Comments on the Species Concept in Some Australian Anisops Spinola (Hemiptera: Notonectidae)

https://doi.org/10.24199/j.mmv.1978.39.08 7 July 1978

COMMENTS ON THE SPECIES CONCEPT IN SOME AUSTRALIAN ANISOPS SPINOLA (HEMIPTERA: NOTONECTIDAE)

By I. Lansbury

Hope Department of Entomology, University Museum, Oxford

Introduction tralian species with reliable k spof characters Twenty-five species of Anisops are recorded are stall Kirkaldy; calcaratus Hale; nasuta Fie- from Australia and Tasmania, 16 are endemic. ber; semita Brooks; and tahitiensis Lundblad. the A few species are taxonomically confusing as Of remainder, thienemanni Lundblad; no- ditlata and canaliculate Brooks are easily they occur in more than one form. Anisops is recognized by rostral compared with Bttenoa Kirkaldy which replaces abnormally shaped prongs. Anisops in the New World. The characters used by Brooks (1951) to The species concept is discussed, especially separate fall of those with an extra-Australian distribution species into two categories (1) of the capsule and the which tend to be superficially distinct at the morphometries head periphery of their range. pronotum; (2) presence of grooves, ridges prolongations of the especially Figures and short descriptions are given of or head capsule the facial tubercle. Secondary sexual charac- little known species from New Guinea which ters with combinations of the primary features may eventually be found in Australia. Finally are reworked to enable species to be keyed a new species is described from Lihir, Bis- marck Archipelago. out. The measurements of the head and pronotum are not easy to make as the structures are Species Concept nearly all convex. The chaetotaxy of the male front leg, number of pegs in the stridulatory Alpha taxonomy row and distribution of spines and setae on To understand the problems posed by vari- the front tibia and tarsus are used. The ros- ous species of Anisops, it is necessary to con- tral prongs which are projections borne either sider the species concept in the genus. With few side of the third rostral segment are also im- exceptions, only the males can be named with portant. reasonable certainty. Attempts by Sweeney (1965) and Lansbury Wing polymorphism (1969) to devise reasonably easy keys to the The terminology used to indicate if an Ani-

Australian Anisops have not been completely sops can fly or not is rather confusing. No successful. The occasional failure of my 1969 problem arises over the term macropterous, key is caused by trying to key out forms of but brachypterous can be misleading. Ail Aus- species which were not available to me when tralian Anisops have fully developed fore the key was prepared. The key couplet re- wings and hind or metathoracie wings. Despite ferring to the comparative width of the head the presence of wings, most species produce to the pronotum has proved unreliable when forms with the wing musculature undeveloped,

dealing with species which vary in the degree in its place parenchymatous tissue. Young of development of the flight musculature. (1962) uses the term 'normal' for individuals There are over one hundred described and capable of llight and 'flightless' for those which seemingly valid species of Anisops, some of although possessing full sized wings, have not them with a widespread distribution. The developed (light muscle. characters used to divide up the genus into The term 4 brachyelytrous' has been sug- manageable proportions vary greatly in their gested, although possibly quite suitable, some reliability. Structural peculiarities of the head confusion may arise as this term is also applied and rarely the pronotum easily distinguish a to the Staphylinidae and Dermaptcra with their small number from most of the genus. Aus- characteristic short elytra, a condition not oc-

101 io: r. LANSBURY earring in Anisops. To avoid confusion, the In B. lirtinocastris Hungerford and other terminology of Young (1962) is followed in species, both sexes of the flightless forms have these notes. the head as wide or wider than the pronotal

