2  to  2/5  of  eye  height.  Third  antennal  article  with  slightly  longer  pubescence  than 
in  farfarae  (but  not  so  long  as  in  hyperborea ). 

Costal  ratio  mg2/mg4  1.7-2. 1 (lower  than  in  both  farfarae  and  hyperborea ).  Wing  length 
2. 1-2.7  mm. 

Head  darker  coloured;  orbits  entirely  dark,  densely  grey-dusted;  centre  of  frons  grey- 
dusted  over  brown  to  reddish  black  ground  colour;  face  largely  dark  brown  or  black;  genae 
brown. 

Male  postabdomen  and  genitalia  very  similar  to  those  of  farfarae,  but  with  some  differ- 
ence in  shape  of  distiphallus  (Fig.  23-24).  Ejaculatory  apodeme  larger  (Fig.  25). 

Puparium  and  third  instar  larva.  — As  in  farfarae.  Spiracles  knob-shaped,  anterior  with 
8-10  bulbs,  posterior  (Fig.  29)  with  7-1 1 bulbs.  Puparium  white,  2. 1-2.7  mm  long  (Fig.  27). 

Mine.  — Larvae  leaf-miners  on  Petasites.  Mine  (Fig.  34)  formed  mainly  on  lower  surface 
of  leaf,  basically  linear  but  usually  convolute  within  restricted  area,  forming  irregular  secon- 
dary blotch,  very  inconspicuous  (virtually  concealed  in  reflected  light  by  dense  pile  of  leaf); 
on  upper  surface  of  leaf  at  most  short  stretches  of  mine  channel  or  area  of  brownish  dis- 
coloration visible;  faeces  deposited  as  very  fine  particles,  in  some  cases  forming  short  strips. 
Puparium  with  its  ventral  surface  adjacent  to  lower  surface  of  leaf,  with  its  anterior  spiracles 
projecting  ventrally  through  epidermis. 

Types.  — Holotype  6;  2 66,  7 99  paratypes  from  larvae  and  puparia  19-26.vii.68  on  Peta- 
sites frigidus  (L.),  Eagle  Summit  (3900  feet  elevation),  Steese  Highway,  Alaska,  emerged 
26.vii-12.viii.68,  leg.  G.  C.  D.  Griffiths.  3 66  paratypes  from  puparia  1 6-25 .viii. 7 1 on  Peta- 
sites hyperboreus  Rydb.,  near  Mount  Cavell  Chalet  (5800-7400  feet  elevation),  Jasper  Na- 
tional Park,  Alberta,  emerged  23.viii-3.ix.71,  leg.  G.  C.  D.  Griffiths. 

Remarks.  — I have  also  bred  a female  from  undersurface  mines  collected  at  Walker  Fork, 
Alaska  (from  larvae  and  puparia  2-3.viii.68  on  Petasites  frigidus  (L.),  emerged  20. viii. 68, 
leg.  G.  C.  D.  Griffiths).  In  this  specimen  the  colour  of  the  head  is  as  in  farfarae,  with  the 
frons  largely  yellow-brown.  In  the  absence  of  associated  males  I cannot  judge  whether  this 
specimen  represents  an  additional  undescribed  species  of  the  robustella  group,  or  a colour 
variant  of  hypophylla. 


Phytomyza  lugentis  new  species 

Adult.  — As  described  for  farfarae,  except  as  follows. 

Costal  ratio  mg2/mg4  1.9-2. 2.  Wing  length  2. 5-2. 8 mm. 

Head  darker  coloured  (compare  also  hypophylla)',  centre  of  frons  brown  or  ochreous; 
face  largely  black;  genae  brown;  antennae  entirely  black;  labella  yellow-brown  or  red-brown. 
Abdomen  black.  Basal  cone  of  ovipositor  (9)  grey-dusted  on  about  basal  third  to  half. 

Male  postabdomen  and  genitalia  similar  to  those  of  farfarae  in  most  respects,  but  with 
clear  differences  in  form  of  aedeagus  (Fig.  17-18).  Distal  section  of  aedeagus  with  larger  Y- 
shaped  distiphallus,  without  paramesophalli;  medial  lobe  with  loop  of  unpigmented  scleroti- 
zation.  Ejaculatory  apodeme  larger  (Fig.  19). 


394 


Griffiths 


Puparium  and  third  instar  larva.  — As  in  farfarae.  Spiracles  knob-shaped,  anterior  with 
9-10  bulbs,  posterior  with  7-10  bulbs.  Puparium  white,  2.3-2.6  mm  long. 

Mine.  — Larvae  leaf-miners  on  Senecio  lugens  Richards  and  S.  sheldonensis  Pors.  Mine 
formed  on  upper  or  lower  surface  of  leaf  (largely  on  lower  surface  in  most  cases),  variable  in 
shape  (more  or  less  linear  throughout,  or  partly  linear  with  irregular  blotchy  areas);  faeces 
deposited  as  discrete  particles  (separated  by  over  1 mm  in  terminal  part  of  mine).  Puparium 
with  its  ventral  surface  adjacent  to  lower  surface  of  leaf,  with  its  anterior  spiracles  project- 
ing ventrally  through  epidermis. 

Types.  — Holotype  <3;  2 66,  4 99  paratypes  from  puparia  3-1  l.viii.70  on  Senecio  lugens 
Richards,  Summit  Lake  Pass  (4200  feet  elevation;  Alaska  Highway  mile  392),  British  Colum- 
bia, emerged  7-15.viii.70,  leg.  G.  C.  D.  Griffiths.  2 66  paratypes  from  puparia  5.viii.70  on 
Senecio  sheldonensis  Pors.,  same  locality  (5000  feet  elevation),  emerged  12.viii.70,  leg. 
G.  C.  D.  Griffiths. 


ACKNOWLEDGEMENTS 

Material  for  study  was  lent  by  K.  A.  Spencer  (London,  England),  M.  Sasakawa  (Kyoto 
Prefectural  University,  Japan),  M.  von  Tschimhaus  (University  of  Kiel,  Germany),  H.  J. 
Hannemann  (Zoologisches  Museum,  Humboldt  University,  Berlin)  and  A.  Kaltenbach  (Nat- 
urhistorisches  Museum,  Vienna).  I am  grateful  to  J.  P.  Dear  and  K.  A.  Spencer  for  informa- 
tion on  the  Hering  mine  herbarium.  My  wife  Deirdre  prepared  the  illustrations  of  leaf  mines 
(Fig.  33-37).  Financial  assistance  for  this  study  was  obtained  from  grant  NRC  A-1399,  held 
by  G.  E.  Ball,  and  from  the  Boreal  Institute  of  the  University  of  Alberta. 

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Boreal  Agromyzidae 


397 


Fig.  1-3.  Phytomyza  alpina  Groschke  (<3),  Lake  Teslin,  Yukon:  1,  aedeagus  and  associated  structures  in  lateral  view 
(AEDAD  aedeagal  apodeme,  AedH  aedeagal  hood,  DPH  distiphallus,  Ml  medial  lobe,  PMPH  paramesophallus,  POG 
postgonite);  2,  distal  section  and  medial  lobe  of  aedeagus  in  anteroventral  view  (lettering  as  Fig.  1);  3,  ejaculatory 
apodeme.  Fig.  4-5.  Phytomyza  alpina  Groschke  (<3),  England:  4,  ejaculatory  apodeme;  5,  aedeagus  in  lateral  view. 


398 


Griffiths 


Fig.  6-8.  Phytomyza  tussilaginis  Hendel  (<3),  England:  6,  aedeagus  in  lateral  view;  7,  distal  section  and  medial  lobe  of 
aedeagus  in  ± ventral  view;  8,  ejaculatory  apodeme.  Fig.  9-10.  Phytomyza  ravasternopleuralis  Sasakawa  (<3):  9,  aedeagus 
in  lateral  view;  10,  ejaculatory  apodeme.  Fig.  11-12.  Phytomyza  senecionis  Kaltenbach  (<5):  11,  aedeagus  in  lateral  view; 
12,  ejaculatory  apodeme. 


Boreal  Agromyzidae 


399 


Fig.  13-14.  Phytomyza  senecionella  Sehgal  (c$).  Alaska:  13,  aedeagus  in  lateral  view;  14,  ejaculatory  apodeme.  Fig. 
15-16.  Phytomyza  buhriella  Spencer  (<3):  15,  aedeagus  in  lateral  view  (after  Spencer,  1969a,  slightly  modified);  16, 
ejaculatory  apodeme.  Fig.  17-19.  Phytomyza  lugentis  n.  sp.,  holotype  S '■  17,  aedeagus  in  lateral  view;  18,  distiphallus 
in  ventral  view;  19,  ejaculatory  apodeme. 


400 


Griffiths 


Fig.  20-22.  Phytomyza  farfarae  Hendel  (<5),  Tirol,  Austria:  20,  aedeagus  in  lateral  view;  21,  distal  section  and  medial 
lobe  of  aedeagus  in  anteroventral  view;  22,  ejaculatory  apodeme.  Fig.  23-25.  Phytomyza  hypophylla  n.  sp.,  holotype  6 : 
23,  aedeagus  and  associated  structures  in  lateral  view;  24,  distal  section  and  medial  lobe  of  aedeagus  in  anteroventral 
view;  25,  ejaculatory  apodeme. 


Boreal  Agromyzidae 


401 


■1 


Fig.  26.  Phytomyza  tussilaginis  Hendel,  puparium  in  lateral  view.  Fig.  27.  Phytomyza  hypophylla  n.  sp.,  puparium  in 
lateral  view.  Fig.  28.  Phytomyza  alpina  Groschke,  posterior  spiracle  of  puparium  in  caudal  view.  Fig.  29.  Phytomyza 
hypophylla  n.  sp.,  posterior  spiracle  of  puparium  in  caudal  view.  Fig.  30.  Phytomyza  hypophylla  n.  sp.,  head  in  left 
lateral  view.  Fig.  31.  Phytomyza  hyperborea  n.  sp.  (holotype  9),  head  in  left  lateral  view.  Fig.  32.  Phytomyza  alpina 
Groschke,  head  in  left  lateral  view. 


402 


Griffiths 


Fig.  33.  Leaf  of  Tussilago  farfara  L.  with  mines  of  Phytomyza  farfarae  Hendel  (A)  and  P.  tussilaginis  Hendel  (B). 


Boreal  Agromyzidae 


403 


Fig.  34.  Leaf  of  Petasites  hyperboreus  Rydb.  (lower  surface),  with  mine  of  Phytomyza  hypophylla  n.  sp. 


404 


Griffiths 


Fig.  35-36.  Leaf  mines  of  Phytomyza  alpina  Groschke:  35,  on  Senecio  lugens  Richards  (Canada);  36,  on  Senecio  jacobaea 
L.  (England).  Fig.  37.  Leaf  mine  of  Phytomyza  senecionis  Kaltenbach  on  Senecio  nemorensis  L. 


Boreal  Agromyzidae 


405 


Fig.  38.  Collection  sites  for  Phytomyza  tussilaginis  Hendel  (■  ssp.  tussilaginis ; • ssp.  petasiti ; ▲ ssp.  kevani).  Fig.  39. 
Collection  sites  for  Phytomyza  alpina  Groschke. 


406 


ANNOUNCEMENT 

Biting  Fly  Control  and  Environmental  Quality  — Proceedings  of  a Symposium 
held  at  the  University  of  Alberta  in  Edmonton,  May  16,  17,  and  18,  1972. 

The  proceedings  of  this  symposium,  which  was  organized  jointly  by  the  Department  of 
Entomology  at  the  University  of  Alberta  and  the  Advisory  Committee  on  Entomology  of 
the  Defence  Research  Board,  are  expected  to  be  published  before  the  end  of  1972.  A 
further  announcement  will  follow. 


<1 


Publication  of  Quaestiones  Entomologicae  was  started  in  1965  as  part 
of  a memorial  project  for  Professor  E.  H.  Strickland,  the  founder  of  the 
Department  of  Entomology  at  the  University  of  Alberta  in  Edmonton 
in  1922. 

It  is  intended  to  provide  prompt  low-cost  publication  for  accounts  of 
entomological  research  of  greater  than  average  length,  with  priority 
given  to  work  in  Professor  Strickland’s  special  fields  of  interest  including 
entomology  in  Alberta,  systematic  work,  and  other  papers  based  on  work 
done  at  the  University  of  Alberta. 

Copy  should  conform  to  the  Style  Manual  for  Biological  Journals 
published  by  the  American  Institute  of  Biological  Sciences,  Second 
Edition,  1964,  except  as  regards  the  abbreviations  of  titles  of  periodicals 
which  should  be  those  given  in  the  World  List  of  Scientific  Periodicals, 
1964  Edition.  The  appropriate  abbreviation  for  this  journal  is  Quaest.  ent. 
An  abstract  of  not  more  than  500  words  is  required.  All  manuscripts  will 
be  reviewed  by  referees. 

Illustrations  and  tables  must  be  suitable  for  reproduction  on  a page 
size  of  93/4x63/4  inches,  text  and  tables  not  more  than  7 %X4%  inches, 
plates  and  figures  not  more  than  8V2  X 5 inches.  Reprints  must  be  ordered 
when  proofs  are  returned,  and  will  be  supplied  at  cost.  Subscription  rates 
are  the  same  for  institutions,  libraries,  and  individuals,  $4.00  per 
volume  of  4 issues,  normally  appearing  at  quarterly  intervals;  single 
issues  $1.00.  An  abstract  edition  is  available,  printed  on  one  or  both 
sides  (according  to  length)  of  3X5  inch  index  cards  (at  $1.00  per 
volume)  or  on  5X8  inch  standard  single  row  punched  cards  ($1.50  per 
volume) . 

Communications  regarding  subscriptions  and  exchanges  should  be 
addressed  to  the  Subscription  Manager  and  regarding  manuscripts  to: 

The  Editor,  Quaestiones  Entomologicae, 
Department  of  Entomology, 

University  of  Alberta,  Edmonton,  Canada. 


Quaestiones 

entomologicae 


:97 


1 vm  V A ••  J 

iVERS]  ry 


ENTOMOLOGY  & EDUCATION 


Proceedings  of  a Symposium  organized  by  the 

Department  of  Entomology,  University  of  Alberta 

on  the  Occasion  of  the  50th  Anniversary 

of  its  Foundation 
May  19,  1972 

A periodical  record  of  entomological  investigations, 
published  at  the  Department  of  Entomology, 
University  of  Alberta,  Edmonton,  Canada. 


VOLUME  VIII 


SUPPLEMENT 


1972 


ENTOMOLOGY  & EDUCATION 


Proceedings  of  a Symposium  organized  by  the 
Department  of  Entomology,  University  of  Alberta 
on  the  Occasion  of  the  50th  Anniversary 
of  its  Foundation 


May  19,  1972 

Lister  Hall,  University  of  Alberta, 
Edmonton,  Alberta. 


11 


ACKNOWLEDGEMENTS 

We  are  grateful  for  financial  support  from  the  University  of  Alberta  and  for  the  enthu- 
siasm and  concern  shown  by  the  speakers,  chairmen,  discussion  leaders,  rapporteurs  and 
participants.  Special  thanks  are  due  to  Brian  Hocking,  Don  Whitehead,  Ron  Gooding,  Ollie 
Frost,  Tony  Thomas,  Doug  Craig,  George  Braybrook,  Brian  Rolseth,  Jack  Scott,  Natalie 
Daviduk,  Marli  Engeland,  Linda  Meissenheimer,  and  Lucille  Queyrane  who  kept  the  sym- 
posium running  smoothly  or  who  assisted  in  the  preparation  of  these  proceedings. 


PREFACE 

To  celebrate  the  50th  anniversary  of  its  foundation  by  E.  H.  Strickland,  the  Department 
of  Entomology  at  the  University  of  Alberta  hosted  two  symposia  during  the  week  of 
May  14.  1972.  The  proceedings  of  the  first  on  “Biting  Fly  Control  and  Environmental 
Quality”  are  to  be  published  by  the  Defence  Research  Board  of  the  Canada  Department  of 
National  Defence.  Those  of  the  second  are  presented  here. 

The  texts  of  the  formal  presentations  appear  here  as  they  did  in  the  authors’  final  type- 
scripts except  for  a few  changes  in  grammar  and  some  adjectival  deletions.  The  discussions, 
as  transcribed  from  tape  recordings  of  the  deliberations,  have  been  left  colloquial  in  the 
hope  that  some  of  the  heat  there  generated  will  come  through  in  the  printed  text.  Much  of 
the  excess  verbiage  typical  of  spontaneous  oral  presentation  has,  however,  been  removed. 

Bruce  Heming 
June,  1972 


Ill 


TABLE  OF  CONTENTS 

Acknowledgements ii 

Preface  ii 

Introduction 

B.  S.  Heming iv 

University  Education  and  High  School  Biology 

Don  N.  Jantzie  1 

A View  of  Biology  and  Entomology  from  Thunder  Bay 

Richard  Freitag 5 

Some  Observations:  Prognostications  on  Trends  in 
Entomological  Curricula  in  U.  S.  Colleges  and  Universities 

A.  R.  Gittins 11 

Discussion  — Morning  Session 

Leader  - D.  H.  Pengelly  18 

Taxonomic  Entomology;  Going,  Going,  - Where? 

W.  R.  M.  Mason 27 

Education  and  the  Professional  Entomologist 

N.  D.  Holmes  33 

Industry  Views  Entomology 

G.  S.  Cooper 39 

Discussion  — Afternoon  Session 

Leader  — D.  H.  Kavanaugh  42 

Summary 

Morning  Session 

Rapporteur  — D.  R.  Whitehead 50 

Afternoon  Session 

Rapporteur  — M.  A.  C.  Chance 52 

Resolutions 

B.  S.  Heming 54 

Entomology  and  Education  - Postscript  to  Symposium 

Brian  Hocking 56 


Participants  in  Entomology  Anniversary  Week 


58 


IV 


INTRODUCTION 

B.  S.  Heming 

Department  of  Entomology 
University  of  Alberta,  Edmonton,  Alberta 

You  might  wonder  why  we  are  having  a symposium  on  this  topic  to  celebrate  our  50 
years  of  operation  as  a Department  of  Entomology  at  the  University  of  Alberta.  As  most 
of  you  are  aware,  and  as  was  mentioned  several  times  during  the  last  three  days,  ento- 
mology, like  science  in  general,  has  entered  a difficult  period  in  its  history.  An  increas- 
ing number  of  graduates  are  unable  to  find  worthwhile  employment  in  a world  of  shrink- 
ing job  opportunities.  Those  who  should  be  the  most  employable  members  in  society 
are  declared  useless  because  of  inexperience  and  “over-qualification.”  At  the  same  time, 
the  public  has  lost  its  faith  in  the  ability  of  entomologists  and  other  scientists  to  solve 
the  problems  now  facing  mankind.  In  addition,  some  of  these  problems,  for  example 
pesticide  pollution,  are  said  to  have  resulted  from  the  activities  of  the  scientists  them- 
selves. As  a result  of  this  “crisis-in-confidence”  taxpayers  are  becoming  increasingly  reluc- 
tant to  have  portions  of  their  incomes  appropriated  for  the  support  of  seemingly  useless 
or  even  dangerous  segments  of  society.  This  feeling  has  been  communicated  to  the  grant- 
ing agencies  and  has  resulted  in  shrinking  financial  support  for  research  in  universities  and 
government  institutes  and  for  education  in  universities. 

Our  responsibility  in  departments  of  entomology  is  to  educate  entomologists.  Thus, 
we  thought  it  worthwhile,  on  our  50th  anniversary,  to  re-evaluate  our  role  in  light  of 
present  and  future  conditions.  We  felt  that  this  could  best  be  accomplished  by  contacting 
practising  entomologists  in  as  many  fields  as  was  practical.  Accordingly,  we  solicited  contri- 
butions from  six  former  students  of  this  department:  a high  school  biology  teacher,  a 
Canadian  biology  teacher  in  a primarily  undergraduate  university,  an  American  entomolo- 
gist in  a U.  S.  land  grant  institution,  a Canadian  government  research  taxonomist,  a Canadian 
government  researcher  in  applied  entomology,  and  the  technical  director  of  a large  indus- 
trial concern.  Each  speaker  was  asked  to  comment  on  the  following  questions:  1.  What 
is  the  proper  education  for  entomologists  of  all  kinds?  2.  What  can  entomology  contribute 
to  education  in  general?  and  3.  What  changes  should  be  made  in  the  curriculum  of  the 
Department  of  Entomology  at  the  University  of  Alberta?  Included  with  these  questions 
were  copies  of  past  curricula,  budgets,  etc.  for  this  department,  selected  at  1 0 year  intervals 
to  illustrate  our  development  through  the  years.  Thus,  all  participants,  regardless  of  when 
they  graduated,  had  a good  idea  of  what  the  department  had  been  doing  in  the  intervening 
years. 

You  will  notice  that  the  program  consists  of  two  parts;  the  speakers  in  the  morning 
session  are  educators  while  those  in  the  afternoon  are  “users”  of  our  graduates.  Each 
group  of  papers  will  be  followed  by  a discussion  period.  In  each  case  we  have  a discussion 
leader,  also  an  alumnus  or  a member  of  the  department,  who  has  received  notice  of  the 
contents  of  the  papers  in  advance.  He  will  open  the  discussion  after  which  we  will  ask 
for  comments  from  the  floor.  We  would  therefore  ask  you  to  hold  onto  your  questions 
until  the  open  discussion  period. 


1 


UNIVERSITY  EDUCATION  AND  HIGH  SCHOOL  BIOLOGY 

Don  N.  Jantzie 
County  Central  High  School 
Vulcan,  Alberta 

First  of  all  I would  like  to  congratulate  Dr.  Hocking  and  his  staff  on  the  unique  manner 
they  have  chosen  to  celebrate  50  years  of  distinguished  service  since  the  founding  of  the 
Department  of  Entomology  at  the  University  of  Alberta.  It  is  an  honor  and  privilege  for 
me  to  have  this  opportunity  to  participate  in  your  celebrations.  I would  like  to  compliment 
you  on  your  choice  of  topic  since  I believe  that  entomology  has  been  almost  totally  by- 
passed in  the  curriculum  of  the  high  schools. 

At  the  outset  I should  point  out  that  the  situation  may  appear  more  critical  to  an  ento- 
mology graduate  teaching  and  administrating  in  a school  system  than  it  would  to  persons 
having  other  specialized  training  in  the  biology  field.  As  a science  and  biology  teacher,  I 
often  found  myself  lamenting  the  fact  that,  because  of  the  structure  and  content  of  the 
courses/  I was  unable  to  utilize  my  special  knowledge  and  skills  to  the  benefit  of  the 
pupils  under  my  charge.  This  was  because  high  school  courses  were  not  structured  toward 
an  in-depth  study  of  the  insect  world  nor  was  there  any  concerted  effort  to  emphasize 
the  environmental  interrelations  of  insects  with  other  organisms.  The  students  of  our  school, 
residing  in  a rural  setting,  actively  involved  in  a battle  of  survival  against  infestations  of 
grasshoppers,  cutworms,  wireworms,  sawflies,  warble  flies,  lice,  mange  and  so  on,  were 
busy  studying  forms  of  life  that  many  of  them  had  never  seen  and  would  never  think 
of  again  in  their  lifetime.  The  biology  courses  were  excellent  according  to  sequential  study 
of  higher  life  forms  in  botany  and  zoology,  but  were  not  particularly  relevant  to  the  experi- 
ences of  the  students  themselves.  Then,  six  years  ago,  our  school  began  a program  that 
was  designed  to  fill  the  needs  of  a large  group  of  these  rural  students  who  planned  to 
remain  in  the  agriculture  field,  and  who  were  negatively  motivated  toward  many  of  the  stan- 
dard options  in  the  high  school  curriculum.  We  developed  an  agriculture  program  that 
encompassed  four  main  areas  of  study:  soils,  horticulture,  field  crops  and  animal  husbandry 
and  in  every  instance,  where  it  was  possible,  included  a comprehensive  study  of  insects.  Our 
experiment  met  with  such  outstanding  success  that  it  has  become  one  of  the  most  impor- 
tant options  offered  in  our  program,  so  much  so  that  my  instruction  time  is  now  totally 
dedicated  to  high  school  agriculture.  My  enthusiasm  has  been  heightened  because  of  the 
fulfillment  I have  received  as  an  entomologist.  It  has  also  made  me  realize  how  much 
more  effective  several  other  high  school  courses  might  be  if  they  too  incorporated  a study 
of  insects  into  their  curricula. 

Unfortunately,  most  high  school  students  have  had  little  exposure  to  the  world  of  insects. 
During  their  1 2 years  of  primary  and  secondary  education  they  have  looked  only  briefly  at 
the  social  habits  of  Hymenoptera,  particularly  at  those  of  bees  and  ants  and  principally  in 
comparison  to  human  social  habits;  they  have  had  a short  overview  of  metamorphosis  with 
particular  emphasis  on  Orthoptera  and  Lepidoptera;  and  they  have  briefly  considered  the 
disease-carrying  ability  of  Diptera  as  a health  hazard.  A brief  introduction  to  rudimentary 
classification  is  made  at  the  high  school  level  in  biology  which  only  affects  those  students 
entered  in  a matriculation  program  (approximately  60%  of  the  student  body).  The  overall 
effect  of  these  references  to  insects  is  minimal.  There  is  little  emphasis  on  creating  a desire 
to  pursue  this  interesting  study  farther,  and  what  we  have  may  even  be  detrimental  to  the 
development  of  future  entomological  interest.  The  average  secondary  school  student  in 
Alberta  would  be  unable  to  contrast  the  eating  habits  or  the  mouth  parts,  for  example,  of 


2 


Jantzie 


the  house  fly,  the  grasshopper,  the  leaf  hopper,  the  bee  and  the  cutworm  and  would 
probably  consider  all  these  insects  very  close  relatives.  These  are  the  same  students  who, 
in  their  social  studies  courses,  are  required  to  make  value  judgements  on  the  ecological  and 
environmental  effects  of  man’s  attempts  to  control  nature,  of  which  one  of  the  prominent 
hot  spots  lies  in  the  chemical  control  of  insects. 

Since  these  courses  do  not  create  an  interest  in  entomology,  how  can  a student  in  high 
school,  going  through  a normal  program,  ever  become  interested  enough  in  insects  to  study 
them  at  post-secondary  institutions?  Another  question  — How  did  each  of  us  become 
interested  in  entomology  to  the  extent  that  we  specialized  in  the  field?  How  much  oppor- 
tunity would  a student  in  the  faculty  of  education  have  of  developing  an  interest  in  entomo- 
logy, should  he  major  in  biology?  These  questions  are  fundamental  to  our  problem  at  the 
secondary  school  level. 

There  are  major  inadequacies  in  our  high  schools  with  respect  to  entomological  education. 
First  and  foremost,  biology  teachers,  for  the  most  part,  are  not  qualified  to  teach  entomo- 
logical concepts,  since  most  of  them  have  had  no  training  in  entomology  at  the  university 
level.  With  a minimum  of  background  in  such  a diverse  and  complicated  field,  it  is  no 
wonder  that  teachers  skirt  the  topic  should  it  arise  during  the  course  of  a lesson.  In  addition, 
it  would  be  extremely  unfair  to  the  student  should  he  be  subjected  to  classroom  instruction 
by  a person  who  has  no  background  experience  in  the  Field.  With  these  points  in  mind,  it 
is  not  surprising  that  our  school  curriculum  covers  only  elementary  concepts  in  the  study  of 
insects. 

Another  major  inadequacy  in  our  educational  system  today,  is  a curriculum  that  side- 
steps any  serious  exploration  of  entomology.  Keeping  in  mind  what  we  have  already  dis- 
cussed regarding  the  teacher,  this  situation  follows  naturally  from  the  first,  for  it  seems  only 
reasonable  that  it  would  be  difficult  to  utilize  a curriculum  weighted  heavily  toward  an  area 
that  could  not  be  adequately  presented.  Although,  at  the  present  time,  biology  is  still 
taught  in  the  standard  pattern,  a concerted  effort  to  improve  old  and  innovate  new  high 
school  and  the  junior  high  school  courses  has  been  undertaken.  With  greater  professional 
freedom,  the  teacher  now  has  the  opportunity  to  stress  parts  of  the  curriculum  that  he  feels 
have  more  meaning  and  more  relevance  to  the  experiences  of  his  class.  There  are  also 
provisions  for  teaching  options  such  as  oceanography,  entomology,  geology,  etc.  at  the 
junior  high  level  by  persons  with  special  interests  and  background  knowledge.  It  is  generally 
agreed  that  our  curriculum  committees  have  made  much  progress  in  developing  research- 
oriented  courses  that  shy  away  from  rote  memorization  (and  regurgitation)  of  facts,  a 
notable  achievement  in  itself.  Despite  these  improvements,  however,  the  basic  problem  of 
including  entomology  with  biology  courses  is  lacking. 

Curriculum  committees  have  an  additional  problem  when  recommending  text  and  refer- 
ence books  at  a reading  level  suitable  for  secondary  schools.  To  my  knowledge  there  are 
no  text  books  available  that  could  be  fully  utilized  in  the  school  system.  The  problem  lies  in 
two  areas.  First  of  all,  since  entomology  has  been  traditionally  a university  discipline  grow- 
ing out  of  the  zoology  pattern,  the  source  books  have  been  written  with  specific  divisions 
in  mind  such  as  in  taxonomy,  morphology  or  physiology.  The  second  area  of  difficulty  lies 
in  the  reading  level  at  which  these  texts  are  written.  In  both  areas,  the  students  would 
encounter  so  much  difficulty  in  utilization  and  comprehension,  that  the  books  would  be 
almost  useless.  Failing  to  find  suitable  text  books  in  print,  it  would  be  necessary  to  write 
our  own,  thus  creating  additional  economic  problems  (high  publication  costs).  School 
boards,  staggering  under  inflationary  increases  in  operating  costs,  have  tightened  the  educa- 
tional belt  to  such  an  extent  that  the  purchase  of  new  text  books,  other  than  those  in 
demand  by  large  numbers  of  jurisdictions,  is  prohibited.  In  addition,  writers  and  publi- 


High  School  Biology 


3 


shers  hesitate  to  develop  a treatise,  unless  it  has  been  specifically  requested  for  a large 
potential  market,  and  particularly  if  it  does  not  measure  up  to  the  professional  standards 
of  the  author.  I believe  the  problem  of  finding  or  producing  a suitable  text  book  to  be  a 
difficult  one  to  overcome. 

Curriculum  committees  have  the  responsibility  of  developing  courses  that  provide  a con- 
cept structure,  suitable  as  a foundation  for  the  students’  future  understanding.  In  the  exer- 
cise of  this  duty  they  must  take  into  consideration  economic  priorities  (very  important  at 
the  present  time);  educational  priorities,  influenced  a great  deal  by  post-secondary  institu- 
tions; and  personnel  priorities,  influenced  by  the  educational  backgrounds  of  the  teachers 
as  well  as  by  those  of  the  committee  members  themselves.  Looking  at  this  situation  we  can 
only  conclude  that,  at  the  present  time,  entomology  must  take  a secondary  position  in  our 
high  school  curriculum. 

Having  looked  at  two  major  inadequacies  in  the  fostering  of  entomological  interests 
in  the  secondary  schools,  let  us  now  take  a brief  look  at  post-secondary  institutions.  How 
many  students  in  the  faculties  of  education  or  science  obtain  their  major  in  entomology?  In 
the  past,  the  Faculty  of  Agriculture  supplied  virtually  the  entire  body  of  graduates  from 
the  Department  of  Entomology.  While  we  must  agree  that  the  study  of  insects  should  be  an 
integral  part  of  the  study  of  agriculture,  it  should  be  closely  linked  to  botany  and  zoology 
as  well.  Traditionally,  an  entomologist  is  an  Aggie  and,  if  we  want  to  enlarge  the  stereotype 
farther,  he  is  also  a somewhat  strange  person,  dressed  in  an  old  fashioned  wool  suit,  and 
holding  the  perennial  butterfly  net  over  his  shoulder.  To  put  it  bluntly,  entomologists  are 
considered  by  the  average  man  in  society  to  be  a bit  weird.  This  negative  attitude  harms 
our  image  and  prevents  expansion  of  the  science  into  the  associated  field  of  education.  I 
consider  myself  fortunate  to  have  graduated  in  agriculture  and  especially  in  entomology. 
My  training  in  the  biological  sciences  — bacteriology,  botany,  zoology,  genetics,  biochemis- 
try, plant  pathology  and  entomology  have  made  my  teaching  in  high  school  a thrilling 
experience.  It  is  unfortunate  that  more  teachers  could  not  have  had  a similar  background. 

