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UBC Theses and Dissertations

Comparative morphology of saldidae and Mesoveliidae (Heteroptera) and its bearing on classification Gupta, Ayodhya Prasad 1961

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COMPARATIVE MORPHOLOGY OF SALDIDAE AND MESOVELIIDAE (HETEROPTERA) AND ITS BEARING ON CLASSIFICATION AYODHYA PRASAD GUPTA B.Sc , Agra U n i v e r s i t y , 1950, M.Sc,, Banaras Hindu U n i v e r s i t y , 1953, F»R.EoS. A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE i n the Department of Zoology We accept t h i s thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA A p r i l , 1961 I n p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f t h e r e q u i r e m e n t s f o r an a d v a n c e d d e g r e e a t t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a , I a g r e e t h a t t h e L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and s t u d y . I f u r t h e r a g r e e t h a t p e r m i s s i o n f o r e x t e n s i v e c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y p u r p o s e s may be g r a n t e d by t h e Head o f my Department o r by h i s r e p r e s e n t a t i v e s . I t i s u n d e r s t o o d t h a t c o p y i n g o r p u b l i c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l n o t be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . D e p a r t m e n t o f ~Z-o o L O Q ^ The U n i v e r s i t y o f B r i t i s h C o l u m b i a , V a n c o u v e r 8, C a n a d a . Date ABSTRACT On the basis of h i s study of the female g e n i t a l i a , Scudder (1959) suggested that i n the Heteroptera-Hemiptera, the fa m i l i e s Saldidae and Mesoveliidae might be c l o s e l y r e l a t e d ; the present morphological study was undertaken to determine whether a study of other characters also supports t h e i r i n c l u s i o n i n a natural group. In these two f a m i l i e s , comparison?! of the head structures revealed that they are quite d i s t i n c t . The thorax revealed two types:-a Saldula type, and a Mesovelia type, and since i t i s shown that the structure of the thorax i s of l i t t l e taxonomic value i n dis t i n g u i s h i n g the s u p r a f a m i l i a l categories, i t was considered that the differences between the Saldidae and the Mesoveliidae need not n e c e s s a r i l y ind i c a t e a fundamental taxonomic d i f f e r e n c e . In the abdomen, the presence of the clasping organ i n the Saldidae completely separates t h i s family from the Mesoveliidae, The present study shows that the Saldidae and the Mesoveliidae are not c l o s e l y r e l a t e d as might be i n f e r r e d from comparisibns of the female g e n i t a l i a ; they are quite d i s t i n c t morphologically. The taxonomic p o s i t i o n of the two f a m i l i e s was also considered. Most a u t h o r i t i e s believe that the Mesoveliidae are appropriately placed i n the Amphibicorisae, and t h i s i s supported II by the present study. The p o s i t i o n of the Saldidae, on the other hand, has heretofore been very uncertain since t h i s f a m i l y shows some Pentatomomorph features as well as some Cimicomorph features. Two a l t e r n a t i v e s have been suggested i n t h i s t h e s i s regarding the systematic p o s i t i o n of the Saldidae. According to the f i r s t , the Saldidae may be considered a Cimicomorph,which branched o f f from the main stem of Cimicomorpha, and subsequently developed Pentatomomorph characters- an assumption which presupposes that p a r a l l e l evolution has occurred. A l t e r n a t i v e l y , the Saldidae may be considered a branch of the Pentatomomorpha, which arose a f t e r the evolution of some Pentatomomorph characters, but before the evolution of the complete Pentatomomorph complex of characters. This l a t t e r a l t e r n a t i v e takes cognisance of the f a c t that the Pentatomomorph complex of characters evolved gradually and not by a single ' s a l t a t i o n ' . I t has been concluded, however, that the data a v a i l a b l e at present are not s u f f i c i e n t to enable one to state which of the two a l t e r n a t i v e s mentioned above i s the c o r r e c t one, although I am i n c l i n e d to consider the former as the more p l a u s i b l e . In addition to the foregoing, two general aspects of the morphology of the Heteroptera were considered, namely the i n t e r p r e t a t i o n of the head s c l e r i t e s and the v a r i a t i o n i n the thoracic structure between apterous and macropterous forms of the two f a m i l i e s . i i r CONTENTS page 1- Introduction,, I 2- Material and Methods. 2 3- Morphology, 3 1- Head: A- Description: 1- Saldula sp, 3 2- Aepophilus bonnairei Sign. 4 3- Mesovelia mu Is a n t i White 4 4- Mesovelia v i t t i g e r a Puton 5 B- Discussion and Interpretation. 5 G- Comparison. I I 2- Thorax: 15 A- Description: 1- Saldula sp. 15 2- Aepophilus bonnairei Sign. 18 3- Mesovelia mulsanti White 19 4- Mesovelia v i t t i g e r a Puton 20 B- Discussion and In t e r p r e t a t i o n . 22 C- Comparison. 27 3- Abdomen: 30 A- Description: 1- Saldula sp. 30 2- Aepophilus bonnairei Sign. 32 3- Mesovelia mulsanti White 33 4- Mesovelia v i t t i g e r a Puton 35 IV B- Discussion and Interpretation. 35 C- Comparison. 36 4- Internal anatomical characters. 38 5- Comparison of alate and apterous (and brachypterous) forms i n Saldidae and Mesoveliidae. 41 6- Relationship between Saldidae and Mesoveliidae. 45 7- Systematic position of Mesoveliidae. 48 8- Systematic position of Saldidae. 49 9- Literature cited. 55 10- Keybto lettering of figures. 59 V LIST OF TABLES I- Table to show differences i n head structure. 13 I I - Table to show i n t e r p r e t a t i o n of veins i n the hind wing. 26 I I I - Table to show differences i n the thorax. 28 17- Table to show differences i n the abdomen. 37 V- Table to show taxonomic characters of Pentatomomorpha, Cimicomorpha, Amphibicorisae and Hydrocorisaeo 39 VI- Table to show differences between alate and apterous forms of Saldidae. 44 VII- Table to show differences between alate and apterous forms of Mesoveliidae. 44 VIII- Table to show taxonomic characters of Saldidae and Mesoveliidae. 46 IX- Table showing i n chronological order the views of various authors on the systematic p o s i t i o n of the S a i d i d a e . 51 VI LIST OF FIGURES F i g s , 1 - 3 Head of Saldula •. ( i ) dorsal ( f r o n t a l ) view ; (2) l a t e r a l view; (3) ventral view. F i g . 4 Head of Aepophilus bonnairei , dorsal view. F i g s , 5 - 7 Head of Mesovelia mulsanti : (5) dorsal view ; (6) l a t e r a l view ; (7) ve n t r a l view. F i g s . 8 - 1 4 Thorax of Saldula : (8) dorsal view ; (9) ventral view ; (IO) l a t e r a l view of prothorax ; ( i i ) l a t e r a l view of pterothorax; (12) i n t e r n a l view of : prothorax showing pleurodema and apodemes ; (13) i n t e r n a l view of mesothorax showing furca , phragmata and pleurodema ; (14) i n t e r n a l view of metathorax showing furoa , phragmata and pleurodema. F i g s . 15 - 17 Thorax of Aepophilus bonnairei : (15) dorsal view ; (16) ventral view ; (17) l a t e r a l view. F i g s . 18 - 20 Thorax of Mesovelia mulsanti: (18) dorsal view ; (19) ventral view ; (20) l a t e r a l view. F i g s . 21 - 23 Legs of Mesovelia mulsanti : (2l) fore leg ; (22) middle leg; (23) hind l e g . F i g s . 24 - 27 Thorax of Mesovelia v i t t i g e r a :(24) dorsal view; (25) ventral view; (26) l a t e r a l view of prothorax ; (27) l a t e r a l view of pterothorax. F i g s , 28 - 29 Wings of Mesovelia v i t t i g e r a : (28) fore wing: (29) hind wing. F i g s . 3© - 31 Wings of Saldula : (30) fore wing ; (31$ hind wing. vrr F i g s . 32 - 34 Legs of Saldula (32) fore l eg ; (33) middle leg j (34) hind leg . F i g s . 35 - 39 Abdomen and female g e n i t a l i a of Saldula :(35) dorsal •view of abdomen of male ; (36) ventral -view of female terminalia; (37) f i r s t gonocoxa and associated parts : (38) second gonocoxa and associated parts ; (39) spermatheca. F i g s . 40 - 44 Abdomen and female g e n i t a l i a of Mesovelia mulsanti : (40) dorsal view of abdomen of female ; (41) ventral view of female terminalia ; (42) f i r s t gonocoxa and associated parts ; (43) second gonocoxa and associated parts; (44) spermatheoa. F i g s . 45 - 46 , 50 Male g e n i t a l i a of Saldula: (45) l a t e r a l view of aedeagus and associated structures ; (46) f r o n t a l view of aedeagus and associated structures; (50) paramere. F i g . 47 Aepophilus bonnairei : l a t e r a l view of aedeagus and associated p a r t s . F i g s . 48 - 49, 51 Male g e n i t a l i a of Mesovelia mulsanti :(48) l a t e r a l view of aedeagus and associated parts ; (49) f r o n t a l view of aedeagus and associated parts; (5l) paramere. F i g s . A and B : showing probable systematic p o s i t i o n of Saldidae and Mesoveliidae, V I I T • ACMOWLEDGMENTS I wish to express my sincere thanks to Dr. G.G.E. Scudder fo r his constant help and encouragement during the progress of the work. I am also indebted to Dr. R.H. Cobben and Mr. I. Lansbury f o r supplying specimens of Mesovelia; the alate specimens of Mesovelia v i t t i g e r a were k i n d l y donated by The Musee Royal de L'Afrique Centrale ,Turvuren, Belgium. The majority of the work was done during the summer of I960 while i n receipt of f i n a n c i a l assistance from the National Research Council of Canada through Dr. G.G.E.Scudder. I I - INTRODUCTION: On the basis of h i s studies of the female g e n i t a l i a of the Heteroptera, Scudder (1959) suggested that the Mesoveliidae might be c l o s e l y r e l a t e d to the Saldidae. The present morphological study was undertaken to determine whether a study of other characters supports the i n c l u s i o n of these two f a m i l i e s together i n a natural group. Both the alate and the apterous forms of the two f a m i l i e s were studied and t h e i r morphology i s compared and discussed • 2 2 - MATERIAL AND METHODS: Saldula sp.* was collected from the beach at Point Grey, Vancouver, British Columbia. Aepophilus bonnairei Sign, from Jersey (Channel Islands) was kindly supplied by Dr. G.G.E. Scudder of the University of British Columbia, and Mesovelia mulsanti White from England and Holland were obtained from Mr. I . Lansbury (Hope Department of Entomology, Oxford) and Dr. R.H. Cobben (Wegeningen, Netherlands). The alate specimens of Mesovelia vittigera Puton were obtained from the Musee Royal de L'Afrique Centrale, Turvuren, Belgium, and were originally collected in the Congo. The external anatomy was studied both from dried and preserved specimens. The material was boiled in 10% potassium hydroxide, was passed through glacial acetic acid, stained in acid fuchsin, and cleared in creosote. Observations on the cephalic muscles of Saldula were made in preserved specimens. This material, which had been preserved in 10% alcohol, was passed through different grades of alcohol, cleared in xylene and examined under polarised light. Specimens of other families of the Heteroptera were collected locally, and were used for comparative study. A l l drawings were made by using a squared graticule eye piece, and are not to the same scale. * The specific name of this species has not yet been decided on by the experts. 