The normal forms with fully developed flight humeral width, normal forms of both sexes musculature are easily recognized by the pig- have the head narrower than the humeral mented scutellum, the clavus and corium also width. Truxal (1953) describing the wing poly- coloured, most commonly black, brown, red morphic forms of limnocastris found that the or yellow, usually a subtle combination of two scutellum of the normal form was longer than or three colours. The metanotum is black or the median length of the pronotum, that of the dark brown. Flightless forms (always with flightless form clearly shorter. These differen- fully developed metathoracic wings in Austra- ces have not been found so far in Anisops. lian species) have hyaline scutellums, the dor- Colour sal pigmentation showing through, likewise the It is the of clavus and corium. The metanotum is not pig- not known how much colour Ani- mented, but occasionally dark brown at the sops species can vary. Young (1962) found a wing bases. The following descriptions are direct correlation between colour and flight taken from Young (1962) and refer to assimi- musculature. There is some evidence that in circumstances Anisops exhibit lis and wakefieldi White, both endemic to New some a pheno- Zealand. The macropterous form of both sexes menon rather better known in the Corixidae have the lateral margins of the pronotum di- (see deanei). Specimens from turbid water with vergent. The males of the brachypterous forms a peaty or dark substrate frequently have a have more-or-less parallel sided pronotums and much larger area of the hemelytra with dark females with slightly divergent pronotums. A brown transverse markings, the pronotum is check of Brooks (1951) and Lansbury (1964) often more heavily pigmented. The same spe- shows that they often described two colour cies of corixid from a sandy or gravel substrate forms of a species but did not comment on the are much paler. The significance of colour in shape of the pronotum. A further complication Anisops where it impinges on food gathering has been found in a number of species which and its role in enabling Anisops to merge with

are isolated from the rest because the head is the background are poorly documented. The as wide or wider than the pronotal humeral differences noted by Young (1962) refer al- width, these are: evansi Brooks (Tasmania) most exclusively to the colour of the scutellum. robusta Hutchinson (E. Africa) lencothea Esa- The pale flightless Anisops with little dorsal ki (Samoa and elsewhere) doris Kirkaldy pigmentation may be difficult for a predator to (Australia) tasmaniaensis Brooks (Tasmania) locate visually, the normal form with a pig- philippinensis Brooks (Philippines and else- mented scutellum and intensified colouring of where) and assimilis. Young's (1962) descrip- the hemelytra may be easier for a predator to tion of assimilis differs from Brooks, the head see. Both forms would be equally at risk viewed

irrespective of the form is always narrower from any position sub-surface when ascending to the surface to than the pronotal humeral width, it seems to renew their gas supply or at risk from predators which use other senses be a general rule in Anisops that the lateral than sight to locate their prey. Bayly, Ebsworth and margins of the pronotum of flightless males are Wan (1975) present data from which it can be more-or-less parallel sided, therefore the taxo- inferred that within closed systems such as nomic value of the head-pronotal width is of exist on Fraser Island, Qld., that fish and limited use in those species which have more water-bugs are mutually exclusive. They pos- voluminous eyes. Series of evansi from Tas- tulate that either the fish eliminate the bugs and the mainland show both forms to mania be by feeding on them, or the fish out-compete present, where females can safely be associated the bugs for the available food supply. Young's with males, the head is always narrower than (1962) data on assimilis shows that both forms the humeral width. were found in almost equal numbers. The num- AUSTRALIAN ANISOPS 103 ber of habitats where only the normal form alike Anisops, one of the main differences be- were found was almost the same as the flight- ing that the male Buenoa has two segmented less form. The total number of habitats with front tarsi, whereas Anisops males have one- both forms present being just over double the segmented tarsus. It is felt that the behaviour of number of those where only one form was Buenoa and Anisops are probably rather simi- present. Anisops wakefieldi differed, the nor- lar. In laboratory experiments it has been mal form was ten times more common than the found that B. limnocastris males will pair with flightless and four times as many habitats were macrotibialis Hungerford females, the Fj gen- populated by the normal form compared with eration are sterile, males morphologically in- the flightless form. Both species were found termediate, their acoustical behaviour similar together in 'swarms', assimilis having a to that of their male parents. Stimson Wilcox greater tolerance of habitat types. In the more (n.d.) (1975). Truxal (1953) descriptions of the stable weedy habitats with little open water stridulatory apparatus of Buenoa species, es- wakefieldi was the dominant species. pecially the number of stridulatory pegs shows that within a species, the variation found is Genitalia less than in Anisops. Young (1962) found that The genitalia of Anisops are of very limited in assimilis males, the shape and number of value at the species level. Brooks (1951) figured stridulatory pegs was fairly constant; the size the left and right parameres but made no re- and arrangement in wakefieldi could be vari- ference to them. Young (1962) did not figure able (28 and 15 pegs respectively). The short or comment on the New Zealand species. Trux- peg row of wakefieldi, rostral prong and ex- al (1953) figured the parameres of seven panded facial tubercle would produce a dif- Buenoa species which closely resemble Ani- ferent sound than the longer peg row of assimi-