What  conclusions  can  we  draw  from  the  obscure  role  of  entomology  in  the  secondary 
school  system?  I believe  that  entomologists  must  become  more  involved  with  education, 
especially  at  the  teaching  and  curriculum-planning  levels.  It  is  imperative  that  high  school 
students,  whether  university  bound  or  otherwise,  be  given  the  opportunity  to  explore  and 
understand  the  fascinating  world  of  insects  and  the  tremendous  impact  that  these  small 
organisms  have  on  other  living  things.  It  is  of  utmost  importance  that  they  learn  to  dis- 
tinguish one  family  from  another;  that  they  comprehend  the  total  meaning  of  biological 
and  chemical  control;  and  that  they  place  this  phase  of  entomology  into  its  proper  ecologi- 
cal perspective.  To  accomplish  this  our  schools  need  teachers  who  have  background  experi- 
ence in  entomology,  and  we  thus  need  university  courses  that  provide  access  to  this  informa- 
tion. At  the  university  level  we  need  greater  enrolments  and  greater  participation  from  facul- 
ty of  education  students  planning  to  work  in  the  biology  and  ecology  fields.  Students 
should  be  able  to  recognize  and  understand  the  interrelationships  that  exist  in  our  living 
world.  The  current  ecological  crisis  demands  that  we  have  knowledgeable  people  in  all 
walks  of  life.  We  need  to  press  for  change  in  the  basic  pattern  of  the  curriculum  at  both 
the  secondary  and  post-secondary  levels  of  education.  We  need  to  shorten  the  educational 
lag  that  is  an  integral  part  of  our  education  system.  We  have  to  provide  a foundation  of 
understanding  through  our  teachers  and  through  our  schools  that  produces  answers,  not 
doubts,  about  our  ability  to  solve  environmental  problems.  Progress  in  the  future  will 
depend  upon  how  we,  the  graduate  entomologists  of  the  past,  influence  the  education 
patterns  of  tomorrow. 


■ 


5 


A VIEW  OF  BIOLOGY  AND  ENTOMOLOGY  FROM  THUNDER  BAY 

Richard  Freitag 
Department  of  Biology 
Lakehead  University,  Thunder  Bay,  Ontario 

“ The  elder  statesmen  sit  on  the  mats, 

And  wrangle  through  half  the  day; 

A hundred  plans  they  have  drafted 
and  dropped, 

And  mine  was  the  only  way.  ” 

— from  the  Book  of  Songs, 

675  B.  C.  by  Confucius 

As  you  probably  know,  during  the  last  five  years  almost  every  university  sector  has  had 
its  share  of  crises.  Most  of  these  have  been  corrected  to  some  extent,  while  others  persist. 
Some  are  growing  worse.  Two  matters  of  concern  to  the  Ontario  university  community  are 
a marked  decline  in  student  enrolment,  and  a scarcity  of  jobs  for  students  following  gradua- 
tion. As  a means  of  offering  information  toward  reaching  the  objectives  of  this  session,  I 
shall  attempt  to  describe  the  two  conditions,  in  particular  as  they  affect  biology  and  ento- 
mology in  Ontario.  Then  I shall  offer  some  possible  remedial  concepts  for  discussion. 

It  is  well  known  now  that  most  universities  in  Ontario  experienced  a shortfall  in  pro- 
jected enrolments  in  September,  1971.  At  first  it  was  looked  upon  as  a brief  dip  in  a clinal 
increase  in  the  numbers  of  students  entering  universities.  The  usual  observations,  analyses 
and  rhetoric  followed  in  the  academic  and  mass  media.  Concurrently,  budgetary  adjust- 
ments were  made  and  by  December,  1971,  the  crisis  had  almost  been  overcome.  However, 
this  year  the  crisis  has  reappeared  with  even  more  vigour  (Table  1). 

According  to  a recent  report  of  the  Committee  on  Statistics  and  Enrolment  Projections, 
of  the  Ontario  Universities’  Council  on  Admissions  (February,  1972),  evidence  from  sub- 
missions of  individual  universities  indicates  that  applications  from  students  in  Grade  13 
are  down  by  5%  compared  to  those  at  the  same  time  a year  ago.  Also,  the  number  of  non- 
grade 13  applicants  into  first  year  is  less  than  half  of  what  it  was  a year  ago.  Thus,  the  total 
number  of  applications  to  Ontario  universities  is  down  by  7 to  8%  from  what  it  was  a year 
ago.  There  are  also  major  shifts  in  program  preferences  of  applicants  as  indicated  in  Table  2. 

Applicants  for  combined  Arts  and  Science  have  declined  by  16%,  for  Arts  by  14%  and 
for  Engineering  by  10%,  while  applicants  for  Science  have  increased  by  9%  and  for  other 
programs  by  4%.  Part  of  the  expected  9%  increase  in  the  sciences  is  due  to  slightly  in- 
creased enrolments  in  the  Life  Sciences  programs,  which  is  probably  because  of  ecologi- 
cal charisma.  Increased  enrolments  in  the  life  sciences  are  taking  place  in  other  Canadian 
universities  as  indicated  by  a survey  conducted  by  Dr.  Von  Borstel,  Chairman  of  the  Depart- 
ment of  Genetics  at  the  University  of  Alberta. 

The  Minister  of  Colleges  and  Universities  of  Ontario,  Mr.  George  Kerr,  recently  announced 
to  the  legislature  that  Ontario  universities  are  expecting  a 5%  drop  in  enrolment  next  fall, 
so  there  is  no  reason  to  believe  that  the  situation  will  improve  by  September.  The  decline 
in  enrolment  last  year  and  possibly  next  year  may  indicate  a trend.  If  so,  it  precedes  by 
approximately  20  years  a projected  decline  in  enrolment  in  universities  in  Ontario  provided 
by  the  Draft  Report  of  the  Commission  on  Post-secondary  Education  in  Ontario  (Queen’s 
Printer,  Toronto,  1971).  The  commission  notes  this  1972  discrepancy  but  does  not  elabo- 


6 


Freitag 


rate  on  its  occurrence. 

In  addition  to  the  problem  of  enrolment,  students  are  experiencing  increasing  difficulty 
in  finding  employment.  Some  statistics  from  student  placement  offices  and  unemployment 
centres  show  this  to  be  a general  trend.  Because  employment  for  biology  and  entomology 
students  is  in  part  what  concerns  us,  I have  selected  the  following  table  from  a report  of 
the  committee  investigating  employment  problems  of  entomologists  in  Ontario,  by  Drs. 
R.  L.  Edwards  and  P.  S.  Corbet  (1970),  as  a more  specific  example  (Table  3). 


Table  1*.  Total  applicants  to  full-time  programs  in  the  Ontario  Universities. 


UNIVERSITY 

1971 

1972 

DIFFERENCE 

TOTAL 

DIFFERENCE 

Brock 

2,512 

1,883 

- 629 

- 8,862 

Carleton 

5,962 

5,265 

- 697 

Guelph 

6,670 

6,475 

- 195 

Lakehead 

1,590 

1,684 

+ 94 

Laurentian 

2,190 

2,100 

90 

McMaster 

10,299 

10,758 

+ 459 

Ottawa 

4,941 

4,942 

+ 1 

Queen’s 

9,258 

8,653 

- 605 

Toronto 

18,034 

17,410 

- 624 

Trent 

3,525 

2,535 

- 990 

Waterloo 

14,517 

12,533 

- 1,984 

Western 

17,648 

15,600 

- 2,048 

Windsor 

4,796 

4,124 

- 672 

York 

11,164 

10,441 

- 723 

W.  L.  U. 

2,631 

2,472 

- 159 

Table  2*.  Applicants  to  first  year  full-time  programs  in  the  Ontario  Universities  on  or  about 
February  15,  1972. 

PROGRAM  1971  TOTAL  1972  TOTAL 


Arts  & Science 

13,970 

Arts 

28,756 

Science 

10,560 

Engineering 

5,412 

Other 

12,155 

11,688 

24,708 

11,507 

4,856 

12,652 


* Tables  1 and  2 adapted  from  Report  of  the  Committee  on  Statistics  and  Enrolment  Pro- 
jections of  the  Ontario  Universities  Council  on  Admissions.  Meeting  of  committee  held 
February  24,  1972. 


Biology  - Thunder  Bay 


7 


Table  3*.  Numbers  of  graduate  students  in  entomology  at  Ontario  Universities  in  1970-71, 
and  the  number  of  existing  positions  for  entomologists  in  Ontario  from  1969  to  1971. 

Number  of  graduate  students  in  entomology  in  Ontario  Universities: 

M.Sc.  33 

Ph.D.  21 

Total  54 

Number  of  existing  positions  for  entomologists  in  Ontario: 


1969 

1970 

1971 

Federal  Government  Service 

120 

115 

115 

Ontario  Government  Service 

8 

8 

4 

Industry 

2 

2 

2 

Ontario  Universities 

38 

40 

40 

Total 

168 

165 

161 

* Report  of  the  committee  investigating  employment  problems  of  entomologists  in  Ontario. 
Proc.  ent.  Soc.  Ont.  101:89-92,  1971. 


The  authors  conclude  that  “if  the  rate  of  turnover  is  3%  per  annum,  we  can  expect 
vacancies  for  entomologists  to  become  available  at  the  rate  of  approximately  five  per  year. 
However,  the  total  number  of  existing  positions  for  entomologists  is  expected  to  decline 
at  about  the  same  rate,  from  168  in  1969  to  a projected  161  in  1971.”  So  opportunities 
for  employment  in  universities,  government  and  industry  do  not  look  too  promising  for 
the  next  few  years.  The  situation  is  probably  not  any  better  for  graduate  students  in  other 
fields  of  biology.  The  drop  in  student  numbers  and  the  decline  in  employment  opportuni- 
ties are  probably  real  and  obvious  while  the  underlying  reasons  seem  more  elusive.  The 
classes  at  Lakehead  University  tend  to  be  small,  (no,  an  optimal  size),  and  teacher  and 
student  usually  communicate  at  the  individual  level.  According  to  some  of  my  students, 
many  high  school  graduates  no  longer  come  to  university  because  they  feel  taking  an 
undergraduate  degree  is  a wasted  effort  if  it  does  not  help  them  in  getting  a good  job. 
Others  tell  me  that  many  students  drop  out  of  university  for  the  same  reason.  One  of  our 
top  graduating  students  expressed  the  view  that  the  increasing  drop  out  rate  in  universi- 
ties is  mainly  due  to  uninteresting  courses  and  programs,  not  because  students  are  unable 
to  do  the  work.  He  suggested  that  the  three-lecture,  one-lab-per-week  course  becomes 
a ‘drag’  by  the  middle  of  the  second  year,  and  that  project  courses  dealing  with  up-to-date 
problems  should  replace  some  of  the  others.  He  was  very  enthusiastic  about  a full-year 
chemistry  course  in  which  the  entire  second  half  was  devoted  to  a laboratory  project  on 
the  DNA  molecule. 

Some  students  believe  that  job-oriented  community  colleges  have  attracted  high  school 
graduates  who  would  otherwise  have  gone  to  university.  In  order  to  understand  more 
quantitatively  the  priorities  of  gaining  qualification  for  work  as  opposed  to  an  education, 
in  the  minds  of  biology  students,  I conducted  a brief  survey  in  two  of  my  classes.  Of  course 
both  processes  are  hardly  separable  in  a biology  degree  program  but  I asked  them  to  give 


8 


Freitag 


priority  to  one  or  the  other.  Thirty-nine  students  stated  that  they  attended  university 
primarily  to  obtain  qualifications  for  employment  and  14  claimed  that  they  were  taking 
the  program  because  they  enjoyed  biology. 

The  relationship  between  job  opportunities  and  education  is  much  more  difficult  to 
define  than  causes  of  decline  in  student  enrolment.  This  is  clearly  expressed  in  the  Draft 
Report  (1971,  p.  28)  as  follows:  “An  important  aspect  of  post-secondary  education  is  its 
relationship  to  the  labour  market.  Paradoxically  enough  this  relationship  is  in  most  cases 
fuzzy.  ...  We  apparently  do  not  have  sufficient  data  even  to  describe  or  evaluate  such  a 
relationship.  Under  such  conditions  it  would  be  difficult  and  irresponsible  to  forecast  future 
linkages  of  education  and  manpower  needs.  It  is  often  said  that  the  majority  of  jobs  our 
current  generation  of  students  will  hold  in  their  lifetime  have  not  yet  been  invented.” 

Although  student  enrolments  are  increasing  in  biology  and  entomology  programs,  there 
is  no  evidence  that  job  opportunities  are  improving  for  biology  graduates.  Indeed,  increas- 
ing numbers  of  our  graduates  find  employment  in  fields  not  related  to  biology.  I was 
surprised  to  find  two  of  our  biology  graduates  working  as  permanent  employees  in  an 
Ontario  liquor  store.  Hardly  a place  for  biologists,  but  they  seemed  to  enjoy  their  work. 

What  then  can  we  do  to  improve  the  situation  for  our  students  while  remaining  commit- 
ted to  our  subjects  and  maintaining  the  good  programs  we  now  have?  I suggest  that  we 
attempt  to  increase  the  development  of  biology  and  entomology  in  a lateral  dimension. 
Surely  in  our  changing  society,  new  and  different  occupations  are  developing  in  which  an 
education  in  biology  or  entomology  is  essential,  but  not  to  the  extent  provided  by  the 
programs  we  now  offer.  The  following  recommendations  are  a list  of  personal  views  which 
have  developed  from  my  experiences  at  Lakehead  University.  These  recommendations  are 
not  intended  to  replace  or  change  any  traditional  activities  or  programs.  They  are  pro- 
posed rather  as  possible  supplementary  avenues  of  development.  I shall  give  the  recom- 
mendations and  a brief  rationale  for  each. 

Recommendations 

1 . That  university  biology  and  entomology  departments  conduct  annual  surveys  with  a 
view  to  determining  the  present  scope  of  entomology  and  biology. 

A few  weeks  ago  Dr.  S.  Madras,  Director  of  the  Liberal  Science  program  at  York 
University,  visited  Lakehead  University  to  discuss  Liberal  Science  with  us.  Dr.  Madras 
served  on  several  education  advisory  committees  in  Quebec  of  which  part  of  their  terms 
of  reference  was  to  determine  the  role  of  educational  institutions  for  the  government.  He 
remarked  that  the  conceptual  profile  of  a university  changes  when  the  university  is  observed 
from  varying  distances  and  positions.  His  experience  was  that  the  interactions  of  univer- 
sities and  society  became  clear  when  he  communicated  with  people  who  were  considerably 
removed  from  the  university  milieu.  Now  this  kind  of  experience  is  not  new  to  field  biolo- 
gists who  in  fact  use  the  process  as  a means  of  studying  a single  species  in  its  natural 
environment,  i.e.  autecology.  In  my  opinion,  a carefully  designed  and  executed  autecology 
of  biology  and  entomology  would  provide  a greater  understanding  of  our  relationships 
to  a changing  society.  It  would  also  reveal  possible  interactions  of  our  disciplines  with 
academic  and  other  sectors  of  society  which  could  benefit  biology,  entomology  and  our 
students.  Perhaps  the  survey  could  be  conducted  on  a larger  scale,  that  is  by  all  life  sciences 
departments  in  a university  or  even  on  a provincial  or  national  scale.  The  survey  could  be 
conducted  at  two  levels.  First,  an  assessment  could  be  made  on  the  relationships  of  the 
classical  programs,  courses  and  course  content  to  the  professional  occupations  of  biolo- 
gists and  entomologists.  Second,  and  probably  more  important  at  this  time,  information 
could  be  obtained  on  occupations  which  require  or  should  require  graduates  with  some 


Biology  — Thunder  Bay 


9 


knowledge  of  biology  and  entomology.  We  could  also  determine  adaptations  necessary  to 
ensure  that  our  subjects  successfully  evolve.  Among  the  many  probable  advantages  deriving 
fj*om  such  a study  would  be  ideas  for  establishing  new  course  programs.  Information 
would  accrue  that  would  be  valuable  in  counselling  first  year  students  in  determining  a 
course  program  for  a specific  goal.  The  data  obtained  would  be  of  benefit  to  all  biologists 
across  the  country. 

2.  That  major  (biology,  entomology)  - minor  (business  administration,  economics,  law, 
political  science,  chemistry,  physics,  etc.)  programs  of  study  be  developed  for  uncom- 
mitted students. 

By  uncommitted  students  I mean  those  who  are  taking  a degree  in  biology  or  entomology 
but  who  also  have  special  interests  in  other  fields  of  study.  There  can  be  a definite  advantage 
to  having  a strong  training  in  two  subjects  with  respect  to  getting  a job.  One  of  our  students 
has  a diploma  in  forestry  and  a B.Sc.  degree  in  biology.  He  was  hired  to  teach  vocational 
subjects  and  academic  courses  in  a vocational  high  school  in  Swan  Lake,  Manitoba.  He  was 
told  that  he  would  not  have  been  offered  the  job  if  he  had  had  only  a degree  in  biology  or 
in  forestry.  Another  one  of  our  biology  education  graduates  was  just  hired  to  teach  science 
in  a high  school  in  North  Bay,  Ontario.  He  completed  a chemistry  program  in  a commu- 
nity college  before  coming  to  Lakehead  University.  The  principal  of  the  high  school  in- 
formed him  that  he  was  hired  primarily  because  he  had  a good  background  in  both  biology 
and  chemistry.  These  are  two  examples  of  a possible  growing  demand  for  university  gradu- 
ates with  such  qualifications.  Major-minor  programs  would  clearly  serve  this  need  not 
only  in  education  but  in  other  occupations  as  well. 

3.  That  honours  students’  research  topics  be  multidisciplinary.  A greater  effort  should  be 
made  to  hire  undergraduates  for  research  being  conducted  by  biology  or  entomology  facul- 
ty, especially  for  multidisciplinary  research. 

Multidisciplinary  research  provides  a good  groundwork  for  more  specialized  research.  I 
feel  that  the  third  undergraduate  year  or  the  honours  year  is  a good  level  for  such  research. 
The  student,  while  completing  a program  composed  of  subjects  often  apparently  related, 
can  obtain  an  appreciation  of  how  they  actually  do  relate.  In  addition,  undergraduate 
students  would  greatly  benefit  from  supervised  research,  particularly  if  they  plan  to  con- 
tinue graduate  studies  in  a specific  biology.  Also,  if  a student  plans  to  leave  university  this 
experience  provides  potential  employers  with  a more  complete  picture  of  the  student’s 
abilities. 

4.  That  a major  (biology,  entomology)  — minor  (business  administration,  economics,  law, 
chemistry,  physics,  etc.)  M.Sc.,  Ph.D.,  program  be  developed. 

This  kind  of  graduate  project  would  require  increased  cooperation  among  university 
departments  which  is  one  of  the  main  reasons  for  the  recommendation.  In  my  view  the 
greatest  failing  in  our  society  is  the  colossal  human  sound  barrier  among  its  components. 
This  problem  is  well  illustrated  by  the  present  environmental  issue  on  which  our  political 
leaders  and  biologists  seem  to  maintain  an  impasse.  What  better  way  to  bridge  this  gap  than 
with  a graduate  ‘biobarrister’  or  ‘entopolitician.’ 


SOME  OBSERVATIONS:  PROGNOSTICATIONS  ON  TRENDS  IN 
ENTOMOLOGICAL  CURRICULA  IN  U.  S.  COLLEGES  AND  UNIVERSITIES 


A.  R.  Gittins 

Department  of  Entomology 
University  of  Idaho,  Moscow,  Idaho 

Before  proceeding  into  the  topic  for  discussion,  I would  like  to  extend  my  sincere  appre- 
ciation to  the  committee  for  the  opportunity  of  appearing  on  this  symposium.  This  allowed 
me  to  make  a pilgrimage  back  to  the  old  Alma  Mater  after  many  years  of  absence.  While 
change  is  the  nature  of  all  living  things,  and  that  includes  man’s  institutions,  and  the 
University  of  Alberta  is  certainly  far  from  an  exception,  I have  pleasant  twinges  of  nostalgia 
when  picking  out  some  old  landmarks  and  reminiscing  about  my  experiences  here. 

In  my  years  subsequent  to  the  baccalaureate  at  Alberta,  I’ve  retained  a very  fond  memory 
of  Prof.  Strickland  and  have  never  waivered  in  the  opinion  that  he  was  a great  teacher,  a fine 
entomologist  and  one  of  the  most  sincere,  dedicated  and  human  individuals  I have  met. 
Many  times  I’ve  had  cause  to  reflect  on  how  fortunate  I was  in  being  exposed  to  his  parti- 
cular style  of  teaching. 

And  of  course  we  are  gathered  here  today,  in  part  at  least,  to  pay  homage  to  Dr. 
Strickland,  for  his  establishment  of  this  department  in  1922  has  provided  reason  for  our 
celebration  these  50  years  later.  But  if  we  are  to  honor  Dr.  Strickland  for  “creating”  and 
nurturing  the  Department  of  Entomology  through  the  earlier  years,  it  would  indeed  be 
ungracious  of  me  not  to  honor  equally  Brian  Hocking,  under  whose  vision  and  capable 
direction  this  department  has  developed  into  the  significant  and  prestigious  position  it  now 
holds  among  departments  of  entomology  in  institutions  of  higher  learning  throughout  Cana- 
da and  the  United  States.  Indeed,  as  an  alumus  I extend  my  gratitude  to  you,  Dr.  Hocking, 
and  to  the  other  members  of  the  faculty  of  the  Department  of  Entomology  for  your  out- 
standing and  successful  efforts  in  building  this  proud  and  productive  entomological  unit. 

Moving  on  to  the  subject  matter  of  the  symposium,  and  referring  to  Dr.  Heming’s  outline, 
I view  his  charge  to  the  symposial  participants  as  a three-part  directive.  First  he  asks  that  we 
consider  the  subject  of  a “proper  education  for  entomologists  of  all  kinds.”  My,  what  far 
reaching  import  lies  in  that  word  “proper”!  Secondly,  we’re  asked  to  direct  some  thought  as 
to  what  entomology  can  contribute  to  education  in  general.  And  finally,  if  appropriate,  to 
provide  some  feedback  on  entomological  activities  at  the  U.  of  A.  over  the  preceding  50 
years  and  perhaps  direct  some  constructive  comments  as  to  the  future.  This  is  a substantial 
responsibility. 

In  public  speaking  one  is  cautioned  never  to  provide  excuses  for  the  comments  or  direc- 
tions he  takes  in  formally  presenting  his  views  on  a given  subject.  However,  today  I choose 
to  avoid  such  caution  and  beg  your  indulgence  regarding  what  might  be  claimed  to  be  a 
degree  of  “tunnel  vision,”  insofar  as  my  direct  association  with  entomological  education 
over  the  past  20  years  has  been  confined  to  institutions  in  the  United  States  and  even  more 
specifically  to  land  grant  institutions.  As  most  of  you  know,  the  land  grant  institution  in  the 
United  States  was  created  primarily  for  the  purpose  of  meeting  obligations  related  to  the 
practical  needs  of  the  agricultural  and  mechanical  arts.  And  while  these  institutions  have 
embraced  the  concept  of  a total  university,  practical  aspects  of  our  field  nevertheless 
commonly  dominate  in  our  activities. 

I believe  that  before  we  can  truly  attend  to  the  matter  of  education  in  entomology,  we 
have  to  outline  some  evaluations  regarding  the  contemporary  position  of  the  scientific 
community  — at  least  as  it  appears  in  the  public  mind.  Liberally  paraphrasing  some  leading 


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educators  in  science,  immediately  and  regrettably  we  note  that,  at  a time  in  our  history 
when  we  most  need  the  talents  of  specialized  intellectuals,  they  are  becoming  a drug  on  the 
market.  Primarily,  of  course,  I am  referring  to  Ph.D.’s  who,  throughout  the  1950’s  and  60’s, 
were  in  such  short  supply  and  long  demand  that  they  could  almost  name  their  own  salaries 
as  they  emerged  from  graduate  school.  During  that  decade,  incidentally,  Ph.D.  production 
rates  soared  from  9,400  in  the  U.  S.  in  1959  to  more  than  26,000  in  1969.  And  entomology 
generally  kept  pace. 

But  now,  suddenly,  the  stampede  is  over.  Education  — by  far  the  largest  consumer  of 
learned  specialists  — has  moved  to  a period  of  contracting  activity.  Government  and  indus- 
try, caught  in  a recession,  have  curtailed  those  activities  that  used  the  greatest  shares  of  such 
talent.  Such  fields  of  particular  interest  to  us  as  environmental  quality  control  and  modem 
pest  management  are  just  two  areas  which  would  seem  to  cry  for  the  intellect  and  training 
of  Ph.D.’s  who  are  now  being  turned  aside  as  surplus. 

It  is  proposed  today  that  we  must  retreat  and  guide  this  stream  of  extraordinary  man- 
power into  more  productive  channels  and  call  for  coordinated  placement  efforts  by  govern- 
ment and  private  industry  on  the  one  hand,  and  for  greater  discipline  and  imagination  by 
universities  on  the  other.  “Should  universities  judiciously  curtail  their  production  in  certain 
areas  and  steer  into  new  avenues  of  greater  need,  government  and  industry  could  come 
together  to  insure  better  opportunities  for  the  numerical  surplus  of  Ph.D.’s.” 

This,  however,  presents  us  with  our  first  of  a series  of  dilemmas.  On  the  one  hand,  ento- 
mological educators  are  almost  totally  committed  time-wise  to  redirection  into  the  future, 
replete  with  all  the  time-consuming  activities  of  re-organization,  re-evaluation,  re-education 
and  development  of  an  enthusiasm  for  broader  interdisciplinary  involvements.  Little  time  is 
left,  therefore,  for  political  involvement  which  appears  to  be  becoming  increasingly  neces- 
sary to  assure  adequate  consideration  by  the  body  politic  and  the  body  public  of  entomolo- 
gy and  attendance  to  entomological  needs  within  the  aegis  of  “Environmental  Issues.”  While 
the  issue  of  insecticides  has  gained  substantial  public  exposure,  the  vast  research  efforts 
needed  to  develop  alternate  means  of  keeping  pestiferous  species  at  non-economic  levels 
have  been  dramatically  over-simplified,  undersold,  or  even  ignored.  And  in  this  regard  we, 
the  entomologists,  may  be  substantially  to  blame.  Now  is  an  ideal  time  for  us  to  emphasize 
paramount  need  for  research  on  long  term  insect  control  and  to  see  that  public  interest  is 
translated  into  fiscal  support. 

In  effect  then,  entomologists  in  the  field  of  education  must  not  only  be  active  in  modern- 
izing and  redirecting  the  total  curriculum,  but  must  also  involve  themselves  in  providing  the 
public  and  government  with  a clear  awareness  of  the  overall  needs  for  entomological  exper- 
tise. We  must  be  pragmatic  enough  to  learn  that  it’s  not  sufficient  to  train  and  educate 
people  in  what  we  deem  a viable  entomological  curriculum  today.  If  we  are  remiss  in 
educating  the  public,  government  and  industry  to  a need  for  our  product,  then  we  are 
failing  our  students,  ourselves,  our  profession  and  society  at  large. 

In  backing  our  attitude  towards  political  activism  we,  in  entomology,  must  assume  some 
responsibility  in  educating  the  whole  man.  Granted,  we  cannot  be  expected  to  teach  all  the 
courses  that  contribute  to  his  wholeness,  but  “the  implications  of  our  field  for  the  well 
being  of  mankind  are  too  important  to  be  placed  in  the  hands  of  leaders  who  lack  the 
capacity  for  wide  judgement.”  “The  role  of  the  biologist  has  assumed  new  importance  as  he 
has  become  able  to  manipulate  growth  processes  and  control  living  organisms  in  his  environ- 
ment. But  to  provide  this  knowledge  for  biological  manipulation  without  developing  an 
accompanying  awareness  in  the  area  of  human  affairs  and  moral  responsibility  will  render 
our  products  ill-prepared  for  the  new  role  that  will  surely  be  thrust  upon  them.”  These  new 
professionals  must  be  provided  with  a breadth  of  exposure  and  understanding,  for,  if  our 


U.  S.  Colleges  and  Universities 


13 


premise  be  correct  relative  to  the  importance  of  our  field,  they  will  constitute  our  primary 
agents  in  the  re-education  of  the  “establishment”  to  the  vital  role  of  entomology.  As  our 
banquet  speaker,  Mr.  Coffey,  stated  the  other  evening  . . . : “We  are  no  longer  society’s 
decision  makers”  — accordingly  then,  we  must  maximize  communication  between  our- 
selves and  those  who  make  the  final  decisions  so  that  their  judgements  are  based  upon  at 
least  adequate  information. 

In  reflection,  it’s  ironic  that  just  as  scientists  are  on  the  crest  of  a wave  of  achievement, 
we  must  plead  increasingly  strongly  for  financial  support.  Just  a short  time  ago,  national 
prestige  and  well-being  seemed  to  be  clearly  dependent  upon  the  ever  increasing  growth  of 
science  and  technology.  Today,  in  spite  of  the  tremendous  impact  scientific  and  technologi- 
cal advances  have  made  on  our  lives,  the  layman  generally  doubts  that  scientists  are  doing 
what  they  should  for  mankind.  And  a lack  of  concert  among  ourselves  adds  to  this  confu- 
sion. The  average  man  on  the  street  sees  a dialectic  ...  on  the  one  hand  he  is  deeply  im- 
pressed by  the  ability  of  scientists  to  “create,”  but  on  the  other  hand  he  is  critical  that 
scientists  have  allowed  technological  advances  to  proceed  in  a way  that  creates  profound 
environmental  and  societal  problems.  Public  disappointments  have  led  to  cutbacks  in  finan- 
cial support  for  scientific  research  in  many  areas.  And  so  it  is  up  to  scientists  themselves  to 
bridge  some  so-called  “credibility  gaps”  and  to  convince  the  public  and  government  that 
science  is  indeed  very  relevant. 

Moving  on  now  to  some  specifics  relating  to  entomological  education  . . . first,  I believe 
it  self-evident  that  no  single  entomology  department  can  be  all  things  to  all  people.  No 
department  can  provide  all  the  courses  and  training  experiences  we  deem  necessary  in  pre- 
paring students  for  all  kinds  of  entomological  activities.  Obviously,  there  are  too  many 
limits  to  our  resources,  both  financial  and  expertise-wise.  But  to  go  one  step  further,  while 
we  often  point  to  a need  for  a certain  “critical  mass”  in  terms  of  faculty  numbers  for  co- 
ordinated departmental  operation,  a department  geared  in  an  attempt  to  do  all  things  might 
in  fact  exceed  a “maximum  desirable  mass.”  One  might  paraphrase  such  a situation  as  “the 
law  of  increasing  negative  returns.”  Another  trap  we  sometimes  fall  into  without  realization 
is  the  unconscious  acceptance  of  the  concept  that  there  is  indeed  a perfect  entomological 
curriculum.  Perhaps  this  subtly  emerges  as  a result  of  conditioning  where  we  think  in  terms 
of  the  individual  when,  in  reality,  we  must  think  of  a collective  group  of  individuals  — 
students  of  various  interests,  ambitions,  abilities,  for  whom  we  must  provide  mosaics  of 
educational  patterns.  If  we  fix  our  focus  firmly  in  terms  of  the  collective  rather  than  the 
individual  then  we  are  more  inclined  to  present  flexible  programs  within  which  we  design  or 
at  least  accommodate  for  a number  of  directions  of  emphasis. 

