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Adv. Odomtol. 1: Review 251-265 of December reproductive 31, isolating barriers 1982 in Odonata K.J. 1949 Hickory Premating and habitat, ences tive in a as researched, and tactile tandem type ence of Groups mental on idae, and mating has not been dragonflies which not exist Polythoridae, the isolation are is need to be effec- sense of in interpreted some barriers have not been have been reported. actual data. Studies and genitalic expected Gomphidae, classical on for functions of the which sense. experi- Macromiidae, various are the exist- nongenitalic but Aeshnidae, and Megapodagrionidae, Pseudostigmatidae, Protoneuridae. The exact still been stimuli in Anisoptera isolation in both and Separation of potential isolating barriers far outweighs ethological important in the demonstrated. has Differ- exchange. gene most isolation, (nongenitalic union) importance of tactile studies do duliidae, in on mechanical in the comprise several hybrid although the stimuli Odonata involve temporal, interspecific of mechanical isolation. Postmating Speculation needed to United States 35630, barriers in isolating order. Mechanical genitalia, during groups Florence, A1 barriers within the incompatible mates reproductive ethological visual barriers Avenue, Tennessen Cor- Platystict- structures used clarified. INTRODUCTION The essence interbreeding such groups of the biological populations (MAYR are 1970), prompts a by was devised term, reproductive cause it of anism. Knowledge closely related species pered gene isolating does not imply derstanding animal isolation The term by DOBZHANSKY (1937) which interspecific an the of exchange a is major to is function barriers exact animals groups from how is encompass the the does the isolating does not exist. especially stumbling more used in this as in block of other isolating mechan- prevented; barriers (RIBs) evolved communication, studies; isolated fundamental question: interbreeding between species prevented? ism that concept, species reproductively the ways current paper be- term mech- majority of Difficulties in un- insects, have hamhas been how iso- 252 K.J. lating barriers arise of (modes Tennessen and speciation) the roles of relative natural selection and chance in these processes. Given taxonomic status of the Odonata, the advanced size and atively large ease of recognition in the field, a their rel- fair knowledge of RIBs in dragonflies would be expected. Despite considerable infor- mation reproductive on needed, was there of this review poses odonate RIBs, to gaps in briefly to are identify in guidelines some JOHNSON’S review (1962) of which he pointed out large are and behavior, isolation in reproductive of the group. The pur- summarize present knowledge is where study areas the approaching where research areas knowledge of study needed, RIBs in and to on give dragonflies. STATE OF KNOWLEDGE All RIBs. in published Almost studies nothing I reviewed Odonata, probably because dragonflies and will (1965) The not show promise study of where groups crosses are of ity mate when readily that RIBs postmating could postmating from yet barriers eggs known be to interspecific copulations two species of be especially in produced rewarding occur, 100 percent Odonata. WAAGE (1975) operative as mortal- interspecific by in copul- demorsa, the only example of based on his assumed observations of and the absence of obvious hybrids between Calopteryx. Discoveries of possible hybrid I) further point to the need for studies in this (Table techniques frequently JOHNSON (1975) reported hatching mortality hybrid difficult to hold captive JOHNSON’S attempts ations between Ischnura damula and I. premating damselflies may be cultured. some tandem with postmating mechanisms are confined. at least interspecific possible. larvae concerned were is known of possible specimens area. PREMATING ISOLATING BARRIERS The cates dominance of its potential mates, and barriers in vision among importance indeed this addition to a as appears to optical JOHNSON (1962). In the of RIBs, the examples are senses operative. are also following are of dragonflies of recognizing not intended ation of the literature, but rather erate. means be stimuli nised by given the On conspecific the other hand, operating, as discussion of the as a indi- recog- types complete compil- used to show how the RIBs op- Reproductive isolation in Table A list of Kiauta dragonfly species 1967; Asahina reported to have 253 Odonata I individuals hybrid produced (main sources: 1974) GOMPHIDAE GOMPHIDAE Gomphus fraternus fraternus X G. G. externus extemus 9 Calvert Calvert 1901 1901 Gomphus X G. graslinellus G. graslinellus (J � Williamson Williamson 