sops in general plan. The female genitalia of lis drawn across a larger rostral prong. The Anisops are of less taxonomic value than the expanded facial tubercle of wakefieldi would male, the same seems to apply to Buenoa al- greatly hinder their attempts to pair with assi- though hungerjordi Truxal differs from the milis females. Presumably in New Zealand others in the shape and distribution of large where Anisops has been geographically isolated setae on the first gonapophysis, hungerfordi for a long period and the number of species oviposits in rock crevices rather than plant small, stable populations would develop. Sound tissue as do most other species of Buenoa and, production in Buenoa is assisted by the body as far as is known, all Anisops. The figures of which in whole or part acts as a frequency gen- the male genitalia (Figs. 43-58) are slightly erator. Experimental work with Buenoa shows misleading as they tend to show differences that acoustical behaviour is important in repro- which do not exist, this is because some diffi- ductive isolation amongst coexistent species, culty was experienced in placing the parameres Stimson Wilcox (n.d.), (1975). No information planes the slides leading to on identical on is available on Anisops. Acoustical behaviour as distortion. apex of the left paramere some The an isolating mechanism in mixed populations is variable, the tip sometimes being is rather well known in other groups. Ragge (1965) curved inwardly, thus when viewed from the describing British Acrididae (Orthoptera) likely side, the tip is not visible. The right paramere is to be found in the same general habitat states a simple flat plate. that Chorthippus brunneus (Thunberg) has Isolating mechanisms 50-90 stridulatory pegs on each hind femora; C. parallelus (Zetterstedt) 70-130 and Myr- As it seems that the genitalia and the dis- meleotettix maculatus (Thunberg) 1 30-1 80 tinctions contrived by the taxonomists using the morphometries of the head and pronotum pegs etc. Ragge's diagrams of the songs of are of limited value, the problem arises of what these and other species shows that irrespective in certain circumstances constitutes a species of the number of pegs, the songs of the males in Anisops. Since Buenoa are so very much of each species remain consistent in volume H 104 I. LANSBURY

and duration. It is reasonable to assume from ditions in Tasmania tend to be rather different this that Anisops males only utilize sufficient from those on the mainland with its greater di- pegs to produce a consistent species signal. versity of species. The carrying capacity of This factor also tends to reduce the taxonomic habitats, especially species diversity rather value of minor variations in the number, size than the total population of water-bugs etc.

and arrangement of the stridulatory comb. is not too well known. In Tasmania, out This variation seems to be a feature of a num- of 175 samples, only one habitat had eight ber of nondescript species with an extensive species of water-bug present (Blackmans geographical distribution. Lagoon). Three habitats with six species Assuming that part of any population of an and six with five. The great majority of Anisops species living in ponds, dams and samples composed of two-three species. These ephemeral pools contains a percentage of in- figures may reflect vagaries of sampling rather dividuals with fully developed musculature, it than an accurate reflection of these habitats. is essential that should conditions deteriorate There are 24 species of water-bugs in Tas-