3 3- MORPHOLOGY: I- HEAD:* A- DESCRIPTION: I- Saldula sp. ( F i g s . 1-3) Head short, but broad; vertex well developed; coronal and ecdysial cleavage l i n e s d i s t i n c t i n nymph, but not i n adult; frons not extensive; frontoclypeal sulcus absent; two invaginations present on each side of vertex; c l y p e a l region r e l a t i v e l y extensive, and d i f f e r e n t i a t e d i n t o postclypeus, anteclypeus and paraclypeus; cephalic p o r t i o n of postclypeus with two l a t e r a l unpigmented lobes (absent i n nymph)j paraclypeal lobes with unpigmented areas; bucculae well developed; postgenal bridge short; mandibular plate absent; mandibular lever well developed and t r i a n g u l a r ( i n nymph and ad u l t ) ; postocciput i n the form of t h i n r i n g around o c c i p i t a l foramen, and apparently d i f f e r e n t i a t e d i n t o dorsal and l a t e r a l elements, the l a t e r a l parts bearing paired condyles; labrum broad and f l a p - l i k e , reaching d i s t a l end of second l a b i a l segment; epipharyngeal process absent; labium f o u r - j o i n t e d , f i r s t segment the t h i c k e s t , t h i r d the longest, being swollen proximally and tapering d i s t a l l y ; antennae four-segmented with small intersegmental s c l e r i t e s , f i r s t segment much thicker than r e s t , the whole beset with h a i r s , the t h i r d and fourth segments also bearing stout b r i s t l e s ; eight to ten p a i r s of t r i c h o b o t h r i a present, scattered over vertex, frons, and postclypeus(nymph with four p a i r s , one p a i r on frons, two p a i r s on the postclypeus, and the fourth p a i r * The terminology of parts follows that of Snodgrass (i960). 4 on the anteclypeus); compound eyes oonspic^us; two o c e l l i present; two unpigmented areas present l a t e r a d of the o c e l l i . 2- Aepophilus bonnairei S i g n . ( F i g . 4) Similar to Saldula i n e s s e n t i a l parts, but d i f f e r i n g i n the following features: Cephalic p o r t i o n of the postclypeus without two l a t e r a l unpigmented areas; paraclypeal region not as well defined as i n Saldula, and r e s t r i c t e d to the upper two t h i r d s of the anteclypeus; m a x i l l a r y plate area well developed; bucculae not well developed; labrum reaching the d i s t a l end of the f i r s t l a b i a l segment, and not the second as i n Saldula; oompound eyes not conspicuous; o c e l l i absent; postoociput not divided i n t o dorsal and l a t e r a l elements; postgenal bridge longer than i n Saldula; four p a i r s of t r i c h o b o t h r i a -one p a i r on frons, two p a i r s on postclypeus and one p a i r on anteclypeus. 3 - Mesovelia mulsanti Tfhite.(Figs. 5 -7) Head longer than i n Saldula; vertex well developed, and overlapped by prothorax; coronal and f r o n t a l ecdysial cleavage l i n e s i n d i s t i n c t i n nymph and adult; fronto-clypeal sulcus absent; cl y p e a l region well developed, and d i f f e r e n t i a t e d i n t o postclypeus, anteclypeus and paraclypeus; maxillary plate region well developed and separated from the paraclypeal region by the short 5 genal s u l c u s ; lower l i m i t of m a x i l l a r y p l a t e area d e l i m i t e d by an i n d i s t i n c t l i n e ; bucculae not w e l l developed; mandibular p l a t e absent; mandibular l e v e r w e l l developed and roughly r e c t a n g u l a r ; labrum f l a p - l i k e , w i t h an epipharyngeal process extending almost t o the d i s t a l end of the second l a b i a l segment; postrenal bridge longer than i n S a l d u l a ; p o s t o c c i p u t i n d i s t i n g u i s h a b l e from o c c i p u t , and bearing two d o r s o - l a t e r a l condyles; labium f o u r - j o i n t e d , f i r s t segment the t h i c k e s t , t h i r d the l o n g e s t , being swollen on the inner side p r o x i m a l l y and t a p e r i n g d i s t a l l y ; antennae f o u r - j o i n t e d w i t h small intersegmental s c l e r i t e s ; " f i r s t antennal segment t h i c k e r than r e s t , and bearing a stout b r i s t l e l a t e r o - m e s a l l y ; three p a i r s of t r i c h o b o t h r i a , one p a i r on the frons l a t e r a d of the o c e l l i , and two p a i r s on the p o s t c l y p e u s ( i n the nymph one p a i r on the anteclypeus as w e l l ) ; compound eyes not very conspicuous; o c e l l i rudimentary. ^~ Mesovelia v i t t i g e r a Puton. S i m i l a r to Mesovelia mulsanti White but o c e l l i w e l l developed. B- DISGUSSIOH AND INTERPRETATION: S a l d u l a sp.: I n the- a d u l t i n s e c t , on the antero-dorsal p a r t of the head capsule, the p o s i t i o n of the f r o n t a l e c d y s i a l cleavage l i n e 6 can be recognized by the p o s i t i o n of two pits-; these are found on each side of the vertex along the f r o n t a l ecdysial cleavage l i n e i n the nymph. These two l a t e r a l p i t s are not homologous with the p r e t e n t o r i a of Homoptera (Cica d e l l i d a e ) (Spooner,1938). They have no counterparts i n other Heteroptera as f a r as i s known, and the name e p i c r a n i a l p i t s i s here suggested for these structures. I t should be mentioned that the frons i s not always delimited l a t e r a l l y by the ecdysial cleavage l i n e , f o r Snodgrass (i960) states that, " they vary g r e a t l y i n t h e i r extent and p o s i t i o n i n d i f f e r e n t i n s e c t s . " He suggests the name cephalic apotome fo r the part cut out at ecdysis by the ecdysial cleavage l i n e . In Saldula the f r o n t o - c l y p e a l sulcus i s absent, and consequently there i s no external demarcation between the frons and the clypeus. The p o s i t i o n s of the antafossae or the mandibular levers are often used as landmarks to delimit the f r o n t a l and the c l y p e a l areas (Spooner, 1938). However, i n Saldula sp., they are situated cephalad and so are perhaps u n r e l i a b l e . These areas can be distinguished, however, by muscle attachment; the d i l a t o r muscles of the sucking-pump are attached to the clypeus i n t e r n a l l y (Snodgrass,1935). Snodgrass(l960) states that the cibarium has often been c a l l e d the 'pharynx' although i t l i e s outside the mouth. He further states that t h i s cibarium has become the sucking-pump of the l i q u i d - f e e d i n g i n s e c t s . I t i s evident, therefore, that the term 'pharyngeal pump' i s inaccurate. In Saldula the c r i t e r i o n of the muscle attachment has been used to delimit the f r o n t a l and the c l y p e a l areas, although i t i s r e a l i z e d that t h i s 7 procedure i s open to some c r i t i c i s m . For example, F e r r i s (1944), DuPorte (1946) and Parsons (1959) have c r i t i c i s e d the c r i t e r i o n of muscle attachment f o r the i d e n t i f i c a t i o n of the s c l e r i t e s , the l a t t e r i n her account of Gelastoooris used the f r o n t a l ganglion as the landmark to del i m i t the point of attachment f o r the d i l a t o r muscle of the food-pump. She recognizes two sets of muscles - c i b a r i a l , which l i e a n t e r i or to the f r o n t a l ganglion and attach on the clypeus, and pharyngeal muscles, which l i e p o s t e r i o r to the f r o n t a l ganglion and attach on the frons. In t h i s she follows Marks (1959). Parsons (1959), however, suggests," that the muscle po s t e r i o r to the f r o n t a l ganglion could come to i n s e r t on a c i b a r i a l p o r t i o n of the food pump, or that c i b a r i a l muscles might s h i f t t h e i r • i n s e r t i o n s to the pharyngeal portion." She further quotes Marks (1959) and states that the p o s i t i o n of the f r o n t a l ganglion also varies r e l a t i v e to the muscles from one species to another. I t i s evident then that an i n t e r p r e t a t i o n based on the p o s i t i o n of the f r o n t a l ganglion i s also not r e l i a b l e . In the present study, therefore, i n the absence of any a l t e r n a t i v e , the c r i t e r i o n of muscle attachment has been adopted to d e l i m i t the f r o n t a l and the cly p e a l portions of the head capsule's Examination shows that the d i l a t o r muscles of the sucking-pump attach to a d e f i n i t e part of the head capsule, and so i t i s probably correct to i n t e r p r e t t h i s area as the clypeus. Ekblom (1926) states that i n Saldula s a l t a t o r i a ( L a t . ) , " the forehead forms i n front a transverse ridge where i t l i m i t s the clypeus." This ridge would appear to be merely the pos t e r i o r margins of the unpigmented postclypeal areas. In Saldula sp,, therefore, as i n other Heteroptera, 8 the clypeus appears to be d i f f e r e n t i a t e d Into a postclypeus, which i s united with the frons, a d i s t i n c t anteclypeus, and two paraclypeal lobes. The anteclypeus i s a c l e a r l y defined area, and appears e x t e r n a l l y as a convex lobe, i t s i n t e r n a l concavity together with i t s l a t e r a l invaginations forms a supporting base f o r the food pump. To the cephalic margin of the anteclypeus i s attached the f l a p - l i k e labrum. The paraclypeal lobes are well defined areas on each side of the anteclypeus, and extend caudad upto h a l f the length of the l a t t e r . They are c l e a r l y v i s i b l e i n both the nymph and the adult, and appear to have no i n t e r n a l muscle attachment. The homology of the paraclypeal lobes i n the Hemiptera i s very c o n t r o v e r s i a l , and has been the subject of much debate. Smith (1892) and Weber (1929) considered them to be homologous with the mandibular p l a t e s . Snodgrass (1935) also mentions, " that these paraclypeal lobes appear to be the mandibular plates of the Homoptera," but he himself doubts t h e i r mandibular o r i g i n i n the ^ omoptera i n the absence of any embryological evidence. Ekblom (1926) and Cobben (i960) designate these lobes i n Saldula -s a l t a t o r i a (Lat.) as ' laminae maxillare ' and 'maxillare p l a t t e s ' r e s p e c t i v e l y . Parsons (1959) states that the paraolypeus i n Gelastocoris i s wholly i n f l e c t e d within the head. However, most authors consider the paraclypeal lobes as parts of the clypeus. Muir and Kershaw ( I 9 H , I9l2) regard them as ' extensions of the clypeus', 9 and confirm that," they have no r e l a t i