1903 Aeshna confusa confusa X A. A. <J � Calvert Calvert 1956 19S6 Anax imperator X A. A. parthenope parthenope 6 � Bilek BUek 1955 Anax nigrofasciatus Anax nigrofasciatus X A. parthenope A. parthenope julius julius 66 <J<3 Hiura Hiura 1968 1968 Anax nigrofasciatus nigrofasciatus X A. parthenope julius julius 99 Hiura 1971 Anax nigrofasciatus nigrofasciatus X X A. parthenope julius �� julius lividus lividus AESHNIDAE AESHNIDAE diffinis diffinis 66 Asahina Asahina 1974 1974 LIBELLULIDAE L1BELLULIDAE X L. intacta Intacta � 6 Tennessen Tennessen 1981 1981 X X L. luctuosa luctuosa larva Wilson Wilson 1920 elatum elatum X S.e. eroticum 66 9 Asahina Asahina 1974 pedemontanum elatum Sympetrum pedemontanum elatum X S.e. 5. e. eroticum 9 Miyazaki 1972 Sympetrum Sympetrum baccha matutinum X X S.e. S.e. eroticum eroticum 6 � Yamamoto 1965 Sympetrum Sympetrum risi risi X S.e. eroticum eroticum 5 Asahina Asahina 1974 1974 Bilek BUek 1963 1963 Leucorrhinia glacialis Libellula pule helia pulchella pedemontanum Sympetrum <sd9 <J<5V COENAGRIONIDAE 6 � Coenagrion pulchellum X C. puella puella civile Enallagma Enallagma civile X X E. E. carunculatum carunculatum 12 different SUMMARY: SUMMARY: species species pairs involved, 24+ 66 �� Williamson 1906 1906 specimens. Temporal isolation. The two modes of temporal isolation, i.e. differences in and son diel adult mating reproductive An can lier example overlap appear be to disappear seasonal related species few cases would have of G. where , wherein adults of G. antilope they (unpublished differences through time and been carried out, the North Ameri- furcillata appear flight antilope adults G. of The literature do ear- sea- not furcillata is replete with relevance to Data relative abundance maturation, including frequencies ignored. on conspecific and heterospecific encounters, are inter- Odonata. although species, reproductive prevent adults before observations). between has on mature G. sea- during the species 1940). Although sympatric, are reproductively isolation studies a effectively flight the time of documented in been (GLOYD in nonoverlap of seasonal isolation may exist in ductive of a Gomphaeschna than those sons of However, genus involve readiness. Obviously the absence of mating activity breeding. periods, the importance of are season repro- needed. Until such in Odonata repro- 254 K.J. Tennessen ductive isolation remains unknown. On diel a basis, an species of Corduliidae agrionid by congeners though of E. females two ity periods of were in the be day The isolated partly (TENNESSEN coen- its from al- 1975), observed being taken into tandem by males MAY (1980) pollutum. to appears earlier mating isolation between two offered by PAULSON (1973). dubium Enallagma of temporal example was differences in the daily activ- reported four species of Micrathyria, although much overlap oc- curred and (1974) showed distinct temporal differences in three damselflies, but the exact species isolation Temporal mating is probably lead changes to in widespread not Variation overlap function secondarily appear to have evolved not to be a Odonata, in breeding directly RIBs in as such as per- amongst individuals another barrier would be necessary. Temporal case probably the isolation appeared study the smaller species. relatively inefficient. environmental which in it is and on pressures MIZUTA’s given. not were congeneric predation haps because and not were response to times mating in times, differences some groups, (PAULSON but 1973). Habitat isolation Often to referred ulation of sympatric differences may be be swarm-feeding of tinctly species separate is initiated; exceedingly (CORBET habitat. or microgeographic isolation, differences in the microhabitats where cop- simultaneously present types ecological as habitat isolation involves small. during in terms of certain even though adult microhabitats, may occur larvae behavior species such activities, 1963), but mating Conversely, these distance, Adults of related in may may as may mixed different live in dis- obliterate the difference. Isolation an due of recognition ethological specific males effected in via a habitat differences results barrier. differences involve placed to which prompted and in because However, behavior responses to habitat a special classify it hetero- (i.e. isolation and is as does not between case stimuli), extrinsic to isolation encounters during females, it represents specific from adult behavior, GARRISON (1979) is therefore distinct category. Groups in which evidence of habitat isolation has been found include phus Enallagma (BYERS (JOHNSON 1962), (TENNESSEN 1939), 1966, 1975, GARRISON Tetragoneuria (TENNESSEN VERDONK Somatochlora (TAKETO isolation is an inefficient