sufficiently to induce flight, the migrant popu- mania, 28 if the Saldidae, Ochteridae and Ge- lation should produce and recognize acoustic lastocoridae are included. signals from the same species in the habitat Fraser Is., Qld. has been studied in some de- emigrated to, otherwise discrete populations tail (Bayly et al.> 1975). Four species is the would form in the same habitat. Exceptional maximum recorded from Boomerang South habitats such as Lake Tengano, Rennell Is- Lake, 13 lakes sampled. Bensink and Bur- land only seem to produce flightless forms ton (1975) on the littoral fauna of Blue see 'capitata'. and Brown Lakes on Stradbroke Is., Qld. found five and nine species respectively. Brown Anisops elstoni, deanei and philippinensis Brooks Lake being sampled more often than Blue Lake. Some of the Anisops listed in Bayly et Under elstoni material from various locali- al. (1975) and Bensink and Burton (1975) are ties in different countries is compared. With discussed in the taxonomic section of these one or two exceptions nothing is known about notes. the type of habitat, population density, species The problem arises of deciding whether the diversity or basic chemical and physical pro- material from Tasmania is conspecific with perties of the habitats where elstoni was col- those from mainland localities in Australia and lected. The genetic aspects remain totally un- further north in New Guinea etc. known. As more material becomes available it has become increasingly difficult to confine the The data can be interpreted in one of two ways, elstoni species to its original concept which, lacking may be a complex of races secondary sexual characters, reproductive or- which because of geographical isolation are in gans indistinguishable from other species and the process of evolving into distinct species or where the morphometries are complicated by it may be a widespread 'plastic' species which structural changes caused by wing polymor- has spread from south east Asia into Austra- lasia, each population still linked phism. The discontinuous distribution of el- by common genetic stoni and other species suggest that the species factors. concept is not fully understood or the term Anisops deanei is confined to Australia and 'super species' might be appropriate. a closely allied form or species is rather com- In Tasmania which has five species of Ani- mon in Tasmania, much more so than elstoni. sops, unpublished data shows that where two Non-Tasmanian deanei varies rather more or more species are found in the same habitat, than elstoni. In Tasmania they are both some- one at least is easily recognized by its larger times found in the same habitat, but do not size and/or secondary sexual characters leav- seem to be so on the mainland. ing the remaining species to be easily recog- Where two or more species occur in the nized by their lack of secondary features. Con- same habitat, acoustic behaviour, phased life —

AUSTRALIAN AN1S0PS 105 cycles and a subtle niche preference not so far revealed by sampling seem to enable Ani- sops species to coexist and avoid competition for the available food supply. |

Habitat and distribution

Lack of data makes it difficult to be pre- cise about the habitat preferences of Austra- lian Anisops species. Sweeney (1965) describes elstoni as rare and confined to the south east, the most westerly record being Narrandera, N.S.W. Most Australian Anisops are found along the eastern coastal zone, a few are confined to the more arid interior. Some species typified by thienemanni, stali and Hyperion Kirkaldy are found across the continent. The coastal zone from Brisbane northwards has a number of species with prominent secondary sexual characters and not related to the New Guinea fauna. Species of the more temperate areas are less well endowed with such features. Anisops philippinensis appears to be a relative newcomer to the Queensland fauna rather like tahitiensis Sweeney (1965). Taxonomy Anisops elstoni Brooks, 1951

Figures 1-15, 43-50 and 59-61 Figures 1-3 Anisops elstoni Brooks

S.A.: 1, front leg. 2, head from side. This species was described from Myponga, 3, head and pronotum from above. South Australia; The Dorrigo, 3,00(y, New South Wales; Brisbane, Queensland and Szech- scutellum which is carmine. Metanotum dark uan, Suifu, China. Brooks compared elstoni reddish brown, tergites black, posteriorly mar- with exigera Horvath (New Guinea). His gined with yellow. Sternites other than keel figures of the male front leg are misleading as black. he omits to include the large spine on the A small series mostly females from Caddies inner proximal margin of the front tarsus Creek, N.S.W. vary between the colour of the (Figs. 1, 4, 7, 10 and 13) and he overlooked it Charters Towers male through to the pale yel- in his description. Fortunately, he mentioned it lowish form. The hemelytra are hyaline, met- in the key to species. He does mention the pro- anotum pale yellow. Pronotal shape the same notal depression which is helpful in isolating regardless of colouration. A series from Valley elstoni from related species (Figs. 3, 6, 9, 11 Heights near Katoomba are a uniform grey- and 14). The material studied has been found ish black. The lack of pigment may be be- to vary from various localities in Tasmania, cause they were kept in alcohol for an un- Australia, New Guinea, Vietnam and possibly known period. The eyes appear to be rather Rennell Island. more elongate than usual and give the im- A male from Charters Towers Qld., Coll. pression of being wider than the humeral Sedlacek (Bishop Museum) has a bright red width, they are consistently narrower, a ratio scutellum. Eyes orange-brown and pronotum of 41:43. A similar phenomenon is found in ( straw yellow. The hemelytra are hyaline ex- the Fraser Island population from Lake AB' cept for the inner claval margin adjacent to the (Fig. 9). The rostral prong of the Fraser Is- — —