o n to the mandibles". Spooner(l938) states that the paraclypeal lobes are undoubtedly parts of the clypeus. However,' MacG-ill (1947) refe r s to these two lobes i n Dysdercus-intermedius Distant as juga. I t i s u s u a l l y not possible to trace the mandibular plates i n the Heteroptera since there i s no sulcus between the mandibular plate area and the gena. I t i s , therefore, advisable to consider the whole area of the head capsule between the eye and the points of attachment of the mouthparts as the genal area; i f mandibular plate areas need to be recognized, i t i s suggested that they be defined as the ventro-anterior area of the gena to which the mandible a r t i c u l a t e s . In mandibulate insects the maxillae are u s u a l l y attached to the ventral area of the occiput and the labium to the v e n t r a l area of the postocciput (Snodgrass, 1935). I t does not appear to be necessary to recognize a maxillary plate area despite the f a c t that such an area i s u s u a l l y described i n the hemipterous head; maxillary plates are absent i n the p r i m i t i v e orthopteran type of the head, and they cannot u s u a l l y be defined by s u l c i i n the hemipterous head(MacGill,l947). The o r i g i n of what i s herein termed the postgenal bridge has been a problem i n the past. Many authors claim that the v e n t r a l region, of the head i s formed by the f u s i o n v e n t r a l l y of the maxillary p l a t e s , the area considered to be equivalent to the postero-ventral \ TO part of the gena anterior to the point of a r t i c u l a t i o n of the labium, MacGill (1947) r e f e r s to the ventral area of the head i n Dysderous -intermedius Distant as "a large median v e n t r a l s c l e r i t e i n t e r p o l a t e d between the labium and the foramen magnum". However, she makes no comment regarding i t s o r i g i n . Parsons (1959), l i k e most e a r l i e r authors, adopts the term, 'gula* but notes that perhaps i t i s not a true gula. Snodgrass (i960) has r e c e n t l y considered the ve n t r a l s c l e r i t e s of the head i n i n s e c t s , and has concluded that they are not homologous i n a l l groups, and thus cannot i n a l l be termed a *gula». He states that there are three d i s t i n c t processes which may r e s u l t i n the formation of vent r a l s c l e r i t e s i n the head of i n s e c t s : i n the f i r s t , a hypostomal bridge may be formed between the o c c i p i t a l foramen and the base of the labium by the v e n t r a l fusion of two hypostomal lohes as i n the Diptera. The hypostomal bridge, he states i s continuous d o r s a l l y with the post-occiput. The second modification, according to Snodgrass, i s the ventral f u s i o n of two postgenal lobes to form a postgenal bridge between the o c c i p i t a l foramen and the base of the labium, as i n Vespula maculata. When t h i s i s the case, the hypostomal bridge i s replaced by the postgenal bridge,whioh d i f f e r s from the former i n not being continuous d o r s a l l y with the postocciput. He jbhus regards the ve n t r a l plate i n Notonecta and Uauooris as the postgenal bridge since i t i s continuous with the postgenae and not with the postocciput. The t h i r d process described by Snodgrass i s the ve n t r a l f u s i o n of the lower ends of the postocciput to form a median plate,which may become extended d i s t a l l y as i n the Coleoptera. T h i s median plate i s the true gula, and i s continuous proximally with the postocciput. IT In Saldula sp., as i n Notonecta and Nauooris, the ventral plate seems to he formed by the f u s i o n v e n t r a l l y of the postgenae. However, a developmental study of t h i s region i s badly needed to determine whether or not the homology accepted i n t h i s t h e s i s i s r e a l l y the correct one. This study should include a consideration of the o r i g i n of the bucculae. These structures,which apparently serve to support the rostrum during feeding, are interpreted as the ven t r a l extensions of the areas anterior to the point of attachment of the labium. Since they often extend p o s t e r i o r to the rostrum, t h i s i n t e r p r e t a t i o n i s perhaps i n c o r r e c t . -C- COMPARISON*!: On oomparing the head structures of the two f a m i l i e s , i t i s found that Saldula and Mesovelia resemble each other i n a number of features,namely, the well developed vertex, the i n d i s t i n c t coronal and f r o n t a l ecdysial cleavage l i n e s i n the adults, well developed clypeal regions, the absence of the fronto-clypeal s u l c i , the fo u r - j o i n t e d labium, the antennae, and i n possessing four p a i r s of cephalic t r i c h o b o t h r i a i n the nymphs. Hovrever, these s i m i l a r i t i e s are not i n characters of great taxonomic importance i n the group. There are a number of important features i n which they are quite d i s t i n c t (Table I ). I t should be mentioned here that although Aepophilus bonnairei appears to resemble Mesovelia more than Saldula, i n respect of the most important taxonomic characters ( i e , the shape of the mandibular l e v e r , presence of e p i c r a n i a l p i t s , and the absence of epipharyngeal process), Aepophilus j2 i a exactly l i k e Saldula. I t i s evident that the two f a m i l i e s , the Saldidae and the Mesoveliidae, show more differences than resemblances i n t h e i r head structures, and thus i t i s concluded that i n the head structure they are d i s t i n c t taxonomically. The e p i c r a n i a l p i t s are a feature which appear to be confined to the Saldidae and can thus be regarded as a character by which t h i s family can be separated from a l l other Heteroptera. 13 TABLE I DIFFERENCES IN BEAD STRUCTURES Parts of Head Saldidae Mesoveliidae I - Postclypeus, 2- Paraclypeal region. 3- Bucculae. 4- Postgenal bridge. 5- Mandibular l e v e r . 6- Postocciput, 7- M a x i l l a r y plate No such unpigmented areas present. 1- Cephalic p o r t i o n of I -postclypeus with two l a t e r a l unpigmented areas(absent i n nymph); absent i n Aepophilus. 2- Well defined i n Saldula; 2- Hot well defined. 3- Not well developed. area. not so i n Aepophilus. 3- YiTell developed i n Saldula; not so i n Aepophilus. 4- Short i n Saldula; longer i n Aepophilus. 5- Triangular both i n Saldula and Aepophilus. 6- D i f f e r e n t i a t e d i n t o dorsal and l a t e r a l portions i n Saldula, not so i n Aepophilus. 7- Not developed d o r s a l l y 7- Developed d o r s a l l y . i n Saldula;developed 4- Longer than i n Saldula, 5- Roughly quadrangular. 6- Not so d i f f e r e n t i a t e d . d o r s a l l y i n Aepophilus, 1-4 Table I continued.... Parts of Head Saldidae Mesoveliidae 8- Labrum. 9- Cephalic t r i c h o b o t h r i a . 10- Compound eyes. I I - O c e l l i , Without epipharyngeal 8 - With epipharyngeal process both i n Saldula process, and Aepophilus. 9- Three pai r s i n the adult« 9- Eight to ten p a i r s i n adult Saldula; four p a i r s i n Aepophilus, 10- Conspicuous i n Saldula; 10- Hot so conspicuous as i n Saldula. not so i n Aepophilus. I I - Present i n Saldula; absent i n Aepophilus. I I - Rudimentary i n Mesovelia mulsant; present i n M . v i t t i g e r a . 12- E p i c r a n i a l p i t s . 12- Present both i n Saldula and Aepophilus. 12- Absent. 1 5 2- THORAX: A- DESCRIPTION: I - Saldula sp. ( F i g s . 8-13, 30-34) PROTHORAX: pronotum large, with anterior c o l l a r , and p o s t e r i o r l y overlapping base of forewings; pronotum with a wide dome-shaped c a l l a l area; the l a t t e r t r i a n g u l a r l y depressed i n middle; epimeron broader than episternum, with part of l a t t e r forming a precoxal shelf; p l e u r a l sulcus and pleurodema d i s t i n c t but short; trochantin present, and very d i s t i n c t i n nymph; sternum d i f f e r e n t i a t e d i n t o a transversely elongated presternum, a tr i a n g u l a r basisternum, and a sternellum; f u r c a l arms rather elongate, extending l a t e r a l l y to meet pleurodema. MESOTHORAX: Mesonotum d i f f e r e n t i a t e d i n t o scutum and scutellum, with the d i v i d i n g suture incomplete; scutum secondarily divided i n t o a median and two l a t e r a l areas; scutellum extending p o s t e r i o r l y over second abdominal segment, and a p i c a l l y pointed; parascutellum also present, extending a n t e r o - l a t e r a l l y i n t o souto-scutellar region, the l a t t e r being unequally s c l e r o t i s e d ; postscutellum present and v i s i b l e l a t e r a l l y ; a n t e r ior notal wing process l y i n g a n t e r o - l a t e r a l l y to p o s t e r i o r notal wing process; p l e u r a l sulcus d i s t i n c t but short; pleurodema short and pointing a n t e r i o r l y ; trochantin short; episternum v e n t r a l l y fused with basisternum, the po s t e r i o r part of the former forming precoxal shelves; epimeron divided i n t o a dorsal anepimeron 16 and a ve n t r a l katepimeron, the l a t t e r produced into a point at the lower l i m i t of coxal c l e f t ; p l e u r a l wing process very d i s t i n c t ; basalar and subalar s c l e r i t e s not dist i n g u i s h a b l e ; sternum well developed, and divided i n t o a presternum, a basisternum, and a sternellum, the l a t t e r extending p o s t e r i o r l y over metasternum; furca well developed, with f u r c a l arms extending l a t e r a l l y and meeting pleurodema; phragma well developed. METATHORAX: Metanotum i n the form of fused metascutum and scutellum; postscutellum well defined; p l e u r a l sulcus h o r i z o n t a l , and dorsal i n p o s i t i o n ; pleurodema short; trochantin comparatively long; episternum broad, forming a large precoxal shelf, the two precoxal shelves being approximated mesially; epimeron small and dorsal i n po s i t i o n ; o r i f i c e s of scent-apparatus located l a t e r a l l y ; basalar and subalar s c l e r i t e s not distinguishable; metapleural wing process l y i n g forward i n the region of mes-epimeron, and rei n f o r c e d by processes from the postscutellum and epimeron of mesothorax and the postscutellum of the metathorax; metasternum reduced to a small plate underneath the mesoscutellum; f u r c a l arms short, not reaching pleurodema; phragma very l a r g e . THORACIC APPENDAGES: WINGS: (F i g s . 