Wav E for M an D Gonopo D Shap E in Sympatric Specie
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WAVE FORM AND GONOPOD SHAPE IN SYMPATRIC SPECIES OF UCA B. TROPICAL EASTERN PACIFIC NAME AND GONOPOD WAVE TYPE WAVE DURATION U.(Minucq) oerstedi (LATERAL) U.(Minucq) deichmanni (LATERAL) U.(Mmucq) beebei [LATERAL) U.(Minuca) batuenta (LATERAL) U.(Minuca) saltitanta (LATERAL) U.fMinuca) lirnicoia (LATERAL) |0 j, SECONDS FIG. 93. Wave form and gonopod shape in sympatric species of Uca. From films made "closely sympatric." When the cheliped is held in place briefly at a wave's peak, the in the tropical eastern Pacific, near Old Panama, R.P. Since all these species are mem pause is indicated in the diagram by a plateau. "VIB": vibration, the strokes being bers of the subgenus Celuca, they are rather closely related. Only three, oerstedi, too rapid to show individually on motion picture film, exposed at speed of 1/48 sec. batuenta, and saltitanta, share so many characteristics that they merit the term (24 frames per sec). (Pp. 528ff.) FIG. 94. Activity phases of Uca {Uca) maracoani maracoani in an outdoor crabbery. Phase sequences in behavior of six adult males. The position of each dot indicates the highest types of activity attained by an individual on a particular date, when these types are arranged in a series from least social to most social. The complete natural sequence appears to range from uninterrupted inactivity underground, through simple maintenance (feeding and digging) activities, wandering, aggressive wandering, territoriality, and, finally, display. Gaps in the diagrams represent days when observations were missing or inadequate. The graph of each individual's activity ends with the day before its death. These specimens were selected for illustration because of their longevity. The shorter records of fifteen other individuals showed similar characteristics. (Pp. 505ff.) rCDKUMrt T MARCH APRIL 10 II 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 UCA SOCIAL BEHAVIOR PATTERNS 95 CLASSES OF CLASSES OF COURTSHIP AGONISTIC COMPONENTS COMPONENTS PRE-MATING COURTSHIP DISPLAY THREAT a SUBMISSION FIG. 95. Diagram indicating the extent of ambivalence in social behavior patterns in Uca. (Pp. 517ff.) °«PONENT 404 7 UCA 6.AFRUCA 4. AMPHIUCA FIG. 96. Dendrogram: subgenera of Uca. Key to under scoring of names: black, occurs in the Indo-Pacinc; pale gray, eastern Pacific; dark gray, western Atlantic. (Pp. 18, 531.) AUSTRALUCA seismella b.longidigita polita b.bellator b.signata FIG. 97. Dendrogram: subgenus Australuca. Key to underscoring as in Fig. 96. (Pp. 63, 531.) 405 arhizophorae a. acuta THALASSUCA d.typhoni \ c.coarctata d. australiae ' d. demani FIG. 98. Dendrogram: subgenus Deltuca. Black bands, superspecies. Hollow bands, sympatric forms; excep tion: demani typhoni and demani demani, which re place each other allopatrically. Key to underscoring as in Fig. 96. (Pp. 24, 531.) 406 v.pacificensis v.vomeris v.dampieri v.hesperiae v.vocans v. borealis formosensis tangeri tetragonon c.chlorophthalmus heteropleura c.crassipes styl if era j.inversa pprinceps i.sindensis P rnonilifera t.thayeri t.umbratila ornata m. insignis FIG. 99. Dendrogram: subgenera Thalassuca, Amphi- uca, Boboruca, Afruca, Uca. Key to underscoring as in Fig. 96. Black bands, superspecies. Hollow bands, sym- patric forms. Key to underscoring as in Fig. 96. (Pp. 76, 97, 111, 117, 127, 532.) 407 mm ax brevifrons mordax galapagensis v.ecuadoriensis subcylindrica FIG. 100. Dendrogram: subgenus Minuca. Black bands, superspecies. Hollow bands, sympatric forms. Key to underscoring as in Fig. 96. (Pp. 156, 533.) tenuipedis stenodactylus ^.perpkxa inaequalis. \ tomentosa l.lactea tallanica beebei oerstedi festae deichmanni m.terpsichores t .triangularis t.bengali argillicola FIG. 101. Dendrogram: subgenus Celuca. Black bands, superspecies. Hollow bands, sympatric forms. Key to underscoring as in Fig. 96. (Pp. 217, 533.) Maps INTRODUCTION The 21 maps that follow consist of 17 in black-and- that I have personally examined were collected. On white and four (nos. 18-21) partly in color. maps where a number of such localities are very The first illustrates the distribution of the genus close together, as on the northwest coast of the Gulf Uca, along with degrees of concentration of its of Davao, a single dot represents two or more sites. species. In some older museum specimens the label gives only Numbers 2-17 show the distribution of most of the a general locality, such as "Japan" or "Madagascar." species, usually arranged in the sequence of sub If I did not find material to examine from particular genera employed in this contribution. Because of the localities in the same area, I used an open diamond prevalence of sympatry, however, and the exigencies symbol on the map; otherwise the more general name of photoreproduction, many species are not presented is not represented. An arrow indicates an extension in taxonomic sequence. In particular, the