106 I. LANSBURY

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Figures 4-6 A. elstoni upper reaches of

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13-15— 1 Figures 10-12—.4. elstoni

elstoni, especially the form from Fraser Island. Limnogonus rennellensis Brown (Gerridae) to In Brooks (1951) key it keys out with rigoen- a subspecies of L. fossarum F. a widespread sis and biroi Brooks, but does not seem to be Indo-Australian and Pacific species, capitata related to them. In Lansbury (1969) it keys is almost certainly at least in the same category. out with evansi and doris because the head is wider than the pronotal humeral width. Like Anisops philippinensis Brooks, 1951 elstoni, the pronotal disk is depressed. The Figures 19-29, 53 and 54 specimens collected by E. S. Brown from Lake Tengano are rather more yellowish than those Described from a series of 90 specimens obtained by Torben Wolff from the same from the Philippine Islands, Mindanao, Lake general habitat which are relatively paler, the Linao, North Slope, Mt Apo Davao Province, hemelytra are hyaline, the costal margin near 7800'. H. Hoogstraal and F. G. Werner. Most the pronotum being infuscated smokey brown. of type series in Field Museum, Chicago. The colour approaches that of the deanei popu- Brooks compared philippinensis with tahitien- lation from North Stradbroke Island. sis, the latter is easily recognized by the carin- Although treated as a distinct species, it is ate frons. Brooks commented that the scutel- very probable that capitata is a brachypterous lum was either testaceous or black with an- form of elstoni. Andersen (1975) relegated terior margin testaceous. Because of the varia- — —

108 I. LANSBURY

hemelytra hyaline. There seems no doubt that this male is a flightless form of philippinensis. The minor variations in the chaetotaxy of the front leg are not considered significant, especially as the paratype(s) do not agree with the figure by Brooks. A pair from N.E. New Guinea, Wau, Mo- robe District, 1100 m., 9.ix.l961, J. Scdlacek (Bishop Museum) (Figs. 22-24) are rather more like the form from the Philippines although the chaetotaxy of the front tarsus is dissimilar. The head is 7-5x anterior width of the vertex, Brooks gives the head width as lOx anterior width of vertex.

Anisops deanei Brooks, 1951 Figures 30-32

A small series from North Stradbroke Island, Brown Lake, September, 1972, G. Monteith are quite unlike the usual form of deanei (Lansbury, 1964). The eyes are dark brown- black. The head between the eyes, facial tu-

Figures 16-18 A. 'capitata' Lansbury

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110 I. LANSBURY

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Figures 25-29—A. philippinensis <£ Fraser Island,

Qd. } Boomerang South: 25, front fe- mur. 26, tibia. 27, tarsus. 28, head and pronotum from above. 29, head from side.

Figures 30-32 A. deanei Brooks

StftI; Gerridae Rhagodotarsus sp., Limnogonus specifically excludes biroi and rigoensis from the re- fossarum (Fabr.). With the exception of Micro- maining species by the absence of setae on the front tarsus. The rostral prong of rigoensis is rather vari- necta, all the water bugs listed are entirely pre- able (Figs. 34, 36). Chaetotaxy of the male front leg datory. Bensink and Burton (1975) comment (Fig. 33). that deanei readily fed on Chaoborus in the laboratory as do the fish in Blue Lake which has a low species diversity. Anisops biroi Brooks, 1951 The colour of deanei from Brown Lake Figures 37-39 which is described as humic brown due to ac- Small species (Brooks states males 4-2-4-3 cumulated dissolved organic matter does sup- mm long, maximum width 1-2-1-3 mm). port the suggestion that the colour of the water Original description based on 3 & 2 9 from and substrate influences the colouration of the New Guinea, Seleo Berlinhafen. '96 (Biro). Anisops species present fish although predators Type series in the Snow Entomological Collec- were absent. Potential predators of Anisops tions, Lawrence Kansas. would include nepids, other notonectids, Co- Material examined; lc? 1$ paratype; male 5 mm leoptera and Odonata nymphs. long, female 4-9 mm long. By courtesy of the Snow Ent. Coll.