barrier, 1979), 1960). as the 1979), Progom1977), Ischnura Leucorrhinia In these potential (PAJUNEN examples, habitat for individuals to Reproductive transgress habitat encounters. The “boundaries” is great, making likely interspecific results of JOHNSON (1966) experimental that habitat suggest 255 isolation in Odonata changes could cause a strongly breakdown of segregation. This breakdown necessitates the presence of another RIB. Ethological isolation Several elucidated. mates) acts In meet that (based examples of ethological on (1) these but elicit mating is A wide Visual stimuli. found to evoke others described in 1963; CONSIGLIO HOLTZ they ing are in cused displayed on their role been and are groups, but (CORBET in which and ways in one pair-form- study has fo- isolation reproductive BUCH- ( Hemiphlebiidae critical only HEY- 1961; Platycnemididae Color patterns in has been Calopterygidae 1973), Chlorocyphidae 1966), and dragonflies, families: to the female many of these in the means be achieved. Courtship JOHNSON 1955; 1974), 1913). can in response discovered. HEYMER 1956; (TILLYARD “recognition” Zygopteran 1951, 1973; WAAGE four basic are variety of visual stimuli have to be four (BUCHHOLTZ MER There the mating bound are (potential individuals intrinsic differences by response. by which species isolation have been behavioral prevented the copulatory the senses) or heterospecific barriers, (WAAGE 1975). Studies and nition” ITO 1960; of only large, dimorphism and the one of acquisition shown a mate that in important are PARR & PARR courtship “recog- sex (JACOBS 1974). However, I (PAJUNEN family 1964). He 1955; am demonstrating visual isolation study diverse have Libellulidae on sexual aware in this concluded that males of two similarly colored Leucorrhinia species females by recognize females appear sight stimuli libellulid to prior genera and color-pattern differences in abdominal recognize to but they by come to mind differences in exist which striking amongst color the species Libellula, Micrathyria, Rhyo- have not been studied. JOHNSON (1975) proposed that male-like andro- morphs within Ischnura offer increased reproductive lation, as he and tandem attempts. Numerous other ( Celithemis, Erythrodiplax, themis), males size, undetermined found male mating preferences in isotwo 256 K.J. species. Many barrier. in likely to be and UV and eyes experimental ocelli of differ- as discrimination, are patterns of ultravioseveral species receptors work such cues, form have been found in 1979), compound for and species. for study species visual detect not Enallagma two discovered. Distinctive reflections need behavior flight BERGLIED in Other did (1979) in males by contain polymorphic groups possible ences let GARRISON However, preferences morph this Tennessen have (MENZEL (SIL- been found The 1979). these various possible on modes of visual isolation is obvious. (2) Tactile uli stimuli. function Experimental RIBs as males of Lestes having and Ischnura experimentally NESSEN (1975) by was havior in ERSON (1982), in the altered of males by These using from respect of experimental on from a under 1970). female. in Tactile tial mates. the differences contacting a stimuli series of signals step In the The in study to (BARTH differences some in tac- or arise rejection groups are just exchanged between poten- Odonata studied appear one of the organ the behavior, which evokes acceptance the shape. organ in particular species one with In studies of other insects, the in ap- mechanism described DOBZHANSKY similar bethat differ in isolation the shape be 1968; approach superior appendage tactile the tactile even and ROBERTSON & PAT- though manner, TEN- provided of Enallagma stimuli consist of males “using” specific copulate appendages. evidence of tactile recognition females based examples Fewith shape of these appendages an other groups of insects. may LOIBL by release copulatory the species. found strong Enallagma, by refused to females of two species specific differences tile tactile stim- concluded that tactile stimuli pear to be isolating a presented the male superior appendages basic that and KRIEGER & KRIEGER-LOIBL (1958). (1958) males evidence first arise thus far, from the however, shape of the organ(s), which is detected and discriminated by females. This hypothesis involves behavior and merits classification appears to serve as be the as an very ethological effective, ultimate barrier. Tactile isolation and premating in some RIB odonates may should other de- Reproductive isolation in 257 Odonata vices break down. Olfactory (3) (4) and substances sound communicative signals in effort these stimuli. Auditory nor waves in is areas have Odonata, almost visual acuteness of