However,  when  one  discusses  the  total  curriculum  offered  by  a department,  of  necessity 
it  must  be  recognized  that  we  are  talking  about  two  tracks:  that  of  the  undergraduate  and 
that  of  the  graduate.  And  within  each  are  a number  of  sub-tracks.  While  I feel  that  a viable 
entomology  department  does  need  both  of  these  programs  (because  of  the  cross-fertiliza- 
tion which  is  evident  for  both  undergraduate  students  and  graduate  students),  it  is  nigh  im- 
possible to  embrace  both  programs  within  a single,  common  philosophy.  Therefore,  I believe 
there  is  need  to  consider  undergraduate  and  graduate  programs  independently. 

Let  us  discuss  for  the  moment  the  undergraduate  program.  The  premise  I offer  is  a 
simple  one  and  paraphrases  that  advanced  earlier  by  Dr.  Edward  H.  Smith.  Entomology  is  a 
branch  of  biology  and  training  at  the  undergraduate  level  in  entomology  fares  best  when 
closely  allied  with  biological  sciences.  Specialization  is  left  to  the  graduate  level.  Entomo- 
logy departments,  by  their  very  existence  with  esprit  de  corps  and  offered  capacity  for 
growth,  tend  to  proliferate  and  fragment  a biological  curriculum.  It’s  perhaps  inevitable  that 
such  units  become  the  victims  of  scientific  isolation,  replete  with  all  the  dangers  of  reduced 


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capacity  for  interdisciplinary  effort.  Therefore,  I agree  with  Smith  when  he  states  that  there 
is  little  justification  for  an  elaborate  undergraduate  major  in  entomology  for  those  students 
planning  graduate  training.  There  are  some  exceptions  to  this,  to  which  I would  like  to  turn 
in  a moment,  but  they  still  relate  primarily  to  training  for  a terminal  degree  at  the  bachelor 
level.  In  order  for  the  student  to  acquire  a firm  foundation  for  an  advanced  degree  program, 
many  hours  must  be  devoted  to  generalized  courses  in  the  biological,  physical  and  social 
sciences.  Consequently,  little  time  is  left  for  specialization  in  entomology.  Courses  in  mathe- 
matics, chemistry,  physics,  zoology  and  botany  are  prerequisites  taking  precedence  over 
a field  of  specialization.  Specialization  then  would  logically  occur  at  the  graduate  level. 
Admittedly,  we  should  retain  a minimal  core  of  courses  at  the  undergraduate  level.  These 
may  be  modified  in  part,  both  to  serve  a general  service  function  and  to  accommodate  the 
student  who  enters  college  with  a career  in  entomology  already  firmly  fixed  in  mind. 
Alternately,  the  university  curricular  regulations  must  be  flexible  enough  to  allow  the  com- 
mitted student  access  to  upper  divisional  and  so-called  graduate  courses. 

The  point  I make  should  not  be  construed  as  a proposal  to  eliminate  entomology  as  an 
undergraduate  offering.  It  is  more  a plea  to  reduce  and  ideally  eliminate  specialization  or 
course  proliferation  at  the  undergraduate  level. 

However,  today  we  must  realistically  face  what  appears  to  be  the  rapidly-emerging  field 
of  pest  management.  Inherent  in  the  implementation  of  many  pest  management  programs  is 
the  implication  that  there  will  be  considerable  manpower  needs  at  probably  the  bachelor’s 
level  of  education  to  conduct  much  of  the  observation  and  scouting  required  for  constant 
surveillance  of  various  units  within  a given  agro-ecosystem.  Admittedly,  this  is  a very  prag- 
matic approach,  but  it  appears  to  me  that  we  will  be  remiss  in  our  responsibilities  to  society 
and  agriculture  if  we  do  not  design  undergraduate  curricula  to  best  prepare  individuals  for 
this  particular  vocation.  I envision  such  a curriculum  with  the  first  two  years  devoted  to 
general  courses  in  the  physical  and  biological  sciences  (much  as  we  have  now)  but  with  the 
last  two  years  specializing  in  applied  ecology  with  the  added  ingredients  of  economic  ento- 
mology, weeds,  plant  diseases,  climatology  and  the  like.  In  addition,  a most  desirable  adden- 
dum would  be  some  kind  of  practicum  course  (admittedly  a harsh  term  to  use  in  the  ivory 
halls  of  learning).  And  should  we  not  be  giving  greater  emphasis  to  in-service  training  pro- 
grams with  government  and  industry?  From  a mercenary  point  of  view  alone,  development 
of  new  in-service  training  programs  at  times  constitutes  a “job-creating”  maneuver. 

Obviously,  we  are  again  faced  with  a difficult  decision  in  determining  what  blend  of 
applied  training  should  be  built  into  a framework  of  academic  study  dealing  with  biological 
principles.  Considering  the  transient  nature  of  technical  knowledge  as  opposed  to  funda- 
mental knowledge,  I hope  that  even  here  we  would  tend  to  stress  the  latter.  Regardless  of 
which  is  emphasized,  we  must  firmly  imbue  every  student  with  the  realization  that  continu- 
ous self-renewal  is  essential  for  professional  survival. 

Incidentally,  that  constitutes  another  basic  consideration  we  sometimes  allow  to  slip 
away  from  us  when  focussing  our  attention  on  what  might  constitute  a complete  and  com- 
prehensive curriculum.  We  must  clearly  remind  ourselves  and  instill  in  the  minds  of  our 
students  that  academia  can  only  hope  to  provide  an  embryogenesis  to  the  profession. 
Growth,  development  and  even  the  metamorphosis  of  an  individual  in  his  profession  must 
be  carefully  and  continuously  nurtured  throughout  his  career. 

Finally,  let  me  phrase  a few  succinct  and  pertinent  questions  being  asked  today  on 
undergraduate  education  in  general.  Are  we  giving  sufficient  attention  to  the  content  and 
approach  being  offered  in  high  school  biology  courses  today  and  accordingly  upgrading  our 
own  lower  division  college  courses  in  light  of  increasing  sophistication  of  the  high  school 
programs?  Are  we  evaluating  and  considering  the  need  for  students,  at  the  upper  levels  par- 


U.  S.  Colleges  and  Universities 


15 


ticularly,  to  exercise  some  voice  in  determining  their  own  curricula  and  setting  their  own 
goals  within  specific  courses?  Regardless  of  the  eventual  outcome  of  discussions  along  these 
lines,  exchanges-of-position-views  by  itself  often  promotes  mutual  enthusiasm  and  has  an 
inspirational  effect. 

Turning  now  to  the  matter  of  graduate  education,  we  seem  faced  with  a kind  of  collage 
incorporating  both  clear  and  obscure  elements.  Obviously,  a graduate  curriculum  must  be 
built  largely  upon  the  areas  of  expertise  of  the  faculty.  This  immediately  places  a substantial 
burden  on  an  existing  faculty  and  their  departmental  administrator  to  plan  carefully  for 
additional  and  replacement  faculty  positions  as  well  as  curricular  programs.  This  is  asking 
both  faculty  and  administration  to  exercise  a considerable  degree  of  clairvoyance  in  deter- 
mining the  nature  of  the  various  specializations  within  the  profession  as  they  relate  to  the 
future. 

There  are,  however,  a number  of  remedial  steps  which  I feel  should  be  taken  today  in 
graduate  educational  programs  regardless  of  the  direction  or  redirection  of  the  curriculum. 
The  first  of  these  concerns  a critical  need  for  the  exposure  of  students  to  the  continuously 
newer  technologies  as  they  become  available.  Even  the  traditional  systematist  must  be  com- 
petent in  the  employment  of  modern  physiological  tools;  the  anatomist  with  the  scanning 
electron  microscope;  the  ecologist  with  infrared  photometry;  and  all  of  us  with  the  language 
and  use  of  the  computer.  These  are  but  a few  examples,  but  ones  which  all  too  frequently 
are  relatively  ignored  or  given  only  vague  consideration  by  teachers  through  inattention, 
preoccupation,  or  sometimes  even  downright  laziness.  It  is  perhaps  fitting  at  this  point, 
since  I have  not  made  reference  elsewhere  to  the  variety  of  graduate  degrees  offered  by 
institutions,  to  pose  a few  additional,  pertinent  questions.  Most  of  the  questions  outlined 
are  under  serious  consideration  at  a number  of  U.  S.  institutions. 

1.  Does  the  research-oriented  Master’s  degree  program  have  a viable  contemporary  pur- 
pose? Or  does  it  simply  serve  as  a second  prize  and  a more  palatable  means  of  “weeding 
out”  Ph.D.  candidates?  Some  educators  refer  to  this  degree  as  a “dinosaur.” 

2.  Should  we  be  instituting  special  graduate  degree  programs  which  differ  in  scope  and 
purpose  from  the  traditional  M.Sc.  and  Ph.D.  programs?  What  about  a non-thesis  M.Sc.  for 
those  going  into  extension,  industrial  sales,  etc.?  What  about  a special  doctorate  (i.e.  Doctor 
of  Arts)  wherein  we  de-emphasize  the  research  aspects  and  emphasize  preparation  for  teach- 
ing or  other  largely  non-research  careers? 

3.  Have  our  language  requirements  become  passe?  Do  these  so-called  “translation  compe- 
tency tests”  represent  simply  another  archaic  “pseudo-intellectual  hoop”  through  which 
students  must  leap  to  obtain  their  parchment?  We  well  know  that  at  many  institutions  the 
language  exam  evaluates  vocabulary  proficiency  and  preparation  for  the  examination  does 
not  often  develop  any  true  proficiency  in  the  language,  including  a knowledge  or  “feeling” 
for  the  culture  or  ways  of  thinking  of  the  people  for  whom  that  language  constitutes 
their  native  tongue.  It  is  being  suggested  that  either  we  require  intensive  study  in  a foreign 
language  or  allow  substitution  of  other  proficiencies.  Computer  courses  have  been  recom- 
mended strongly  as  one  alternative. 

But  back  to  the  central  theme!  While,  of  necessity,  we  must  immerse  a student  in  the 
details  of  a particular  specialization,  we  must  be  careful  not  to  do  him  the  disservice  of  over- 
channelling  him  whereby  he  has  neither  time  nor  opportunity  to  explore  other  fields  which 
philosophically  or  practically  might  relate  directly  to  his  own  area  of  study.  Let  us  use  care 
in  not  over-emphasizing  how  entomology  relates  to  other  fields,  but  rather  to  commence 
emphasis  centering  on  how  such  areas  as  biochemistry,  cellular  biology,  development  biolo- 
gy, evolutionary  biology,  etc.  relate  to  our  own  field.  We  must  develop  our  curriculum  with 
a focus  on  process  rather  than  organism,  even  though  we  realize  this  does  impose  consider- 


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able  stress  on  traditional  departmental  structure.  Fortunately,  many  of  our  research  pro- 
grams today  (at  least  those  receiving  substantial  portion  of  total  funding)  are  both  mission- 
oriented  and  tuned  to  interdisciplinary  team  approaches.  Consequently  and  traditionally,  as 
goes  research,  so  often  goes  the  graduate  academic  program,  unless  we’ve  withdrawn  so  far 
as  to  consider  the  Ph.D.  degree  to  be  largely  a matter  of  formal  course  hurdles,  language 
requirements  and  preliminary  examinations.  And,  I’m  afraid  there  has  been  a tendency  to 
move  in  this  direction  at  the  expense  of  maximizing  opportunity  for  both  directed  and  inde- 
pendent research  effort. 

I realize  I’m  speaking  philosophically  and  in  generalities  but  today  can  one  be  expected  to 
do  much  more  outside  of  one’s  institution,  what  with  each  institution  exhibiting  unique- 
ness relative  to  philosophy,  faculty  interests  and  abilities,  fiscal  and  physical  resources,  etc. 

In  spite  of  all  the  “soul  searching”  we  are  engaged  in  today,  I believe  we’re  very  fortunate 
to  be  in  the  field  of  entomology,  a discipline  which  provides  for  considerable  latitude  and 
opportunity  for  delving  into  numerous  phases  such  as  physiology,  morphology,  behavior 
and  the  like.  I’m  frequently  impressed  with  entomologists  in  general  because  many  have 
interest  in  numerous  subdisciplines.  I honestly  feel  that  the  heterosis  one  finds  in  our  field 
is  what  makes  it  “tick.”  I’m  always  concerned  that  the  alternative  to  general  interest  and 
broad  involvement  is  narrow  specialization  which  frequently  breeds  professional  extinction. 

And  so,  again,  we  are  faced  at  the  graduate  level  with  the  need  for  what  might  first 
appear  to  be  two  antagonistic  responsibilities  — training  the  individual  in  an  area  of  special- 
ization, but  also  providing  him  with  a continual  expansion  of  his  total  education.  To  accom- 
plish the  latter  does  not  necessitate  saddling  the  student  with  a vast  array  of  courses  at  the 
graduate  level,  but  only  requires  that  there  be  a significant  cross-fertilization  between  all 
members  of  the  department  be  they  staff  or  student  and  with  additional  input  from  with- 
out. What  we  are  endeavoring  to  do  this  week  is  a prime  example  of  a very  viable  way  of 
approaching  this  objective.  Symposia,  seminars,  discussion  groups,  coffee  sessions,  reports 
on  meetings,  presentations  of  research  results  and  the  like,  are  means  by  which  this  can  be 
accomplished.  We  should  also  look  into  the  matter  of  involving  those  from  without  the  insti- 
tution in  special  lectures,  programs,  etc.  This  has  apparently  been  successfully  used  in  Law 
and  Medicine.  So  I would  charge  all  university  entomologists  to  greater  involvement  in  these 
kinds  of  activities  even  at  the  expense  of  a few  “precious”  hours  lost  from  their  own  re- 
search laboratory.  To  ask  a student  to  “dig  it  out  for  yourself’  is  commonly  a good  educa- 
tional procedure,  particularly  if  it  generates  some  frustration  followed  by  satisfaction  in 
having  learned  by  self-instruction  (and  how  better  can  one  really  learn?).  From  such  activity 
comes  increased  motivation.  But  to  fail  to  provide  the  individual  with  subsequent  opportu- 
nity to  expound  and  bounce  his  philosophies  and  ideas  off  his  teachers,  constitutes  a 
breakdown  in  the  educational  process. 

Turning  now  to  the  matter  of  how  entomology  can  serve  education,  I’m  perhaps  being 
somewhat  naive  when  I state  that  the  picture  seems  more  clear-cut.  The  insect  is  an  almost 
ideal  critter  for  illustrating  a vast  array  of  biological  principles  being  introduced  at  all 
grade  levels  within  our  educational  system.  Personally,  I’ve  had  considerable  success  in 
teaching  a rather  practical  course  along  these  lines  to  erstwhile  grade  school  and  high  school 
biology  teachers.  Many  insect  species  provide  most  convenient  laboratory  animals,  and 
field  projects  involving  these  same  insects  can  be  easily  carried  out  even  under  minimal 
supervision. 

The  field  of  entomology  provides  both  the  information  and  the  biological  tools  for 
studies  in  many  related  disciplines,  be  they  biochemistry,  genetics,  gerentology,  or  cancer 
research.  May  I take  the  liberty  of  suggesting  perhaps  that  some  of  the  other  participants 
on  this  symposium  can  provide  a far  more  comprehensive  outline  of  this  area  of  our  dis- 


U.  S.  Colleges  and  Universities 


17 


cussion. 

Finally,  what  can  I offer  the  Department  of  Entomology  here  in  both  reflection  and 
prediction?  Very  little  I’m  afraid,  particularly  in  the  area  of  the  future  over  and  above  what 
I’ve  already  covered.  As  to  the  past  how  can  I,  one  of  your  alums,  be  so  modest,  indeed 
be  so  callous  as  to  level  any  large  degree  of  criticism  for  past  things  done  or  undone.  In 
truth  I have  only  two  points  of  reference  — the  first  being  my  experiences  here  some 
20  years  ago,  which  I felt  were  well  tuned  to  my  needs,  being  on  the  one  hand  designed 
to  provide  me  with  sufficient  breadth  as  to  be  aware  of,  if  indeed  not  capable  of,  becom- 
ing educated.  And  on  the  other  hand  providing  me  with  a fundamental  array  of  courses 
in  botany,  zoology,  basic  and  applied  entomology,  chemistry  and  the  like  so  that  I would 
experience  at  least  some  degree  of  confidence  in  my  ability  to  handle  a position  in  entomo- 
logy or  to  be  prepared  (and  it  so  happened  I was)  for  continuing  my  education  at  the 
graduate  level. 

My  second  point  of  reference  is  not  a firm  one  but  revolves  around  my  occasional  con- 
tact with  currently-enrolled  or  recently  graduated  students  of  this  department.  I have  been 
impressed  with  both  their  competency  and  with  their  enthusiasm  and  I could  say  no  more 
that  would  constitute  a greater  endorsement. 


18 


DISCUSSION  - MORNING  SESSION 

Leader  — D.  H.  Pengelly 
Department  of  Environmental  Biology 
University  of  Guelph,  Guelph,  Ontario 

Upon  learning  that  I had  been  designated  a discussion  leader,  I tried  to  find  some  kind  of 
pattern  within  previous  symposia  experienced  that  would  shed  some  light  on  my  respon- 
sibilities in  this  one.  There  isn’t  one.  Nevertheless,  I do  thank  the  committee  for  the  oppor- 
tunity of  being  part  of  this  program  held  to  pay  respect  to  Dr.  Strickland  and  to  the 
department  that  he  started. 

The  speakers  have  presented  a wide  variety  of  ideas  on  Entomology  and  Education. 
Mr.  Jantzie  has  placed  a challenge  squarely  in  front  of  us  as  entomologists.  We  must  be 
represented  on  the  committees  that  formulate  curricula  for  senior  public  and  high  schools 
and  convince  those  responsible  that  the  insect  is  far  too  important  to  be  ignored  at  any 
level  in  an  educational  program.  Our  representatives  will  have  the  assurance  that  most, 
if  not  all,  of  the  entomologists  in  this  country  are  solidly  behind  them.  Some  competent 
soul  will  have  to  write  a book  that  uses  insects  to  illustrate  basic  biological  principles.  We 
should  be  encouraged  on  hearing  that  Dr.  Gittins  has  such  a book  in  preparation  and  we 
are  looking  forward  to  seeing  it. 

It  is  encouraging  too,  to  see  that  Mr.  Jantzie  and  his  staff  have  stepped  over  the  time- 
honoured  traces  by  introducing  detailed  studies  of  insects  and  equating  these  with  local  and 
world  problems  in  food  production,  health,  and  other  aspects  of  an  environmental  nature. 
If  the  rest  of  us  sit  back  and  do  nothing,  there  will  be  little  or  no  progress  and  there  will 
be  a relapse  back  to  the  traditional. 

The  problems  of  using  insects  have  been  covered.  In  many  ways  this  is  a difficult  group 
but  the  challenge  of  the  difficulties  associated  with  these  aliens  of  Planet  Earth  should 
attract  those  with  the  keenest  minds.  Making  students  aware  of  insects  as  early  in  their  life 
as  possible  will  do  much  toward  getting  entomology  into  its  proper  place  in  schools,  colleges, 
universities,  and  back  to  the  layman.  It  must  not  be  left  in  the  hands  of  a chosen  few.  It  is  a 
changing  world  and  we  must  do  our  part  in  bringing  about  change  if  it  will  be  to  the  advan- 
tage of  the  fair-haired  youth  who  someday  must  pass  this  way. 

Our  second  speaker  exposed  a different  set  of  problems  associated  with  declining  student 
numbers.  Our  concern  here  should  perhaps  be  directed  toward  the  administrators  who  in 
their  “crystal  balling”  have  assumed  that  there  are  populations  having  a predictable,  conti- 
nual, and  prolonged  growth  increase.  Under  the  present  system  of  sponsorship  that  demands 
speculation,  admonishes  the  seers,  and  punishes  the  academic  community  by  limiting  funds 
that  they  were  forced  to  anticipate,  I can  see  little  else  other  than  problems.  Dr.  Freitag 
said  that  the  present  problems  in  under-enrolment  were  overcome  by  “budgetary  adjust- 
ments.” Perhaps  he  would  enlarge  on  this  and  tell  us  what  adjustments  were  made.  I suspect 
that  the  academic  who  survives  on  a diet  of  Full-time  Teaching  Equivalents  will  be  the  first 
to  go  hungry. 

The  decline  may  well  continue  and  in  all  likelihood  be  attributable  to  the  lack  of  employ- 
ment opportunities  for  graduates.  It  does  seem  a bit  much  to  expect  that  there  will  be 
jobs  for  everyone  who  graduates  from  the  vast  array  of  specialties  available.  Here  we  see 
the  conflicting  points-of-view  as  to  the  function  or  role  of  a university.  To  some,  one 
goes  to  university  for  the  pure  joy  of  learning.  To  most,  it  is  because  specialized  training 
will  provide  greater  returns  in  a chosen  field  of  endeavour.  The  latter  is  the  basic  philosophy 
of  parents  and  children  as  to  why  an  education  is  necessary.  The  former  reason  is  a dodging 


Discussion 


19 


of  responsibilities  when  it  is  offered  as  an  explanation  to  job-seeking,  fourth-year  students. 
If  this  is  the  basic  function  of  a university  then  it  must  be  made  clear  as  early  as  grade  one 
that  education  and  employment  are  in  no  way  related.  The  initiative  for  going  to  university, 
whether  we  wish  it  to  be  or  not,  is  associated  with  the  monetary  system  of  a materialistic 
society.  If  community  colleges  with  job-oriented  programs  are  attracting  or  just  starting 
to  attract  students,  the  university  must  do  some  soul  searching.  Student-supporting  agencies 
will  soon  decide  where  to  invest  their  monies. 

Dr.  Freitag  has  made  a number  of  recommendations.  Within  the  next  15  years  the  majo- 
rity of  the  entomologists  presently  employed  in  Canada  will  have  retired.  What  are  we 
doing  to  ensure  that  there  will  be  Canadian  students  in  the  “mill”?  It  is  going  to  take 
9 or  10  years  to  produce  the  kind  of  product  that  is  being  sought  occasionally  today. 
An  annual  survey  does  not  seem  to  be  a very  useful  undertaking. 

Our  speaker  has  illustrated  a rather  strange  phenomenon.  He  refers  to  entomology  and 
biology.  Why  do  entomologists  accept  so  passively,  their  categorization  by  “zoologists” 
as  being  neither  biologists  nor  zoologists?  Correcting  this  would  serve  a very  useful  purpose. 
For  what  it  is  worth  from  one  who  has  been  “there  and  back,”  so  to  speak,  I am  convinced 
that  for  entomology  to  maintain  and  increase  its  position  in  the  academic  community, 
which  it  surely  must  do,  the  name  “Entomology”  must  be  retained.  To  be  swallowed  by 
the  “Zoological  Octopus”  that  does  not  accept  entomologists  as  being  either  biologists 
or  zoologists,  is  sheer  folly.  How  many  universities  offering  degrees  in  biology  or  zoology 
have  any  extensive  courses  in  entomology  as  requirements? 

To  establish  programs  embracing  biology  and  law,  or  economics,  chemistry,  physics, 
etc.,  is  an  admirable  idea  but  somewhat  ethereal.  Entomology  is  ignored  by  such  closely 
associated  programs  as  animal  science,  crop  science,  soil  science,  and  landscape  architecture, 
to  mention  a few. 

I envy  those  whose  vantage  point  is  within  a smaller  university.  Research  projects  in 
the  final  year  are  indeed  desirable.  In  crowded  institutions,  especially  in  the  three-term 
system,  this  is  all  but  impossible.  As  far  as  having  M.Sc.  and  Ph.D.  programs  involving 
the  arts,  physical  sciences,  and  biology  I am  again  pessimistic.  Students  in  these  areas  in 
universities  have  the  choice  right  now  to  elect  courses  in  biology  but  they  choose  to  do 
otherwise. 

Dr.  Gittins  has  provided  much  food  for  thought.  One  thought  was  that  there  be  a chan- 
nelling of  Ph.D.’s  into  more  productive  or  needed  areas.  Why  has  this  obvious  fact  been 
ignored  by  both  government  and  university?  Why  have  they  been  immune  to  planned 
programming,  especially  in  this  age  of  computers?  Education  for  education’s  sake  may 
have  a place  but  this  should  not  have  priority  over  education  for  the  sake  of  service  to  man- 
kind and  to  this  fragile  planet  as  a whole.  During  the  war  years  men  and  women  needed 
for  particular  jobs,  even  the  “tail-end-Charlies,”  were  obtained;  not  by  the  free  choice  of 
individuals  but  by  directed  training  programs. 

The  need  to  educate  the  government  and  the  public  has  been  stressed.  The  education  of 
the  student  was  referred  to  by  Dr.  Gittins  as  being  associated  with  modernizing  and  redirect- 
ing the  total  curriculum.  I have  often  wondered  what  it  would  be  like  if  we  reverted  to  the 
courses  required  and  recommended  by  Dr.  Strickland  25  years  ago.  Frankly,  I think  the 
end  product  would  fare  very  well  indeed.  Often  we  are  preoccupied  with  changing  and 
revising  (mostly  downward)  because  of  the  system  of  government  support  and  the  need  for 
self-preservation  in  an  F.T.E.-  (Full-time  Teaching  Equivalents)  dominated  system. 

The  need  to  provide  an  awareness  in  the  areas  of  human  affairs  and  moral  responsibility 
is  a pressing  one.  I do  not  agree  that  this  should  be  left  to  the  university.  If  so,  there  will 
be  no  visible  improvement.  It  must  occur  long  before  this,  preferably  in  the  home. 


20 


Discussion 


Perhaps  biologists  should  shoulder  some  of  the  responsibilities  for  the  adverse  effects 
of  technological  progress  but  there  are  others  who  need  a jolting  with  the  proverbial  “frozen 
boot.”  For  the  lawyers  who  make  laws,  chemists  and  engineers  who  design  processes,  equip- 
ment and  buildings,  architects  who  plan  streets,  gardens,  parks  and  even  universities,  the 
establishment  of  at  least  one  ecological  ethic  would  be  a godsend. 

As  to  leaving  specialization  to  the  graduate  school,  I am  in  complete  disagreement.  Had 
this  been  the  policy  in  the  past,  most  of  us  would  not  be  here  today.  My  interest  in  entomo- 
logy was  aroused  and  maintained  by  Dr.  Strickland.  Had  I not  been  able  to  specialize  in 
this  area,  undoubtedly  I would  have  ended  up  elsewhere.  Learning  to  be  an  entomologist 
in  the  graduate  school  is  next  to  impossible.  By  then,  it  is  much  too  late.  Students  with 
the  so-called  broad  and  varied  background  are  seldom  a match  for  those  who  have  been 
reading,  observing  and  experimenting  with  insects,  or  any  other  animals,  since  childhood. 
A student  who  is  neither  fish  nor  fowl  and  tries  to  become  one  or  the  other  in  graduate 
school  finds  himself  a student  of  the  lecture  room  and  the  text  book  and  not  of  the  library 
and  laboratory. 

Dr.  Gittins  has  cautioned  us  about  relating  entomology  to  other  fields  such  as  biochemis- 
try, cellular,  developmental  and  evolutionary  biology.  To  me  these  are  as  much  a part  of 
entomology  as  they  are  of  any  other  branch  of  zoology.  There  has  been,  and  still  is,  a 
tendency  to  call  all  entomologists  “specialists”  and  to  accept  without  question  the  idea 
that  only  zoologists  have  the  so-called  broad  background.  I am  of  the  opinion  that  if  a 
student  entered  university  and  studied  insects  only,  as  representatives  of  the  animal  king- 
dom, and  studied  these  with  respect  to  morphology,  anatomy,  histology,  embryology, 
genetics,  physiology,  behaviour,  taxonomy,  zoogeography  and  evolution,  he  would  be  at 
least  as  good  a biologist  as  those  who  now  graduate  in  Honours  Programs  in  zoology 
without  so  much  as  looking  at  a single  member  of  the  Class  Insecta. 

Looking  back  over  the  last  20-odd  years  I do  not  remember  there  being  an  abundance  of 
jobs.  There  wasn’t  when  I graduated.  I think  that  if  we  assess  the  past  we  will  see  that  our 
training  stood  us  in  good  stead  by  providing  us  not  only  with  a solid  base  in  entomology 
but  also  with  an  attitude  that  we  could  tackle  almost  anything.  If  we  can  come  close  to 
instilling  the  enthusiasm  and  confidence  in  our  students  that  Dr.  Strickland  did  in  us,  we 
will  have  made  a significant  contribution.  We  must  convince  the  public  and  government  that 
entomology  is  academically  respectable  and  in  some  way  establish  the  study  of  insects  in 
public  and  high  schools.  We  must  provide  the  tools  and  competent,  qualified  teachers.  The 
rest  should  follow. 

Ladies  and  Gentlemen,  your  comments  and  questions,  please. 

Gittins  - Moscow  — When  I spoke  of  leaving  specialization  to  the  graduate  level,  I did  not 
mean  to  infer  that  we  are  leaving  out  specialization  in  entomology  per  se.  I am  in  no  way 
convinced  that  entomology  isn’t  a vital  part  of  an  undergraduate  curriculum,  but  speciali- 
zation in  general  entomology  rather  than  in  systematics,  morphology  or  something  like  that. 
One  point  Dave  made  is  very  important  and  hits  us  right  now.  We  must  resist  this  full-time 
equivalent  business  with  all  the  vehemence  that  we  can  muster.  I presented  the  budget  for 
our  department  to  our  central  administration  a week  or  so  ago  and  suggested  that  we 
cease  worrying  about  student  credit  hours  generated,  at  least  within  shop.  I can  think  of 
no  more  detrimental  activity  for  the  university.  What  would  result  of  course,  if  this  con- 
cept were  carried  to  extremes,  is  a single  lecture  course  with  400  students.  Such  a course 
would  generate  credit  hours  but  at  the  expense  of  innovation,  of  special  programs,  of 
seminars,  of  these  kinds  of  things  that  don’t  generate  that  kind  of  hour.  This  is  one  thing 
we  do  have  to  fight  and  fight  hard. 

Freitag  - Thunder  Bay  — There  is  just  one  point  about  budgetary  adjustments  at  Lakehead 


Discussion 


21 


University  that  I’d  like  to  mention.  I believe  there  were  about  30  people  on  staff  last  year 
whose  term  appointments  will  not  be  renewed  this  summer  because  of  the  drop  in  student 
enrolment  in  1971.  There  were  a number  of  staff  released  in  the  summer  as  well.  I have 
two  questions  from  Dr.  Hocking  regarding  my  talk.  First,  “How  were  1972  enrolment 
predictions  arrived  at?”  These  were  based  on  grade  13  and  non-grade  13  applications 
to  first  year  in  all  Ontario  universities.  They,  in  turn,  passed  this  information  on  to  the 
Ontario  Universities  Council  of  Admissions. 