30, 31 ) Forewings d i f f e r e n t i a t e d i n t o clavus, corium, I'7 embolium, and a membranous area; p o s t e r i o r end of clavus produced mesially i n t o a narrow t r i a n g u l a r area along inner margin of membrane; membranous area with four c e l l s , Hindwings with d i s t i n c t vanal and jugal folds;Costa, Subcosta and base of Radius fused ; Radius and Media d i s t a l l y fused; base of Media and Cubitus i n contact with d i s t a l median pla t e ; two Vanal veins i n the vanal area; 2V with a thickened base; jugal lobe with single Jugal vein; humeral plate well developed; f i r s t and second a x i l l a r i e s reduced; t h i r d a x i l l a r y a r t i c u l a t e d both with proximal median p l a t e and second a x i l l a r y ; two median p l a t e s 1 (proximal and d i s t a l ) present. L E G S : ( F i g s , 32-34) Coxae, e s p e c i a l l y those of hind l e g , well developed, and having only p l e u r a l and trochantinal a r t i c u l a t i o n s ; a r t i c u l a r surface of hind coxa i n f l e c t e d probably to give leverage to muscles; a coxal suture present on proximal h a l f of outer surface of hind coxa, the i n t e r n a l ridge of the suture being continuous with i n f l e c t i o n of a r t i c u l a r surface; coxal suture absent i n fore and middle coxae; d i s t a l l y coxae bearing anterior and p o s t e r i o r a r t i c u l a r surfaces f o r a r t i c u l a t i o n with trochanter; trochanter immovably a r t i c u l a t e d with base of femur; femora f l a t t e n e d l a t e r a l l y ; hind t i b i a longer than hind femora, and bears stout b r i s t l e s ; proximal end of t i b i a with a d i s t i n c t head bent toward femur; t a r s i with three tarsomeres, the basa?the smallest; tarsomeres with b r i s t l e s ; pretarsus i n the form of two claws. 18 2- Aepophilus bonnairei Sign, ( F i g s . 15-17) PROTHORAX: Pronotum large with a pronotal c o l l a r ; epimeron broader than episternum, precoxal shelf formed by both; p l e u r a l sulcus short; pleurodema short; coxal c l e f t prominent; trochantin present; sternum d i f f e r e n t i a t e d i n t o a basisternum and sternellum; f u r e a l arms short. ME60TH0RAX: Mesonotum a t r i a n g u l a r piece; p l e u r a l sulcus short (shorter than that of prothorax); trochantin present; sternum d i f f e r e n t i a t e d i n t o a presternum, basisternum and sternellum, l a t t e r extending over metasternum (as i n Saldula); furca not well developed. METATHORAX: Metanotum u n d i f f e r e n t i a t e d , and shorter than pronotum and mesonotum; p l e u r a l sulcus dorsal i n p o s i t i o n ; epimeron dorsal; pleurodema very short; episternum broad, forming a large precoxal s h e l f . THORACIC APPENDAGES: LEGS: Coxae more or les s l i k e those of Saldula; no outer suture and i n f l e c t i o n ; a n t e r ior and pos t e r i o r a r t i c u l a r processes present i n coxae (as i n Saldula); femora f l a t t e n e d l a t e r a l l y ; hind t i b i a longer than hind femora(as i n Saldula); t i b i a e with stout b r i s t l e s at 19 t h e i r d i s t a l ends; t a r s i with three tarsomeres. Two rudimentary mesothoracic wings present. 3- Mesovelia mulsanti White. (Figs. 18-23) PROTHORAX: Pronotum large, with a short c o l l a r ; epimeron broader than episternum; precoxal shelf formed by both episternum and epimeron; p l e u r a l sulcus not v i s i b l e ; a very short pleurodema present; trochantin present, and very d i s t i n c t i n nymph; sternum d i f f e r e n t i a t e d i n t o a basisternum and a sternellum; f u r c a l arms (apophyseal apodemes) small, and pointing v e n t r o - p o s t e r i o r l y . MESOTHORAX: Mesonotum un d i f f e r e n t i a t e d ; p l e u r a l sulcus not v i s i b l e ; very short pleurodema present; trochantin present; episternum mesially fused basisternum; precoxal shelf formed by both the episternum and epimeron; sternum d i f f e r e n t i a t e d into basisternum and sternellum; f u r c a l arms (apophyseal apodemes) short, and pointing p o s t e r i o r l y . METATHORAX: Metanotum un d i f f e r e n t i a t e d ; p l e u r a l sulcus not v i s i b l e ; coxal c l e f t absent (present i n prothorax and mesothorax); apophyseal apodemes short and pointing l a t e r a l l y ; episternum fused with basisternum, and forming a part of precoxal shelf; metasternum 2 0 d i f f e r e n t i a t e d into basisternum and sternellum, the l a t t e r fused with the f i r s t abdominal sternum; single median o r i f i c e of scent-apparatus present,, THORACIC APPENDAGES: Coxae well developed, the fore and middle ones being swollen; a r t i c u l a t i o n both p l e u r a l and tro c h a n t i n a l ; hind coxa without coxal suture on i t s outer surface; trochanter immovably attached to base of femur; femora f l a t t e n e d l a t e r a l l y ; fore and middle femora with more stout b r i s t l e s than hind femora; hind t i b i a e with stout b r i s t l e s ; middle t i b i a e with combs at t h e i r d i s t a l ends; three tarsomeres; pretarsus i n the form of claws, l a t t e r provided with pseudo-aroliae (parempodium). 4 - Mesovelia v i t t i g e r a Puton. ( F i g s . 2 4 - 2 9 ) PROTHORAX: Pronotum with anterior c o l l a r , and p o s t e r i o r l y overlapping the bases of fore and hind wings; pronotum with a wide c a l l a l area, the l a t t e r with two l a t e r a l l y located depressions; epimeron broader than episternum; p l e u r a l sulcus short; pleurodema very short; trochantin present; sternum as i n Mesovelia mulsanti,. MESOTHORAX: Mesonotum d i f f e r e n t i a t e d i n t o scutum and 21 scutellum, the l a t t e r not extending p o s t e r i o r l y over the second abdominal segment as i n Saldula; anterior n o t a l wing process l y i n g a n t e r o - l a t e r a l l y to p o s t e r i o r notal wing process; postscutellum present; sterno-pleural region same as i n Mesovelia mulsanti. METATKORAX: Metascutum and scutellum fused, the median part extending over the second abdominal segment; postscutellum di s t i n g u i s h a b l e ; pleuro-sternal region same as i n Mesovelia mulsanti. THORACIC APPENDAGES: WINGSJ ( F i g s . 28, 29 ) Fore wings with clavus, corium and an i n d i s t i n c t embolium; bases of Costa, Subcosta and Radius fused; d i s t a l ends of fused Costa, Subcosta plus Radius, and Media and Cubitus form a stigma; r e s t of wing membranous, but without c e l l s . Hind wings without jugal f o l d ; vanal f o l d present; bases of Costa, Subcosta, Radius and Media fused; Radius and Media d i s t a l l y fused; two vanal veins present. LEGS: As i n Mesovelia mulsanti. 22 B- DISCUSSION AND INTERPRETATION: The prothorax i n Saldula i s more or l e s s of a generalised type seen i n other Heteroptera i n that i t shows no d i f f e r e n t i a t i o n i n t o separate s c l e r i t e s . The presence of the dome-shaped c a l l a l area i s not a constant feature i n the Saldidae, f o r Drake and Chapman (1958) state that a c a l l u s i s absent i n the genus Saldoida, Of the three thoracic segments, the mesothorax i n Saldula i s the most developed. This agrees withWebe^s (1930) thesis that i n the Hemiptera the fore wings are the p r i n c i p a l organs of f l i g h t . Both the mesoscutellum and mesosterhum are well developed. Taylor (1918) states that i n the Heteroptera the mesothoracic sternum i s i n d i s t i n g u i s h a b l y fused with the pleura. This i s true i n Saldula, and i t i s because of t h i s fusion that the l i m i t s of the p l e u r a l and sternal s c l e r i t e s cannot be c l e a r l y defined. The f u s i o n of the sterno-pleural s c l e r o t i z a t i o n s i s also seen i n the metathorax, Brindley (1934) figures the thorax of Saldula p i l o s e l l a (Thomson), and l a b e l s the anterior and p o s t e r i o r areas of the mesosternum BS2 (basisternum of mesothorax) and BS3 ( basisternum of metathorax), and leaves the middle area unnamed. I f the l o c a t i o n of the apophyseal p i t s i s taken as the landmark i n d e l i m i t i n g the sternal p l a t e s , her i n t e r p r e t a t i o n appears to be inaccurate with respect to the mesosternum. According to the i n t e r p r e t a t i o n given, the area which she has l e f t unnamed i s the basisternum of the mesothorax, and the areas .which she c a l l s BS2 and BS3 are merely the mesothoracic presternum and the sternellum r e s p e c t i v e l y . The basisternum of the metathorax i n 23 Saldula Is a small p l a t e , and l i e s underneath the mesosternellum,with the metasternal apophyseal p i t s l y i n g on each side of it„ This can be seen i f the mesothorax and the metathorax are p u l l e d apart. This structure i s most c l e a r i n the nymph. Thus what Brindley considers as the basisternum of the metathorax i s evidently the sternellum of the mesothorax. The p l e u r a l sulcus of the metathorax i s h o r i z o n t a l , and l i e s on the upper margin of the pleura, so much so that the epimeron occupies a dorsal p o s i t i o n , and i s attached to the l a t e r a l margin of the metascutellum. According to Taylor (1918), the hor i z o n t a l p o s i t i o n of the p l e u r a l sulcus seems to be a general feature i n the Heteroptera. Brindley (1934) states that t h i s h o r i z o n t a l p o s i t i o n i s c h a r a c t e r i s t i c of the aquatic bugs because of the enlarged coxae, which extend behind rather than from beneath the thorax. Larsen (1945) mentions a horizontal p l e u r a l ridge i n Salda muelleri (G-melin). The upper forward margin of the metapleuron extends forwards beneath the p o s t e r i o r margin of the mesopleuron, and terminates i n the metapleural wing process i n the region of the mesepimeron. Taylor (1918) mentions that s i m i l a r condition i s seen i n the Nabidae, G-erridae and Berytidae. However, the metapleural wing process i n Saldula i s p e c u l i a r i n that i t i s also supported by processes from the mesepimeron and the metascutellum, and l i e s i n the region of the mesothorax. The large precoxal shelves of the metapleuron seem to be associated with the greatest development of the hind coxae. 24 In the fore wing, the corium i s secondarily divided i n t o a marginal embolium. Drake and Chapman (1958) also mention an embolium i n the genus Saldoida. In the i n t e r p r e t a t i o n of the veins of the hind wing of Saldula the c r i t e r i o n of the a x i l l a r i e s and t h e i r a s s o c i a t i o n with p a r t i c u l a r veins has been adopted. Thus the veins i n as s o c i a t i o n with the d i s t a l median plate have been designated as the Media and Cubitus, the former together with the Radius seems to be very much approximated toward the Costa and Subcosta. I t may be mentioned here that d i f f e r e n t degrees of f u s i o n of the Costa, Subcosta, and Radius occur also i n the Miridae, Lygaeidae, Phymatidae, Mesoveliidae, and Piesmidae (Hoke, 1926). Drake and Davis (1958) have figured the hind wing of the piesmid Miespa splendida Drake, and have shown the Cubitus i n the same p o s i t i o n as i n S a l d u l a . Hoke (1926) has studied the venation of the hind wings of the Heteroptera i n d e t a i l . She follows: the Comstock-Needham system, and divides the wing in t o four areas- the c o s t a l area with Costa, Subcosta, Radius and Media I and 2; medial area with Media 3 and 4; c u b i t a l area with Cubitus and f i r s t Anal, and the anal area with the remaining anal veins. On comparision, i t becomes obvious that her C u b i t a l area i n Salda b o u o h e r v i l l e i Prov.