Brook's description is brief but adequate to distinguish biroi from rigoensis. The synthlipsis Anisops rigoensis Brooks, 1951 of biroi is wider than that of rigoensis and the Figures 33-36 male examined lacks a carinate vertex. The chaetotaxy of the front leg (Fig. 37) hardly Small species 4-4-5 mm long, maximum differs from rigoensis. The rostral prong is a width 1-1-3 mm. little longer, Brooks states that the prong of Originally, description based on specimens biroi is at least as long as the third segment from New Guinea: N.G., Rigo, Luglio [July], whereas in rigoensis the prong is shorter than 1889, L. Loria; N.G., Lemian Berlinhafen the third rostral segment. Comparison of figure 1896, Biro and N.G. Friedrich-Wilh.-hafen, 38 with figures 34 and 36 do not wholly sup- 1896, Biro in the Snow Entomological Collec- port Brooks diagnosis. The eyes of rigoensis tions, Lawrence Kansas. are larger (Figures 34-36) than those of biroi (Figs. 38, 39). Material examined, lc? 1? paratype N.G. Rigo, Anisops rigoensis and biroi are similar in Luglio, 1889 and lie? 1? New Guinea: Neth. Genjam 40 km west of Hollandia, 100-200 m; l-10.iii. T.C. general appearance, chaetotaxy of the front Man by courtesy of the Snow Ent. Coll. and Bishop legs and shape of the rostral prong. The only Museum respectively. real difference is that rigoensis has a feebly

The description given by Brooks is rather confus- carinate vertex and a very narrow synthlipsis. ing. The two paratypes do not agree with the original Because of a lack of data on other species, it description. The synthlipsis of the male is not 1/5- is not possible to comment on the differences in 1/3 anterior width of vertex as stated by Brooks, eye size between these two species. but 1/9-1/12 anterior width of vertex. The vertex is feebly carinate, commencing at the synthlipsis and extending about 1/3 median head length between the Anisops lihiriensis sp. n. eyes. The development of the carina is rather variable and difficult to see. Brooks gives a muddled descrip- (Figures 40-42 and 51, 52) tion of the morphometries of the head and prono- Males, 5 5 long, maximum width 1-75 tum. The ratios in the series examined have the mm head length about the same as the median pronotal mm. length, pronotal humeral width just under twice the Colour, eyes grey, vertex, pronotal fovea and median length. Brooks comparing biroi with rigoensis pronotum yellowish white. Scutellum basally states that rigoensis has three small setae on the in- broadly brown, apex yellowish white. Hemely- ner surface of the male front tarsus, they are not shown in his figure and in his key, couplet 93 he tra hyaline, claval margins and hemelytral com- —

112 I. LANSBURY

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Figures 33-36—,4. rigoensis Brooks 6*, 33-35, paratype. 36, Genjam, N.G.: 33, front leg. 34, head from side. 35 and 36, head and pronotum from above.

Figures 37-39 A. biroi Brooks

AUSTRALIAN ANISOPS 113

Figures 40-42 A. lihiriensis sp. n.

missure pale red. Abdomen dark brown to On the same island on the 13.vi.1965, A. W. black, legs pale yellow. Originally preserved in Sweeney collected a long series of tahitiensis. alcohol. This species has spread over much of the Indo- Australian Pacific region. Originally described Structure, greatest width of head between from Tahiti. It is common in the Hebri- ten and twelve times the anterior width of New des, Guinea, Solomons, etc. the vertex, narrower than pronotal humeral New Malaya Sweeney 1 recorded tahitiensis from width. Synthlipsis about half the anterior width ( 965) Queensland. Anisops nasuta Fieber has a of vertex. Median head length slightly shorter than the pronotum. Humeral width about two similar distribution although it does not extend so far across the Pacific. Known from Tonga and a half times median pronotal length. Fa- Is. (Nova Pou Is.) Hebrides, Cale- cial tubercle prominent. New New donia, New Guinea, Solomons, Malaya, Java, to couplet in Keys out 53 Brooks (1951) Bismarck Archipelago and northern Australia. similar to philippinensis and windi, lihiriensis The first gonapophysis of the female (Fig. 62) differs having an inflated facial from them by drawn to the same scale as figures 59-61 shows tubercle, the rostral prong is rather stouter that although nasuta is larger than elstoni 6- tibia have and the front does not the setae char- 7*5 mm compared with 5 mm, the gonapophy- acteristic of these species. In Lansbury (1969) sis is not significantly larger. lihiriensis keys out with deanei, the same differences apply. Parameres (Figs. 51, 52). Acknowledgements Holotype 3 , 2 $ paratypes: Bismarck Ar- chipelago, Lihir, Put Put, pig wallows, 1(X dia- I wish to thank Dr P. S. Lake, Zoology De- meter, bottom muddy, water opaque 3" deep, partment, Monash University, Victoria for the 30.vi.1965, A. W. Sweeney (Oxford). loan of many samples of water bugs from Tas- 114 I. LANSBURY