The  other  question  was,  “What  is  wrong  with  a liquor  store  as  an  environment  for  a 
biologist?”  Answer  — They  were  holding  jobs  for  which  entomologists  were  better  qualified. 
Downer  - Waterloo  — Several  years  ago  the  Entomological  Society  of  Canada  recognized 
many  of  the  problems  which  we  have  been  discussing  this  morning  and,  to  find  means  of 
alleviating  some  of  them,  established  a Committee  on  Student  Encouragement.  I thought 
it  might  be  appropriate  to  mention  some  of  the  activities  which  are  presently  being  pursued 
by  this  committee.  Like  Dr.  Gittins,  this  committee  has  recognized  the  need  for  suitable 
resource  material  for  teachers.  We  felt  that  teachers  must  be  convinced  that  working  with 
insects  is  quite  feasible  in  the  classroom.  To  this  end  the  committee  is  preparing  an  experi- 
mental manual  containing  information  on  methods  of  rearing  and  maintaining  insect  cultures 
in  the  classroom  and  some  simple  experiments  that  can  be  used  by  teachers  and  students. 
A note  to  this  effect  was  placed  in  the  bulletin  of  the  Society  a few  months  ago  in  which  we 
invited  interested  people  to  submit  experiments  which  could  be  included  in  this  manual. 
We  are  still  looking  for  contributions  and  they  will  be  gratefully  received.  We  are  also 
preparing  a list  of  audio-visual  aids,  films  and  other  material  which  might  be  of  use  in  the 
classroom.  If  anyone  knows  of  any  films  which  might  be  suitable  for  this  purpose,  we  would 
be  very  pleased  to  hear  comments  from  them.  Thirdly,  we  have  established  liaison  with 
several  provincial  science  teacher’s  associations  in  Ontario  with  the  hope  that,  by  providing 
articles  of  interest  to  these  teachers,  it  will  be  possible  to  instill  greater  interest  in  things 
entomological  among  them.  The  final  approach  has  been  to  attempt  infiltration  of  the 
various  provincial  committees  which  formulate  school  curricula.  We  have  met  with  mixed 
success  in  this  regard.  I think  Doug  Craig  established  some  contact  with  the  Alberta  authori- 
ties and  we  have  been  successful  in  Ontario  in  getting  membership  on  one  of  these  commit- 
tees. In  Ontario  we  have  been  helped  by  the  fact  that  provincial  Bill  145  has  limited  to  a 
considerable  extent  the  use  of  non-human  vertebrates  in  the  classroom.  Teachers  are  there- 
fore looking  desperately  for  alternative  animals  to  use  in  traditional  experiments. 

Pepper  — Bozeman  — The  problems  arising  out  of  our  current  methods  of  insect  control 
should  have  been  foreseen  by  many  of  us  at  least  15  years  ago.  The  indiscriminate  use 
of  insecticides  and  the  resulting  environmental  problems  is  only  one  of  these.  Our  biggest 
problem  has  been  our  failure  to  recognize  that  we  are  dealing  with  evolutionary  units, 
that  we  are  artificially  selecting  for  undesirable  characteristics  in  pest  populations.  Do  we 
teach  this  in  our  courses?  Hopefully  we  do  now. 

I used  to  teach  a very  specialized  course  in  insect  physiology.  One  of  my  students  asked, 
“What  relevance  has  this  course  to  the  problems  I will  be  confronted  with  on  graduation?” 
At  that  time  my  answer  was,  “None,  but  I will  give  you  a grade  in  it  anyway.”  Entomologi- 
cal education  today  seems  to  be  comprised  of  a bunch  of  fragments:  histology,  morphology, 
taxonomy,  toxicology,  etc.  Students  graduate  with  these  fragments.  Their  education  started 
no  place,  led  no  place,  and  ended  no  place.  Yet  we  send  these  same  students  out  to  be  con- 
fronted on  the  job  with  completely  new  situations.  What  are  they  going  to  do?  What  back- 
ground do  they  have  to  do  it?  Who  is  going  to  give  them  this  background?  Entomology, 
as  we  all  know,  started  out  as  a hobby  with  scattered  practitioners  all  over  the  world. 
They  collected,  studied  life  histories,  and  in  some  instances  made  great  contributions.  Even- 


22 


Discussion 


tually  there  came  a time  when  these  people  were  replaced  by  those  whose  approach  was 
more  scientific.  We  started  using  tools,  every  tool  we  could  lay  our  hands  on.  All  you  had  to 
have  was  something  to  measure,  and  by  God,  you  measured  it.  You  measured  it  so  that  you 
could  write  a paper  on  it  and  if  you  wrote  enough  papers  you  got  an  increase  in  salary. 
This  was,  and  unfortunately  still  is,  true.  Eventually,  we  became  a group  of  gadgeteers. 
This  sort  of  thing  continued.  Then,  along  came  mathematics.  To  use  mathematics  in  ento- 
mology, we  must  quantify.  We  now  have  at  our  disposal  computers  that  can  do  anything  — 
can  even  put  a man  on  the  moon.  But,  can  you  quantify  a qualitative  property  of  an 
entity?  Can  you  put  a qualitative  characteristic  into  a computer  and  have  it  analyzed?  That 
well  known  phrase,  “garbage  in,  garbage  out”  answers  that  question. 

All  entomology  students  must  take  math,  physics,  chemistry,  and  so  on  during  their 
undergraduate  years.  But  what  kind  of  math?  They  get  the  math  being  taught  to  those 
who  want  to  go  into  industry.  Statistics?  Statistics  is  said  to  be  based  on  logic.  The  best 
definition  I’ve  heard  for  logic  is  “an  organized  procedure  for  going  wrong  with  competence 
and  certainty.”  Chemistry?  Since  I received  my  undergraduate  degree  in  chemistry,  I feel 
that  I can  criticize  chemical  education  legitimately.  The  chemistry  taught  in  university  is 
industrial  chemistry.  The  biochemistry  taught  in  universities  is  the  application  of  organic 
chemistry  to  some  industrial  product.  Physics?  We  learn  about  solids.  Living  systems  are 
liquid  ones.  Biologists  of  all  kinds  should  demand  from  these  departments  that  they  teach  a 
type  of  math,  chemistry  and  physics  suited  to  biological  thinking. 

Once  I visited  each  of  my  staff  and  asked,  “How  do  you  use  your  knowledge  of  math?” 
“Well,  I don’t  use  it  myself  but  everyone  else  seems  to.”  I eventually  discovered  that  no 
one  in  my  department  used  math.  I found  the  same  to  be  true  of  chemistry  and  physics. 
Then  why  in  the  world  do  we  require  that  our  students  take  these  subjects?  If  a student 
asks  this  question  we  usually  answer,  “It’s  in  the  calendar  as  a requirement.”  What  I’ve  been 
trying  to  say  is  that  the  whole  educational  bag  must  be  re-evaluated  as  a whole,  not  frag- 
ment by  fragment. 

Jantzie  — Vulcan  — I hope  I didn’t  leave  you  thinking  that  the  current  high  school  biology 
courses  here  in  Alberta  are  poor  ones.  I agree  with  Dr.  Gittins  that  when  students  go  to 
university,  their  high  school  biology  is  usually  adequate  for  their  first  year.  In  fact,  first 
year  biology  in  Alberta  is  almost  a complete  repeat  of  high  school  biology.  I have  some- 
times found  the  high  school  course,  if  anything,  to  be  overly  sophisticated  for  the  majority 
of  students.  They  seemed  to  enjoy  it  but  I sometimes  felt  that  they  went  into  some  things 
a little  too  deeply  for  that  age  level.  I think  they  would  have  appreciated  a look  at  some  of 
the  more  simple  things  in  life.  We  could  have  spent  more  time  on  insects,  on  plant  patho- 
logy, etc.  and  less  on  DNA.  You  can  talk  about  fungus  all  you  want,  but  unless  you 
then  go  and  say,  “Okay,  let’s  look  at  some  of  the  plant  diseases  of  cereal  crops,”  these 
students  aren’t  going  to  relate  it  to  anything  they  already  know. 

As  I think  back  to  the  broom  closets  that  comprised  the  entomology  department  of  the 
40’s,  I see  now  that  entomology  occupied  a lesser  position  in  the  eyes  of  the  administration 
than  did  the  departments  of  zoology,  botany,  etc.  I hope  that  things  are  different  now,  but 
I’m  not  so  sure  that  they  are.  This  may  be  because  administrators  have  not  appreciated  the 
ubiquitous  role  that  insects  have  in  the  living  world. 

Mr.  McGregor  has  asked  about  the  agriculture  program  that  we  have  in  Alberta  high 
schools.  There  is  no  specific  program  in  agriculture.  If  a person  wants  to  teach  it,  he  must 
get  special  permission  from  the  Department  of  Education.  He  must  have  the  qualifications 
to  teach  it  and  he  must  make  up  his  own  syllabus.  This  is  nice  because  he  can  structure  it  to 
his  own  particular  specifications.  That  is  why  I am  so  happy  with  our  program. 

Following  graduation  from  the  University  of  Alberta,  I worked  in  the  seismograph  field 


Discussion 


23 


for  several  years  where  I didn’t  use  much  of  my  background  in  entomology.  But  I had  a 
wonderful  time  until  I took  up  farming  at  Vulcan.  I farmed  there  for  almost  10  years 
until  I decided  to  try  to  do  something  for  our  school  system  which  had,  at  that  time,  been 
plagued  by  a rapid  turnover  in  teachers.  That’s  how  I got  involved  in  education,  and  I 
must  say  I have  enjoyed  every  second  of  it  since. 

Hocking  — Edmonton  - I have  one  or  two  comments  that  I would  like  to  add  on  this  ques- 
tion of  predictions.  I was  glad  to  learn  from  Rick  Freitag  that  predictions  in  Ontario  are 
made  from  high  school  enrolments.  I have  been  doing  this,  on  behalf  of  biology  and  science 
at  this  university,  for  a number  of  years  and  the  predictions  I come  up  with  usually  seem 
to  be  closer  to  what  actually  transpires  than  the  predictions  of  our  office  of  institution  • 
research  and  planning.  Much  of  the  devastating  effect  of  declining  enrolments  is  not  so 
much  the  fact  that  they  are  declining  but  that  they  are  declining  in  the  face  of  growing 
predictions.  To  illustrate  this  point  and  possibly  add  a note  of  optimism,  I have  three 
slides  here  illustrating  percentages  of  enrolments  in  grades  11,  12  and  first  year  university, 
taking  the  three  sciences:  chemistry,  physics  and  biology  since  1963-64.  Grade  11:  Green 
is  biology,  red  is  chemistry,  blue  is  physics,  white  is  science.  You  will  note  that  until  1968 
biology  was  not  a science.  Grade  12:  My  only  comment  here  is  that  the  dotted  green  line 
is  the  old  biology  32  going  down  as  the  new  biology  30  comes  up  — to  a higher  position 
in  relation  to  the  other  sciences  than  32.  The  next  one  is  first  year  university  which  I 
think  is  perhaps  the  most  encouraging  of  the  three. 

I would  like  to  answer  two  rhetorical  questions  which  the  speakers  asked.  Mr.  Jantzie 
asked  how  many  students  in  education,  agriculture  and  science  get  their  major  in  entomo- 
logy. I haven’t  precise  figures  but  I would  guess  that  over  the  last  10  years,  there  were  very 
few  altogether.  Science  would  have  the  largest  number,  agriculture  next,  and  education 
very  few.  Dr.  Gittins  asked  about  the  role  of  the  Master’s  degree.  I would  like  to  give  you  my 
interpretation  of  the  role  of  the  Master’s  degree.  I consider  it  a stepping  stone  between  a 
doubtfully  adequate  undergraduate  program  and  a Ph.D.,  or  if  you  like,  a trial  run  for  a 
Ph.D.  — specifically  for  students  with  a dubious  background  in  language,  writing,  research 
and  research  procedures. 

Gurba  - Edmonton  — One  of  the  problems  facing  entomology  seems  to  be  the  same  as  that 
confronting  other  disciplines  in  university.  Too  many  of  the  teaching  programs  there  are 
still  research-oriented.  This  was  alright  in  the  days  when  graduates  were  filling  positions  in 
government  research  and  university  teaching,  but  these  positions  have  now  been  filled  and 
there  is  now  a surplus  of  students  with  this  type  of  background  being  graduated.  Why 
haven’t  these  curricula  been  changed?  There  are  interesting  opportunities  in  fields  other 
than  research  and  teaching.  There  are  openings  in  extension,  in  agriculture,  in  forestry,  in 
industry  and  in  parks  and  recreation.  Many  of  these  jobs  involve  helping  other  people  and 
are  therefore  very  satisfying. 

Dr.  Gittins  mentioned  the  need  for  work  and  training  in  pest  management.  This  is  some- 
thing we,  in  the  Department  of  Agriculture,  have  been  discussing  for  a long  time.  What  is 
our  entomology  department  doing  about  training  people  in  this  area?  Those  of  us  who 
came  through  the  University  of  Alberta  in  the  late  40’s  and  in  the  early  50’s  got  this  sort 
of  thing  from  Dr.  Strickland  in  his  courses  in  general  and  economic  entomology.  This 
sort  of  approach  is  not  available  today  and  I am  wondering  why. 

Pengelly  — The  answer  could  lie  in  consideration  of  administration.  My  point  is  this:  if  they 
turned  out  one  apiculturist  a year  and  he  found  a job  open  somewhere  in  Canada,  they 
would  have  served  Canada  well.  Unfortunately,  this  does  not  suit  present-day  systems. 
You  cannot  institute  a new  course  without  assuring  administration  that  you  will  have  rela- 
tively large  enrolments  in  it.  Three  students  might  want  to  take  a course  in  pest  manage- 


24 


Discussion 


ment.  Well,  you’ll  have  to  be  sure  of  more  students  than  that  or  you  can’t  offer  it.  It 
brings  us  back  to  our  old  problem  of  full-time  teaching  equivalents. 

Hocking  — I entirely  agree  with  what  David  has  just  said.  I would  like  to  indicate  what  we 
are  doing  about  this  question  here.  There  is  one  way  of  getting  a course  accepted  and 
allowing  it  to  persist  in  spite  of  apparently  inadequate  enrolments.  We  have  a project  course 
in  applied  entomology.  The  only  way  we  have  been  able  to  keep  this  course  in  existence 
is  by  having  it  a catch-all.  Anybody  wanting  pest  management  can  enrol  in  this  course 
and  get  it.  Unfortunately,  the  university  is  now  looking  very  hard  at  any  course  with  an 
enrolment  of  10  or  less.  They  are  trying  to  weed  these  courses  out  to  cut  costs.  If  we 
don’t  have  a demand,  there  is  nothing  we  can  do  about  it. 

Kavanaugh  - Edmonton  — My  question  concerns  the  possibility  of  channelling  students 
into  specific  areas  of  interest.  I would  like  to  ask  those  here  with  considerable  experience 
in  teaching  at  what  point  in  the  education  of  the  individual  we  can  begin  this  channelling. 
More  importantly,  how  late  in  the  process  can  we  accomplish  this  sort  of  thing?  I ask  this 
question  as  a student  realizing  that  motivation  is  of  prime  importance  and  I wonder  where 
in  my  own  education  I would  have  been  susceptible  to  rechannelling.  I might  add  that  I 
am  interested  in  taxonomy  which  should  put  things  in  perspective  for  you.  Certainly,  it  had 
to  be  pretty  early  in  my  case.  I would  say,  in  very  early  grade  school.  In  my  experience  how- 
ever, students  in  entomology  are  generally  pretty  turned-on  people.  I think  in  biology  that 
to  rechannel  these  people  would  mean  to  redirect  or  change  the  object  of  their  enthusiasm 
and  I don’t  think  this  is  going  to  be  too  easy,  especially  later  on.  You  have  to  remember 
that  we  are  dealing  with  people  that  have  motives  and  are  motivated.  They  are  not  objects 
sitting  on  a conveyor  belt  that  we  can  manipulate. 

Pengelly  — I wasn’t  suggesting  manipulation.  Dave.  What  I was  suggesting  in  agreeing  with 
Dr.  Gittins  on  channelling  is  this:  most  kinds  of  financial  support  require  that  some  knowl- 
edgeable professor  who  really  knows  the  art  of  grantsmanship  apply  for  support  from  our 
friendly  uncle  in  Ottawa.  The  group  of  people  making  decisions  on  research  grants  will 
simply  say  that  because  there  are  no  job  opportunities  in  taxonomy  in  the  foreseeable 
10  years,  we  are  not  going  to  sponsor  55  taxonomists  in  Canadian  universities.  This  is 
the  kind  of  channelling  that  I was  envisioning.  Funds  would  be  directed  into  areas  of  need. 
Enthusiastic  students  who  followed  the  dollar  sign  a little  bit  might  find  problems  that 
fitted  into  these  areas.  I wasn’t  saying  that  you  are  not  going  to  be  what  you  want  to  be. 
Kavanaugh  - If  we  can  accept  the  fact  that  we  do  have  some  capacity  to  direct  where  the 
funds  go  and  that  we  are  not  completely  at  the  mercy  of  public  opinion,  I would  be 
reasonably  satisfied.  I just  wanted  to  know  at  what  point  we  can  really  influence  a student  — 
interest  him  in  a given  field. 

Pengelly  - In  Ontario  there  is  a committee  in  existence  that  is  going  around  and  visiting 
various  departments  in  various  universities.  Members  of  this  committee  are  asking  that  each 
professor  justify  what  he  is  teaching  in  relation  to  those  courses  offered  elsewhere  in  his 
university.  If  he  can't,  if  his  entomology  course  is  too  specialized  for  a general  biology 
program,  he  could  be  asked  to  move  to  another  university  with  strength  in  entomology.  I 
suspect  that  we  will  see.  within  Ontario  at  least,  different  campuses  specializing  in  different 
disciplines  such  as  engineering  or  food  science  or  entomology.  There  will  be  only  one  or 
two  “centres  of  excellence”  for  entomology.  You.  as  a student,  will  have  to  be  very  careful 
when  choosing  an  institution  to  further  your  education  because  all  will  be  specialized. 
Gittins  - Let’s  not  be  too  gloomy  about  the  future  of  entomology.  I think  it  looks  pretty 
good.  Also,  let’s  eliminate  the  use  of  those  two  terms  “basic”  and  “applied”  - they  are 
meaningless  today.  In  Idaho  we  are  now  looking  at  insect  control  in  terms  of  population 
suppression.  No  longer  are  we  going  to  rely  solely  on  insecticides.  We  are  investigating 


Discussion 


25 


instead  the  fundamental  biology  of  pest  insects  and  seeing  how  we  can  manipulate  various 
aspects  of  these.  The  20  years  we  have  just  completed  were  atypical,  not  typical.  As  pest 
management  programs  develop  we  are  going  to  require  improved  insect  keys  and  a fuller 
understanding  of  basic  biology.  Whether  we  are  interested  in  physiology  or  taxonomy  or 
toxicology  we  are  all  still  interested  in  insects;  interested  in  all  of  their  aspects.  Surely  we 
can  redirect  our  activities  without  instituting  a bunch  of  new  courses.  Our  primary  interest 
may  still  be  systematics  but  we  can  practise  it  with  a different  emphasis.  We  are  not  just 
taxonomists  or  physiologists,  we  are  entomologists,  all  of  us,  and  we  should  be  proud  of  it. 
Morrison  — Waterloo  — Is  the  government  of  Alberta  funding  the  university  here  in  terms 
of  the  number  of  students  enrolled?  Is  the  funding  reaching  down  to  the  department  or 
faculty  level  as  it  has  in  Ontario?  This  creates  very  serious  problems  in  the  teaching  pro- 
grams that  a department  tries  to  implement.  It  is  devastating  in  many  many  ways.  When 
Guelph  talks  of  their  problems  they  talk  of  them  in  this  light,  and  when  we  talk  of  our  prob- 
lems we  talk  of  them  in  this  light.  Could  you  please  enlighten  us? 

Hocking  — A large  part  of  the  funding  at  the  University  of  Alberta  is  geared  through  weekly 
student  hours  but,  hitherto,  this  has  been  on  the  basis  of  predicted  weekly  student  hours. 
Initially,  a correction  factor  applied  if  enrolment  did  not  meet  predictions  and  this  is  per- 
haps one  reason  why  predictions  are  high.  It  is,  I agree,  a terrible  system  but  what  is  the 
alternative?  I wish  there  were  some  way  of  getting  away  from  this.  We  resisted  it  to  our  cost 
as  a department  just  about  as  far  as  we  could  go.  I think  there  is  an  increasing  tendency  to 
resist  this  and  hopefully  somebody  will,  sooner  or  later,  really  come  to  grips  with  it  and  get 
something  done  about  it. 

Pengelly  — You  are  really  fighting  Mother  when  you  go  after  these  people  because  they  are 
the  ones  with  the  purse  strings  and,  as  Dr.  Hocking  suggests,  this  might  be  to  your  cost  if 
you  challenge  them  too  hard. 

W.  E.  Heming  — Guelph  — For  many  years  I have  had  the  feeling  that  a gifted  popularizer  of 
science  is  making  as  great  a contribution  as  many  of  the  research  scientists,  and  I feel  maybe 
we  have  reached  the  point  where  a gifted  popularizer  of  entomology  could  reach  the  public, 
could  reach  the  granting  agencies  and  make  a very  real  contribution.  I wonder  what  some  of 
the  rest  of  you  feel? 

Schwab  - Edmonton  - My  name  is  Betty  Schwab  and  I am  a psychiatrist.  I like  Dr. 
Heming’s  comments.  Those  I am  going  to  make  are  probably  a little  philosophical.  I am 
speaking  as  a person  who  now  realizes  that  the  two  years  that  I spent  in  entomology  were, 
to  a degree,  wasted  because  I did  not  realize  how  valuable  they  were  going  to  be  to  me  as  a 
mother.  I think  as  teachers,  that  we  are  waiting  much  too  long  to  interest  children  in  the 
fascinating  field  of  entomology.  I heard  Mr.  Jantzie  say  that  he  was  dealing  with  a turned 
off  group  by  the  time  that  they  reached  high  school.  This  struck  me  as  being  very  sad 
because  I think  that  if  those  children  had  been  motivated  to  look  at  insects  as  kindergarten 
students,  in  grade  school  and  high  school  they  would  have  been  a different  population 
of  young  people.  I would  suggest  that  even  at  the  university  level,  it  would  be  to  the 
advantage  of  the  general  public,  to  the  children  and  to  entomology  to  develop  simple 
programs  for  kindergarten  students. 

Pengelly  — That’s  certainly  endorsement  of  Mr.  Jantzie’s  idea  that  we  start  teaching  ento- 
mology a little  earlier. 

Craig  - Edmonton  - I would  like  to  speak  as  a father.  We  have  a daughter  four  years  old 
and  a son  two.  Our  daughter  has  a number  of  friends  around  the  place  who  come  and  play. 
Last  year  we  made  a breakthrough  when  our  daughter  lost  her  fear  of  mosquitos.  We  have 
had  great  difficulty,  however,  in  stopping  her  friends  from  squashing  every  bug  that  they 
see.  Thus,  this  business  of  education  must  be  started  very  early.  Our  2-year  old  son  is  just 


26 


Discussion 


the  opposite  — he  loves  everything.  He  will  often  lie  on  the  foot  path  and  say  “Hi!”  to 
every  ant  that  comes  along.  In  the  next  few  years  he  is  going  to  see  his  friends  stomping 
on  insects.  Perhaps  we  should  be  looking  at  this  age  group.  I am  directing  these  comments 
to  Dr.  Downer  and  also  to  myself  and  perhaps  to  the  Student  Encouragement  Committee 
of  the  Entomological  Society  of  Canada.  Perhaps  we  should  be  attempting  kindergarten 
encouragement  rather  than  student  encouragement. 

Corbet  — Waterloo  — I think  the  last  few  comments  have  raised  in  my  mind  very  clearly, 
two  questions.  One  relates  to  communication  and  one  relates  to  the  reason  why  we  recog- 
nize entomology  as  a separate  science.  I think  there  is  a reason  why  we  recognize  entomolo- 
gy as  a separate  science  and  it  is  largely  one  of  convenience.  It  is  particularly  related  to 
training  in  the  direction  in  which  we  may  expect  to  get  professional  employment  afterwards. 
I think  there  is  another  reason  why  we  should  recognize  entomology  as  part  of  and  not 
separate  from  biology  and,  speaking  now  to  the  question  of  communication,  why  we  should 
teach  entomology.  I think  one  of  the  most  compelling  reasons  we  teach  entomology  is  to 
illustrate  biological  principles.  In  this  connection  it  matters  not  one  whit  as  to  whether  the 
person  who  learns  entomology  later  on  becomes  a professional  entomologist.  It  is  very  im- 
portant however  that  he  become  a knowledgeable  biologist.  The  people  here  who  don’t 
already  know  it,  might  be  interested  to  hear  of  one  of  the  small  number  of  resolutions  that 
the  founding  meeting  of  the  Canadian  Committee  of  University  Biology  Chairmen  made 
when  it  met  in  November  (1971)  in  Montebello.  This  resolution  was  that  members  of 
the  biological  disciplines  or  subdisciplines,  and  of  course  that  includes  entomology,  should 
refer  to  themselves  first  as  biologists  and  only  second,  if  necessary  and  appropriate,  as 
adherents  of  a subdiscipline. 

Pengelly  — Thanks,  Dr.  Corbet.  Maybe  my  comments  were  not  exactly  parallel  to  yours 
but  I still  feel  the  name  entomology  has  to  hang  around  someplace.  I sat  and  watched  it 
going  down  the  drain  once. 


27 


TAXONOMIC  ENTOMOLOGY;  GOING,  GOING,  - WHERE? 

W.  R.  M.  Mason 
Entomology  Research  Institute 
Canada  Department  of  Agriculture,  Ottawa,  Ontario 

From  looking  at  the  program  I suspect  that  my  views  are  supposed  to  represent  the 
government  employment  ivory  tower  segment  of  entomology.  Accordingly,  I take  it  that  I 
can  discuss  any  reasonable  aspect  of  entomology  and  education.  Because  of  my  background, 
the  viewpoint  that  will  come  through  loud  and  clear  will  be  that  of  the  basic  research 
worker  in  a government  department.  Thus,  I will  start  with  a description  of  the  condition  of 
systematic  entomology  at  the  present  time. 

Much  of  what  I have  to  say  about  systematic  entomology  has  been  stated  in  a report 
recently  published  by  the  National  Science  Foundation  in  the  United  States  (Steere  et  al. , 
1971).  This  report  emphasizes  that  the  great  systematic  biology  collections  are  an  irre- 
placeable and  priceless  national  resource,  functioning  as  the  key  to  all  filing  and  retrieval 
systems  for  information  on  the  diversity  of  living  creatures  of  the  world.  “These  great  col- 
lections are  now  in  a situation  that  can  only  be  called  desperate.  They  are  suffering  from: 
(1)  a great  influx  of  new  specimens  resulting  largely  from  government  financed  research 
projects;  (2)  stepped-up  use  of  collections  in  response  to  specific  practical  questions  as  well 
as  intensified  research;  (3)  sky-rocketing  inflation  on  top  of  a static  income  base  leading  to  a 
merciless  financial  crunch.  The  result  has  been  substandard  salaries,  deteriorating  personnel 
strength,  reduced  activities  and  deferred  expansion.  Deficits  have  been  reflected  more  in 
deterioration  than  in  actual  dollar  figures  but  soon  actual  deficits  will  be  inevitable.”  The 
publication  closes  with  an  urgent  plea  for  financial  assistance  from  the  federal  government 
but  the  sum  is  surprisingly  modest  considering  the  means  of  our  neighbours  to  the  south: 
they  want  only  198  million  dollars  over  the  next  10  years. 

The  above  publication  covers  conditions  in  the  great  systematic  collections  (exclusive  of 
the  U.  S.  National  Museum)  in  the  United  States  but,  nevertheless,  much  of  its  content 
applies  equally  to  the  great  collections  in  Canada.  Let  me  tell  you  what  has  happened  to  the 
Entomology  Research  Institute,  undoubtedly  the  largest  and  best  financed  in  the  country. 
Since  1968  our  budget  has  been  cut  to  less  than  half  the  number  of  dollars  that  were  avail- 
able in  1968.  Our  staff  of  professionals  has  suffered  six  resignations  or  retirements  and  only 
three  have  been  replaced.  During  the  same  four  years  we  have  had  to  increase  our  identifica- 
tion service  from  43,000  to  77,000  identifications  per  year.  And  we  have  had  no  additional 
space,  although  the  collections  continue  to  grow  at  an  alarming  rate.  Fortunately,  our  Insti- 
tute experienced  rapid  growth  of  staff  and  facilities  during  the  1950’s  and  early  1960’sand 
thus  things  are  not  as  bleak  as  my  figures  might  indicate. 

The  more  entomological  museums  one  visits  the  stronger  becomes  the  pervading  impres- 
sion of  overcrowding,  under-staffing  and  financial  stringency.  This,  sad  to  say,  is  the  atmos- 
phere into  which  the  aspiring  taxonomist  will  be  heading  and  it  is  not  likely  to  change  until 
taxonomists  themselves  are  better  able  to  compete  for  the  budget  dollar. 

If  present  trends  continue  unaltered  one  can  only  foresee  a continuing  deterioration,  not 
only  of  the  great  collections  themselves,  but  also  of  the  associated  services  such  as  identifi- 
cation and  bibliographic  advice.  The  time  may  well  come  when  identifications  will  become 
very  slow  and  difficult  to  make  in  some  groups  and  completely  impossible  in  others.  Then 
perhaps,  biologists  in  general,  and  especially  ecologists,  will  finally  awaken  to  the  plight  of 
museums  and  bring  pressure  to  bear  upon  governments  and  granting  institutions  to  make 
available  enough  funds  to  put  life  back  into  the  museum  segment  of  biological  sciences. 


28 


Mason 


Even  at  that  it  will  take  many  years  to  train  a new  generation  of  taxonomists  to  revive 
ancient  and  moribund  institutions  and  to  modernize  their  methodology.  For  some  institu- 
tions a revival  could  come  too  late  because  museum  pests  and  other  natural  agencies  may 
have  already  destroyed  large,  neglected  parts  of  the  collections. 

Now  this  is  the  kind  of  thing  that  might  happen  and  I hope  it  represents  a pessimistic  pole 
for  the  probabilities  available  to  us.  What  is  more  likely,  I expect,  is  that  the  cries  for  help, 
such  as  the  one  from  the  U.  S.  National  Science  Foundation  that  I referred  to  earlier,  will  be 
heeded  in  time  and  the  museums  will  be  kept  going  even  though  on  a more  stringent  scale 
than  they  would  like.  There  are  plenty  of  intelligent  people  in  high  places  who  can  see  the 
need  and  urgency  and  we  can  only  hope  that  their  priorities  will  include  museums.  At  any 
rate  none  of  us  dare  sit  back  complacently  and  wait  for  “George”  to  do  it.  It  is  up  to  every- 
one who  has  an  interest  in  the  maintenance  of  museums  in  good  financial  and  physical 
health  to  do  whatever  he  can  in  the  way  of  agitation  for  their  better  maintenance. 

Another  great  trouble  bothering  taxonomy  is  the  conservative  nature  of  taxonomists 
themselves.  We  tend  far  too  much  to  cling  to  old,  time-consuming  methods  of  working, 
necessitated  by  rules  that  were  originally  drawn  up  in  the  days  when  taxonomy  was  the 
exclusive  field  of  amateur  dilettantes  of  independent  wealth.  Consider  the  amount  of  time 
and  effort  that  goes  into  searching  through  ancient  literature  trying  to  dig  out  the  oldest 
name  and  the  first  combination  etc.  etc.  This  is  not  zoology;  it  is  bibliographic  research 
done  by  the  most  old  fashioned  and  time-consuming  methods.  Furthermore,  it  is  only  of 
historical  interest  and  has  very  little  place  in  science.  Think  where  the  science  of  chemistry 
would  be  today  if  they  had  a code  of  nomenclature  that  rigorously  demanded  that  the  earli- 
est name  for  every  chemical  compound  be  exhumed  from  the  literature  and  used  regardless 
of  all  logic.  Think  also  where  they  would  be  if  all  papers  had  equal  value  no  matter  how 
good  or  poor  the  work  was.  In  other  branches  of  science  a poor  piece  of  work  is  quickly 
read  and  thrown  aside  to  be  forgotten  if  it  is  of  no  value,  whereas  in  taxonomy  we  are  stuck 
with  it  forever  no  matter  how  ambiguous  or  unrecognizable  or  even  crazy  the  work  happens 
to  be.  And  what  a horrible  condition  our  cataloguing  is  in  when  the  chief  cataloguing  journal 
for  taxonomists,  Zoological  Record,  is  six  years  behind  the  literature  and  steadily  losing 
place.  If  all  taxonomic  novelties  such  as  new  names  and  new  combinations  had  to  be  vali- 
dated by  filing  with  an  international  bureau  (why  not  the  Zoological  Record?)  there  would 
be  little  trouble  in  keeping  the  catalogues  up  to  date.  Some  of  the  money  now  going  into 
grants  for  biological  research  should  certainly  be  put  into  cataloguing  and  bibliographic 
services,  fully  computerized  and  internationally  available  for  all  zoologists  or  botanists  to 
draw  upon.  This  kind  of  thing  ought  to  be  a function  of  a large  world  organization  such  as 
the  United  Nations.  The  ancient  literature  should  be  fully  catalogued  once  and  for  all  and 
the  data  filed  in  computer  banks.  Then  the  old  rare  and  expensive  books  need  never  again 
be  consulted  except  for  historical  interest.  Anything  missed  in  that  cataloguing  process 
(after  a waiting  period  during  which  biologists  at  large  could  note  deficiencies)  should  sim- 
ply be  written  off  because  only  a rare  and  valueless  publication  would  be  omitted. 