(=coriacea Uhler) and Saldula p a l l i p e s (F.)(= separata Uhler) should r e a l l y be interpreted as the vanal area, and thus i t appears that the v e i n which she designates as Cubitus i s probably one of the vanal veins (Table I I ) . Although she has omitted the consideration of the a x i l l a r i e s i n her i n t e r p r e t a t i o n , she has fi g u r e d them i n the two saldids she studied, and i t i s evident that her Cubitus; i s not i n a s s o c i a t i o n with the d i s t a l median pl a t e , which i t should be according to the i n t e r p r e t a t i o n (Snodgrass,1935) adopted i n t h i s t h e s i s . Since many of the veins of the adult wing 25 i n Heteroptera are without a corresponding trachea (Hoke,1926), the i n t e r p r e t a t i o n of the adult veins seems to be more r e l i a b l e i f based on the associated a x i l l a r i e s . Very l i t t l e information i s obtained by studying the p o s i t i o n of the trachea i n the nymph. A r e i n t e r p r e t a t i o n of the wing venation of the Heteroptera based on the asso c i a t i o n of veins with a x i l l a r i e s i s thus required. 26 TABLE II INTERPRETATION OF THE VEINS OF HIND WING Hoke Gupta 1- Costa r- Costa 2- Subcosta 2- Subcosta 3- Radius. 3- Media 4- r-m 4- m-ou 5- Media I plus 2 5- Cubitus 6- Cubitus 6- Vanal I 7- Anal I 7- Vanal 2 8- Anal 2 8- Jugal I 2 7 ' O COMPARISON : on comparing the two f a m i l i e s , we find" that they show c e r t a i n resemblances, p a r t i c u l a r l y i n the pronotal c o l l a r , c a l l a l area with i t s median or l a t e r a l depressions, precoxal s h e l f , trochantin, and fused metascutum and scutellum, well developed coxae, f l a t t e n e d femora, and three tarsomeres , e t e „ , but they e x h i b i t d i s t i n c t differences (Table I I I ) : these differences outweigh the resemblances,, A preliminary examination of some of the fa m i l i e s of Pentatomomorpha, Cimicomorpha, Amphibicorisae and Hydrocorisae revealed that the slructure of the mesonotum and the metasternum i n the various f a m i l i e s are of either Saldula-type ( the mesosteimum pro j e c t s over the metasternum) or Mesovelia-type ( the mesosternum not projecting over the metasternum). Examination also revealed that, with the exception of Amphibicorisae, both the Pentatomomorpha and the Cimicomorpha as well as the Hydrocorisae possess both types (Table T V l ) . This indicates that i f the above mentioned taxa are natural groups, the structure of the thorax i s of l i t t l e value i n d i s t i n g u i s h i n g the suprafamilial categories. 28 TABLE III: DIFFERENCES ,IN THE THORAX Parts of thorax Saldidae Mesoveliidae I- Scutellum(mesoth.) 2- Parascutellum (mesothorax) 3- P l e u r a l sulcus-(mesothorax) 4- Epimeron(mesoth.) 5- Sternum(mesotho) 6- Coxal c l e f t (metathorax) 7- P l e u r a l sulcus (metathorax) 8- O r i f i c e of S c ent-apparatus, I - Extends over second abdominal segment i n Saldula; I - Does not extend over the abdomen. not so i n Aepophilus. 2- Present i n Saldula; 2- Absent. absent i n Aepophilus. 3- D i s t i n c t but short i n both, 4- Divided i n t o dorsal anepimeron and ve n t r a l katepimeron, 5- Sternellum extending over metasternum i n both, 6- Present i n both. 7- Present and dorsal i n both, 8- Two, located l a t e r a l l y . 3- Absent i n both(not v i s i b l e externally) 4- Not so divided i n both, 5- Sternellum not extending over metasternum. 6- Absent i n both. 7- Absent In both. 8- One, median i n p o s i t i o n . 29 Table III continued....... Parts of thorax Saldidae Mesoveliidae 9- Fore wings 10- Hind wing 11- Base of media 12- T i b i a l comb 13- Pseudo-aroliae (par-empodium) 9- D i f f e r e n t i a t e d i n t o clavus,embolium and membranous part, 10- With jugal f o l d . 11- Not fused to Costa, Subcosta and Radius-. 12- Absent i n both. 13- Absent, 9- Mostly membranous. 10- Without jugal f o l d , 11- Fused to C o s t a , Subcosta and ^adius, 12- Fore and middle t i b i a e with combs at t h e i r d i s t a l ends, 13- Present. 30 3- ABDOMEN: A* DESCRIPTION: I- Saldula sp. ( F i g s . 35-39,45,46,50) Abdomen Y i d t h ten segments. TERGA: Terga of segments two to eight d i f f e r e n t i a t e d i n t o dorsal median p l a t e s , and l a t e r a l p aratergites; tergum of f i r s t segment not complete medially; i n the male, pos t e r i o r margin of second paratergite modified i n t o a granulated convex lobe; conjuctiva between second and t h i r d p aratergite forming, underneath the anterior margin of t h i r d p a r a t e r g i t e , a f o l d with a concave anterior margin, the lattenbeing beset with stout setae; t h i s together with the convex lobe of second paratergite forms,during copulation, the grasping mechanism i n the male; remnants of o r i f i c e s of abdominal scent-glands present at posterior margin of t h i r d t e r g i t e . STERNA: Sternum of f i r s t segment rudimentary and i n d i s t i n g u i s h a b l y fused with the p a r t i a l l y membranous sternum of second segment; seven p a i r s of s p i r a c l e s present on l a t e r a l margins of the sterna two to eight, FEMALE GENITALIA: ( F i g s . 36-39) Previous Description: Ekblom (1926); Leston (1956); Scudder (1959). 31 Firsttgonapophyses joined by membrane, elongate,tapering and s p l i t l o n g i t u d i n a l l y with serrate dorsal t i p s ; rami s c l e r o t i z e d ; f i r s t gonocoxa fused with v e n t r a l part of eighth p a r a t e r g i t e ; ninth tergum without separate p a r a t e r g i t e ; gonangulum t r i a n g u l a r , i t s anterior limbs being fused with f i r s t ramus, and the p o s t e r i o r side with ninth tergum; ve n t r a l angle of p o s t e r i o r side a r t i c u l a t i n g i n a notbh on mid-dorsal side of second gonocoxa, thus forming a fulcrum on which the l a t t e r pivots'; second gonapophyses broader than f i r s t , s c l e r o t i z e d and united except at the apices, the l a t t e r being truncate; second gonocoxa elongate and thickened d o r s a l l y ; gonoplacs broad, curved and free d i s t a l l y , and united proximally by p a r t i a l l y s c l e r o t i z e d membrane; spaermatheca s i n g l e , with arybval bulb or receptacle, and an elongated spermathecal tube, the l a t t e r communicating with the vagina; a muscular pump with a single flange present between the receptacle and the main part of the duct; wall of vagina l i n e d i n t e r n a l l y with wrinkled chitinous intima, and strenghtened by a s c l e r o t i z e d r i n g 0 MALE GENITALIA: ( F i g s . 45,46,50) Previous Description: Ekblom (1926); Marks(l95l); P r u t h i (1925). Aedeagus d i f f e r e n t i a t e d i n t o phallosoma and endosoma, the" l a t t e r being further d i v i s i b l e i n t o conjuctiva and v e s i c a ; proximal part of phallosoma wide and membranous, the d i s t a l part being narrow, h e a v i l y s c l e r o t i z e d and bent over the proximal part; mouth of phallosoma located v e n t r o - l a t e r a l l y and confined to i t s d i s t a l h a l f ; two minute 32 p o s t e r i o r l y d i r e c t e d appendages located one on each side of the anterior end of the mouthj base of phallosoma supported by a inverted^ Y-shaped s c l e r i t e , with i t s upper limb fused with the junction of the basal p l a t e s ; dorsal h a l f of the conjuctiva s c l e r o t i z e d , forming a curved s c l e r i t e with i t s two ends produced i n t o l a t e r a l 'wings' on either side of mid-dorsal l i n e ; d i s t a l end of conjuctiva containing two p a i r s of d o r s o - l a t e r a l l y located appendages, the anterior p a i r smaller than the p o s t e r i o r one; ventral wall of conjuctiva containing, near the mouth of phallosoma, a V-shaped structure with two p a i r s of processes p r o j e c t i n g a n t e r i o r l y from the inner angle of V, the inner p a i r being smaller than the outer; vesica narrow, c o i l e d , and i n close contact with the ejaculatory duct; ejaculatory r e s e r v o i r located at the junction of conjuctiva and vesica; basal plates completely fused i n the middle l i n e , forming a horse-shoe-shaped structure; capitate processes attached on to the ends of basal p l a t e s ; parameres long, sickle-shaped, and pointed at apices, the proximal ends being broader and curved f o r muscle attachment, 2- Aepophilus bonnairei: ( F i g . 47) Abdomen with ten segments. TERG-A: Terga two to eight d i f f e r e n t i a t e d i n t o median t e r g i t e s and l a t e r a l p aratergites; f i r s t tergum fused with second; clasping organ present i n the second and t h i r d segments. 33 STERNA: F i r s t sternum rudimentary and i n d i s t i n g u i s h a b l y fused with p a r t i a l l y membranous second sternum: seven p a i r s of s p i r a c l e s on segments two to eight. FEMALE GENITALIA: More or l e s s as i n Saldula. MALE GENITALIA: ( F i g . 47) Aedeagus d i f f e r e n t i a t e d i n t o a phallosoma and endosoma, l a t t e r with two appendages pr o j e c t i n g out of the mouth of phallosomaj base of phallosoma membranous, d i s t a l part narrow and h e a v i l y s c l e r o t i z e d and bent over the proximal part; base of phallosoma supported by an L-shaped s c l e r o t i z e d structure, the h o r i z o n t a l limb being fused with the junction of the basal p l a t e s ; dorsal part of endosoma presents a s c l e r o t i z e d strvicture; basal plates completely fused and l i k e those i n Saldula. 3- Mesovelia mulsanti : ( F i g s . 40-44,48,49,51) Abdomen with ten segments. TERGA: Terga of segments two to eight i n the female and two to' seven i n the male d i f f e r e n t i a t e d i n t o dorsal median t e r g i t e s and l a t e r a l p a r a t e r g i t e s ; o r i f i c e of abdominal scent gland i n the middle of fourth tergum; clasping organ i n the male absent. 34 STERNA: F i r s t sternum i n d i s t i n g u i s h a b l y fused a n t e r i o r l y with metasternellum and p o s t e r i o r l y with second sternum; seven p a i r s of sp&fcac.lesspresent on segments two to eight; anterior margin of the seventh sternum produced a n t e r o - l a t e r a l l y , i n mid-line, as a long apodeme; two c i r c u l a r patches of black setae present on the eighth sternum i n the male. FEMALE GENITALIA} (F i g s . 