dragonflies to the approaches long-range ity of fact study females detecting material may bond. Is material contain this be a on the these determined. problems proaches, To my such summary, interspecific between superior has (see ata BICK & may exchange, related easily BICK the be isolated by there and could or the could it are of Enallagma function has devised, and may be other ap- of sound detection in may few not solving be rewarding. investigated. thoroughly to for procedures possibility been not species thought 1981). their chemical analysis, as that the tandem fluid, appendages 1975) but not a that appear to be secretory ethological isolating barriers gene closely are knowledge, dragonflies In Cells seminal Experimental as reported mesostigmal plates phenomenon displaced males (TENNESSEN been female function in strengthening dried, in rule out chemo- the chemical and producing the common a not species of Argia. He postulated pheromone? found were be weeks (any phero- even WILLIAMSON (1906) it. left males of several the or dissipated) does males may white substance by attention. The deserves attempt tandem when presented females would have as of the dominated unlikely, the possibil- seems that have been dead for days reception as research Recognition has obviously chemoreception that males mone nil. implicated although of their biology. And although chemoreception contact Neither chemical been effectively prevent reports of copulation ethologically isolated The majority of cognate species of Odon- ethological means, with visual and tactile dif- ferences playing the major roles. Mechanical isolation Also called the “lock-and-key” under much criticism, withstood strated scrutiny. convincingly ated that in as hypothesis, few supposed examples this in Incompatibility of genitalia has in any odonate idea has come the Insecta have not been demon- species. WATSON (1966) specul- Tramea male hamules interlock with the female genitalia; JOHNSON’S study (1972) showed that these structures in Hagenius 258 K.J. Tennessen support the penis probably on lock a and key how the genitalia during and held engaged are in tandem because appendages of CORBET (1963) dence of this type of mental evidence of supportive the Based I studies, isolation species, on genera into tandem upon specific males, performed “refusal motions” and had no 1975). When presented “difficulties female from a from a them. PAULSON interpreted isolation. on to the “difficulty” inappropriate stim- Separation of such pairs gives the contact stimuli mechanical barrier when different type of mechanical ap- behavior and isolation may be using WAAGE’s discovery (1979a) that males the female’s would work but bursa as copulatrix follows; a A, sp. B; barring sperm of the sp. A remove transferred. The hypothesis the previous remove sperm fertilizing The least of eggs hypothesis tandem previous is taken of isolation, hypothesized sperm stored in remove The matings. A mates with into tandem by copulation is the (see also following some species last male; no sperm would not male’s sperm apply the facilitate sperm productive a barrier male of male of or length from the sp. B male if a are male does not have in order to transfer his own be- male to mate appear to have precedence WAAGE 1979a for would not hold for copulation. If this discussion and those species hypothesis is in references). ovipositing in for at correct pairs, it would fit the classical definition of mech- anical isolation in that genitalic incompatibility much of a attempted; the intromittent organ is not of the correct shape however, cause she other type any from female of species before oviposition, sp. to the However, involved. A to hetero- released (TEN- were fitting of the male appendages female discrimination based heterospecific male. pearance of are mechanical as Females by heterospecific females, males dead clasping associated with the thorax” could arise uli in securely difficulty and has been inter- tactile stimuli. being taken experi- observations what that involve may presented that he considered own my believe provide evi- to considered coenagrionid is preceding union JOHNSON (1962) isolation. PAULSON (1974) hypothesis. mechanical NESSEN on how do hamules females with anal clasp differences). cited several studies other of two Enallagma securely to anatomical from several of understanding as needed in Anisoptera linkage, the nongenitalic copulation (i.e. inability of males preted is together (i.e. Another mechanical barrier which has been proposed function?). incompatibility and translocation, casting doubt sperm More work operation. is involved. However, morphology of the odonate penis probably evolved displacement isolation may be (WAAGE