A number  of  computerized  systems  for  use  by  taxonomists  and  other  biologists  are  at 
present  being  developed  in  Washington  and  other  centers.  Working  out  suitable  techniques 
for  the  manipulation  of  a large  body  of  systematic  data  is  not  an  easy  task,  but  in  the  last 
few  years  computer  technology  has  evolved  to  the  point  where  direct  application  to  the 
management  of  large  amounts  of  systematic  data  is  feasible.  The  Smithsonian  Information 
Retrieval  system,  the  Flora  of  North  America  Program  and  the  Museum  Computer  Network 
system  are  at  present  under  development.  Lack  of  funds  and  staff  very  seriously  limit  the 
participation  of  many  museums  in  these  programs  but  the  outlook  for  computerization  of 
museum  technology  is  bright. 


Taxonomic  Entomology 


29 


Computerization  of  taxonomic  data  has  recently  achieved  a rather  bad  reputation  among 
taxonomists  because  of  early  misapplications  toward  the  study  of  phylogeny  and  other  rela- 
tionships. However,  in  proper  applications  the  computer  could  revolutionize  the  abilities  of 
curators  to  retrieve  data.  As  antipathy  toward  modernization  and  fear  of  the  machine  give 
way  to  the  realization  of  the  computers’  enormous  ability  to  manipulate  data,  the  advances 
necessary  for  the  maximum  utilization  of  these  great  collections  will  soon  follow.  Compu- 
ters can  also  be  very  important  in  making  identifications  of  specimens  in  reasonably  well 
known  groups.  Not  only  can  a computer  construct  a key  but  it  can  also  run  a specimen 
through  the  key,  needing  only  an  operator  to  select  key  characters  from  observation  of  the 
specimen.  The  computer  can  even  tell  the  observer  what  characters  to  read.  This  will  release 
highly  trained  research  workers  from  the  routine  part  of  identification  activities  and  enable 
the  latter  to  be  carried  on  by  technicians. 

Will  the  research  taxonomist  be  needed  at  all  in  this  idyllic  future?  Yes,  indeed  he  will, 
because  the  computer  cannot  replace  the  research  brain  and  it  cannot  extract  new  data  from 
unworked  specimens.  Fortunately  for  insect  taxonomists  the  Insecta  are  not  only  the  most 
numerous  of  all  living  creatures  but  also  the  least  known.  Many  groups  of  Insecta  are  five  or 
10  percent  known  on  a world  basis.  Some  are  that  poorly  known  even  within  such  reason- 
ably well  studied  areas  as  North  America. 

Dr.  H.  K.  Townes  has  recently  compiled  figures  (Townes,  1969)  for  the  world  fauna  of 
Ichneumonidae  (Table  1).  Note  his  estimate  that  only  35%  of  the  North  American  species 
are  presently  known.  Note  also  his  estimate  that  between  85  and  90%  of  all  tropical  species 
are  unknown  and  that  in  the  world  as  a whole  there  are  approximately  four  undescribed 
species  for  every  described  one.  I might  also  add  out  of  personal  knowledge  that  at  least  half 
the  described  species  are  known  from  only  one  or  a very  few  specimens  with  no  associated 
ecological  data.  To  put  it  in  other  words  our  acquaintance  with  at  least  90%  of  the  species 
of  Ichneumonidae  is  limited  to  a few  museum  specimens  only.  Since  a modern  taxonomist 
can  revise  at  best  only  a few  hundred  species  a year  even  at  the  alpha  level,  and  since 
the  Ichneumonidae  are  typical  of  large  parts  of  the  lesser  known  sections  of  the  Insecta,  it  is 
easy  to  see  that  there  is  absolutely  no  danger  of  the  entomological  taxonomist  running  out 
of  work  for  generations  to  come.  The  real  danger  is  that  large  parts  of  the  Insecta  will  be 
extinct  before  they  are  even  described. 

Now  I move  from  the  needs  for,  and  opportunities  in,  insect  taxonomy  to  the  demand 
for  taxonomists  — a very  different  subject. 

This  graph  (Figure  1)  is  adapted  from  one  published  by  Kelly  (1971).  I think  it  might 
serve  very  well  as  the  symbol  of  the  symposium,  showing  as  it  does  the  really  outstanding 
problem  facing  entomology  departments  and  graduate  students  today.  The  contrast  in  the 
actual  numbers  and  also  the  trends  of  the  output  of  Ph.D.’s  and  the  positions  available  for 
them  is  too  obvious  to  need  any  further  comment,  but  the  plight  of  the  unfortunate 
individuals  who  are  caught  in  the  gap  calls  for  a lot  of  hard  thinking  and  action.  The  reason 
I show  this  graph  is  to  serve  as  background  for  a story  illustrating  how  badly  wrong  the 
common  wisdom  may  be  and  how  desperately  we  really  need  accurate  forecasts  of  a type 
not  available  to  us  in  the  past  (nor  at  present,  according  to  Kelly). 

During  1967,  the  editor  of  the  Canadian  Entomologist,  Dr.  D.  P.  Pielou  asked  me  if  I 
would  write  an  article,  preferably  controversial  enough  to  start  some  correspondence  and 
stimulate  interest,  for  the  first  number  of  a popular  publication  he  planned  to  start  for  the 
entomologists  of  Canada.  The  publication  has  since  appeared  and  is  now  established  as  The 
Bulletin  of  the  Entomological  Society  of  Canada.  It  sounds  strange  to  say  it  now,  but  at  that 
time,  in  1967,  I was  trying  very  hard  to  recruit  Canadian  taxonomists  for  positions  in  the 
Entomology  Research  Institute.  Quite  a number  of  well  qualified  foreign  students  were 


30 


Mason 


Table  1.  World  species  of  Ichneumonidae 


Valid  described 

Estimated  % 

Estimated  total 

species 

described 

species 

Nearctic 

2809 

35 

8026 

W.  Palaearctic 

4023 

70 

5747 

E.  Palaearctic 

2400 

20 

12000 

Indo-Australian 

2424 

15 

16160 

Ethiopian 

1618 

15 

10787 

Neotropical 

1718 

10 

17180 

14992 

69900 

Allowance  for  duplication  (13.5%) 

-2025 

-9400 

Corrected  Total 

12967 

21 

60500 

-2000 


75  67  68  70  7\  ?2 


Figure  1.  Supply  and  demand  for  Ph.D.’s  in  science  and  engineering  in  Canada.  ABD  - all  but  dissertation. 


available  and  anxious  to  take  the  positions  but  the  executive  insisted  on  having  Canadian 
citizens  for  the  positions  if  at  all  possible.  There  were  very  few  students  qualified  to  work  in 
taxonomy  and  the  few  who  were  qualified  seemed  disinterested.  So  in  an  attempt,  which  I 
should  in  honesty  now  admit  was  a little  contrived,  to  bait  a few  university  professors  I 
wrote  an  article  (Mason,  1969)  complaining  of  the  lack  of  Canadian  students  trained  in 
taxonomy  and  laid  the  blame  on  the  shoulders  of  the  Canadian  universities.  Well,  I certainly 
got  a reaction  (Kevan,  1969;  Scudder,  1969).  However,  before  I elaborate,  examine  the 
graph  once  more  and  see  when  the  article  was  written.  In  1 967  the  production  of  graduate 
students  was  well  below  the  accumulation  of  new  positions.  Allow  a lapse  of  almost  18 


Taxonomic  Entomology 


31 


months  while  the  article  was  in  press  and  then  see  the  result  in  late  1 969  when  the  letters 
from  the  professors  came  in  to  the  editor.  By  that  time  new  positions  were  declining  and 
the  output  of  Ph.D.’s  was  still  rising.  This  time  no  one  needed  experts  to  tell  them  what 
the  trends  were.  Unemployment  was  screaming  at  us  from  every  university  campus  in  the 
country  and  the  federal  government  was  in  the  midst  of  reducing  its  staff  by  not  replacing 
retirements,  by  forcing  early  retirements,  and  even  by  a little  outright  firing. 

Well  at  least  I was  not  alone  in  my  ignorance  back  in  the  middle  sixties.  In  those  days  we 
were  living  in  an  era  when  all  graphs  went  upwards  and  upwards  forever.  The  catastrophe  of 
1929  seemed  far  away  and  we  were  secure  in  our  knowledge  that  modern  economics  had 
now  made  such  an  occurrence  impossible.  We  heard  no  academic  voices  crying  that  the  pro- 
duction of  graduate  students  should  be  curtailed  because  the  young  hopefuls  who  were  then 
enrolling  in  graduate  studies  would  not  be  able  to  find  jobs  in  three  or  four  years  when  they 
launched  themselves  upon  the  market.  No,  the  cry  was  for  Canadian  universities  to  enlarge 
their  graduate  departments,  train  our  own  Canadian  students  at  home  and  generally  more 
and  more  of  everything.  But  the  bubble  burst,  not  just  as  it  did  in  1 929  and  not  with  the 
same  drastic  results.  Nevertheless,  burst  it  did,  and  we  are  now  living  with  the  results.  Those 
who  should  be  the  most  eligible  and  employable  members  of  our  society  are  declared  useless 
because  of  inexperience  and  overqualification.  Those  with  experience  are  declared  obsolete, 
overaged  and  unemployable  because  of  pension  fund  computations.  It  seems  that  no  matter 
who  you  are  or  what  your  background  the  personnel  department  has  a stainless-steel-clad 
reason  for  declaring  you  ineligible  for  any  vacancy. 

Well  all  this  gloomy  talk  serves  to  indicate  that  the  demand  for  taxonomists  in  the  near 
future  is  likely  to  be  far  below  what  taxonomists  themselves  believe  is  the  need. 

But  what  is  a need?  Certainly  everybody  knows  there  is  a difference  between  demand  and 
need  but  what  determines  it?  Well  as  I see  it  needs  are  what  you  believe  you  wish  to  have 
and  demand  is  what  you  are  willing  to  pay  for.  Needs  may  be  divided  into  necessities  and 
luxuries  and  of  course  the  dividing  line  is  always  a nebulous  zone  that  varies  from  one  indi- 
vidual to  another  and  from  one  community  to  another.  For  instance,  I have  travelled  in 
primitive  parts  of  the  world  where  the  population  considers  a man  who  suffers  only  from 
malnutrition,  intestinal  parasites  and  endemic  malaria  to  be  healthy  and  where  they  can  see 
no  need  for  any  measures  to  prevent  these  conditions.  In  other  words,  if  you  are  poor 
enough  and  ignorant  enough,  malaria  control  and  sanitation  are  luxuries,  not  necessities. 

Obviously,  if  the  public  and  its  representatives  are  to  hire  entomologists  they  must  be 
sold  on  the  idea  that  the  services  of  entomologists  are  a necessity.  To  some  extent  this  job 
has  been  done  but  it  seems  that,  at  the  moment,  entomologists  above  a certain  number  in 
the  Public  Service  are  regarded  as  a luxury.  In  fact  the  federal  government,  through  its  top 
executives,  regards  the  Public  Service  of  Canada  as  being  presently  overloaded  with  entomo- 
logists. Hence,  the  lack  of  demand  for  entomologists  in  the  Public  Service.  It  is  quite  clearly 
a disastrous  policy  for  a department  such  as  this  one  in  the  University  of  Alberta  to  continue 
to  train  entomological  graduates  in  numbers  larger  than  the  market  place  will  absorb.  Only 
four  courses  of  action  seem  open:  (1)  cut  down  the  production  of  the  department;  (2)  con- 
vince employing  agencies,  chiefly  the  governmental  and  university  authorities,  that  more 
entomologists  are  needed;  (3)  direct  the  graduates  into  non-entomological  employment;  or 
(4)  do  nothing.  The  last  two  alternatives  seem  silly.  The  first  course  of  action  is  a passive 
one  of  simply  trying  to  foresee  the  demand  and  filling  it  as  closely  as  possible.  The  present 
situation  demonstrates  how  well  universities  have  been  succeeding  recently.  They  are  not 
entirely  to  blame,  admittedly,  for  there,  is  no  source  of  information  on  which  to  make 
reasonable  forecasts  (Kelly,  1971).  The  second  course  of  action  involves  salesmanship  and  is 
the  kind  of  thing  that  has  made  business  expand  in  a free  enterprise  system.  An  effective 


32 


Mason 


enough  advertising  campaign  can  convert  a luxury  into  a necessity  in  the  minds  of  people 
with  enough  money  to  pay  for  it.  The  present  popularity  enjoyed  by  environmentalists  and 
conservationists  should  be  the  key  to  a sales  approach  by  entomologists.  After  all,  insects 
are  by  far  the  most  numerous  forms  of  animal  life  in  Canada  or  in  any  other  part  of  the 
terrestrial  world  and  are  the  least  known. 

Should  the  university  be  in  the  advertising  business?  Well,  that  is  a decision  that  is  up  to 
the  university,  but  it  seems  to  me  that  there  is  already  tacit  admission  that  something  along 
these  lines  is  necessary  when  universities  maintain  placement  services  to  assist  their  students 
in  obtaining  employment.  Furthermore;  there  seems  little  point  in  using  the  university 
budget  to  build  larger  facilities  and  improved  labs  and  hire  more  staff  if  no  one  will  buy  the 
product.  Any  factory  that  was  glutting  the  market  with  an  unsalable  commodity  would  very 
quickly  close  its  production  lines  and  pump  increased  funds  into  advertising  in  order  to 
move  the  stock.  And  if  universities  will  not  indulge  in  this  messy  business  of  advertising, 
what  then?  Well  clearly  young  people  are  going  to  look  elsewhere  and  the  great  facilities  so 
painstakingly  built  up  in  the  universities  will  be  running  at  half  capacity.  Maybe  empty  class- 
rooms and  empty  lab  space  will  be  a very  pleasant  novelty  for  a while  but  it  might  also  mean 
reductions  in  staff  (horrible  thought)  and  reductions  in  budget  (more  horrible  thoughts). 
Probably  the  apparently  inevitable  population  growth  will  catch  up  with  the  empty  spaces 
and  fill  them  again  but  by  that  time  the  facilities  may  well  be  obsolete. 

In  spite  of  all  these  words  of  gloom,  I can  foresee  a very  bright  future  for  entomologists 
in  general  and  also  for  taxonomists  if  the  opportunities  inherent  in  our  present  situation  are 
correctly  handled.  Entomology  can  even  be  made  a popular  study  with  the  public.  As  proof 
of  that  one  need  only  examine  the  number  of  entomological  clubs  existing  in  Japan.  I am 
told  that  butterfly  and  dragonfly  collecting  in  Japan  are  about  as  popular  as  snowmobiling 
is  in  Canada. 

I think  the  fundamental  problem  is  that  the  public  must  be  made  aware  of  insects  in  a 
pleasant  and  positive  way  and  thus  be  made  interested  in  the  encouragement  of  entomolo- 
gists. At  the  moment  our  average  fellow  Canadian  thinks  of  an  insect  as  almost  synonymous 
with  a house  fly  or  a mosquito,  in  other  words  an  unmitigated  nuisance  — the  only  good  one 
being  a dead  one.  With  that  kind  of  attitude  the  good  entomologist  is  the  one  who  can  kill 
insects  with  the  greatest  efficiency.  But  you  and  I know  that  the  vast  majority  of  insects  go 
about  minding  their  own  business  in  ways  that  are  really  beneficial  to  humanity  even  though 
humanity  does  not  appreciate  it.  I think  it  behooves  all  entomologists  to  think  very  hard 
and  very  seriously  about  improving  our  public  image  as  a group  and  thereby  improving  our 
lot  and  our  share  of  the  national  budget. 

REFERENCES 

Kelly,  F.  1971 . Prospects  for  scientists  and  engineers  in  Canada.  Science  Council  of  Canada, 
Special  Study  20. 

Kevan.  D.  K.  McE.  1969.  Insect  taxonomy  in  Canada.  Bui.  Ent.  Soc.  Can.  2:13-15. 

Mason.  W.  R.  M.  1969.  Insect  taxonomy  - Is  it  dying  in  Canada?  Bui.  Ent.  Soc.  Can.  1:15-17. 
Scudder.  G.  G.  E.  1969.  Insect  taxonomy  training  in  Canada.  Why  not  help  instead  of  blame 
the  universities?  Bui.  Ent.  Soc.  Can.  2:16-18. 

Steere.  W.  C..  et  al.  1971.  The  systematic  biology  collections  of  the  United  States  — an 
essential  resource.  Part  I.  II.  A report  to  the  National  Science  Foundation  by  the  Confer- 
ence of  Directors  of  Systematic  Collections.  New  York  Botanical  Garden.  Bronx.  N.  Y. 
Townes,  H.  K.  1969.  The  genera  of  Ichneumonidae,  Part  I.  Size  and  complexity  of  the 
Ichneumonidae.  Mem.  Amer.  Ent.  Inst.  11:3-7. 


33 


EDUCATION  AND  THE  PROFESSIONAL  ENTOMOLOGIST 

N.  D.  Holmes 
Research  Station 

Canada  Department  of  Agriculture,  Lethbridge,  Alberta 

Although  entomologists  occupy  various  positions  as  in  teaching,  extension,  or  industry, 
I wish  to  limit  my  discussion  to  those  employed  primarily  to  produce  new  knowledge  that 
will  be  applied  for  the  direct  benefit  of  society.  I will  concentrate  mostly  on  those  associ- 
ated with  research  in  agriculture,  because  they  constitute  the  largest  group  of  professional 
entomologists  in  Canada. 

Entomologists  have  found  relatively  few  positions  in  most  fields  in  recent  years  and  there 
is  little  evidence  of  much  immediate  improvement.  Better  long-term  prospects,  however, 
make  this  an  appropriate  time  to  examine  their  education.  As  the  present  generation  in  uni- 
versities and  elsewhere  retire,  new  entomologists  will  undoubtedly  be  hired.  It  is  likely, 
however,  that  the  largest  number  of  replacements  will  be  required  in  departments  of  the 
federal  government  either  in  Agriculture  or  Forestry,  providing  that  drastic  changes  are 
not  made  as  a result  of  advice  from  the  new  Ministry  of  State  for  Science  and  Technology. 

Prospects  in  Agriculture 

The  Canada  Department  of  Agriculture  employed  about  140  entomologists  in  1944, 
rising  to  over  350  in  1956  (2). 

By  1964  this  number  had  fallen  to  173,  which  still  amounted  to  over  half  the  entomolo- 
gical force  employed  in  Canada  (4).  Of  the  remainder,  about  40  were  employed  by  univer- 
sities and  provincial  governments,  and  about  96  by  the  Department  of  Forestry  (now  in 
Environment  Canada). 

In  1972  the  Canadian  Forestry  Service  employed  87  entomologists  plus  15  administra- 
tors, some  of  whom  may  not  be  entomologists.  The  Research  Branch,  CDA,  employed 
162  entomologists;  five  in  stored  products  research,  1 1 in  veterinary-medical  entomology, 
33  in  taxonomy,  and  1 13  mostly  in  crop  entomology. 

Most  entomologists  in  the  Research  Branch  were  probably  hired  between  1945  and  1950. 
Because  early  retirement  after  30  or  35  years  of  service  is  encouraged  by  the  federal  govern- 
ment, it  is  likely  that  by  1980  many  will  be  retired  or  close  to  retirement.  Allowing  for 
some  attrition  of  positions,  the  Branch  will  require  60  to  80  entomologists  within  the 
next  eight  to  13  years.  Undoubtedly  the  Forestry  Service  will  also  require  some  in  the  same 
period,  bringing  the  probable  total  to  well  over  100. 

The  demand  will  depend  on  entomologists  explaining  the  importance  of  their  research. 
The  belief  that  the  need  for  entomology  was  self-evident  became  obsolete  with  the  advent 
of  tight  budgets  and  cost-benefit  analyses  of  research.  Data  to  show  the  need  for  entomolo- 
gical research  are  scarce.  The  decision  of  the  Canadian  Entomological  Society  to  remain 
separate  from  organizations  such  as  the  Agricultural  Institute  of  Canada  removed  us  from 
the  places  of  influence.  Until  data  are  obtained  to  show  the  potential  hazards  from  insect 
pests  or  until  we  experience  more  dramatic  outbreaks,  such  as  that  of  the  Bertha  army- 
worm  in  1971,  we  will  continue  to  find  it  difficult  to  change  the  current  impression  that 
there  are  too  many  entomologists. 

Another  factor  has  been  the  success  of  the  control  measures  that  we  developed,  primarily 
centering  around  the  use  of  insecticides.  Insecticide  use  resulted  in  the  expansion  of  regula- 
tory agencies  and  increased  the  numbers  of  pesticide  analysts,  but  reduced  the  apparent 
need  for  entomological  research.  Even  though  insecticides  have  detrimental  side  effects 


34 


Holmes 


and  have  not  provided  permanent  insect  control,  their  immediate  and  evident  effective- 
ness makes  it  difficult  for  non-entomologists  to  understand  why  entomologists  advocate 
the  continued  search  for  better  controls.  Some  support  and  understanding  for  our  position 
has  been  generated  by  public  reaction  in  recent  years. 

Even  if  we  had  sufficient  knowledge  to  solve  all  our  present  problems  satisfactorily, 
the  need  for  entomological  research  would  continue.  New  insect  outbreaks  will  inevitably 
follow  the  introduction  of  new  crops  and  cultural  practices.  Environmental  changes,  the 
development  of  resistant  strains  of  insects,  the  necessity  for  opening  up  new  agricultural 
areas,  and  the  increasing  demand  for  food  will  all  have  this  result.  Unless  a sufficient  num- 
ber of  trained  entomologists  are  available  to  meet  these  new  problems,  Canada  could  be  in 
serious  trouble  in  the  future. 

The  Entomologist  in  Agricultural  Research 

Most  entomologists  in  the  Research  Branch  are  engaged  in  applied  research.  Although 
some  are  in  relatively  pure  research,  their  numbers  are  decreasing.  At  Lethbridge,  the  work 
on  insect  cold-hardiness  ended  when  R.  W.  Salt  retired.  The  Belleville  Institute  will  be 
closed  and  its  personnel  transferred.  The  taxonomic  work  of  the  Entomology  Institute  at 
Ottawa  has  been  more  closely  related  to  its  service  for  field  entomologists. 

The  research  of  the  Branch  has  been  reorganized  into  programs,  most  of  which  are  aimed 
at  the  production  of  agricultural  commodities.  Objectives  are  established  and  goals  are  set 
for  attainment  within  a specified  period  of  time.  The  programs  generally  include  a wide 
range  of  disciplines  particularly  at  the  larger  research  centres.  The  scientists  have  thus  been 
drawn  into  a true  interdisciplinary  approach  in  planning  and  conducting  research.  Despite 
early  misgivings,  we  find  that  we  are  obtaining  a better  appreciation  of  the  problems  in- 
volved, thus  providing  a more  rational  approach  to  research  planning. 

The  research  of  the  applied  entomologist  is  best  described  by  a few  examples.  In  the  pro- 
ject on  cutworms  at  our  Research  Station  at  Lethbridge,  work  is  proceeding  to  develop 
suitable  artificial  diets  and  environmental  conditions  for  mass  rearing  of  moths  for  phero- 
mone studies.  Concurrently,  the  testing  of  synthetic  pheromones  is  being  advanced  in  con- 
junction with  analyses  with  the  gas  chromatograph  and  mass  spectrometer  and  electro- 
antennal  response  tests.  Studies  of  the  pea  aphid  on  alfalfa  involve  the  collection  and  identi- 
fication of  the  species  complex  in  alfalfa  fields,  determination  of  the  roles  of  the  species 
involved,  and  the  effects  of  insecticides  on  parasites  and  predators.  Research  on  the  tech- 
nology of  the  manipulation  of  the  alfalfa  leafcutter  bee  is  accompanied  by  studies  on  behav- 
ior of  bumblebees  and  of  their  enemies,  the  cuckoo  bees.  The  resistance  of  new  wheats  to 
the  wheat  stem  sawfly  is  being  examined  along  with  determination  of  the  influence  of 
plant  hormones  and  quality  and  quantity  of  light  on  host  plant  resistance  and  larval  behav- 
ior. 

Although  I have  described  our  research  as  applied,  it  could  as  well  be  classified  as  research 
and  development,  or  even  as  applied  and  background.  Perhaps  it  defies  classification  because 
it  is  so  varied.  The  entomologist  in  agricultural  research  inevitably  becomes  involved  in 
activities  that  range  from  extension,  preparation  of  popular  articles,  training  post-doctoral 
fellows  and  undergraduates,  developing  ad  hoc  controls  for  unexpected  outbreaks  of  new 
pests,  participating  in  advisory  committees  on  insecticides,  conducting  field  surveys,  and 
developing  computer  programs,  to  keeping  up  with  the  scientific  literature,  designing  new 
experiments,  and  publishing  results  in  scientific  journals. 

The  primary  aim  of  entomologists  in  applied  research  is  to  develop  the  knowledge  and 
techniques  required  for  the  management  of  insect  populations.  To  accomplish  this  aim 
the  ideal  entomologist  should  possess,  in  addition  to  an  interest  in  research  and  insects,  the 


The  Professional  Entomologist 


35 


ability  to  determine  the  significant  questions,  the  ingenuity  and  imagination  to  design  expe- 
riments that  will  answer  these  questions,  the  willingness  to  discard  hypotheses  proven  un- 
tenable, and  the  intelligence  to  interpret  the  results  of  the  experiments. 

Educating  the  Entomologist  for  Applied  Research 

Now  that  I have  described  some  of  the  work  and  characteristics  of  the  entomologist  in 
applied  research,  all  that  remains  is  to  decide  how  to  educate  him.  Seriously  though,  his 
formal  education  should  enable  a reduction  in  the  amount  of  experience  required  before 
he  can  make  a significant  contribution  and  it  should  provide  a base  of  knowledge  that 
will  enable  a broad  selection  of  strategies  for  solving  research  problems. 

It  would  be  difficult  for  anyone  to  provide  a list  of  the  subjects  necessary  without  spend- 
ing a great  deal  more  time  than  I have.  Research  in  applied  entomology  involves  a range  of 
sciences  and  of  scientific  fields  that  is  probably  broader  than  that  for  any  other  single 
modern  scientific  specialty.  The  main  branches  of  the  seven  major  sciences  total  88  (3). 
Applied  entomology  involves  a knowledge  of,  or  at  least  an  association  with,  32  of  them. 

The  research  entomologist  should  have  a basic  grounding  in  the  classical  fields  of  ento- 
mology, with  a general  knowledge  of  insect  behavior  and  ecology  probably  ranking  first. 
He  must  be  able  to  identify  rapidly  many  of  the  major  families  of  insects  and  possess 
instant  recognition  of  a large  number  of  the  more  common  species.  He  should  be  knowledge- 
able in  the  properties  of  insecticides  and  with  the  methods  and  equipment  of  their  applica- 
tion. He  must  have  a thorough  knowledge  of  how  to  find  and  search  the  relevant  scientific 
literature. 

Knowledge  of  statistical  design  and  analysis  of  experiments  is  essential  and  a thorough 
grounding  in  mathematics  related  to  the  study  of  ecology  would  be  valuable  if  not  essen- 
tial. He  should  take  chemistry,  physics,  and  economics.  Plant  taxonomy  and  ecology,  verte- 
brate and  invertebrate  zoology,  and  animal  ecology  would  be  important  additions.  A course 
in  instrumentation  would  be  excellent  and  computer  science  would  be  valuable.  Exposure 
to  most  of  the  other  agricultural  sciences  should  be  required  for  the  agricultural  entomolo- 
gist, with  a similar  requirement  in  forestry  for  the  forest  entomologist. 

Unless  he  has  already  taken  French  or  a foreign  language,  the  student  will  usually  benefit 
little  from  such  training  in  his  graduate  program.  A foreign  language  should  not  be  auto- 
matically required  for  an  advanced  degree.  It  would  be  far  better  to  concentrate  on  English 
composition  and  grammar.  Only  the  rare  scientist  can  write  with  clarity  and  simplicity. 
Most  spend  long  and  painful  hours  attempting  to  write  good  scientific  papers  and  presenta- 
tions, and  few  are  able  to  write  decent  popular  articles.  The  theses  that  I have  read  convince 
me  that  professors  who  act  as  major  advisors  deserve  hardship  pay  on  the  one  hand,  and  on 
the  other,  demerit  in  their  pay  for  allowing  the  production  of  what  is  all  too  often  a dull, 
verbose,  and  pretentious  composition  of  interminable  length.  But  perhaps  times  have 
changed  since  I wrote  my  thesis.  It  would  be  preferable  if  the  theses  were  written  as  a 
scientific  paper  that  had  to  be  published  in  a properly  refereed  journal.  Although  the  schools 
have  failed,  the  universities  share  the  failure  by  allowing  its  students  to  graduate  with  only 
a rudiment  of  the  skill  of  communication.  If  the  proper  courses  for  writing  and  editing 
scientific  and  popular  articles  do  not  exist,  the  department  should  insist  that  they  be 
created. 

The  Future  Entomologist 

Entomological  research  of  the  next  two  or  three  decades  will  be  similar  to  that  of  today 
because  our  knowledge  of  the  behavior  and  ecology  of  many  insects  of  economic  signifi- 
cance is  still  insufficient.  For  example,  the  flea  beetles  that  annually  damage  a variety  of 


36 


Holmes 


our  crops  remain  unidentified  even  to  species,  and  their  behavior  and  habits  are  almost  un- 
known. Bierne  ( 1 ) has  listed  314  species  or  species-groups  of  insects  that  have  been  recorded 
as  damaging  annual  crops  in  Canada.  He  covered  only  three  of  the  27  orders  of  insects  and 
did  not  deal  with  perennial  crops,  ornamentals,  stored  products,  animals,  households,  and 
forests,  nor  the  beneficial  species.  It  is  obvious  that  the  list  of  all  insects  of  direct  importance 
to  Canadians  includes  many  hundreds  more.  The  five  species  that  appear  annually  as  new 
pests  of  crops  also  add  to  the  list.  Even  with  many  more  entomologists  it  would  take  years 
before  we  could  expect  to  know  all  that  is  required  of  even  50  percent  of  these  species. 

Although  cotton  insects  in  the  U.  S.  have  received  a great  deal  of  attention  over  the 
years,  only  seven  years  ago  an  investigative  committee  decided  that  little  of  the  essential 
research  had  been  done  on  them.  In  Canada,  orchard  pests  have  been  studied  intensively, 
yet  a recent  research  planning  meeting  on  these  pests  listed  1 1 major  research  areas  still  re- 
quiring attention  including  sampling  techniques,  rearing  methods,  methodology  for  use  of 
pheromones,  determination  of  crop  losses,  and  the  search  for  parasites  and  predators. 