41-44) Previous Description: Ekblom (1926); Neering (1954); Pendergrast (1957); Scudder (1959). F i r s t gonapophyses joined by membrane, elongate, tapering and s p l i t l o n g i t u d i n a l l y , with serrate t i p s ; rami s c l e r o t i z e d and i n t e r l o c k i n g ; f i r s t gonocoxa fused with eighth p a r a t e r g i t e ; gonangulum t r i a n g u l a r , Its p o s t e r i o r side also fused with an i n f l e c t i o n between the eighth and ninth terga; second gonapophyses elongate, s c l e r o t i z e d , and l a o i n a t e , united except at the t i p s ; second gonocoxa elongate; gonoplacs t r i a n g u l a r , s c l e r o t i z e d , curved and attached to second gonocoxae; spermatheca single with an accessory fecundation canal* MALE GENITALIA: ( F i g s . 48,49, 51)• Previous Description: Ekblom (1926); P r u t h i (1925). Aedeagus d i f f e r e n t i a t e d i n t o phallosoma and endosoma, the l a t t e r being further d i v i s i b l e i n t o conjuctiva and vesica; proximal part of phallosoma membranous, d i s t a l part h e a v i l y s c l e r o t i z e d and bent over the proximal part; a t r i a n g u l a r s c l e r o t i z e d area present i n the mid-dorsal part of the conjuctiva; proximally conjuctiva produced i n t o 35 s c l e r o t i z e d appendages projecting from the mouth of the phallosoma; vesica aarrow and short and i n close contact with ejaculatory duct; two ejaculatory r e s e r v o i r s d i s t i n g u i s h a b l e ; basal plates completely fused i n the middle l i n e , forming a horse-shoe-shaped structure; capitate processes attached l a t e r a l l y ; parameres hook-like, pointed at the apices, the proximal ends being broader f o r muscle attachment„ 4- Mesovelia v i t t i g e r a : Abdomen very s i m i l a r to that of Mesovelia mulsanti. B- DISCUSSION AND INTERPRETATION: Drake and Hottes ( l 9 5 l ) state that the " hardened and p l a t e - l i k e and roughened lobe of the f i r s t paratergite together with the f p e g - l i k e ' or ' s p i n e - l i k e ' organs" of the second paratergite form the s t r i d u l a t o r y organ i n the Saldidae. Leston (1957), however, established that the granulated plate and the pegs were a c t u a l l y situated on the second and t h i r d paratergites r e s - p e c t i v e l y , and not on the f i r s t and second as indicated by Drake and Hottes 0 Leston also states that the organ functions not as a s t r i d u l a t o r but as aclasping mechanism i n the male during copulationo Examination of the organ i n Saldula has revealed, however, that one small modification i s needed i n Leston's description; the pegs are lodated a c t u a l l y on a f o l d of the conjuctiva underneath the anterior margin of the t h i r d paratergite and not on the anterior margin of the paratergite i t s e l f . 56 In the male g e n i t a l i a , the s c l e r o t i z e d curved structure i n the base of the conjuctiva probably acts as a guiding mechanism for the v e s i c a , which passes along the ventral surface of t h i s s c l e r i t e j the ves i c a i s then directed on to the V-shaped structure i n the ventral wall of the conjuctiva and i s thus everted out. P r u t h i (1925), i n h i s account of the male g e n i t a l i a of Chiloxanthus p i l o s u s ( F a l l . ) and Salda 1 i t t o r a l i s ( L . ) does not mention the c o n j u c t i v a l appendages, the ejaculatory r e s e r v o i r , andjthe capitate processes, the l a t t e r according to Marks (l95l()), are secondary developments i n the Heteroptera, and mark the o r i g i n a l point of attachment of the parameres to the basal p l a t e s . Ekblom (1926) also f a i l e d to notice the ejaculatory r e s e r v o i r and the capitate processes. P r u t h i (1925) mentions that the aedeagus i n Mesovelia i s not d i f f e r e n t i a t e d i n t o phallosoma and endosoma, but examination of Mesovelia mulsanti has shown that the phallosoma, conjuctiva and vesica are d i s t i n g u i s h a b l e . Ekblom (1926) figures a drawn-out endosoma with i t s two appendages i n his account of Mesovelia furoata. C- COMPARISON!:": On comparing the structure of the female g e n i t a l i a , the aedeagus, the presence of ejaculatory r e s e r v o i r , p a r a t e r g i t e s , and the seven p a i r s o f abdominal s p i r a c l e s , the two fa m i l i e s are found to be very s i m i l a r , but i t i s evident that other characters i n d i c a t e that they are taxonomically d i s t i n c t (Table I V ) . 37 TABLE IV DIFFERENCES IN THE ABDOMEN Parts of abdomen Saldidae Mesoveliidae I - Apodeme i n sternum V I I . I - Absent. I - Present. 2 - Clasping organ. 2- Present. 2- Absent. 3- Aedeagus. 3- D i f f e r e n t i a t e d 3- D i f f e r e n t i a t e d i n t o i n t o phallosoma, phallosoma, conjuctiva conjuctiva and and v e s i c a . v e s i c a i n Saldula; i n Aepophilus,into phallosoma and endosoma only* 4- Base of phallosoma,, 4- Supported by an 4- Absent. inverted Y-shaped structure i n Saldula; by L-shaped structure i n Aepophilus, 5- Conjuctiva. 5- Dorsal h a l f s c l e r o - 5- Present. t i z e d i n t o a curved structure i n both. 6- E j a c u l a t o r y r e s e r v o i r . 6- Present(one). 6- Present (two). 7- Parameres. 7- sickle-shaped. 7- Hook-like. 4- INTERNAL ANATOMICAL CHARACTERS: To supplement the characters obtained from a study of the external morphology, information on the i n t e r n a l anatomy and other important features of the insects has been obtained from the literature» Studies on the mandibular lever (Spooner, 1938), accessory s a l i v a r y glands (Baptist,1941; Southwood,l954), wing venation (Hoke,1926), testes (Pendergrast,l956), ovary (Garayon,l950j Miyamoto,1957), sex-determining mechanism (Makino,l950), accessory'^fecundation canal (Pendergrast, 1957), eggs (Southwood,l956) were consulted, and the information obtained i s included i n Table V« 39 TABLE V TAXONOMIC CHARACTERS OF PENTATOMOMORPHA,CIMICOMORPIiA,AMPHIBICORISAE AND HYDROCORISAE Parts Pentatomomorpha Cimicomorpha Amphibicorisae Hydrocorisae I- Labrum (Spooner,I938), 2 - Mandibular lever (Ekblom , I 9 2 6 ) . 3- Accessory s a l i -vary gland $Baptist,I94l; Southwood,l955) 4- Mesoscutellum (person, observ,) 5- Mesosternum (person, observ.) 6- O r i f i c e of Scent-gland(pers. obs 0) 7- Metacoxal c l e f t (person, observ.) 8- R and M i n hind wing(Leston,Pen-der grast,South-wood, 1954; pers. I- Not broad, 1onger. 2 - Triangular. 3- Tubular. 1- Broad,flap- I- Broad,flap- I- Broad, f l a p -l i k e or l i k e , w i t h l i k e , longer. epipharynge-a l process. 2 - Triangular. 2 - Quadrangular . 2 - Three branched or t r i a n g u l a r . 3- V e s i c u l a r . 3- Vesi c u l a r 3- V e s i c u l a r , (Gerridae), 4- Projects or 4- Does not does not project project over abdomen, 5- Saldula plus Mesovelia type. 6- L a t e r a l . 4- Does not pro-4- Does or does ject over not project over abdomen. abdomen. 5- Saldula plus- 5- Mesovelia Mesovelia type. type, 6- L a t e r a l . 6- Median or 7- Present. 8- D i a - t a l l y separated. 7- Present or absent, 8- D i s t a l l y fused. absent, 7- Absent. 8- D i s t a l l y fused (Gerridae) over abdomen, 5- Saldula plus Mesovelia type, 6- L a t e r a l or absent, 7- Present or absent. 8- D i s t a l l y fused. 40 Table V continued,„ ........ Parts Pentatomomorpha Cimicomorpha Amphibioorisae Hydrocorisae 9- Male g e n i t a l i a 2- Pentatomid type 9- Reduvid 9- Reduvid type. 9- Reduvid (Pruthi,I925), or r e l a t e d thereto. 10- Female g e n i t a l i a 10- Pentatomid (Scudder,l959). type. type or type, r e l a t e d thereto. 10- CimicomorphlO- Cimicomorph 10- Cimicomoph type type, I I - T e s t i c u l a r f o i l - I I - Usually seven. I I - Usually type or r e l - or r e l a t e d ated thereto, thereto. I I - Usually one. I I - Usually seven. icles(Pender-grast,I956). 12- Male sex Chrom. 12- XY or XO type, 12- XY or XO 12- XO type. (l£akino,I950). 13- Eggs 13- With microp-(Southwood, y l a r processes. 1956) type. 13- With micr- 13-pylar appa-ratus. seven. 12- XY or XO type. . 13 14- Accessory fecu- 14- Absent, ndation canal (Pendergrast, 1957) 14- Absent, 14- Present except i n Ochteridae, ^4- Absent, 4r 5- COMPARISON.! OF ALATE AND APTEROUS (AND BRACHYPTEROUS) FORMS IN SALDIDAE AND MESOVELIIDAE t From comparative morphology of the alate and the apterous forms i n both the Saldidae and the Mesoveliidae, i t i s evident that these two forms show d i s t i n c t s t r u c t u r a l d i f f e r e n c e s , p a r t i c u l a r l y i n the thorax (Tables VI and V i i ) . Insects with f l i g h t possess well developed f l i g h t muscles and corr e l a t e d with t h i s an elaborate thoracic structure, while those with l i m i t e d or no f l i g h t have reduced f l i g h t muscles, and consequently le s s developed thoracic s t r u c t u r e . F l i g h t i n insects Is effe c t e d by two sets of muscles, the d i r e c t and the i n d i r e c t . The i n d i r e c t muscles include the dorsal l o n g i t u d i n a l muscles and the dorso-ventral muscles. The d i r e c t muscles are attached to wing bases or wing s c l e r i t e s and include p r i n c i p a l l y the basalar, the subalar and the muscles of the a x i l l a r i e s . According to Larsen (1945), the p r i n c i p a l muscles are present i n Saldula, and accordingly, as one would expect, the apodemes and the i n t e r n a l margin of the pronotal c o l l a r are well developed i n the prothorax. The l o n g i t u d i n a l muscles, running from the f i r s t phragma to the second phragma i n the pterothorax, mainly produce the arching of the nota, and thus r a i s i n g the notal processes r e l a t i v e to the p l e u r a l processes, act as depressors of the wings. And since these muscles are important i n f l i g h t , the phragmata i n Saldula are well developed. S i m i l a r l y the development of the furca seems to be co r r e l a t e d with the development of the d i r e c t muscle, M. f u r c a - p l e u r a l i s (of Larsan). 