only an 1982), and any resulting occasional consequence. to re- isolation in Reproductive 259 Odonata BREAKDOWN AND REINFORCEMENT The above premating how interspecific riers fail ing in gene is unknown. sequence, RIBs is habitat differences may isolate two sympatric poral or most part; if visually and avoid mating. inefficient, a tactile mechanical or these barriers would result in mating RIBs, zygote or such hybrid bridization. increase in specific mate. foregoing selection example, males distinguishable. and sites, to attempt in these tandem chance of obtaining wered questions speciate?). isms riers arise into come little or are supportive possible tropic may benefit The biggest anisopteran are also be female, such been evidence has a favor females are those versus this as furtive (i.e. unans- how do organ- that isolating bar- incipient species 1978), been found (PATERSON 1978). other Further gene exchange evolved incidental or very It of studies of is pleio- functions (ROBERTSON advancement population-genetic are for the increasing stem from or near 1975, WHITE to an con- advantageous more thereby postulated 1970, BUSH & speciation theory isolating barriers 1981). gap in our knowledge and effectiveness families Aeshnidae, needed females it (MAYR to heterospecific how RIBs originate has securing seem via natural selection after of prevent hy- phenomenon in to perfected from importance would may CONTINUING THE QUEST the may rare Although 1982. (TEMPLETON a devoted Nondescript mate. Problems a consequences post- case different groups, females may with any that many barriers PATERSON, or female. However, owing to dif- groups it concerning contact absent Failure of sequence, there appears to be and energy ferent sexual selection pressures in breeding inferiority or discriminate against visually are 1981). time of For stimuli interspecific mating, in which those that attempt to mate with any be tem- species for the intervene. may mating is apparently in the wastage If visual RIB mortality, sterility Interspecific With each step not operat- chromosome incompatibility (no fertilization), as Odonata (BICK & BICK males that as bar- heterospecific individuals meet, they may “recognize” differences may should supposed reinforcing another. Accordingly, one and 100% effective, not prevented Nevertheless, RIBs have been viewed series, or probably are exchange on of - STUDY NEEDS of RIBs ethological Gomphidae Cordulegasteridae, in Odonata barriers in concerns the large and Corduliidae; studies Macromiidae and Synthemist- 260 K.J. Observations idae. these groups ly has to the entire mating due to tandem is achieved. these general, hyaline uli and is a belong made. once tandem pairs involving (BICK & BICK groups in to which the 1981). wings the species, RIBs might and in these much be are stimonce aeshnids, indicating that tactile the effective macromiids barriers. Male uni- remarkably are in libellulid genera, and discover- as is groups number of large a shaped, uniquely are cordulegasterids form amongst in appendages corduliids of appendages the of prior unreliable, and that isolation is effected or Male mechanical differences and/or number sexes from the site courtship behavior is unknown, evidence that visual gomphids and ing However, in species nonexistent. Rare- within these groups flight, and disappearance species nonexistent are contact are for any species sequence individuals have been collected heterospecific In closely-related data the unknown whereabouts of the their rapid meeting, of behavor mating on scanty, and experimental are been observed, Tennessen especially Much challenging. more study of Libellulidae is needed. Very ferences exist coveries await in the well as world. Acanthagrion, is knowledge undertake studies of large and Argia available. Epallagidae of in genera Ischnura neo- color dif- and dis- Polythoridae, Platystictidae and Protoneu- and Chlorocyphidae are and Intriguing problems Pseudostigmatidae, the There species. who investigators as behavior and isolation of on which many unusual structural between related Megapodagrionidae, ridae, published little has been tropical Zygoptera, for other parts of Coenagrionidae, the which little very such as behavioral Much remains to be learned in the Caloptery" gidae. ' The terms used by phological be structures directed minimize clarify and in not uniform terminology difficulties. importance of each and mor- and confusion needed document to T. Takeuchi on are being groups. Fe- possible barriers contact published work studies although two are possibility of deserves attention. No appeared, Obana and Mr. prevent is barrier in certain sperm removal further study. The has will that Research and 1972) and effort