last maps, of a range beyond the confines of the map. nos. 18-21, follow the American series, but show, Of special interest on Maps 18-21 are the colored with the aid of color, the complex distribution of symbols falling outside the usual boundaries of the certain Indo-Pacific forms that are taxonomically species or subspecies, within the range of an allo- scattered. patric form. These distributions are discussed under To aid rapid comparison of ranges, an alphabeti the species headings in the systematic section, in cal list of species with corresponding map numbers Chapter 1 and in Chapter 7; see also Tables 3, 6, follows this introduction. and 22. No symbols appear on any American map, since Screens. Throughout the series the extent of each they proved in some cases to be impractical because screen represents my conclusion on the probable dis of extensive sympatry and in all to be of doubtful tribution of the species at the present time, based on value. These numerous American forms are usually information that appears to me to be reliable. This characterized by short or narrow ranges in uncom information consists first of published records which plicated patterns, while their taxonomic histories are, seem to me trustworthy, either because the contribu with a few famous exceptions, less tortuous than tion is the work of an experienced specialist, or the those of most Indo-Pacific species. It seemed, there record of a species in a part of its range where it is fore, that on American shores the precise localiza otherwise known to occur, or because I have exam tion on a crowded map of the origins of examined ined the material in the museum where it is deposited. specimens would not be very helpful, either in clari Second, the ranges are based on additional specimens fying allopatric situations or in evaluating evidence in museums which have never been recorded in print, for particular distributions. or for which references to the published accounts All of the records of material examined, both pre were not certainly determined, if at all. The third cisely and imprecisely localized, are listed in Appen source of information is provided by specimens I dix A. Ranges of the species and subspecies appear collected in the field (Table 24). under appropriate headings in the systematic section. Records that seem to me wholly erroneous, such The chapter dealing with zoogeography starts on p. as the occurrence of a species in Odessa or Yugo 431. slavia, have been omitted from the maps, although Excluded from the maps and from Appendix A they are discussed under the species concerned and is museum material I examined early in the course listed in the section on questionable geographic rec of the study before I had attained sufficient knowl ords (p. 326). Where boundaries are uncertain be edge of the forms involved to refer them with confi cause of the absence of recent records, as in Chile, dence to species or subspecies. This limitation applies the screen ends with an irregular edge; where records particularly to the collection of Australian material in are wholly lacking throughout a wide area between Sydney. The indicated ranges of some species occur populations, the sections of screen are rounded off. ring in Australia are, therefore, almost certainly al Symbols. For all the Indo-Pacific species dots and ready in need of extension. other symbols mark the localities where specimens 410 ALPHABETICAL LIST OF SPECIES OF THE GENUS Uca, WITH NUMBER OF MAP ON WHICH EACH APPEARS Species Number Species Number Species Number Species Number Name of Map Name of Map Name of Map Name of Map acuta 2 festae 16 mordax 12 stenodactylus 16 arcuata 19 forcipata 19 musica 17 stylifera 10 argillicola 15 jormosensis 4 oerstedi 15 subcylindrica 11 batuenta 15 galapagensis 14 ornata 10 tallanica 15 beebei 16 helleri 15 panamensis 12 tangeri 8 bellator 3 heteropleura 10 polita 4 tenuipedis 15 brevijrons 13 inaequalis 15 pnnceps 9 tetragonon 4 burgersi 12 inversa 7 pugilator 16 thayeri 11 chlorophthalmus 5, 6 lactea 21 pugnax 10, 14 tomentosa 15 coarctata 19 latimanus 17 pygmaea 11 triangularis 7 crenulata 15 leptochela 15 rapax 14 uruguayensis 15 cumulanta 15 leptodactyla 17 rosea 2 urvillei 19 deichmanni 17 limicola 17 saltitanta 15 vocans 20 demani 2 major 10 seismella 4 vocator 13 dorotheae 15 maracoani 9 speciosa 15 zacae 14 dussumieri 18 minax 12 MAP 1. Distribution of the genus Uca, showing relative concentrations of species in different parts of the range. See p. 409ff. "" llllll ff: MAP 2. Distribution of the subgenus Deltuca (part). (General explanation: p. 409.) MAP 3. Distribution of the subgenus Australuca (part). {General explanation: p. 409.) • —polita formosensis 45 60 75 90 105 1_ _i_ _L_ _L_ _J MAP 4. Distribution of the subgenera Australuca (con cluded) and Thalassuca (part). (General explanation: p. 409.) chlorophthalmus crassipes chlorophthalmus chlorophthalmus 30 60 90 120 150 180 150 120 90 _i J _J ? i _j _J J it MAP 5.