4 2 By the same token, the absence of another d i r e c t muscle,M.coxa-subalaris (of Larsen) both i n the mesothorax and the metathorax i s c o r r e l a t e d w i t h the absence of the subalar s c l e r i t e i n the pterothorax. The l a t e r a l oblique muscle, M.mesonoti secundus (of Larsen) i s w e l l developed i n the mesothorax of S a l d u l a , but i s absent i n the metathorax. This can be explained on the b a s i s of Weber's t h e s i s that i n the Heteroptera the f o r e wings are the p r i n c i p a l organs of f l i g h t , and thus the mesothorax i s more developed than the metathorax. I t i s evident from the fofcegoing t h a t the morphological d i f f e r e n c e s are the r e f l e c t i o n s of the f u n c t i o n a l d i f f e r e n c e s i n the a l a t e and the apterous forms. U n f o r t u n a t e l y , no account of the musculature of Mesovelia i s a v a i l a b l e f o r comparision, but the s t r u c t u r a l d i f f e r e n c e s i n the thorax of Mesovelia v i t t i g e r a and Mesovelia mulsanti could a l s o be explained on a f u n c t i o n a l b a s i s . This study has a l s o revealed t h a t the s t e r n a l r e g i o n i n the a l a t e and the apterous forms shows very l i t t l e d i f f e r e n c e , and thus perhaps the dor s o - v e n t r a l muscles are not of great importance i n the f l i g h t of these i n s e c t s . On comparing the a l a t e and the apterous forms of both f a m i l i e s , i t i s found that the o c e l l i are rudimentary or absent i n the apterous forms, but are present i n the a l a t e forms. Accompanied w i t h t h i s presence of the o c e l l i i n the a l a t e forms, are the w e l l developed compound eyes, which are not so conspicuous I n the apterous forms. Such c o r r e l a t e d presence or absence of c e r t a i n s t r u c t u r e s has a l s o been reported i n the Lygaeidae (Scudder, personal communication). I t i s . l i k e l y that perhaps an a l a t e i n s e c t needs more p e r f e c t v i s u a l 43 apparatus than an apterous one, and probably the o c e l l i supplement the compound' eyes i n the l a t t e r ' s v i s u a l perception. I t i s also possible that the presence of the o c e l l i i n the alate forms and i t s absence i n the apterous- ones may be due i n part to a genetic linkage with some other character effected by l o s s of f l i g h t . 44 TABLE VI DIFFERENCES IN ALATE AND APTEROUS FORMS IN THE SALDIDAE Parts Saldula sp. Aepophilus bonnairei 1- C a l l a l area. 2 - F u r c a l arms. 3- Mesoscutellum. 4- O c e l l i . 1- Present. 2 - Well developed. 3- Extends over the abdomen. 4- Present. 1- Absent. 2 - Not well developed, 3- Does not extend over the abdomen. 4- Absent, TABLE VII DIFFERENCES IN ALATE AND APTEROUS FORMS IN THE MESOVELIIDAE Parts Mesovelia v i t t i g e r a Mesovelia mulsanti 1- C a l l a l area, 2 - Mesonotum, 3 - Metanotum, 4- O c e l l i . 1- Present. I - Rudimentary, 2 - D i f f e r e n t i a t e d into meso- 2- Mesonotum scutum and scutellumj, the fo- und i f f e r e n t i a t e d , rmer being overlapped by the p o s t e r i o r part of pronotum. 3- Median part extending 3- Does not extend over abdomen, over abdomen. 4- Present. 4- Rudimentary. 45 6- RELATIONSHIP BETWEEN SALDIDAE AND MESOVELIIDAE: From the foregoing study i t i s evident that the Saldidae and the Mesoveliidae are not c l o s e l y r e l a t e d as suggested by the comparative morphological study of the female g e n i t a l i a ; t h e y are quite d i s t i n c t i n gross morphology. In order to a s c e r t a i n or check the systematic p o s i t i o n of the two f a m i l i e s , i t has been necessary to compile a table showing the differences between the four higher taxonomic groups u s u a l l y recognized i n the Heteroptera. This table (Table V) l i s t s only the characters considered i n recent works to be of r e a l importance i n the higher c l a s s i f i c a t i o n of the group. I f we also l i s t these same characters i n the Saldidae and Mesoveliidae (Table V I I I ) , and compare Table V and Table V I I I , we can work out the possible systematic p o s i t i o n of the two f a m i l i e s under consideration. 46 TABLE VIII TAXONOMIC CHARACTERS OF THE SALDIDAE AND MESOVELIIDAE Parts Saldidae Mesoveliidae 1- Labrum. 2- Mandibular l e v e r . 3- Accessory s a l i -vary gland. 4- Mesoscutum. 5- Mesosternum. 6- O r i f i c e of scent gland. 7- Metacoxal c l e f t . 8- R and M of hind wing. 9- Male g e n i t a l i a . 10- Female g e n i t a l i a . 11- T e s t i c u l a r f o l l i c l e s . 1- Broad, f l a p - l i k e . 2- Triangular. 3- V e s i c u l a r . 4- Projects over abd-omen(exception i n C imic omorpha). 5- Projects over meta-sternum(Saldula type). 6- L a t e r a l o 7- Present. 8- D i s t a l l y fused. 9- Pantatomid type. 10- Cimicomorph type. 11- Seven. 1- Broad,flap-like,with epipharyngeal process. 2- Quadrangular. 3 4- Does not project over abdomen. 5- Does not project over metasternum (Mesovelia type). 6- Median. 7- Absent. 8- D i s t a l l y fused. 9- Reduvid type. 10- Cimicomorph type, with gonoplac. 11- One, 47 TableVT.il continued......... Parts Saldidae Mesoveliidae 12- Male sex- 12- 12- XI type. chromosomes, 13- Eggs, 13- Operculum absent, 13-egg burster present, pdeudo-micropyle absent, 14- Accessory 14- Absent, 14- Present, fecundation canal. 48 7- SYSTEMATIC POSITION OF THE MESOVELIIDAE: Pruthi(1925) and China (1933) included Mesoveliidae and Hydrometridae i n Gerridae. Ekblom (1926) also maintained that," Mesoveliidae shows perfect conformity with Hydrometra group". Handlirsch (1908) included both Mesoveliidae and Saldidae i n Gymnocerata. He probably thought that Aepophilidae arose from Mesoveliidae, Reuter (1910), however, doubted i f Mesoveliidae i s a branch of Gerroidea, Leston, Pendergrast and Southwood (1954) placed Mesoveliidae i n Amphibicorisae. I t i s thus evident from the foregoing that most of the authors agree that Mesoveliidae belongs to the present group of Amphibicorisae. The present study of the morphology of Mesoveliidae shows that t h i s family i s c o r r e c t l y placed i n the Amphibicorisae, On the basis of the epipharyngeal process, mandibular l e v e r , mesoscutum, mesosternum, median scent-gland o r i f i c e , absence of metacoxal c l e f t , number of t e s t i c u l a r f o l l i c l e s and the presence of accessory fecundation canal, Mesoveliidae d i s t i n c t l y belongs to Amphibicorisae. However, Scudder (1959) showed that the female g e n i t a l i a of Mesoveliidae are of a p r i m i t i v e Cimicomorph type with a well developed gonoplacj gonoplacs are absent i n a l l other Amphibicorisae as f a r as i s known. I t i s probable that Mesoveliidae represents a p r i m i t i v e stage or separate branch of the Amphibicorisae stem-a branch which separated before the gonoplac was l o s t i n the phylogeny(Fig.A)o 49 8- SYSTEMATIC POSITION OF SALDIDAE: There has been l i t t l e agreement among the heteropteristas on the probable p o s i t i o n of the Saldidae i n the higher c l a s s i f i c a t i o n of the Heteroptera, and t h i s family, more than any other, has been v a r i o u s l y moved from one group to another. Table IX shows In chronological order the views of d i f f e r e n t authors on the p o s i t i o n of the Saldidae, and i t i s evident that the Saldidae has no recognized p o s i t i o n to date. I t i s evident that on the basis of the labrum, mandibular l e v e r , mesoscutum, mesosternum, l a t e r a l scent-gland o r i f i c e , metacoxal c l e f t , the male g e n i t a l i a and t e s t i c u l a r f o l l i c l e s , the Saldidae cannot be included i n the Amphibicorisae. The Hydrocorisae i n general are almost i d e n t i c a l with the Cimicomorph section of the Geocorisae, and" so these groups can be considered as one. The p o s i t i o n of the Saldidae i s to be sought e i t h e r among Pentatomomorpha or Cimicomorpha. On the basis of the labrum, accessory s a l i v a r y gland and d i s t a l l y fused Radius and Media of the hind wing, Saldidae show a f f i n i t y with Cimicomorpha, but i n the structure of male g e n i t a l i a and the eggs i t c l e a r l y belongs to the Pentatomomorpha. Two a l t e r n a t i v e i n t e r p r e t a t i o n s of t h i s s i t u a t i o n are p o s s i b l e , and these are p i c t o r i a l l y i l l u s t r a t e d i n F i g s , A and B. I f we consider the Cimicomorph characters as the most important, and i f i t is- accepted that p a r a l l e l evolution can occur i n the egg and male g e n i t a l i a , i t can be argued that the Saldidae i s a Cimicomorph,which has branched o f f the main Cimicomorph stem and has evolved p a r a l l e l to the Pentatomomorph l i n e , having subsequently developed the Pentatomomorph type of male g e n i t a l i a 50 and eggs, However, i t should be noted that there i s l i t t l e evidence for p a r a l l e l evolution. But i t s p o s s i b i l i t y cannot be discounted since so many cases are known elswhere In the animal kingdom, The a l t e r n a t i v e systematic p o s i t i o n can be determined by assuming that p a r a l l e l evolution has not occurred. One must also state i n t h i s a l t e r n a t i v e scheme that the Pentatomomorph complex of characters has evolved gradually and not by a single ' s a l t a t i o n 1 . The a l t e r n a t i v e scheme shows the Saldidae as a side branch of the main Pentatomomorph stem, a branch which arose a f t e r the evolution of the male g e n i t a l i a and eggs but before the evolution of the r e s t of the Pentatomomorph characters. The data that we have available at the present time are not s u f f i c i e n t to enable one bo state which of the two a l t e r n a t i v e s mentioned i n t h i s t h e s i s i s the correct one although I am i n c l i n e d to consider the Saldidae as a branch of the Cimicomorph l i n e , which probably subsequently developed the Pentatomomorph male g e n i t a l i a and eggs. 5r TABLE IX VIEWS OF VARIOUS AUTHORS ON THE SYSTEMATIC POSITION OF THE SALDIDAE Authors Dates Views 1- L a t r e i l l e 2- Amyot & S e r v i l l e 3- Osborn 4- Kirkaldy 5- Handlirsch 6- Distant 7ruReuter 8- Pruthi 9- Ekblom 10- Esaki & China 11- Borner 1825 1843 1898 1908 1908 I902-I9I8 1912 1925 1926 1927 I2- Spooner 1934 1938 Grouped Saldidae i n Oculatae(Geocorisae). Included Saldidae i n Nudirostres (Geocorisae). A l l i e d Saldidae with Gerridae. included Saldidae i n the superfamily Notonectoidea. Included Saldidae and Mesoveliidae i n Gymnocerata. Associated Saldidae with Reduviidae. Stated that Saldidae are c l o s e l y a l l i e d to Nabidae* Put Saldidae i n Pentatoma. Stated that Saldidae have a f f i n i t i e s with Nabidae. Put Saldidae and Leptopodidae i n Hydrocorisae, Included Saldidae i n superfamily Reduvioidea. Grouped Saldidae, Anthocoridae, and Cimicidae together on the basis of f l a p - l i k e labrum. Table IX continued Authors Dates Views 13- Larsen 14- Leston 1945 1953 15- Leston, Pendergrast, 1954 & Southwood 16- China 17- Pendergrast 18- Leston & Scudder 19- Scudder 1955 1957 1957 1959 D i f f e r e d i n p l a c i n g Saldidae intermediate between Cryptocerata and Gymnoceratao ind i c a t e d a Trichophoran a f f i n i t y of Saldidae on the basis of wing venation. Included S a i d o i d e a i n Pentatomomorpha, although they stated that" Saldoidea i s f a r removed from the main Pentatomomorph stem. Put Saldidae at the base of Amphibicorisae on the basis of three p a i r s of cephalic t r i c h o b o t h r i a . Stated that Saldidae have the type of spermatheca found i n most Trichophora. Included Saldidae i n the Geocorisae. Included ^aldidae i n Cimicomorpha on the basis of the female g e n i t a l i a . 