Advances  have  been  made  in  the  laboratory  in  the  identification  and  synthesis  of  phero- 
mones and  other  compounds  such  as  hormones.  But,  as  with  other  new  or  potential  methods 
of  insect  control,  the  entomologist  is  the  only  one  who  can  test  these  advances  in  the 
field  and  develop  the  proper  methods  for  their  application.  The  lack  of  adequate  research 
by  entomologists  and  others  led  us  into  the  difficulties  associated  with  the  application  of 
insecticides,  and  we  should  try  to  avoid  a similar  omission  with  these  new  techniques. 

The  professional  entomologist  of  the  future  will  work  more  closely  with  specialists  in 
other  scientific  disciplines  in  his  attempts  to  develop  integrated  controls.  He  will  be  re- 
quired to  work  with  economists  to  generate  more  data  on  the  losses  caused  by  insects, 
and  to  determine  the  cost-benefit  ratios  of  his  research  and  the  control  measures  he  recom- 
mends. He  will  have  to  develop  methods  for  accurately  forecasting  outbreaks  as  well  as  for 
determining  the  levels  of  population  that  require  control.  He  will  have  to  cooperate  in 
studies  to  determine  the  effects  of  his  control  measures  on  the  biosphere. 

The  University  Department 

The  department  of  entomology  has  a responsibility  as  a professional  department  as  well 
as  in  the  liberal  arts  context.  At  the  University  of  Alberta  this  is  acknowledged  by  placing 
the  department  in  the  Faculty  of  Agriculture  and  Forestry  and  by  listing  entomology  as  a 
major  in  the  Faculty  of  Science.  In  view  of  this  dual  role  I would  suggest  that  the  depart- 
ment re-examine  its  1970  objectives. 

Because  of  its  responsibility  in  applied  science,  the  department  should  ensure  that  its 
staff  will  continue  to  include  members  that  have  had  professional  experience  as  entomolo- 
gists. This  opinion  is  based  on  the  hypothesis  that  such  members  are  more  likely  to  know 
the  courses  that  are  best  suited  for  positions  outside  the  university.  Professional  experience 
enables  the  professor  to  draw  on  his  own  experience  for  counselling  students  and  for  the 
provision  of  personal  anecdotes  to  embellish  his  lectures.  Additional  benefit  will  be  derived 
from  the  addition  to  the  department  of  a point  of  view  not  usually  acquired  in  the  univer- 
sity environment.  The  university  is  such  a distinctly  separate  subculture  that  selection  of 
faculty  solely  from  those  who  have  spent  all  of  their  life  in  the  academic  environment  is 
likely  to  make  it  difficult  for  the  university  to  understand  the  needs  of  society  and  vice 
versa. 

The  department  should  encourage  the  student  who  is  interested  in  research  to  seek 
work  at  a research  establishment  during  the  summer.  This  would  provide  the  student  with 
a sample  of  applied  research,  and  exposure  to  different  attitudes  concerning  the  objectives 
of  research.  The  student  who  intends  to  pursue  graduate  studies  with  the  purpose  of  getting 


The  Professional  Entomologist 


37 


a job  in  applied  research  would  be  well  advised  to  take  one  or  two  years  off  to  work  in  such 
research  before  proceeding  to  his  terminal  degree.  This  should  assist  him  in  selecting  the 
appropriate  courses  and  in  determining  whether  or  not  he  is  suited  to  the  job. 

Students  may  wish  to  pursue  a field  of  entomology  primarily  of  academic  interest  or  of 
limited  potential  for  future  employment.  The  professor  concerned  should  discover  the  ulti- 
mate goal  of  the  student  and  advise  him  accordingly.  It  is  a sincere  tribute  to  any  profes- 
sor that  he  can  make  his  own  field  so  interesting  that  many  students  wish  to  emulate  him. 
Unfortunately  for  the  student  interested  in  a career,  not  all  of  the  interesting  fields  are 
needed  outside  the  university. 

Despite  the  importance  of  ecology  in  entomology,  it  is  my  impression  that  there  are  few 
if  any  universities  in  Canada  noted  for  their  expertise  in  this  field.  The  field  of  insect  cold- 
hardiness is  now  vacant  even  though  it  should  be  an  important  area  of  study  in  a country 
with  our  climate.  Another  orphan,  although  certainly  less  appealing  scientifically,  is  the 
study  of  insects  that  inhabit  households. 

Canada  will  require  substantial  numbers  of  entomologists  in  the  relatively  near  future.  In 
the  past  we  relied  on  the  U.  S.  to  provide  most  of  the  graduate  training  of  entomologists 
but  Canadian  universities  should  be  able  to  carry  more  of  this  load.  Departments  that  are 
part  of  an  applied  science  faculty  should  be  best  suited  to  fulfill  this  role. 

In  Summary 

The  Department  of  Entomology  of  the  University  of  Alberta  has  a long  history  of  excel- 
lence and  experience  in  the  education  of  entomologists,  most  of  whom  probably  ended  up 
in  applied  research.  Indeed  most  of  the  entomologists  at  the  Lethbridge  Research  Station, 
including  myself,  are  graduates  of  the  Department.  The  Crop  Entomology  Section  and  the 
Department  had  a common  progenitor  in  E.  H.  Strickland,  and  close  contact  has  been  main- 
tained for  the  past  50  years.  A recently  retired  member  of  the  Section  was  the  acting  Head 
of  the  Department  for  two  winters.  It  would  not  be  surprising  therefore,  if  the  ideas  that 
I have  presented  differ  little  from  those  already  considered  by  the  Department.  Perhaps 
the  major  difference,  if  any  exists,  is  in  the  attitude  toward  the  purposes  and  goals  of 
entomology. 

Research  is  becoming  more  difficult.  Many  of  the  easier  problems  have  been  solved. 
Entomologists  of  the  future  will  have  to  be  better  educated.  They  will  have  to  work  close- 
ly with  scientists  of  other  disciplines,  and  they  will  have  to  be  more  imaginative  in  their 
approach  to  research.  If  entomology  is  to  continue  as  a strong  discipline,  entomologists 
will  have  to  educate  people  outside  of  the  discipline  as  to  the  importance  of  our  research. 
The  department  is  in  the  ideal  position  to  perform  this  role. 

The  public  is  becoming  increasingly  reluctant  to  support  science  without  the  promise  of 
social  benefit.  The  measurable  benefit  from  entomology  is  likely  to  be  demonstrated  only 
by  its  application.  Unless  the  department  makes  the  error  of  embarking  on  an  extensive 
program  of  applied  research  on  its  own,  it  will  have  to  rely  on  the  success  of  its  graduates  in 
order  to  obtain  public  support.  The  success  of  applied  entomology  in  turn  relies  on  the 
education  provided  by  a strong  department  that  continues  to  maintain  the  basic  fields  of 
entomology.  Applied  entomology  has  been  and  will  continue  to  be  important  to  society. 
Departments,  to  serve  society  and  to  ensure  their  own  strengths,  have  to  be  concerned 
with  providing  the  most  suitable  education  for  applied  entomologists. 

Departments,  as  well  as  entomologists  in  general,  will  also  have  to  concern  themselves 
with  the  new  Ministry  of  State  for  Science  and  Technology.  The  Ministry  is  likely  to  have 
an  important  voice  in  the  agreements  on  financing  universities  that  will  be  re-negotiated 
within  two  years  between  the  federal  and  provincial  governments.  Currently,  the  federal 


38 


Holmes 


government  provides  50%  of  the  operating  costs  and  over  70%  of  the  research  grants  to 
the  universities.  The  Ministry  has  already  indicated  that  it  expects  to  play  a major  role  in 
decisions  of  government  departments  and  in  planning  the  science  programs  of  universities 
(Science  Forum  4:16-18,  1971).  The  practical  course  will  be  to  determine  what  we  think 
the  future  needs  for  entomology  are  likely  to  be  and  to  ensure  that  our  scientific  and  pro- 
fessional associations  present  these  views  to  the  Ministry.  This  should  be  done  within  the 
next  two  years. 

The  Department  of  Entomology  of  the  University  of  Alberta  deserves  credit  for  taking 
the  highly  unusual  step  of  asking  entomologists  from  outside  of  the  University  to  offer 
suggestions  and  criticisms  on  the  education  of  the  entomologist.  While  major  changes  may 
not  be  indicated  or  made,  an  exchange  of  ideas  between  the  Department  and  outside  sources 
at  this  symposium  should  benefit  all  concerned. 

REFERENCES 

1.  Bieme,  B.  P.  1971.  Pest  insects  of  annual  plants  in  Canada.  Memoir  No.  78,  Entomol. 

Soc.  Canad. 

2.  Glen,  Robert.  1956.  Entomology  in  Canada  up  to  1956.  Canad.  Entomol.  85:290-371. 

3.  Margenau,  Henry  et  al.  1964.  The  Scientist.  Time  Inc.,  New  York. 

4.  Willis,  T.  G.  1964.  Agricultural  Research  and  Development  Survey.  Canad.  Dept.  Agric. 

Admin.  Branch. 


39 


INDUSTRY  VIEWS  ENTOMOLOGY 

G.  S.  Cooper 
Cyanamid  of  Canada  Ltd. 

Rex  dale,  Ontario 

Everyone  has  experienced  events  in  early  childhood,  that  remain  forever  imprinted  in 
his  mind.  Living  through  the  grasshopper  invasions  around  Medicine  Hat  and  Lethbridge 
during  1920,  *21,  and  ’22  was  such  an  experience  for  me.  It  was  my  first  major  encounter 
with  an  insect  pest,  and  the  sight  of  crops  disappearing  almost  overnight  will  never  be  for- 
gotten. Grasshoppers  were  everywhere,  and  were  even  eating  the  cotton  clothing  hanging 
on  clothes-lines  and  the  bark  from  the  unpeeled  fence  posts  of  barbed  wire  fences.  In 
1922,  about  the  only  green  material  that  survived  in  some  areas  was  Russian  thistle,  and  in 
many  cases  this  was  harvested  for  feed  for  the  coming  winter.  It  was  during  this  period  that 
I had  the  good  fortune  to  meet  my  first  entomologist,  E.  H.  Strickland.  I was  very  young 
at  that  time,  and  was  very  impressed  with  the  meeting,  even  though  I am  certain  that  this 
feeling  was  not  mutual.  At  a much  later  date  that  first  meeting  and  its  impact  on  me  were 
discussed.  It  was  through  Professor  Strickland  that  the  first,  large  chemical  control  program 
in  Alberta  was  started.  I was  to  spend  many  hours  spreading  poisoned  bait  for  grasshopper 
control,  and  well  remember  the  instructions  on  how  to  handle  the  bran  bait. 

Professor  Strickland  was,  to  me,  a practical,  economic  entomologist.  I have  always  felt 
that  it  was  his  approach  to  insect  control  that  provided  the  encouragement  necessary  to 
establish  a sound  pesticide  industry  in  Western  Canada.  The  wide-scale  application  of  arseni- 
cals  and  strychnine  provided  incentive  to  the  suppliers  of  pesticides.  During  this  period, 
two  other  economic  entomologists  also  gave  support  to  the  young  industry.  Both  Mr.  R. 
(Dick)  Painter  and  Dr.  H.  E.  Gray  had  a marked  impact  on  the  industry  through  then- 
work  on  chemical  control  of  insect  pests. 

Following  the  war,  I enrolled  in  the  Faculty  of  Agriculture  of  the  University  of  Alberta 
rather  than  in  that  of  Education,  a field  which  I had  been  following  prior  to  the  war.  In 
my  first  year  I had  the  privilege  of  taking  my  first  course  in  economic  entomology  from 
Professor  Strickland.  To  those  of  you  who  did  not  have  the  good  fortune  of  attending  his 
classes  may  I state  that  you  missed  a great  and  inspiring  experience.  He  was  a dedicated  and 
superb  teacher,  a first-rate  practical  economic  entomologist,  and  had  developed  to  a very 
high  degree  that  rare  commodity  — a sense  of  humour. 

Lack  of  a sense  of  humour  could  be  responsible  for  some  of  the  problems  we  presently 
face.  If  one  recalls  the  highly  successful  economic  entomologists  of  the  past,  this  one  trait 
is  found  in  all  of  them.  As  the  years  have  passed,  a marked  change  in  entomologists  and  in 
entomology,  has  occurred.  Polarization  now  exists  in  both.  The  economic  entomologist  no 
longer  has  anything  in  common  with  the  taxonomist,  the  physiologist,  etc.  and  today,  it  is 
difficult  at  times  to  realize  they  are  all  part  of  entomology.  There  has  been  so  little  interest, 
cooperation,  and  communication  between  the  entomologist  interested  in  the  biological 
control  of  insects  and  the  entomologist  working  with  chemicals,  that  at  times  each  has  pro- 
claimed that  his  was  the  only  meaningful  work.  Rarely  has  there  been  any  attempt  to 
consider  both  areas  in  proper  perspective.  This  fragmentation  of  entomology  by  entomolo- 
gists has  been  detrimental  to  all. 

For  too  long  entomology  has  not  been  an  active,  integrated  part  of  biology.  Little  interest 
has  been  shown  by  entomologists  in  considering  their  contributions  in  relation  to  those  of 
other  biologists  and  little  interest  in  relating  these  to  the  needs  of  agriculture.  Entomology 
and  entomologists  have  tended  to  remain  aloof  from  other  associated  disciplines  and  as  a 


40 


Cooper 


result,  must  now  make  a concerted  effort  to  break  out  of  the  web  in  which  they  find  them- 
selves trapped.  In  the  world  of  today  and  tomorrow,  the  entomologist  must  become  an 
active  participant  in  applied  biology  if  we  are  to  protect  our  environment,  economy,  and 
social  customs. 

Just  as  happened  in  the  period  prior  to  1950  when  chemical  control  became  almost  the 
only  love  of  the  economic  entomologist,  so  the  period  following  the  early  1 950’s  initiated 
the  trend  to  basic  research.  While  basic  research  is  essential  and  will  always  be  so,  it  is 
unfortunate  that  it  is  now  implemented  to  such  an  extent  that  it  deters  the  applied  entomo- 
logist. Today,  industry  employs  very  few  entomologists  as  there  are  very  few  good,  applied 
entomologists  available.  At  the  same  time,  there  is  little  opportunity  for  the  employment 
of  basic  research  entomologists  in  the  pesticide  industry  in  Canada.  There  is,  in  fact,  a mark- 
ed shortage  of  applied  entomologists  employed  by  governments  too,  and  I am  aware  of  two 
positions  that  are  not  being  filled  because  of  this  shortage.  Indications  are  that  this  shortage 
will  become  more  acute  in  the  next  few  years,  and  this  can  only  result  in  serious  problems 
for  the  future.  More  and  more  major  insect  problems  are  developing  as  we  move  into  the 
“Green  Revolution,”  and  yet  we  are  not  solving  these  problems.  It  is  time  we  returned  to 
the  concept  of  cooperation,  and  worked  together  in  developing  a well-balanced,  integrated 
program  of  insect  control  that  will  benefit  our  entire  society.  The  time  for  “class  distinc- 
tion” between  entomologists  is  over;  each  segment  is  essential,  each  must  be  kept  in  balance, 
and  each  must  contribute  and  be  given  equal  recognition  for  the  part  it  plays  in  producing 
programs  that  are  economically  sound  and  give  maximum  protection  to  our  environment. 

In  the  early  days  of  entomology,  the  universities  offered  courses  that  both  introduced 
the  subject  broadly  and  emphasized  practical  implications.  One  has  only  to  review  the 
names  of  noted  entomologists  occupying  important  positions  in  Canada’s  past,  to  find 
that  in  most  instances  their  basic  training  was  in  economic  entomology.  In  the  past,  gradu- 
ating entomologists  had  far  more  exposure  to  other  areas  of  biology  than  many  get  today. 
It  may  be  true  that  most  of  these  courses  were  then  offered  by  the  Faculty  of  Agriculture, 
but  faculties  of  science  were  involved  as  well.  Today,  in  many  universities,  “entomology”  is 
no  longer  being  taught. 

During  the  past  four  or  five  years,  I have  interviewed  many  graduates  applying  for  posi- 
tions in  our  company.  All  too  often  I find  that  their  training  has  been  restricted  to  one 
small  phase  of  entomology  and  even,  at  times,  to  one  specific  aspect  of  one  specific  insect. 
Although  they  had  taken  a course  in  general  entomology,  they  admitted  having  little  or  no 
interest  in  insects  other  than  the  ones  on  which  they  had  specialized.  This  type  of  speciali- 
zation might  have  been  acceptable,  provided  the  applicant  had  received  an  adequate  ground- 
ing in  basic  entomology,  but  this  was  apparently  lacking.  In  several  instances,  I have  indi- 
cated to  the  applicant  that  a broader  knowledge  of  biology  and  chemistry  is  essential  if 
employment  with  industry  is  expected.  A closer  check  on  the  quality  of  graduating  students 
also  indicated  that  many  of  our  universities  have,  as  their  main  interest,  the  number  of  stu- 
dents enrolled,  rather  than  the  quality  of  the  students  or  of  the  courses  they  teach.  In 
attempting  to  take  maximum  advantage  of  grants  available  for  student  support,  universities 
have  ceased  being  selective  when  enrolling  students.  Instances  are  available  of  universities 
encouraging  every  student  to  continue  on  to  higher  degrees,  regardless  of  the  ability  they 
have  shown.  It  would  appear  that  little  attempt  has  been  made  to  encourage  students  to 
enter  fields  in  which  employment  opportunities  are  available,  although  this  may  be  partially 
due  to  a lack  of  understanding  of  such  needs  or  a failure,  on  the  part  of  student  advisors,  to 
be  far-sighted  when  counselling.  Many  recent  graduates  are  inflexible  and  seem  to  think  that 
a university  degree  is  the  key  to  instant  success.  The  phrase:  “I  was  encouraged  by  my 
professor  to  accept  the  fact  that  an  advanced  degree  would  assure  me  of  employment”  has 


Industry  Views  Entomology 


41 


been  too  often  repeated  by  job  applicants.  Graduating  students  should  certainly  be  made 
aware  of  the  significance  and  need  of  advanced  training,  but  it  should  also  be  emphasized 
to  them  that  obtaining  such  a degree  alone  is  not  the  complete  answer.  They  must  be  versa- 
tile, adaptable,  tolerant,  and  realistic  in  their  approach  to  employment.  The  entomological 
departments  of  universities  and  the  entomological  sections  of  biological  departments  have  a 
very  important  role  to  play  in  the  future  training  of  entomologists.  There  must  be  broader 
exposure  to  other  biological  sciences,  greater  selectivity  in  accepting  candidates  for  advanced 
degrees,  and  more  awareness  of  the  immediate  and  long-range  needs  for  entomologists 
trained  in  the  various  facets  of  entomology.  Only  in  this  way  can  we  keep  the  supply  of  and 
the  demand  for  entomologists  in  balance,  and  avoid  the  serious  employment  problem  that  is 
facing  us  today. 

Entomology  should  be  taught  in  public  school,  in  high  school,  and  in  university  at  the 
undergraduate  level,  to  all  students.  A quick  check  of  schools  in  the  Province  of  Ontario  last 
year  revealed  a disturbing  ignorance  of  insects  on  the  part  of  teachers,  a failure  of  teachers 
to  use  insects  in  biological  studies  and  in  fact,  a tendency  of  teachers  to  create  an  adverse 
reaction  to  anything  connected  with  insects.  Entomophobia  seems  to  be  increasing  because 
of  this  widespread  ignorance  of  insects.  Lately,  there  has  been  a revival  in  superstitions  and 
fables  regarding  insects  that  date  back  to  the  middle  ages.  A larger  percentage  of  young 
people  seem  to  fear  insects  than  was  true  a few  years  ago.  One  contributing  factor  to  this 
present  ignorance  and  fear  of  insects  is  the  failure  of  entomologists  to  communicate  with 
the  public. 

When  one  reviews  today’s  curricula  in  universities,  elementary  schools,  and  high  schools, 
it  is  apparent  that  economic  entomology  and  general  entomology  are  no  longer  present. 
Compare  the  programs  of  the  Department  of  Entomology,  University  of  Alberta,  from  1922 
to  1936  with  those  of  1970,  and  the  changes  that  have  been  made  become  apparent. 

I have  been  very  critical  these  last  few  minutes,  and  this  has  been  deliberate.  While  we  in 
industry  realize  the  important  role  that  the  entomologist  has  played  in  the  success  of  the 
chemical  industry  in  the  past,  we  also  feel  that  there  is  a need  for  change  in  the  future.  More 
economic  entomologists  must  be  .graduated  if  the  major  agricultural  problems  of  the  imme- 
diate future  are  to  be  solved.  Chemical  control  of  pest  insects  is  essential  now  and  will 
remain  so  for  many  years.  The  answer  to  our  insect  pest  problems  surely  lies  in  our  accep- 
tance, development  and  implementation  of  integrated  control  programs.  Full  cooperation 
by  all  entomologists  is  essential.  Let  us  get  back  to  teaching  more  general  and  economic 
entomology  before  specialization  begins. 


42 


DISCUSSION  - AFTERNOON  SESSION 

Leader  — D.  H.  Kavanaugh 
Department  of  Entomology 
University  of  Alberta,  Edmonton,  Alberta 

First  of  all  I would  like  to  thank  our  speakers  on  behalf  of  all  of  us.  I would  especially 
thank  Dr.  Mason  for  discriminating  between  the  terms  “need,”  “demand,”  “luxury”  and 
“necessity.”  It  is  clear  from  what  has  been  said  that  our  profession  today  is  diseased.  The 
outstanding  symptom  of  the  affliction  has  been  repeatedly  described  today  in  graphs  and 
words:  namely,  that  entomologists  are,  using  Dr.  Mason’s  terminology  again,  “necessary” 
and  yet  not  in  demand.  We  are  confronted  today  then  with  the  job  of  diagnosing  the 
disease.  Once  we  have  succeeded  in  doing  this  we  should  begin  to  formulate  some  treat- 
ment for  it.  If  we  succeed  in  this,  then  perhaps  we’ll  have  something  which  will  assist 
entomology  departments  in  Canada  and  the  U.  S.,  and  particularly  our  department  here, 
in  understanding  the  present  situation  and  possibly  even  in  planning  for  the  future.  The 
papers  presented  today  covered  a great  variety  of  points  all,  I believe,  very  relevant  to 
the  problems  at  hand.  Numerous  suggestions  have  been  put  forth  which  should  stimulate 
discussion.  In  my  attempts  to  relate  the  various  topics  discussed  to  one  another,  I felt 
the  same  frustration  that  others  have  mentioned  throughout  the  week.  I found  it  useful 
to  jot  the  main  points  down  on  a flow  chart  (Fig.  1)  and  I would  like  to  use  this  to  outline 
what  I think  we  are  facing.  I realize  that  all  of  you  would  probably  organize  your  thoughts 
a little  differently  but  I hope  that  this  approach  will  promote  rather  than  stifle  the  discus- 
sion to  follow. 

Roman  numeral  I is  the  Entomology  Department,  the  focal  point  for  this  symposium. 

Roman  numeral  II  is  financial  support.  I think  we  will  include  in  this  category  our  symp- 
toms — the  fact  that  we  have  lots  of  entomologists  coming  along,  that  we  have  a need  for 
them  but  no  place  for  them  to  go.  Related  to  this  topic  under  Roman  numeral  II  we  have 
the  sources  of  support.  Under  support  I would  include  not  only  funds  but  jobs.  We  have 
three  sources  of  these:  the  public,  government  and  industry  which  narrow  down  to  one,  the 
public  at  large.  Dr.  Mason  has  aptly  presented  the  position  of  some  of  us  with  regard  to  job 
support.  The  other  speakers  today  have  provided  us  with  a good  deal  of  insight  into  what 
kind  of  support  we  can  expect  and  into  what  we  will  have  to  do  if  we  want  to  get  more. 

Under  Roman  numeral  III,  I have  listed  what  I consider  to  be  three  goals  of  an  entomol- 
ogy department.  Their  first  responsibility  is  to  educate  professionals.  Here,  a big  problem 
seems  to  be  relevance  — I use  that  term  in  the  Madison  Avenue  sense  — getting  the  back- 
ground knowledge  necessary  to  engage  in  interdisciplinary  research.  Dr.  Cooper’s  talk  had 
surprises  for  me  since  I was  given  a little  indication  of  what  he  was  to  say  beforehand.  He 
changed  much  of  it  — for  the  better  I think.  A lot  of  what  he  had  to  say  is  directly 
applicable  to  this  responsibility  of  educating  professionals. 

The  second  goal  of  an  entomology  department  is  to  educate  teachers  - not  only  univer- 
sity educators  but  primary  and  secondary  school  teachers  too.  Special  attention  was  paid 
to  this  topic  by  Mr.  Jantzie  and  especially  by  Dr.  Gittins.  I hope  we  will  hit  the  topic 
hard  in  our  discussion  because  it  is  related  to  many  of  our  problems. 

Lastly,  we  have  a responsibility  to  educate  the  general  public.  I would  suggest  that  we 
have  two  problems  here.  First,  we  have  to  educate  young  people.  We  can  do  this  by  taking 
care  of  some  of  our  other  goals,  namely  by  getting  some  good  teachers  into  the  school  sys- 
tem at  all  levels.  Secondly  and  perhaps  most  difficult,  will  be  to  reach  the  people  who 
control  the  money;  I call  them  the  “post-schoolers.” 


Discussion 


43 


HI 


Flow  chart  to  and  from  a department  of  entomology. 


Roman  numeral  IV  I have  headed  Programs  and  Curricula.  The  goals  I indicated  for  an 
entomology  department  are  only  achieved  through  some  sort  of  plan  — through  specific 
programs  and  curricula  as  emphasized  by  Dr.  Freitag  and  Dr.  Gittins.  The  distinction  made 
by  Dr.  Gittins  between  the  goals  of  graduate  and  undergraduate  programs  is  very  impor- 
tant. 

My  fifth  point  doesn’t  fit  into  my  scheme  very  well.  It  involves  channelling  students 
into  different  specialties  during  their  educational  process.  Here,  I include  the  screening  of 
students  to  find  out  whether  or  not  they  are  really  dedicated  and  to  learn  to  what  they  are 
dedicated.  We  may  have  to  resort  to  these  procedures  to  handle  the  imbalance  now  existing 
between  students  available  and  jobs  available. 

Next,  I would  like  to  consider  faculty,  a topic  pretty  well  skirted  in  this  symposium  so 
far.  In  this  context  I would  like  to  pose  a few  questions.  First,  what  sort  of  staff  should 
an  entomology  department  have  in  light  of  its  goals?  At  present  faculty  members  must 
be  teachers,  researchers,  and  administrators.  Have  we  left  room  for  our  best  teachers?  We 
certainly  have  left  lots  of  room  for  our  best  researchers!  Can  an  individual  whose  forte 
is  teaching  keep  a job  in  a university  today?  The  system  requires  that  he  carry  out  research 
at  the  same  time  that  he  is  teaching.  What  does  this  drain  on  his  time  do  to  his  teaching 


44 


Discussion 


potential?  Dr.  Cooper  suggested  that  there  are  other  aspects  of  teaching  that  could  be 
better  investigated,  i.e.  that  there  are  activities  outside  the  classroom  just  as  or  more  impor- 
tant than  those  going  on  inside. 

Finally,  is  the  public  really  getting  the  education  it  needs  from  the  entomology  depart- 
ment? Here  I mean  the  public;  not  industry  nor  government.  To  link  this  all  together,  and 
I have  indicated  this  on  my  chart  by  dotted  arrows,  we  have  to  communicate.  Communi- 
cation is  a problem  in  any  group  situation.  Looking  at  the  different  parts  of  my  scheme, 
and  we  all  know  this  from  our  own  experience,  we  can  see  the  problem:  there  is  little 
communication  between  sections.  Perhaps  this  is  really  the  disease  I referred  to  earlier. 
This  lack  of  communication  is  the  disease  we  have  to  cure.  All  day  we  have  been  hearing 
about  this  lack  of  communication  between  entomology  departments,  and  entomologists, 
and  the  public,  government  and  industry  but  this  communication  must  be  a two-way  street. 
Hopefully,  this  symposium  has  been  a step  in  the  right  direction.  Intrauniversity  and  inter- 
departmental communication  needs  to  be  increased  and  encouraged  as  well. 

Before  opening  the  floor  for  discussion,  I would  like  to  say  a few  things  about  the 
public  as  a separate  entity.  The  most  challenging  problem  we  face  as  scientists  and  educators 
is  communicating  effectively  with  the  public.  Those  who  supply  the  funds  and  jobs  for 
entomologists  are  not  available  to  educators  as  students;  you  can’t  get  them  into  a class- 
room to  have  at  them.  They  are  not  likely  to  pick  up  our  point  of  view  from  television 
either,  since,  even  if  we  had  presentations  on  T.V.,  they  would  be  watching  “All  in  the 
Family”  instead.  Maybe,  as  Dr.  Mason  suggested,  we  will  have  to  adopt  Madison  Avenue 
techniques  to  sell  ourselves  to  the  public.  For  is  it  not  pathetic  that  Madison  Avenue  can 
sell  luxuries  ...  as  necessities  when  we  can’t  even  sell  necessities  as  necessities? 

Until  Dr.  Cooper  did  so,  no  one  had  mentioned  one  additional  barrier  to  our  communi- 
cating with  the  public.  I refer  here  to  the  increasing  problem  of  entomophobia.  Entomo- 
phobia  isn’t  just  squeamishness,  it’s  a real  psychiatric  problem  and  I’m  sorry  to  see  that 
our  resident  psychiatrist,  Dr.  Schwab  has  left  for  the  day.  Entomophobia  is  an  irrational 
fear  of  insects.  I was  an  extension  entomologist  for  five  minutes  a few  days  ago  when 
Bruce  Stewart,  our  extension  expert,  was  out  somewhere.  A woman  came  in  with  a tiny 
little  insect  which  she  insisted  was  infesting  her  bed.  She  had  broken  out  in  a rash  and  was 
itching  all  over  the  place.  The  insect  she  showed  me  was  a collembolan,  yet  I could  not 
convince  her  that  it  was  harmless.  I’m  afraid  that  the  problem  of  entomophobia  will  not 
be  solved  by  education  alone.  It  will  require  the  integrated  efforts  of  psychologists,  sociolo- 
gists, psychiatrists  and  maybe  even  advertising  agencies. 

I would  now  like  to  open  the  floor  to  a general  discussion.  Dr.  Cooper  made  some 
stiff  accusations.  Let’s  answer  them. 

Craig  — Edmonton  — Dr.  Holmes  indicated  the  need  for  research  on  cold-hardiness  and 
household  pests.  Who’s  going  to  provide  the  funds? 

Holmes  - Lethbridge  — This  problem  of  obtaining  support  should  be  considered  clearly. 
We  can  talk  all  we  like  about  communicating  with  the  public;  but  the  public  does  not 
provide  us  with  funds,  at  least  not  directly.  In  the  federal  government  money  is  first  appor- 
tioned to  the  various  agencies.  From  there  it  is  passed  down  through  various  levels  of 
authority.  Eventually,  we  have  to  compete  with  people  in  other  disciplines  for  a certain 
amount  of  the  money.  In  other  words  priorities  are  established.  We  are  not,  as  entomolo- 
gists, able  to  increase  the  total  amount  of  money  coming  to  the  government;  but  we  can 
increase  the  percentage  of  this  amount  that  comes  to  us.  You  have  to  sell  your  research 
proposal  to  the  individuals  controlling  the  purse  strings. 

Gittins  — Moscow  — There  are  ways  of  getting  funds  without  involving  the  government  at 
all.  In  California  there  are  many  organizations  called  commodity  groups.  Many  of  these 


Discussion 


45 


organizations  have  funds  available  for  research  on  the  commodity  of  their  concern.  I am 
sure,  for  example,  that  Gord  Hobbs  at  Lethbridge  has  received  money  from  the  alfalfa 
seed  association.  The  sugar  beet  industry  in  southern  Alberta,  I am  sure,  has  such  a fund. 
Many  of  these  agencies  provide  money  for  research  in  insect  control.  Very  often  the  results 
can  be  lucrative  for  all  concerned. 