should (BICK difference between tactile and mechanical isolation, polymorphism Zygoptera riers the defining the quiring a interpretive and male toward are ' for behavioral acts various workers re- in chemoreception on bar- postmating conducted by Dr. Sympetrum in Japan (K. Inoue, S. pers. comm.). The exact male accessory of genitalia and need of clarification. There ture of these the functions parts within different how is a the they are structures used large degree order, and in comprising mating are the still in of variation in the struc- different functional roles isolation Reproductive for structures homologous JOHNSON of the understanding mechanical cover reconstruct tions and (1972), in exist. may WAAGE Studies (1979a, sperm-transfer 261 Odonata is rarely possible, and as rarely un- from having arise relationships of male and female parts, during copulation (1971), furthered have Such research may process. barriers. Difficulties isolating by PFAU 1982) to direct observa- do in pairs stay the wheel position after being collected. PROBLEM-SOLVING GUIDELINES Temporal and under study. and useful. dicates through time, between Any overlap either initially in the group maturation periods, considered. Marking tech- described by as for heterospecific potential a changes movement must be such analyses, particularly of isolation afforded by Population mating frequencies and niques isolation should be investigated ecological to determine the extent GARRISON (1978), in time species or are space in- males and females to meet, and other barriers must be sought. In such ethologically-isolated questions ponse in males? of the different fishing-line PAULSON do males visually species? Models technique there is model, on a nothing in favor tandem by males and/or dead sented achieve dence and sult their a from & with copulation males models in tandem be the be observed. If be be be females into and taken though even perform males are pre- able experimental of the male with difficult to to evi- appendages fe- conspecific control. Evi- (1) separation of heterospeachieved (this could also re- however); (2) separation of refusal motions by the female; (3) without sperm transfer. can broken; (2) when attempts lat- problematical. Further shape may 1958; the Evi- species, type of tactile requirement, lack can cooperate them. can With in tandem, the females tandem orven- male’s attempting a (1) tandem is hold techniques secure can of different altering success not using the 1952), 1975). res- females mechanical barrier includes: fly answer KRIEGER-LOIBL or two do and securely such a MOORE WAAGE mechanical barrier would be: some pair isolation can tandem a to males presented 1934; tactile a motions and obtained by pairs before of the in be and although dence of cific female observing males, be interfere with to latter the them clasp refusal tandem can tactile heterospecific able to are escape of models details of tandem behavior so differentiating presumed; of use discriminate between (KRIEGER visual isolation is found lacking, dence can 1974; TENNESSEN 1975; technique, to land (2) the species, aspects of the female elicit what technique (ST. QUENTIN trally-supported ter (1) as a break 262 K.J. If interspecific the possibility that vious conspecific artificially or using species may be every Hybrids fertilized were with used attempt should be result, although may with sperm from in seen a pre- Therefore, attempts from other species sperm previously tandem. The techniques . not be ruled out. can eggs observed, larvae rear the eggs mating fertilize especially is copulation made to obtain eggs and Tennessen to needed, are copulation interspecific by FREMLING (1967) for mayflies applicable. CONCLUDING REMARKS Visual (1) in important needed stimuli color distinctive in in the or isolating uniformly-colored of understanding mechanical morphological of range character displacement of RIBs (WAAGE species; and groups; more in efforts serve to as a work of which some vary research should the importance focus on are within possible of selection in the origin 1979b). be used difficult taxonomic problems; such application solve is Odonata. amongst populations Knowledge from reproductive isolation studies should (3) with species isolation characteristics, isolating species, often geographic barriers patterns, whereas tactile differences may be equally Behavioral and (2) be important clear-winged the important may test of conventional in could taxonomy. ACKNOWLEDGEMENT I thank JONATHAN K. WAAGE for many valuable comments on the manuscript. REFERENCES ASAHINA, S. BARTH, Part B1CK, 1974. R.H. I. G.H. Interspecific 1968. The Mating behaviour 1972. 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