References ' Andersen, N. M., 1975. The Limnogonus and Neo- gerris of the Old World. Entomologica Scan- dinavica. Supplementum 7: 1-93. ' Bayly, I. A. E., E. P. Ebsworth « i and Hang Fong Wan, 1975. Studies on the lakes of Fraser Is- land, Queensland. Aust. J. Mar. Freshwat. Res., 26: 1-13.

Bensink, A. H. A. and H. Burton, 1975. North Stradbroke Island a place for freshwater inverte- brates. Proc. R. Soc. Qd. 86(7): 29-45. Brooks, G. T., 1951. A revision of the genus Anisops ( Notonectidae, Hemiptera). Kans. Univ. Sci. Bull. 34: 301-519.

Lansbury, L, 1962. Notes on the genus Anisops in Bishop Museum. Pacific Insects. 4: 141-151.

, 1964. The genus Anisops in Australia (Hemiptera: Notonectidae). /. ent. Soc. Qd. 3: 52-65.

, 1968. Notonectidae (Hemiptera-Heterop- tera) of Rennell Island. The Natural History of Rennell Island 5 (Zoology), pp. 95-98.

, 1969. The genus Anisops in Australia (Hemiptera-Heteroptera Notonectidae). /. nat. 55 57 Hist. 3: 433-458. 56 58 Ragge, D. R., 1965. Grasshoppers, Crickets and Cockroaches of the British Isles, F. Warae, Lon- Figures 43-58 Anisops parameres. Figures 43-50 el- don. stoni; 43-44 myponga; 45-46 Sorrell Stimson Wilcox, R. (not dated). Abstract Acou- River; 47-48 Stradbroke Island; 49-50 stical behaviour, sound producing-structures and Fraser Island. Figures 51-52 lihirien- biology of Buenoa (Hemiptera, Notonectidae).' sis. Figures 53-54 philippinensis, Fra- Thesis, Department of Zoology, University of ser Island. Figures 55-56 deanei, Michigan, Ann Arbor, Michigan, 271 pages. Stradbroke Island. Figures 57-58 Nasuta, Fraser Island. , 1975. Sound producing mechanisms of Buenoa macrotibialis Hungerford (Hemiptera: Notonectidae), mania. Dr Int. J. Insect Morphol & Embriol. A. Neboiss for the loan of material 4(2): 169-182. from Blackmans Lagoon, Tasmania. The Ber- Sweeney, A. W., 1965. The distribution of the No- nice P. Bishop Museum, Honolulu for the tonectidae (Hemiptera) in south-eastern Austra- loan of important material. Dr G. Byers of the lia. Proc. Linn. Soc. N.S.W. 90(1): 87-94. Snow Entomological Collections, Lawrence, Truxal, F. S., 1953. A revision of the genus Buenoa (Hemiptera Kansas for the loan of specimens from the Notonectidae) . Kans. Univ. Sci Btdl.35: 1351-1523. type series of species described by G. T. Brooks. Young, E. C, 1962. The Corixidae and Notonectidae Lastly Mr A. W. Sweeney for the gift (Hemiptera-Heteroptera) of New Zealand. Rec. of material from the Bismarck Archipelago. Canterbury Mus. 7(V): 327-374. —

AUSTRALIAN ANISOPS 115

59 .

l . A i 61 62

Figures 59-62 Anisops gonapophyses. Figure 59, elstoni, Myponga. Figure 60, elstoni, Sorrel River. Figure 61, elstoni, Fraser Island. Figure 62, nasuta, Fraser Island.