I would  like  to  support  Dr.  Cooper’s  plea  for  entomologists  to  work  together.  We  have 
gone  through  periods  when  “squirt  gun”  entomologists  were  considered  to  be  at  the  bot- 
tom of  the  totem  pole.  Let’s  get  away  from  this  - all  of  us  will  profit. 

One  positive  result  of  the  student  activism  of  the  last  few  years  is  this  word  “relevance.” 
In  many  of  our  universities  this  is  the  call  we  hear  now.  “Publish  or  perish”  is  being  re- 
placed by  “teach  or  travel”.  I think  this  is  great. 

Cooper  — Rexdale  — About  five  years  ago  we  at  Cyanamid  had  money  available  for  research 
on  household  insects  - but  no  one  was  interested  - there  was  no  demand  for  research  of 
this  type.  There  is  more  misuse  of  pesticides  in  the  household  than  anywhere  else.  No  one 
tells  the  housewife  that  a vacuum  cleaner  is  often  more  useful  in  controlling  insects  than 
the  flit  gun.  If  more  people  were  aware  of  the  problems  resulting  from  the  misuse  of  house- 
hold sprays,  funds  could  be  made  available  to  investigate  alternate  methods  for  controlling 
household  pests.  If  I go  to  a university  with  some  suggestions  for  research  on  household 
insects,  pray  tell  me  where  do  I go?  If  I do  find  a university  that  wishes  to  do  the  work,  I 
am  then  told  that  I am  not  allowed  to  put  any  strings  upon  the  work  which  is  to  be  done 
even  though  my  company  is  providing  the  money.  In  addition,  it  has  been  my  experience 
that  university  personnel  seem  to  think  that  a student  being  supported  on  an  industrial 
grant  will  become  contaminated  if  he  undertakes  a project  which  involves  pesticide  evalua- 
tion or  a study  of  termite  habits.  Why  have  the  universities  stigmatized  the  student  interes- 
ted in  applied  entomology? 

Pengelly  — Guelph  — Having  had  our  academic  tails  twisted  slightly,  I,  as  a representative 
of  academe,  would  like  to  clarify  our  position.  Dr.  Cooper  referred  to  the  poor  calibre 
of  student  he  is  getting.  If  he  thinks  the  students  graduating  now  are  mediocre,  he  should 
wait  a couple  of  years.  They  have  just  introduced  a system  at  the  University  of  Guelph 
called  “Pass  by  Course.”  If  you  take  50  courses  while  at  university,  and  pass  40  of  them, 
then  you  get  your  degree.  There  is  nothing  that  says  you  have  to  pass  Chem  1 00  before 
taking  Chem  200  or  400.  Why?  Our  all-wise  Senate  is  composed  primarily  of  “Arts”  types 
because  Wellington  College  (our  Faculty  of  Arts  and  Science)  has  the  largest  student  body 
on  campus.  These  are  the  people  swinging  the  cat  — and  their  current  educational  fad  is 
“doing  your  own  thing.” 

Dr.  Cooper  mentioned  sources  of  support  for  students  doing  research  in  integrated  con- 
trol. In  our  Department  of  Environmental  Biology  we  have  a staff  member  who  is  interested 
in  integrated  control.  He  was  quietly  told  that  research  in  integrated  control  of  apple  pests 
is  now  the  province  of  the  CDA  lab  at  Vineland.  Since  he  has  no  money  he  can’t  take  on 
a student  in  integrated  control. 

At  one  time  it  was  not  academically  respectable  in  most  universities  for  a student  to  sub- 
mit a thesis  on  the  use  of  pesticides.  In  fact  there  were  graduate  schools  that  refused  to 
accept  students  with  this  interest.  We  have  tried  to  get  students  to  work  in  applied  re- 
search but  they  just  said,  “Pooh.”  Dr.  Cooper  mentioned  that  when  he  wanted  an  applied 
entomologist,  he  couldn’t  get  one.  Well,  I am  sympathetic  because  I would  like  to  have 
one  or  two  as  graduate  students.  Our  educational  program  here  at  Alberta  was  a sensible 
one  and,  at  one  time  we  did  have  a sensible  one  at  Guelph.  If  you  came  into  the  depart- 
ment or  into  the  faculty  of  agriculture  you  at  least  got  one  course  called  introductory 
or  general  entomology.  One  or  two  people  took  on  the  challenge  introduced  by  this  course 


46 


Discussion 


and  eventually  became  entomologists.  Well,  at  Guelph,  through  judicious  maneuvering  of 
“zoologists”  we  ended  up  with  entomology  being  required  by  no  one.  Now  how  in  the 
world  can  we  interest  anyone  in  entomology  when  we  never  get  a chance  to  talk  to  him? 
At  Guelph,  if  you  are  enrolled  in  animal  husbandry,  you  don’t  have  to  take  a course  in 
livestock  insects.  The  same  is  true  of  students  in  crop  science,  and  so  on.  A student  must 
have  an  “Introduction  to  Microbiology,”  and  an  “Introduction  to  Zoology”  but  he  doesn’t 
have  to  have  an  “Introduction  to  Entomology.” 

Heming  - Edmonton  - Bob  Dixon,  with  the  Alberta  Department  of  Agriculture  was  won- 
dering why  it  was  we  didn’t  have  a speaker  from  the  provincial  government  presenting  the 
point  of  view  of  the  extension  entomologist.  His  point  is  well  taken.  We  probably  would 
not  have  had  time  to  put  him  on  but  something  that  has  come  through  loud  and  clear 
throughout  this  symposium  is  the  need  to  communicate  — and  an  extension  expert’s  job 
is  communication.  We  are  forewarned  next  time. 

Dixon  — Edmonton  — I created  13  jobs  this  year  for  students.  Five  students  are  working  on 
mosquito  biology  for  the  city  of  Edmonton;  one  at  Fort  Saskatchewan;  four  in  Calgary 
and  two  on  blackflies  for  the  provincial  government.  Three  students  were  turned  down. 
In  1957  there  was  only  one  entomologist  on  staff  in  the  Department  of  Agriculture.  By 
1962  there  were  two  and  by  1971  three  full-time  entomologists  with  the  department.  In 
addition,  one  entomologist  transferred  to  the  Department  of  the  Environment.  This  June 
I am  creating  another  position  for  an  entomologist  to  bring  our  total  to  four.  I suspect  that 
by  1973  there  will  be  need  of  an  additional  entomologist  in  the  Department  of  the  Environ- 
ment. I am  not  an  Albertan.  I received  my  training  in  Manitoba.  Alberta  entomology 
graduates  right  now  are  in  trouble.  The  Department  of  Entomology  at  the  University  of 
Alberta  does  not  have  any  applied  entomology  so  that  their  graduates  are  competing  for 
jobs  with  people  from  outside  the  province  who  do.  I have  had  to  reduce  the  level  of 
one  position  in  the  hope  of  getting  an  Alberta  graduate  to  fill  it. 

Freitag  - Thunder  Bay  — Mr.  Dixon,  are  the  entomologists  you  refer  to  M.Sc.’s,  Ph.D.’s 
or  undergraduates? 

Dixon  — The  students  I placed  were  undergraduates.  In  this  province  not  one  Alberta 
entomology  student  got  any  of  these  positions.  There  were  none  or  they  did  not  apply. 
Where  are  they?  The  other  positions  are  for  M.Sc.’s  and  possibly  Ph.D.’s  if  they  have  an 
applied  bent.  Otherwise  we  don’t  want  them  since  we  can’t  afford  to  train  them. 

Gooding  - Edmonton  - 1 am  not  looking  for  a job,  Bob,  but  I would  love  to  know  where 
these  jobs  are  being  advertised. 

Dixon  — I phoned  your  department  twice,  Ron.  1 got  no  response  from  the  Faculty  of 
Agriculture.  I did  get  responses  from  Physical  Education,  Zoology,  and  Education.  The 
professional  jobs  were  and  are  advertised  in  the  papers.  All  you  have  to  do  is  read  the 
classified  ads. 

Gooding  - We  have  only  one  undergraduate  majoring  in  entomology  and  he  is  employed 
elsewhere  for  the  summer.  Another  is  working  with  mosquitoes  for  the  summer  but  I’m 
not  sure  if  he  is  one  of  your  13.  He  has  four  assistants.  Surely  there  are  lots  of  undergradu- 
ates who  have  had  some  entomology.  There  is  a girl  working  for  the  city  of  Edmonton  at 
$600.00  a month  who  took  one  of  our  courses.  The  student  you  have  working  at  Fort 
Saskatchewan  took  a course  in  medical  and  veterinary  entomology.  Most  of  the  students 
you  have  with  you  have  had  some  entomology.  We  have  few  students  majoring  in  entomol- 
ogy when  undergraduates. 

Dixon  — This  is  the  point  I am  trying  to  make.  There  is  no  requirement  in  the  Faculty  of 
Agriculture  that  students  take  entomology,  even  though  all  agriculture  graduates  should 
have  at  least  an  introduction  to  the  subject.  We  have  60  district  agriculturalists  on  our 


Discussion 


47 


staff,  few  of  whom  have  any  entomological  background.  I credit  you  people  with  teaching 
the  ones  that  do.  This  is  true  too  of  the  students  from  other  faculties  who  we  have  em- 
ployed. Nevertheless,  where  are  the  entomologists?  We  can  create  jobs  but  we  have  to  have 
entomologists  to  fill  them. 

Gooding  — During  his  talk  Dr.  Cooper  referred  to  certain  aspects  of  our  entomology  curri- 
culum that  he  did  not  like.  What  doesn’t  he  like  and  what  would  he  like  to  see  added  or 
substituted? 

Cooper  — Too  many  of  your  courses  are  specialized  and  too  few  cover  introductory  and 
applied  entomology.  When  I was  a student,  everybody  in  agriculture  had  to  take  some  ento- 
mology. What  good  are  agriculture  graduates  to  us  if  they  have  had  no  entomology?  All  of 
our  chemical  salesmen  are  agriculture  graduates  but  we  cannot  find  enough  of  them  that 
have  had  any  entomological  exposure.  We  have  agriculture  graduates  that  specialize  in  engi- 
neering, crop  science,  animal  husbandry  . . . but  not  entomology.  In  looking  over  your 
number  of  professors,  students  and  grad  students,  there  are  few  in  what  I would  call  the 
economic  field.  I would,  therefore,  probably  have  to  take  a very  broad  interpretation  of 
what  you  are  calling  economic  entomology. 

Kavanaugh  — Dr.  Cooper,  you  have  just  told  us  what  you  would  like  to  see  in  an  agriculture 
graduate  with  a Bachelor’s  degree.  What  do  you  want  in  a postgraduate? 

Cooper  — We  want  a student  who  has  had  enough  general  and  applied  entomology  to  be 
flexible.  If  you  talk  to  a man  who  has  taken  nothing  but  taxonomy,  that  is  the  only  field  he 
is  interested  in.  All  I ask  is  that  he  have  a good  general  background  upon  which  he  can  build. 
If  he  has  only  taxonomy  and  physiology  he  is  of  no  use  to  us.  A student  should  have  at  least 
two  years  exposure  to  general  entomology  no  matter  what  his  specialty.  We  have  nothing 
against  taxonomists;  we  have  nothing  against  physiologists;  but  we  do  have  something 
against  the  taxonomist  who  has  so  little  general  background  that  we  must  re-train  him. 
Hocking  — Edmonton  — The  curriculum  of  the  Faculty  of  Agriculture  is  decided  by  demo- 
cratic process.  Many  years  ago  most  courses  required  of  all  students  were  eliminated  and 
with  them  entomology.  There  was  nothing  we  could  or  would  do  about  this  because  we  do 
not  believe  in  teaching  entomology  to  students  who  are  in  class  only  because  they  have  to 
be.  I would  like  to  list  the  applied  courses  we  offer,  since  there  seems  to  be  some  ignorance 
of  this.  There  are  three  courses  in  medical  and  veterinary  entomology,  two  in  forest  ento- 
mology, one  in  general  agricultural  entomology;  one  in  toxicology,  and  a project  course 
which  can  be  used  to  cover  aspects  of  applied  entomology  not  included  in  the  courses  I have 
mentioned.  We  have  six  research  projects  in  progress  on  applied  aspects,  all  of  these  in- 
volving at  least  one  graduate  student  as  well  as  faculty.  During  the  last  10  to  15  years,  one 
quarter  of  the  theses  produced  were  in  applied  fields. 

I would  like  to  thank  Dr.  Holmes  for  repeating  what  I have  been  saying  to  graduate 
students  for  years  about  the  writing  of  theses.  I have  also  said  it  to  the  chairman  of  the 
Department  of  English  here,  and  on  more  than  one  occasion  in  the  hope,  so  far  vain,  of 
getting  his  department  to  do  something  about  this.  All  I have  been  able  to  persuade  them 
to  do  is  to  intensify  their  instruction  in  such  fields  as  the  appreciation  of  the  writings  of 
Geoffery  Chaucer.  Most  students  in  entomology,  however,  already  have  a reasonable  appre- 
ciation of  some  of  his  works.  However,  we  have  had  a real  problem,  especially  with  graduate 
students  who  have  never  had  to  write  anything  other  than  the  yes’s  and  no‘s  required  in 
high  school  examination  papers,  in  getting  these  people  to  the  position  where  they  can  write 
a well  written  paper.  This  problem  we  have  tried  to  solve  ourselves  by  offering,  on  Saturday 
mornings  in  the  fall  term,  a course  in  scientific  writing.  I think  this  is  improving  the  situa- 
tion, but  it  takes  time. 

Dr.  Mason  showed  a graph  illustrating  Ph.D.  production  and  job  openings  in  science  and 


48 


Discussion 


technology  in  Canada.  It  did  not  show  this  information  for  entomology  and  for  the  whole 
world.  It  has  always  been  my  belief  that  entomology  is  international  and  should  be  treated 
as  such.  If  a similar  pair  of  curves  were  drawn  for  the  life  sciences  and  especially  for  ento- 
mology and  on  a world-wide  basis,  the  picture  would  be  a different  one.  It  certainly  was  in 
the  past  and  I predict  that  it  will  be  in  the  future. 

Shemanchuk  — Lethbridge  — Dr.  Hocking  emphasized  that  the  curriculum  of  the  Faculty 
of  Agriculture  here  had  been  derived  through  democratic  process.  I would  suggest  that  the 
curriculum  has  also  become  too  liberal;  that  the  student  is  allowed  too  much  freedom  to 
choose  to  take  only  what  he  feels  like  taking,  not,  necessarily,  what  he  needs.  Everybody 
knows  that  a little  entomological  knowledge  is  of  use  to  all  agriculturalists.  Surely  none  of 
us  would  go  to  a surgeon  for  surgery  if  we  knew  that  he  had  not  taken  surgical  procedure 
because  he  did  not  have  to  take  it  when  in  medical  school.  Therefore,  I refuse  to  endorse 
the  idea  that  a student  need  not  take  a course  because  he  doesn’t  want  to  do  so.  If  he  has 
committed  himself  to  a program  in  agriculture,  or  animal  science  etc.,  an  entomology  course 
should  be  mandatory. 

Evans  — Edmonton  — Dr.  Hocking  listed  the  applied  courses  that  we  offer.  Ecology  is  basic 
science.  The  textbook  I use  in  this  course  is  by  Andre wartha  and  Birch,  two  well  known 
economic  entomologists.  The  whole  book  is  a review  of  the  applied  entomological  literature. 
Morphology  is  often  considered  basic.  Our  extension  entomologist,  Bruce  Stewart,  studied 
the  sense  organs  of  the  red  turnip  beetle  as  a project  in  this  course.  Projects  in  physiology 
often  involve  the  use  of  mosquitos,  grain  beetles  and  cockroaches.  Thus,  a course  appearing 
basic  from  its  description  in  the  calendar  is  not  necessarily  so. 

Nelson  — Lethbridge  — I would  like  to  support  Dr.  Gittins  and  Dr.  Corbet  in  their  emphasis 
on  leaving  specialization  in  entomology  until  the  graduate  level.  I did  this  myself  and  found 
it  very  helpful. 

At  our  last  meeting  of  the  Entomological  Society  of  Alberta  a motion  was  made  that  we 
attempt  to  hold  a joint  meeting  with  the  Canadian  Society  of  Zoologists.  It  was  defeated. 
This  was  a shame  for  reasons  which  are  obvious. 

Mason  — Ottawa  — Dr.  Hocking,  in  answer  to  your  statements  on  my  graph,  I did  not  have 
figures  available  on  life  scientists  or  entomologists,  and  your  remarks  about  internationalism 
are  to  the  point.  In  scientific  research,  nationality  is  irrelevant;  in  employment  it  surely  is 
not.  People  who  hire  entomologists  do  look  at  your  citizenship.  Therefore,  the  fact  that 
there  are  entomological  jobs  in  France  or  Brazil  does  not  help  Canadians  one  bit.  These 
people  demand  French  or  Brazilian  citizenship  of  the  people  they  employ. 

Dolinsky  — Edmonton  — I am  the  latest  asset  of  the  Alberta  Department  of  Agriculture  — 
a newly-hired  supervisor  of  entomology.  I would  suggest  that  the  reason  we  are  not  getting 
the  type  of  people  we  need  out  of  our  universities  is  because  the  people  who  are  teaching 
there  are  not  teachers  but  researchers.  When  they  graduated  with  their  Ph.D.’s,  they  knew 
nothing  about  teaching.  Thus,  it’s  not  surprising  that  they  teach  only  their  own  little  field. 
They  know  nothing  of  cutworms,  flea  beetles,  red  turnip  beetles  or  clover  mites;  in  other 
words,  they  have  no  background  in  general  entomology.  I would  like  to  see  a course  in 
entomology  offered  at  a university  that  a person  could  take  to  learn  about  pest  problems  in 
his  own  country.  What  insects  are  forest  pests?  What  do  these  insects  look  like?  What  are 
their  life  cycles? 

I have  gone  to  university.  I have  taken  all  kinds  of  introductory  courses.  Yet,  when  I 
graduated  a couple  of  months  ago  I knew  nothing  about  the  behaviour  of  grasshoppers, 
about  flea  beetles,  about  Bertha  army  worms.  I came  here  with  a background  in  grain  pests. 
I knew  a little  about  grain  pests  and  household  pests.  But  when  you  get  into  a field  like 
extension  you  are  expected  to  be  a G.  P.  Every  time  you  get  up  to  talk  you  shake  because 


Discussion 


49 


you  are  aware  of  your  limitations.  Nobody  can  know  everything  about  everything.  The  CDA 
in  the  next  five  years  is  going  to  retire  a large  number  of  researchers.  Yet,  there  is  no  one 
coming  up  through  the  ranks  with  experience  of  Canadian  pest  problems.  Last  year  we  had 
an  outbreak  of  Bertha  armyworm  on  rape  but,  because  for  years  there  had  been  no  problem 
with  this  insect,  no  one  knew  what  to  do.  In  a few  years  we  are  not  going  to  have  any  grass- 
hopper experts  in  western  Canada.  Who  is  going  to  control  the  outbreaks  to  come? 
Heming  — Mike,  you  mentioned  earlier  that  we  in  the  universities  are  researchers  not 
teachers.  Since  coming  here  four  years  ago  I have  had  about  10  students  in  my  two  intro- 
ductory courses  who  were  majoring  in  education.  Seven  of  them  told  me  that  the  quality 
of  the  teaching  in  that  faculty  was  lower  than  in  any  other  they  had  experience  of.  Where 
are  we  supposed  to  learn  to  teach  — probably  not  in  the  faculty  of  education. 

Henri  Goulet  — Edmonton  — Taxonomists  are  always  the  ones  singled  out  as  narrow- 
minded. I have  been  a student  both  here  and  at  Macdonald  College.  I would  like  to  work  in 
applied  entomology  when  I graduate.  Students  in  entomology  here  do  get  a broad  back- 
ground. Dr.  Ball  insists  that  his  students  know  all  aspects  of  entomology  and  much  in  other 
fields  as  well.  I have  had  courses  in  basic  and  applied  entomology  and  in  botany,  zoology, 
anthropology,  and  geology  among  others.  I will  agree  with  you  that  there  are  narrow- 
minded entomologists,  but  few  of  them  graduate  from  this  department.  Dr.  Hocking  would 
not  allow  it.  Mike  Dolinsky  complained  about  the  lack  of  courses  on  specifically  Canadian 
pests.  I advise  him  to  get  in  touch  with  Macdonald  College.  They  offer  a superb  course  of 
this  type. 

O'Keefe  — Moscow  — We  are  talking  quite  frankly  to  each  other  today  and  this  is  good,  but 
we  are  also  polarizing  people  in  this  room  who  should  not  be  polarized.  I came  into  ento- 
mology after  having  received  my  B.Sc.  degree  in  agriculture;  I was  not  an  amateur  entomolo- 
gist as  a five-year-old  as  many  of  you  seem  to  have  been.  This  doesn’t  necessarily  mean  that 
I can’t  be  a good  entomologist.  I also  have  five  years  experience  in  extension  entomology  — 
fireline,  talk-to-the-people  type  of  entomology.  Until  about  1965  if  you  asked  an  entomolo- 
gist a question  you  got  an  answer,  right  or  wrong.  Since  then  this  has  ceased  to  be  true. 
Entomologists  won’t  answer  a question  now  unless  they  know  the  answer.  Yet,  there  are 
tasks  to  be  done  that  don’t  require  a lot  of  detailed  knowledge.  In  university  we  should  be 
learning  how  to  pose  questions  and  how  to  look  for  answers.  If  you  learn  to  do  this  properly 
you  should  have  no  problem  fitting  yourself  into  a new  job  or  situation.  Although  I have 
been  working  in  entomology  for  five  years  I have  never  had  a course  in  applied  entomology. 
I have  had  a commitment  to  agriculture  and,  more  recently,  one  to  entomology  and  biology. 
I am  having  no  trouble  in  my  job. 

Dr.  Mason  mentioned  that  we  had  to  adopt  Madison  Avenue  techniques  if  we  were  to  sell 
entomology.  This  isn’t  necessary.  If  you  talk  to  someone  who  is  having  trouble  with  insects 
he  wants  to  listen  to  you.  This  is  why  Edmonton  is  having  trouble  allocating  the  right 
amount  of  money  for  mosquito  control  — because  we  entomologists  are  afraid  to  go  down 
and  talk  to  them  not  knowing  all  the  answers.  If  we  had  the  answers  we  would  be  more  than 
willing  to  talk  to  them  and  they  would  be  willing  to  pay  us  to  listen.  I talk  to  people  about 
insects.  As  a result,  both  my  applied  entomology  and  my  insect  physiology  suffer,  but  I still 
take  the  time  to  talk.  As  a result,  it  helps  other  entomologists  because  I am  considered  to  be 
typical  of  other  entomologists  and  since  I take  the  time  to  talk  to  people,  others  probably 
do  as  well.  If  a person  comes  to  you  with  an  insect  problem  you  have  a chance  to  sell  the 
subject  of  entomology  to  him. 


50 


SUMMARY 
Morning  Session 

Rapporteur  - D.  R.  Whitehead 
Department  of  Entomology 
University  of  Alberta,  Edmonton,  Alberta 

Bruce  Heming  introduced  today’s  discussion  by  pinpointing  what  he  considers  to  be  the 
crisis  in  entomological  education.  Too  many  students  are  graduating  to  fill  too  few  jobs.  In 
the  eyes  of  the  public  our  product  is  overqualified  and  is  therefore,  paradoxically,  useless. 
To  the  modem  taxpayer  it  seems  that  science  creates  rather  than  solves  problems  and  tax- 
payer support  is  therefore  lacking.  We  must  therefore  re-evaluate  what  is  required  of  ento- 
mological education  and  we  must  also  re-educate  the  public  as  to  our  actual  goals.  The  goals 
of  this  meeting  are,  as  was  noted  again  later  by  Dr.  Gittins,  to  answer  these  three  questions. 
First,  what  is  the  proper  education  for  entomologists  of  all  kinds?  Second,  what  can  ento- 
mology contribute  to  education  in  general?  Third,  what  has  been  done  at  the  University  of 
Alberta  and  what  should  be  done? 

Our  first  speaker  this  morning,  Don  Jantzie,  gave  us  his  thoughts  as  a teacher  of  high 
school  biology.  He  lamented  that  entomology,  general  and  environmental,  is  not  treated  in 
sufficient  depth  in  high  schools.  Students  should  not  only  be  stimulated  but  should  have 
sufficient  background  to  make  value  judgements  when  they  become  taxpayers.  They  should 
further,  at  secondary  school  level  if  not  perhaps  much  earlier,  be  instructed  in  effect  in  what 
I would  call  insect  appreciation  or  relevant  entomology. 

Problems  in  entomological  education  at  secondary  levels  are  lack  of  adequately  trained 
teachers  and  lack  of  suitable  texts.  The  second  of  these,  promised  by  Dr.  Gittins,  is  about 
to  be  resolved.  The  first,  however,  seems  to  be  up  to  us.  We  need  lateral  education  at  univer- 
sity level  with  participation  of  faculty  of  education  students  in  entomology  programs.  And, 
entomologists  must  become  more  involved  with  secondary  education  at  teaching  and  curri- 
culum planning  levels. 

Rick  Freitag  gave  us  his  views  from  a base  at  a small,  mainly  undergraduate,  Canadian 
university.  He  modified  the  statement  of  crisis  in  entomological  education  given  earlier  by 
Dr.  Heming  to  one  that  university  enrolments  are  decreasing  and  job  placement  becoming 
more  difficult.  Science  enrolments,  however,  continue  to  increase,  complicating  the  problem 
of  job  placement.  Students  in  science  programs  seem  job  oriented  — only  about  10  to  15% 
seem  interested  in  education  per  se.  Some  students  drop  out  because  of  bleak  employment 
prospects,  some  prospective  students  do  not  even  enrol  for  the  same  reason,  and  some  opt 
to  take  further  training  at  more  vocational-oriented  institutions.  According  to  Rick  we 
should  prepare  the  entomology  or  biology  student  with  a broad  range  of  options  by  (1) 
continuing  surveys  designed  to  acquaint  academia  with  public  requirements,  (2)  offering 
enhanced  opportunities  for  undergraduates  to  experience  multi-disciplinary  research,  and 
(3)  insuring  at  both  undergraduate  and  graduate  levels,  opportunities  for  lateral  education. 
This  is  essentially  the  reverse  of  what  Mr.  Jantzie  suggested  but  completely  complementary. 
Jobs  of  the  future  may  not  now  be  foreseen  and  yet  provision  should  be  made  for  their 
fulfillment. 

Art  Gittins  gave  his  views  from  a base  that  is  somewhat  larger  — a U.  S.  Land  Grant 
University  which  I take  to  be  strong  in  both  undergraduate  and  graduate  programs.  He 
noted  that  while  Ph.D.  production  rates  in  the  U.  S.  nearly  tripled  from  1959  to  1969, 
these  Ph.D.’s,  though  still  urgently  needed  in  such  areas  as  environmental  quality  control 


Summary 


51 


and  pest  management,  are  no  longer  in  demand.  Entomologists  in  education  must  modernize 
and  redirect  the  total  curriculum  and  must  also  provide  both  public  and  government  with  a 
clear  awareness  of  the  need  for  entomologists.  At  university,  broadened  education  is  again 
called  for.  At  undergraduate  levels,  training  should  be  closely  allied  with  biological  science 
with  specialization  left  to  the  graduate  level.  Exceptions  may  be  when  undergraduate  train- 
ing is  terminal  perhaps  as  in  pest  management  entomology.  Courses  should  be  updated  to 
accommodate  today’s  more  sophisticated  high  school  graduate.  Above  all,  students  should 
be  made  aware  that  education  must  continue  after  graduation  through  their  professional 
lives. 

Graduate  level  education  should  be  re-examined.  Faculties  should  be  carefully  structured 
to  meet  modern  needs.  Students  should  be  exposed  to  new  technologies  as  they  become 
available.  Dr.  Gittins  asked  a number  of  interesting  questions.  Should  we  have  research- 
oriented  Master’s  level  degree  programs?  Should  we  not,  in  fact,  have  non-research-oriented 
Ph.D.  programs?  Should  not  superficial  foreign  language  requirements  be  eliminated  or 
replaced  by  requirements  in  computer  languages?  The  student  should  have  an  opportunity 
to  be  broadened  in  experienced,  not  channelled. 

Now,  to  get  at  the  three  main  questions  addressed  to  this  symposium,  I shall  attempt  to 
put  together  remarks  made  by  our  three  morning  speakers,  by  the  discussion  leader  Dr. 
Pengelly  and  by  members  of  the  audience. 

First,  “What  is  the  proper  education  for  entomologists  of  all  kinds?”  As  suggested  by  Dr. 
Pengelly,  entomology  should  be  clearly  defined  for  the  public,  including  the  government. 
This  is  up  to  us  as  entomologists.  Students  should  be  introduced  to  the  subject  at  an  early 
age,  perhaps  even  before  they  become  students.  We  must,  and  it  seems  we  are  about  to, 
produce  suitable  books  that  can  be  read  in  high  school  level  or  by  the  general  public.  Should 
the  orientation  of  training  be  vocational  or  educational?  Again,  there  seems  to  be  some 
disagreement.  It  was  suggested  first,  by  Dr.  Pengelly  I believe,  that  the  current  job  situation 
is  normal,  not  depressed.  Thus,  perhaps  continued  emphasis  on  educational  aspects  may  be 
desirable.  Dr.  Pepper,  however,  implied  that  professional  entomological  training  never  has 
been  relevant. 

Some  members  of  this  symposium  have  suggested  broadened  lateral  education  both  for 
entomologists  and  for  non-entomologists.  I gathered  that  there  is  not  total  agreement  with 
this  idea  though  it  seems  a good  one  to  me.  In  my  opinion,  which  perhaps  is  not  out  of 
place  here,  Rick  Freitag’s  suggestion  — increased  lateral  and  multidisciplinary  education  in 
entomology  — does  seem  appropriate. 

I think  Mr.  Gurba  may  have  alluded  to  the  central  problem,  in  providing  more  emphasis 
in  communication  in  entomological  training,  so  that  our  graduates  need  not  be  solely  re- 
search entomologists. 

The  second  question  was,  “What  can  entomology  contribute  to  education  in  general?” 
This  question,  really,  is  not  easily  distinguished  from  the  first  and  perhaps  should  not  be. 
Entomology  should  indeed  be  a part  of  education  in  general.  The  emphasis  again  is  on  com- 
munication and  we  as  entomologists  are  responsible  for  educating  the  public  and  the  govern- 
ment. 

The  final  question  was,  “What  has  been  done  at  the  University  of  Alberta  and  what 
should  be  done?”  If  my  memory  serves  me  correctly,  there  was  general  agreement  that  in 
Professor  Strickland’s  day  entomology  at  the  university  was  ideal.  It  was  suggested  by  some 
participants  this  morning  that  entomology  at  the  University  of  Alberta  should  return  to  the 
form  it  then  took;  indeed,  perhaps  if  enrolments  continue  to  decline,  entomology  here  may 
do  just  that  — but  as  a discipline  it  would  very  probably  soon  disappear.  Again,  this  question 
is  not  really  separable  from  the  first  and  may  be  similarly  answered.  It  would  seem  desirable 


52 


Summary 


that  future  students  have  broadened,  non-entomological  training,  especially  to  include  expo- 
sure to  and  communication  with  non-entomologists  including  professionals  in  other  fields 
and  with  the  public.  Please  bear  in  mind  that  this  is  this  morning’s  discussion,  however. 

I hope  that  I have  done  reasonable  justice  to  thoughts  given  by  those  involved  in  the 
preparation  of  the  product  and  will  now  turn  this  over  to  Martin  Chance  whose  responsi- 
bility will  be  to  summarize  views  of  the  users  of  the  product. 