5 3 FIGURE A Cimicomorpha & Hydrocorisae E- I- Labrum broad and f l a p - l i k e , 2- Mandibular lever three branched or t r i a n g u l a r , 3 - O r i f i c e of scent-gland l a t e r a l or absent, 4 - Metacoxal c l e f t present or absent, 5 - T e s t i c u l a r f o l l i c l e s u s u a l l y seven. 6- Accessory fecundation canal absent. Pentat omomorpha Mesoveliidae Amnhibicorisae C- I- Labrum not broad, and f l a p - l i k e . 2- Accessory s a l i v a r y gland tubular type, 3- R and M i n hind -wing separate d i s t a l l y , 4 - Male g e n i t a l i a Pent-atomomorph type. 5 - Female g e n i t a l i a Pentatomomorph type. 6- Eggs with micropylar processes. 1- Labrum broad and f l a p - l i k e . 2- Accessory s a l i v a r y gland of v e s i c u l -ar type. 3 - R and M of hind wing fused d i s t a l l y . 4 - Female g e n i t a l i a Cimicomorph type, 5 - Eggs with microp-y l a r apparatus. 1- Labrum,broad, f l a p - l i k e with epipharyngeal process, 2- Mandibular lever quadrangular. 3 - O r i f i c e of scent-gland median or absent, 4 - Metacoxal c l e f t absent, 5 - T e s t i c u l a r f o l l i c l e s one. 6- Accessory fecundation canal present except i n Ochteridae. Cimicomorpha 5 4 FIGURE B Cimicomorpha & Hydrocorisae Cimicomorpha D,E,F as i n F i g . A 55 LITERATURE CITED Amyot, C. 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Some remarks on the phylogeny of the Hemiptera-Heteroptera. Canad. Ent. 40 j 357 - 364. Larsen, 0. 1945. Der Thorax der Heteropteren, Skelett und Muskulatur. Lunds Univ. A r s s k r i f t . N.F. Ad 2, 4 l j 1945. * L a t r e i l l e , P. A. 1825. Families n a t u r e l l e s du regne animal; expoees succinctement et dans un ordre anaTytique, avec 1 * i n d i c a t i o n de leurgenres P a r i s . 570 p. Leston. D, 1956. Systematics of the marine bug- Aepophilus.Nat., 178:427-428, 1957. The s t r i d u l a t o r y mechanism i n t e r r e s t r i a l species of Hemiptera-Heteroptera. Proc. z o o l . Soc. Lond.128(3)•369-386. 5 7 Leston, Do, Pendergrast, J, G. and Southwood, T. R. E. 1954. C l a s s i f i c a t i o n of the t e r r e s t r i a l Heteroptera(Geocorisae). Nature, 174; 91 - 9,2. Leston, D. and Scudder, G. G. E. 1956, A key to larvae of the f a m i l i e s of B r i t i s h Hemiptera-Heteroptera. Entomologist.89; "223- 3 l . MacGi l l , E. I . 1947. The anatomy of head and mouthparts of Dysdercus intermedius D i s t . Proc. z o o l . Soc. Load. 117: 115- 128. Makino, S. 1950. Chromosomer.numbers i n animals. The Iowa State College Press. Marks, E. P. 1951. Comparative studies of male g e n i t a l i a of Hemiptera (Homoptera-Heteroptera), Jn. Kansas ent. Soc. 24; 134-41. 1959. The food pump of Pe l o c o r i s and comparative studies on the and other aquatic liemiptera. Psyche, 64:123-134. Miyamoto, S. 1957. L i s t of ovarioles i n Japanese Heteroptera. S i e b o l d i a , 2 ( l ) j $ 9 - 82. Muir, F. and Kershaw, J. C. 1911. On the homologies and mechanism of the mouthparts of Hemiptera, Psyche, I 8 ( l ) : I . 1912. The development of the mouthparts i n the Plomoptera, with observations on the embryo of Siphanta. I b i d . 19(3):77. Neering, T. 1954. Morphological v a r i a t i o n s i n Mesovelia mulsanti (Hemiptera, Mesoveliidae). Univg Kans, Sc. B u l l . , 36':K5-148, *0sborn, H. 1898, The phylogeny of the Hemiptera. Proc. ent. Soc. Wash. 3:185-90. Parsons, M. C. 1959, Skeleton and musculature of the head of Gelasbcoris o^culatus (Fabr.) Hemiptera-Heteroptera. B u l l . Mus.^ompT^Zool. Harvard College, I 2 2 ( l ) . Pendergrast, J . G. 1956. T e s t i s f o l l i c l e numbers i n the Heteroptera. Ent, mon. Mag. 92 : 275-76. 58 Pendergrast, J. G. 1957. Studies on the reproductive organs of the Heteroptera with a consideration of t h e i r bearing on c l a s s i f i c a t i o n . Trans. R. ent. Soc. Lond. 109 ( l ) : 1 - 63. P r u t h i , H. S. 1925. The morphology of the male g e n i t a l i a i n Rhynchota, Trans. R. ent. S o c Lond. !925 pp. I2I - 267. *Reuter, 0. M. 1912. Bermerkungen uber mein neues Heteropteren System. Qfvers, f i n s k a Vetensk. Soc. Forh 54 A (6) : I - 62. Scudder, G. G. E. 1959. The female g e n i t a l i a of the Heteroptera : morphology and bearing on c l a s s i f i c a t i o n . Trans. R. ent. Soc. Lond. I l l (14) : 405 - 67. ~ Smith, J. B. 1892, Notes on the homology of the Homopterous mouth, Proc. Amer. Assoc. Adv. S c i . 40 : 325. Snodgrass, R. E. 1635, P r i n c i p l e s of insect morphology, McGraw H i l l Book Com. Inc. NTY, ~ 1960. Facts and theories concerning the insect head, Smithson. misc. C o l l , 142 ( i ) . Southwood, T. R. E. 1955. The morphology of the s a l i v a r y glands of t e r r e s t r i a l Heteroptera (Geocorisae) and i t s bearing on c l a s s i f i c a t i o n , T i j d s c h r . Ent. 98 : 77 - 84, 1956, The stucture of the eggs of the t e r r e s t r i a l Heterpptera and i t s r e l a t i o n s h i p to the c l a s s i f i c a t i o n of the group. Trans. R. ent, Soc. L 0nd. 108(g) : I6e - 221. Spooner, C. S. 1938. The phylogeny of Hemiptera based on a study of the head capsule. Univ. I l l i n o i s B u l l . 35 No. 70 : I - 102, Taylor, L. H. 1918, The thoracic s c l e r i t e s of Hemiptera and Heteroptera. Ann, ent. Soc. Amer. II : 225 - 249. *T/eber, H, 1929. Kopf und Thorax von P s y l l a m a l i . Z e i t s c h r . Morph. Okol , 14 : 59 - 165. 1930, Bio l o g i e der Hemipteren. B e r l i n , * not consulted. 5 9 KEY TO LETTERING- OF FIGURES A o o o o o . a o a . a o o . anal "Vein. AG o . o o . o o . o o . o . anteclypeus. AEM • « > < < * t o < * * « anepimeroa. AF oco«o.«oo»o.» antafossae 0 ANP 9 > o o « . « e o « o 9 ant e r i o r notal wing process, AP ............. apodemes, APP o o . o o o . o . o o . apophyseal p i t s . AS ............. antennal tubercle. AT o a o . o . o « . o « o . antenna. 1 .AX . . 0 . . . f i r s t a x i l l a r y . 2 AX . o o . . . . . . . . second a x i l l a r y . 3 AX o o . . . . . . . . . t h i r d a x i l l a r y . B . . . . . . . . o o » . . . spermathecal bulb. BP O . . O O . 0 . . . o o o basal p l a t e s . I BS o . . . . . . . . . . basisternum of prothorax. 2BS . . . . . . . . o s . " " mesothorax. 3 BS ........... " " metathorax. BU ............. bucculae, C . . . . . o o . . . . . « . costa. GA o o « a o . o . o » . o o callus« CE . . . . . . . . . . . . a compound eye. CL . A . . . . . . . . . . . clypeus» CNJ . . . o o . . . o s . . c o n j u c t i v a . 60 A P P «i>oo«ao#tt 0 0 0 0 c o n j u c t i v a l appendages. G O 0 0 0 0 0 0 0 0 0 0 9 0 0 0 0 0 0 0 corium. C P 0 0 9 0 0 0 0 0 0 0 0 9 0 0 6 0 0 0 capitate process, 0 C T 0 9 0 0 0 0 0 0 0 9 0 9 0 0 t 0 O 0 coxal c l e f t o C T Y o o o o e o « o « « « o Q 0 « 0 O coxal c a v i t y . C U 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 cubitus. C V 0 0 0 0 0 0 0 0 0 0 9 9 0 0 B 0 0 0 clavus o Gj£ o o o o o o o o o o t o o o » 0 0 o coxa, " D M P 0 0 0 0 0 0 0 0 0 0 0 0 0 » 0 0 9 d i s t a l median process. T p T > Xjir o o 0 e o o 0 0 0 0 0 0 0 0 » 0 0 0 e p i c r a n i a l p i t . JliDS 0 0 0 0 0 0 0 0 0 0 9 0 0 » * 0 0 endosoma. EDS iU?P « o * t o » o « * > 0 0 0 endosomal appendages. ejaculatory duct. EtTH. 0 0 0 0 0 0 0 0 0 0 0 0 0 1 9 0 0 ejaculatory r e s e r v o i r . X "EM o » « « « « o * * « o o > 0 0 0 epimeron of prothorax. 2 E M o o « a « a « * o e s o ( 1 0 0 0 " " mesothorax. 0 JiiM 0 0 0 0 0 0 0 0 0 0 0 0 » 0 9 9 " " metathorax. E P P 0 O O O 9 O 9 9 0 9 0 O O I k 9 0 0 epipharyngeal process. 9 0 0 episternum of prothorax. 2 E S 0 0 9 9 9 0 0 0 0 0 0 0 c 0 * 0 " " mesothorax. 3 E S • o o a « o 4 « 0 0 * « f l 0 9 0 " " metathorax. E C 0 0 0 0 0 0 9 0 0 0 9 0 0 0 1 0 9 0 fecundation canal. E L 0 * 0 9 0 0 0 0 0 0 0 0 0 0 1 0 9 * flange of pump. E M 0 0 O O 9 O 0 9 0 0 0 0 O 0 Q o a e femur. E H 9 0 O 0 0 a 0 O O 9 9 9 9 9 « 0 0 0 9 frons. 61 FTJ o . e a - e a . a a ' o . a . o . 1*111*0 &. a FWP o . o . fore wing prooess. G .................. gena, GA gonangulum. GP . . . . . . . . . . . . 0 . . . granular plate of clasping organ. GPC .............. concavity of clasping organ. GPL gonoplac. 1 GPO . . . . . . o o . . . . f i r s t gonapophysis. 2 GPO .second " 1 GS f i r s t gonocoxa. 2 GX o o,o.. o. second n HP ......... humeral p l a t e . BWP hind wing process. J « . . . . . . . . . . . o . . o jugal v e i n . JF jugal f o l d . KEM katepimeron. L labium. LB ............... labrum. i M o o . e s . . o . . o . g o . « o medxae ME . 0 0 0 0 . 0 0 0 0 0 0 0 0 0 membrane 0 MP o o . o... o o. o . o . . . median p l a t e . MSN o . . . o . o . . . . . . . mesonotum. MOT metanotunu MXPA a . . « , . . . . . . . . maxillary plate area 0 62 IT 0 0 0 0 0 0 9 9 O 0 0 0 * 9 0 notumo 0 00000**9 0 0 * » O 0 0 ocelluso OC © o o 0 Q 0 ft 0 0 * (•OO* occiput a OCC o*oeo* 0 0 0 1 * 0 0 o c c i p i t a l condyle « OF O O O 0 0 O O 0 0 0 10 9 0 o c c i p i t a l foramen• PAC » • * • « • 0 0 0 » 0 0 0 paraclypeua* postclypeus fl PCS • 0 * 0 * * tt * 0 • 0 0 0 precoxal s h e l f 0 P G 0 0 0 0 0 0.0 » 0 * i » 0 0 9 postgena e postgenal bridge. 2 PH O • O O 0 i o o t 0 9 * phragma of mesothorax,. 3 PH 0 0 O • 0 1 * 0 * 0 0 0 ,T " metathorax* phallosoma,. PHS APP ., » o 0 e o o o phallosomal appendages. PLA • 0 • 0 O 9 C 0 0 4 0 0 0 pleurodema9 P M P O 0 O 0 O * 1 * o a 0 0 0 proximal median plate. PIT 0 O 0 0 9 O O C 0 * 0 0 * 9 pronotum. PltfC o 0 * o e « a 0 0 0 0 o * pronotal c o l l a r 0 PMP * O 0 0 0 0 • 0 0 0 0 * 0 posterior notal wing process• postocciput* P H 0 0 0 0 0 0 0 a 0 0 0 0 0 0 paramere0 X P H S o o o o• « O 9 0 « * presternum of prothorax. 2 PRS 0 0 0 0 4 0 0 0 0 0 0 " " mesothorax. PHSC 0 O 0 O 0 O 0 9 0 0 0 0 prescutum. pleural sulcus• PSCU 0 0 0 0 * 0 o e o O O 0 • parascutellum. 2 PSXJ o o o o o 0 0 0 0 0 9 postscutellum of mesothorax* 63 3 PSL ............. postscutellum of metathorax. P5H . . o . . . . . . . . . . . . postnotum. PT ................ p a r a t e r g i t e . PTAR pretarsus. R . . . . . o o . . . . . . . . . . radius. SO . . . . . . . . . . . . o o . . subcosta. 2 SCL mesoscutellum, 3 SCL . . . . . . » o . o 'metascutellum. 2 SCU 0 . . . . . . . . . . . . mesoscutum. 3 SCU . . . . . . o o . o o . . metascutum. SGO ............... scent-gland o r i f i c e , SPCD .............. spermathecal duct. ST « o « . o o . . . . . . . . o o stigma. 1 STL . . . o o . . . . . . . . sternellum of prothorax. 2 STL , . . . . . . . o o o . , " " mesothorax. T «o .,o o . o . o . o . o o . o . tergum, TAR tarsus. TB . , o o « « « » . , . a o o o , t i b i a . TN o o . o . o o o . . . o . . o o trochantin. TR trochanter, 1 V . o o . . . . . f i r s t vanal vei n . 2 V . o o . . . . . . o o . . . . second vanal v e i n . VF «.«««......os... vanal f o l d . VG ,. vagina. VS . . . . . . . . o . o . o . . . v e s i c a . VX o o . . . . o . , . . . . . . . vertex. WP o o . o o . o . . . . . . . . . p l e u r a l wing process. PNC' 

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