Afternoon  Session 

Rapporteur  - M.  A.  C.  Chance 
Department  of  Entomology 
University  of  Alberta,  Edmonton,  Alberta 

Bill  Mason  began  with  what  he  hoped  was  a pessimistic  view  of  the  future  of  the  great 
systematic  biology  collections  in  Canada  and  the  U.  S.  With  static  financial  support,  system- 
atists  will  be  expected  to  cope  with  an  ever-increasing  inflow  of  new  specimens  and  an 
ever-increasing  number  of  requests  for  identification.  Eventually,  conditions  will  become 
such  that  identification  will  become  slow  or  impossible.  Large  portions  of  valuable  collec- 
tions may  be  destroyed  because  they  were  neglected  and  large  amounts  of  data  will  become 
unretrievable.  Dr.  Mason  points  out  that  to  avoid  this  dismal  future  those  of  us  interested  in 
the  maintenance  of  museums  must  agitate  for  increased  museum  support.  The  second  prob- 
lem, he  tells  us,  is  the  conservative  nature  of  the  taxonomists  themselves.  They  retain  old, 
time-consuming  nomenclatural  and  information  retrieval  methods  which  could  be  replaced 
by  a fast,  computerized  validating  service.  As  well,  with  the  help  of  computers,  much  of  the 
routine  identification  could  be  carried  out  by  technicians.  Will  the  computer  replace  the 
scientist?  Dr.  Mason  thinks  not.  The  computer  cannot  extract  new  data  from  unworked 
specimens  nor  can  it  replace  the  research  brain.  Is  the  taxonomist  likely  to  run  out  of  mate- 
rial? The  Insecta  are  not  only  the  most  numerous  of  all  living  creatures  but  are  also  the  least 
known.  Largely  due  to  expansion  of  Canadian  facilities  in  the  1960’s,  the  production  of 
Canadian  Ph.D.’s  now  far  exceeds  the  demand  for  them.  Continued  overproduction  is  a 
disastrous  policy.  Dr.  Mason  suggested  what  he  feels  are  two  reasonable  courses  of  action. 
First,  lowering  production  and  second,  convincing  employing  agencies  that  more  entomolo- 
gists are  needed.  The  first  has  been  unsuccessful  hitherto  as  can  be  seen  by  the  present 
employment  situation.  The  second  involves  salesmanship  — the  conversion  of  something 
thought  to  be  a luxury  into  a necessity  in  the  minds  of  those  with  enough  money  to  pay  for 
it.  The  popularity  enjoyed  by  environmentalists  and  conservationists  should  be  the  key  to 
this  sales  approach.  Dr.  Mason  thinks  the  university  should  be  in  advertising  but  also  that  all 
entomologists  should  be  involved.  Our  public  image  as  a group  must  be  improved. 

Neil  Holmes  began  by  indicating  that  the  CDA  has,  in  the  past,  been  the  largest  employer 
of  entomologists  in  Canada  and  that,  within  the  next  eight  to  13  years,  it  will  require 
60  to  80  new  entomologists.  This  demand  will  be  modified  by  the  capacity  of  entomolo- 
gists to  explain  the  importance  of  their  research,  by  additional  dramatic  outbreaks  of  pest 
species  such  as  that  of  the  Bertha  armyworm  last  summer,  and  by  public  reaction  to  the  use 
of  insecticides.  Dr.  Holmes  indicated  that  most  entomologists  in  agricultural  research  are  in 
applied  fields.  Those  involved  with  relatively  pure  research  are  decreasing  in  numbers.  There 
is  a general  trend  towards  programs  with  an  interdisciplinary  approach.  Whatever  the  re- 
search, the  agricultural  entomologist  becomes  involved  in  a wide  range  of  activities  directed 
to  the  management  of  insect  populations.  How  should  an  agricultural  entomologist  be 


Summary 


53 


trained?  Dr.  Holmes  says  that  of  the  88  main  branches  of  the  seven  major  sciences,  applied 
entomology  has  associated  with  it  32.  A wide  background  is  thus  essential.  The  study  of 
foreign  languages  should,  perhaps,  be  replaced  by  increased  emphasis  on  English  composi- 
tion and  grammar.  Theses  should  be  written  as  scientific  papers  to  be  published  in  refereed 
journals.  The  skills  of  communication  should  be  increasingly  developed.  The  entomologist  in 
the  next  two  or  three  decades,  Dr.  Holmes  feels,  will  carry  on  research  similar  to  that  of  to- 
day. Our  knowledge  of  behaviour  and  ecology  of  many  pest  insects  is  still  inadequate.  Many 
pest  species  remain  to  be  identified.  New  techniques,  such  as  the  use  of  pheromones,  must 
be  adequately  tested  to  avoid  past  mistakes.  Entomologists  will  integrate  their  efforts  more 
with  those  of  other  specialists.  Dr.  Holmes  feels  that  the  university  entomology  department 
should  include  staff  members  who  have  professional  experience  as  entomologists.  These 
members  might  better  understand  the  needs  of  society.  Students  interested  in  research 
should  be  encouraged  to  seek  work  at  research  establishments  during  the  summer.  Those 
entering  grad  studies  should  work  at  such  establishments  for  one  or  two  years  before  pro- 
ceeding to  terminal  degrees.  Such  experience  might  help  the  beginning  student  in  choosing  a 
research  proposal. 

George  Cooper  believes  that  there  is  a need  to  reintroduce  the  study  of  insects  into  public 
and  high  schools.  Little  is  done  at  these  levels  to  create  a favourable  image,  or  an  under- 
standing, of  insects.  There  is  probably  even  a tendency  to  create  an  adverse  reaction  to  in- 
sects at  this  level.  He  believes  that  entomologists  with  different  specialties  have  become  far 
too  fragmented  — far  too  isolated  from  the  activities  of  other  biologists.  Broad  biological 
background  no  longer  seems  to  be  provided  to  the  entomology  student.  The  trend  toward 
basic  research  is  so  extensive  that  applied  entomologists  are  sometimes  in  short  supply. 
Industry  and  government  need  applied  entomologists.  Dr.  Cooper  feels  that  there  is  a need 
for  a long  term  survey  to  determine  Canada’s  future  needs  for  entomologists.  With  this 
as  a guide,  the  universities  should  become  more  selective  when  taking  on  new  students. 
On-the-job  training  for  university  students  should  be  encouraged.  Dr.  Cooper  believes  that 
there  is  a need  today  and  that  there  will  be  a need  in  the  future  in  industry  for  entomolo- 
gists with  broad  backgrounds  in  biology  and  chemistry.  Today’s  graduates  do  not  fulfill 
industry’s  needs.  Advertisement  and  especially  improved  communication  may  overcome  this 
problem.  Dr.  Cooper  believes  there  is  a need  for  some  consideration  of  psychiatric  problems 
caused  by  insects.  There  is  also  a very  large  need  for  the  personal  counselling  of  students  by 
university  staff.  Dr.  Cooper  is  also  alarmed  by  the  lack  of  interest,  desire,  enthusiasm  and 
salesmanship  evidenced  by  university  staff  themselves. 

I will  try  and  sum  up  what  has  transpired  this  afternoon  in  four  points. 

1.  Both  public  and  government  should  be  made  aware  that  entomologists  are  necessary 
and  not  a luxury.  However,  the  activities  of  entomologists  should  not  be  directed  solely 
towards  solving  entomological  problems  arising  in  a free  enterprise  economic  system,  since 
some  of  the  assumptions  supporting  such  a system  are  questionable. 

2.  A favourable  view  of  insects  should  be  encouraged  at  all  levels  of  our  school  system. 

3.  Input  should  be  provided  for  graduate  schools  so  that  their  graduates  are  of  use  to 
those  hiring  entomologists.  I hope,  in  the  future,  that  we  won’t  again  have  as  much  negative 
feedback  from  employers  as  we  did  today. 

4.  A narrow  education  in  entomology  should  be  avoided  in  this  day  of  problems  requiring 
multidisciplinary  answers.  If  the  languages  of  chemistry,  physics  and  mathematics  have  too 
strong  an  industrial  accent,  then  we  should  develop  entomological  dialects  to  fill  our  re- 
quirements. 


54 


RESOLUTIONS 

B.  S.  Heming 

Department  of  Entomology 
University  of  Alberta,  Edmonton,  Alberta 

This  has  been  a long  day.  I think  you  will  be  surprised  to  see  how  small  a package  every- 
thing has  salted  down  to.  We  have  only  two  resolutions  resulting  from  today’s  activities.  Be- 
fore presenting  them  I’d  like  to  thank  Brian  Hocking  and  Philip  Corbet  for  helping  the  rap- 
porteurs and  me  to  put  them  together.  The  first  resolution  is  entitled  “The  Need  for 
Communication.” 

WHEREAS,  There  is  a lack  of  appreciation  of  the  social  and  biological  significance  of  insects 
and  of  entomology,  especially  among  parents,  educators,  employers,  the  com- 
munications media,  and  governments  at  all  levels,  therefore 
Be  it  Resolved,  That  continuing  and  increasing  efforts  be  devoted  to  remedying  this  lack,  in 
particular  by:  1 . encouraging  children  to  regard  insects  as  friends  rather  than 
foes;  2.  emphasizing  the  increasing  value  of  living  insects  as  classroom  mate- 
rial, especially  for  illustrating  biological  principles;  3.  encouraging  the  im- 
provement of  communication  between  educators  and  employers;  4.  assisting 
the  media  to  present,  insofar  as  possible,  information  about  insects  and  ento- 
mological problems  that  is  authoritative  and  accurate;  and  5.  making  known 
to  governments  the  benefits  and  the  costs  to  man  of  insect  activities,  and 
thus  the  need  for  entomologists,  now  and  in  societies  of  the  future. 

The  second  resolution  pertains  to  the  lack  of  practical  experience  of  today’s  graduates 
in  entomology. 

WHEREAS,  Too  few  graduating  entomologists  have  adequate  background  in  applied  fields 
for  immediate  employment  in  them,  therefore 
Be  it  Resolved,  That  greater  efforts  be  made  by  all  concerned  to:  1.  facilitate  summer  em- 
ployment for  undergraduates  in  research  especially  in  these  fields,  and  2. 
make  additional  material  in  applied  fields  available  in  the  curriculum  both 
as  formal  courses  and  through  appropriate  invited  speakers. 

Are  there  any  questions? 

y/  yy 

Belicek  — Edmonton  — Entomology  should  be  made  compulsory  for  students  in  agriculture 
and  related  fields.  If  you  want  to  be  a physicist  you  have  to  take  math.  This  should  be  true 
also  of  biologists  and  agriculturalists  with  respect  to  entomology.  I suggest  that  we  include 
a resolution  to  the  effect  that  in  agriculture  and  other  pertinent  fields  an  entomology  course 
be  made  compulsory  for  graduation. 

Heming  - I would  like  to  hear  Dr.  Hocking  speak  on  this. 

Hocking  — Edmonton  — We  don’t  have  the  power  to  institute  it.  The  curriculum  in  agri- 
culture is  decided  by  agriculture  faculty  council  and  that  is  what  I meant  when  I said 
curriculum  is  decided  by  democratic  process  — not  by  democratic  process  among  the 
students  — but  by  faculty  council.  Faculty  council  eliminated  a large  proportion  of  required 
courses  some  years  ago  and  I see  little  prospect  of  convincing  them  to  change  their  minds. 
Corbet  — Waterloo  — In  relation  to  that  last  point,  Mr.  Chairman,  we  seem  to  have  identified 
a serious  deficiency  among  agriculture  graduates,  namely  that  they  can  graduate  in  agricul- 


Resolutions 


55 


ture  and  not  have  been  exposed  to  entomology.  Since  the  present  mechanism  within  univer- 
sities is  not  competent  to  correct  this,  I think  it  would  be  appropriate  to  see  how  it  can  be 
corrected  and  I suggest  that  one  way  towards  this  might  be  for  the  employers  to  take  a very 
stern  line  here  and  make  it  clear  to  the  universities  that  they  don’t  consider  in  many  fields  a 
graduate  in  agriculture  as  being  trained  adequately  unless  he  or  she  has  had  entomology. 
Heming  — Do  other  people  here  agree?  I wonder  if  Dr.  Corbet’s  contribution  can  be  re- 
phrased as  a resolution?  Do  you  think  this  would  have  any  impact  on  the  powers  that  be  in 
our  faculty  council? 

Mary  Chance  — Edmonton  — Perhaps  we  should  include  it  as  a resolution  and  send  it  on  to 
the  Faculty  of  Agriculture.  Where  are  these  resolutions  going  anyway? 

Heming  — I have  to  admit  that  I have  given  little  thought  about  where  these  resolutions  are 
going  to  go.  They  are  to  appear  in  the  proceedings  of  this  meeting,  of  course,  but  I am  open 
for  additional  suggestions.  One  target  will  certainly  be  the  Faculty  of  Agriculture. 

Fredeen  — Saskatoon  — The  Agricultural  Institute  of  Canada  and  its  provincial  branches 
have  professional  standards  committees.  If  these  resolutions  are  worth  anything  they  should 
go  to  these  committees. 

Rowes  — Pinawa,  Manitoba  — I agree  with  Dr.  Corbet  that  we  should  get  potential  em- 
ployers involved.  I also  think  that  another  symposium  should  be  organized  involving  profes- 
sors and  employers  that  will  get  down  to  the  problem  of  course  details  etc.  What  does  the 
employer  need?  I suggest  that  an  additional  resolution  should  be  formulated  having  to  do 
with  such  a future  symposium. 

Gittins  — Moscow  — Something  additional  should  be  added  to  the  resolution  soon  to  come 
from  Dr.  Corbet’s  remarks.  Those  education  students  who  are  majoring  in  biology  and  who 
will  be  teaching  biology  at  the  elementary  and  high  school  level  should  be  included.  They 
too  should  be  required  to  take  entomology. 

Hocking  — I have  spent  the  last  few  minutes  framing  a third  resolution  dealing  with  the  last 
question  to  come  up.  I don’t  think  of  itself  it  will  do  much  good;  nevertheless,  if  there  is 
additional  support  brought  forward  from  other  directions  for  this  sort  of  thing,  it  just  could 
bring  about  some  changes. 

WHEREAS,  Knowledge  of  insects  is  considered  to  be  essential  in  the  work  of  all  agricultur- 
alists, foresters,  and  educators  in  the  life  sciences  and  desirable  in  the  work  of 
educators  in  all  sciences,  therefore 

Be  it  Resolved,  That  faculties  of  agriculture  and  forestry  be  requested  to  give  favourable 
reconsideration  to  a requirement  for  entomology  or  at  least  to  facilitate 
increased  enrolment  in  courses  in  this  field  and  faculties  of  education  be 
similarly  requested  in  respect  of  students  in  science  and  especially  biology 
majors. 

Moved  by  Evans,  seconded  by  Holmes  that  the  three  resolutions  be  accepted  in  principle. 
Passed  unanimously. 


56 


ENTOMOLOGY  AND  EDUCATION  - POSTSCRIPT  TO  SYMPOSIUM 

Unquestionably,  this  Symposium  has  achieved  a number  of  the  objectives  for  which  it  was 
organized:  it  has  given  us  a number  of  practical  suggestions  for  here  and  now  action,  the 
more  important  of  which  have  been  drawn  together  in  the  resolutions.  It  is  our  intention  to 
act  on  these  in  the  widest  possible  ways;  directly,  through  contributors  as  intermediaries 
where  appropriate,  and  more  indirectly  through  wide  distribution  of  these  proceedings. 

Some  of  us,  I think  organizers  and  contributors  alike,  had  other  problems  in  our  minds 
which  never  really  found  explicit  expression,  but  which  must  have  been  apparent,  by  impli- 
cation, to  most  sensitive  listeners.  Collectively,  these  may  perhaps  be  covered  by  the  expres- 
sion “Education  Sickness,”  which  would  include  the  current  wave  of  anti-education  feeling, 
declining  budgets  for  education,  declining  enrolments  and  mounting  drop-outs  at  schools 
and  universities.  These  problems  are  difficult  to  focus  on  and  have  so  far  proved  impossible 
to  solve;  Hoar  (1972)  aired  some  of  them  in  his  address  to  the  heads  of  departments  of 
biological  sciences  in  Canada  at  Montebello,  Quebec  in  November,  1971. 

To  me,  this  sickness  shows  some  of  the  symptoms  of  chronic  indigestion:  a constant  sense 
of  fullness  — of  the  head  with  knowledge  rather  than  the  stomach  with  food  — , a feeling  of 
revulsion  when  confronted  with  further  knowledge,  and  relief  obtainable  only  by  abstention 
and  the  passage  of  time  — accelerated  perhaps  by  exercise.  Since  wisdom  may  be  described 
as  digested  knowledge,  an  approach  to  a solution  to  these  problems  at  once  suggests  itself: 
a move  towards  balancing  the  knowledge  we  create  by  research  with  some  wisdom  digested 
from  it,  and  towards  teaching  the  wisdom  along  with  the  knowledge.  This  is  really  saying  no 
more  than  that  we  should  be  putting  more  emphasis  on  education  and  less  on  training. 
Education  - leading  out  — produces  wisdom  from  knowledge,  thus  curing  mental  indiges- 
tion. Training  facilitates  and  improves  the  use  of  knowledge,  mainly  by  repetition,  and  is 
thus  not  far  removed  from  rote  learning.  While  some  training  is  a necessary  part  of  educa- 
tion, training  without  education  is  not  only  possible,  but  perhaps  usual.  No  man  is  made 
wise  merely  by  training  his  memory.  That  we  talk  of  training  Ph.D.’s  is  tragic  if  this  is  really 
what  we  mean,  especially  since  training  has  as  its  only  raison  d'etre,  preparation  for  a 
specific  job. 

As  measured  by  the  printed  pages  recording  it,  scientific  knowledge  has  been  multiplied 
more  than  tenfold  in  the  last  century.  It  is  said  that  ninety  per  cent  of  the  scientists  who 
have  ever  lived  are  alive  today  — busily  producing  more  at  an  ever  increasing  rate.  The 
capacity  of  human  brains  to  absorb  knowledge  has  not  increased  correspondingly;  indeed 
one  may  question  whether  any  selection  pressure  for  this  still  exists.  Attempts,  in  this  situ- 
ation, to  continue  to  teach  all  that  is  known,  even  in  a narrowing  field,  may  make  a reality 
of  that  cynics’  specialist  — the  man  who  knows  everything  about  nothing.  Clearly  there  is  a 
glut  of  scientific  knowledge  and  it  is  no  wonder  its  price  in  the  market  place  has  declined, 
and  the  public  is  disillusioned  with  its  misapplications.  Though  entomology  is  a bit  of  a 
laggard  it  is  no  exception.  The  sins  of  technology  are  being  visited  upon  the  world  of  learn- 
ing and  both  knowledge  and  wisdom  are  threatened.  A man  who  has  taken  every  course  a 
single  university  department  offers  is  less  well  educated  than  one  who  has  taken  the  same 
number  of  introductory  courses  in  different  departments.  Wisdom,  though  not  wealth,  is 
more  readily  distilled  from  breadth  than  from  depth.  Knowledge  is  already  in  disgrace  but 
wisdom,  synthesis,  and  the  broad  view  must,  in  time,  prevail  over  that  self-destructive, 
narrow  nationalism  which  in  the  present  must  favour  insects  over  ourselves  and  in  the  future 
threaten  them  and  us  through  our  common  environment. 

The  insect  world  reflects  in  miniature  so  many  aspects  of  mankind  that  entomology  can- 
not lag  for  long.  In  education  the  insects  will  illustrate  more  biological  principles  for  a dollar 


Postscript 


57 


than  any  other  class  of  animal;  increasingly  so  as  our  pressure  on  the  environment  increases. 
And  after  education?  While  our  population  continues  to  grow,  insects  will  increasingly 
threaten  our  food,  fibre,  and  health;  when  it  shrinks,  as  it  soon  must,  we  shall  have  them  to 
learn  from  still. 

Hoar,  W.  S.  1972.  Educational  patterns  and  manpower  requirements  in  the  biological  sci- 
ences. Canadian  Soc.  of  Zoologists  Newsletter  3(4):  1-9. 


BRIAN  HOCKING 
University  of  Queensland 
Brisbane,  Queensland,  Australia 
June  1972 


58 

PARTICIPANTS  IN  ENTOMOLOGY  ANNIVERSARY  WEEK 
MAY  15-19,  1972 

Albert,  P.  J University  of  Saskatchewan,  Regina 

Anderson,  J University  of  California,  Berkeley 

Ashraff,  M.  A CIBA  - Geigy  (Canada)  Ltd.,  Winnipeg,  Manitoba 

Awram,  J Alberta  Department  of  Agriculture,  Edmonton 

Baldwin,  W.  F Atomic  Energy  of  Canada  Ltd.,  Chalk  River,  Ontario 

Bay,  E University  of  Maryland,  College  Park 

Belicek,  J University  of  Alberta,  Edmonton 

Bellamy,  R.  E Canada  Department  of  Agriculture,  Saskatoon,  Saskatchewan 

Bowden,  B Winnipeg,  Manitoba 

Bradbury,  W.  C Memorial  University  of  Newfoundland,  St.  John’s 

Brust,  R.  A South  Bend,  Indiana 

Burzynski,  A Pinawa,  Manitoba 

Campbell,  J.  B University  of  Nebraska,  North  Platte 

Chance,  M.  A University  of  Alberta,  Edmonton 

Chance,  M.  M University  of  Alberta,  Edmonton 

Chapman,  H.  C United  States  Department  of  Agriculture,  Lake  Charles,  Louisiana 

Cheung,  A University  of  Alberta,  Edmonton 

Clay,  M.  E Ohio  State  University,  Columbus 

Coffey,  M.  F Canadian  Armed  Forces,  Northern  Command,  Yellowknife,  NWT 

Cooper,  G.  S Cyanamid  of  Canada  Ltd.,  Rexdale,  Ontario 

Corbet,  P.  S University  of  Waterloo,  Waterloo,  Ontario 

Craig,  D.  A University  of  Alberta,  Edmonton 

Davids,  D.  E Defence  Research  Establishment  Suffield,  Ralston,  Alberta 

DeFoliart,  G University  of  Wisconsin,  Madison 

Depner,  K.  R Canada  Department  of  Agriculture,  Lethbridge,  Alberta 

Dixon,  R Alberta  Department  of  Agriculture,  Edmonton 

Dobson,  R.  C Purdue  University,  Lafayette,  Indiana 

Dolinsky,  M Alberta  Department  of  Agriculture,  Edmonton 

Downer,  R.  G.  H University  of  Waterloo,  Waterloo,  Ontario 

Downes,  J.  A Canada  Department  of  Agriculture,  Ottawa,  Ontario 

Duffy,  J.  R University  of  Prince  Edward  Island,  Charlottetown 

Dyke,  G.  R Northern  Alberta  Institute  of  Technology,  Edmonton 

Ellis,  R University  of  Manitoba,  Winnipeg 

Evans,  W.  G University  of  Alberta,  Edmonton 

Fallis,  A.  M University  of  Toronto,  Toronto,  Ontario 

Fettes,  J.  J Department  of  the  Environment,  Ottawa,  Ontario 

Fowler,  H.  W.  Jr Walter  Reed  Army  Medical  Center,  Washington,  D.  C. 

Frank,  R University  of  Guelph,  Guelph,  Ontario 

Fredeen,  F.  J.  H Canada  Department  of  Agriculture,  Saskatoon,  Saskatchewan 

Freitag,  R Lakehead  University,  Thunder  Bay,  Ontario 

Friend,  W.  G University  of  Toronto,  Toronto,  Ontario 

Gaunt,  R.  A Defence  Research  Establishment  Suffield,  Ralston,  Alberta 

Gillespie,  J.  M University  of  Idaho,  Moscow 

Gittins,  A.  R University  of  Idaho,  Moscow 

Gooding,  R.  H University  of  Alberta,  Edmonton 

Gorham,  J.  R Arctic  Health  Research  Center,  Fairbanks,  Alaska 


Participants 


59 


Goulet,  F.  E University  of  Alberta,  Edmonton 

Goulet,  H University  of  Alberta,  Edmonton 

Graham,  W.  M Lakehead  University,  Thunder  Bay,  Ontario 

Gray,  D.  P Canada  Department  of  Agriculture,  Sackville,  New  Brunswick 

Griffiths,  G.  C.  D University  of  Alberta,  Edmonton 

Griffiths,  J University  of  Alberta,  Edmonton 

Gurba,  J Alberta  Department  of  Agriculture,  Edmonton 

Hansens,  E.  J Rutgers  University,  New  Brunswick,  New  Jersey 

Harris,  C.  R Canada  Department  of  Agriculture,  London,  Ontario 

Haufe,  W.  O Canada  Department  of  Agriculture,  Lethbridge,  Alberta 

Hergert,  C.  R University  of  Alberta,  Edmonton 

Heming,  B.  S University  of  Alberta,  Edmonton 

Heming,  W.  E Guelph,  Ontario 

Hocking,  B . University  of  Alberta,  Edmonton 

Hollebone,  J.  R University  of  Alberta,  Edmonton 

Holmes,  N.  D Canada  Department  of  Agriculture,  Lethbridge,  Alberta 

Hooper,  R.  L University  of  Alberta,  Edmonton 

Hudson,  A Canada  Department  of  Agriculture,  Ottawa,  Ontario 

Hudson,  J.  E University  of  Alberta,  Edmonton 

Jantzie,  D.  N.  ^ County  Central  High  School,  Vulcan,  Alberta 

lessen,  J.  I University  of  Idaho,  Moscow 

Judson,  C.  L University  of  California,  Davis 

Kavanaugh,  D.  H University  of  Alberta,  Edmonton 

Khan,  M.  A Canada  Department  of  Agriculture,  Lethbridge,  Alberta 

Laird,  M Memorial  University  of  Newfoundland,  St.  John’s 

Lee,  R University  of  Alberta,  Edmonton 

Leech,  R National  and  Historic  Parks,  Ottawa,  Ontario 

Lefkovitch,  L.  P Canada  Department  of  Agriculture,  Ottawa,  Ontario 

Lewis,  D.  J Memorial  University  of  Newfoundland,  St.  John’s 

Lindsay,  I.  S Defence  Research  Establishment,  Ottawa,  Ontario 

MacGregor,  H.  A Edmonton,  Alberta 

Mason,  W.  R.  M Canada  Department  of  Agriculture,  Ottawa,  Ontario 

McClelland,  G.  A.  H University  of  California,  Davis 

McKague,  B British  Columbia  Research  Council,  Vancouver 

Mclver,  S.  B University  of  Toronto,  Toronto,  Ontario 

McLintock,  J.  R Canada  Department  of  Agriculture,  Saskatoon,  Saskatchewan 

Morrison,  P.  E University  of  Waterloo,  Waterloo,  Ontario 

Mulla,  M.  S University  of  California,  Riverside 

Neilson,  C.  L British  Columbia  Department  of  Agriculture,  Victoria 

Nelson,  W.  A Canada  Department  of  Agriculture,  Lethbridge,  Alberta 

Nuss,  J.  R Churchill  Falls  (Labrador)  Corp.  Ltd.,  Montreal,  Quebec 

O’Keeffe,  L.  E University  of  Idaho,  Moscow 

Orr,  D Lakehead  University,  Thunder  Bay,  Ontario 

Osgood,  C.  E Canada  Department  of  Agriculture,  Belleville,  Ontario 

Patterson,  J.  H Canadian  Wildlife  Service,  Edmonton,  Alberta 

Pengelly,  D.  H University  of  Guelph,  Guelph,  Ontario 

Pepper,  J.  H Bozeman,  Montana 

Peterson,  B.  V Canada  Department  of  Agriculture,  Ottawa,  Ontario 

Peterson,  L.  K Alberta  Department  of  the  Environment,  Edmonton 


60 


Participants 


Pilon,  J.  G University  of  Montreal,  Montreal,  Quebec 

Poinar,  G.  Jr University  of  California,  Berkeley 

Provost,  M.  W Entomology  Research  Center,  Vero  Beach,  Florida 

V.  v 

Pucat,  A.  M Champlain  Regional  College,  St.  Lambert,  Quebec 

Rai,  K.  S University  of  Notre  Dame,  Notre  Dame,  Indiana 

Rowes,  K.  D Pinawa,  Manitoba 

Ryan,  J.  K University  of  Alberta,  Edmonton 

Saha,  J.  G Canada  Department  of  Agriculture,  Saskatoon,  Saskatchewan 

Schwab,  E Edmonton,  Alberta 

Service,  M.  W Monks  Wood  Exp.  Stn.,  Abbots  Ripton,  Huntingdon,  England 

Shemanchuk,  J.  A Canada  Department  of  Agriculture,  Lethbridge,  Alberta 

Siddiqui,  F Northern  Alberta  Institute  of  Technology,  Edmonton 

Smallman,  B.  N Queen’s  University,  Kingston,  Ontario 

Smith,  J.  J.  B University  of  Toronto,  Toronto,  Ontario 

Sommerman,  K.  M Arctic  Health  Research  Center,  Fairbanks,  Alaska 

Stewart,  B.  D University  of  Alberta,  Edmonton 

Strong,  F.  E University  of  California,  Davis 

Taylor,  B.  W University  of  Manitoba,  Winnipeg 

Taylor,  D.  C Ottawa,  Ontario 

Thomas,  A.  W University  of  Alberta,  Edmonton 

Tidwell,  M.  A Louisiana  State  University,  Baton  Rouge 

Toblan,  V.  E Calgary,  Alberta 

Washino,  R.  K University  of  California,  Davis 

Watler,  D University  of  Alberta,  Edmonton 

Watson,  G Edmonton,  Alberta 

Weidhaus,  D.  E United  States  Department  of  Agriculture,  Gainesville,  Florida 

West,  A.  S Queen’s  University,  Kingston,  Ontario 

Whitehead,  D.  R University  of  Alberta,  Edmonton 

Wilkinson,  P.  R Canada  Department  of  Agriculture,  Lethbridge,  Alberta 

Zacharuk,  R.  Y University  of  Saskatchewan,  Regina 


Publication  of  Quaestiones  Entomologicae  was  started  in  1965  as  part 
of  a memorial  project  for  Professor  E.  H.  Strickland,  the  founder  of  the 
Department  of  Entomology  at  the  University  of  Alberta  in  Edmonton 
in  1922. 

It  is  intended  to  provide  prompt  low-cost  publication  for  accounts  of 
entomological  research  of  greater  than  average  length,  with  priority 
given  to  work  in  Professor  Strickland’s  special  fields  of  interest  including 
entomology  in  Alberta,  systematic  work,  and  other  papers  based  on  work 
done  at  the  University  of  Alberta. 

Copy  should  conform  to  the  Style  Manual  for  Biological  Journals 
published  by  the  American  Institute  of  Biological  Sciences,  Second 
Edition,  1964,  except  as  regards  the  abbreviations  of  titles  of  periodicals 
which  should  be  those  given  in  the  World  List  of  Scientific  Periodicals, 
1964  Edition.  The  appropriate  abbreviation  for  this  journal  is  Quaest.  ent. 
An  abstract  of  not  more  than  500  words  is  required.  All  manuscripts  will 
be  reviewed  by  referees. 

Illustrations  and  tables  must  be  suitable  for  reproduction  on  a page 
size  of  9%X6%  inches,  text  and  tables  not  more  than  13AX^3A  inches, 
plates  and  figures  not  more  than  8Mj  X5  inches.  Reprints  must  be  ordered 
when  proofs  are  returned,  and  will  be  supplied  at  cost.  Subscription  rates 
are  the  same  for  institutions,  libraries,  and  individuals,  $4.00  per 
volume  of  4 issues,  normally  appearing  at  quarterly  intervals;  single 
issues  $1.00.  An  abstract  edition  is  available,  printed  on  one  or  both 
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Communications  regarding  subscriptions  and  exchanges  should  be 
addressed  to  the  Subscription  Manager  and  regarding  manuscripts  to: 

The  Editor,  Quaestiones  Entomologicae, 
Department  of  Entomology, 

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