THE RELATIONSHIP OF EPIGAMIC DISPLAY TO THE SYSTEMATICS OF JUMPING SPIDERS (ARANEAE: SALTICIDAE)
By
DAVID B. RICHMAN
A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1977 Dedicated to Dr. Joseph C. Bequaert and Dr. H. K. V/allace. ACKNOWLEDGEMENTS
I wish to thank my committee members, Dr. Jonathan Reiskind,
cha i rman, Dr. John F. Anderson, Dr. CI ifford Johnson, Dr. Dale H.
Habeck and Dr. James E. Lloyd for their encouragement and criticism.
I would also like to thank Dr. James T. Giesel, Dr. James L. Nation,
Dr. Howard K. Wallace, and Dr. Henry M. Wallbrunn for suggestions or materials used in this study. Other people contributing specimens, materials, time or information include Dr. Bruce Cutler, G. B. Edwards,
David E. Hill, Dr. Robert R. Jackson, Will Kopachik, Wayne Maddison,
Randy Nicholson and Richard Skinner. 1 am indebted to both the
Departments of Zoology and of Entomology and Nematology, University of Florida, for the use of their facilities
Special thanks are due Dr. W. B. Miller, Dr. W. J. Gertsch and
Mr. Vincent D. Roth for their advice while 1 was a graduate student at the University of Arizona.
Part of the field work, especially that in south Florida, was
supported by a Grant-in-Aid of Research from Sigma Xi . Finally, I wish
to thank Ms. Lynda Coin for her advice and encouragement, without which
much of the writing of this dissertation would have been more difficult V
TABLE OF CONTENTS
Page
ACKNOWLEDEMENTS. iii
LIST OF TABLES vi
LIST OF FIGURES vi i
KEY FOR SYMBOLS USED IN FIGURES 1-17 ix
ABSTRACT xi i
CHAPTER
I INTRODUCTION 1
The Senses involved in Epigamic Display k
1 METHODS AND MATERIALS 20 1
ill COMPARATIVE EPIGAMIC DISPLAYS 26
The Epigamic Display of "Runners" 72 The Epigamic Display of "Intermediates" 79 The Epigamic Display of "Hoppers" 90 Summary 107
IV EPIGAMIC DISPLAY IN JUMPING SPIDERS AND ITS SYSTEMATIC IMPLICATIONS 109
Crane's Behavioral Groups and the Systematics of Jumping Spiders 109 Sexual Dimorphism and Epigamic Display 120 Summary. 126
APPENDICES
A COMPARATIVE DISPLAY DATA IN SEVEN SUBFAMILIES OF
SALTICID SPIDERS ' 128
B A CONTACT SEX PHEROMONE IN mNEMERUS BIVITTATUS 139
C THE GENUS HENTZIA IN FLORIDA 141
D A LIST OF THE SPECIES OF PELLENES FROM FLORIDA ]hh
I E CHARACTERISTIC SALTICIDS iN FLORIDA HABITAT ]hl
LITERATURE CITED 153
BIOGRAPHICAL SKETCH 162
V LIST OF TABLES
Table Page
1 Sources for courtship and agonistic display of salticidspiders 5
2 Collecting areas in Florida 21
3 Courtship, agonistic display, mirror display and
sexual dimorphism observed in North American salticids. . . 27
k Comparison sets of species in which courtship
characters are useful in spec i es- 1 evel systematics 36
5 Subfamilial placement of selected genera of salticids 112
V i LIST OF FIGURES
Figure Page
1 Et hog ram of courtship of Menemerus bivittatus male from Gainesville, Alachua County, Florida 39
2 Ethogram of courtship of Metacyrba undata male from Charlton County, Georgia k\
3 Ethogram of courtship of Hentzia palmavwn male from Way Key, Levy County, Florida A3
k Ethogram of courtship of different Hentzia palmanm male from Way Key, Levy County, Florida 45
5 Ethogram of agonistic display of Hentzia palmaruiv male from V/ay Key, Levy County, Florida kj
6 Ethogram of agonistic display of different Hentzia paluiariori male from Way Key, Levy County, Florida kS
7 Ethogram of courtship of Hentzia grenada male from Archbold Biological Research Station, Highlands County, Florida 5I
8 Ethogram of courtship of Hentzia mitvata. male from Lake Kanapaha, Alachua County, Florida 53
9 Ethogram of courtship of Covytl'ialia. auvata male from the River Styx, Alachua County, Florida 55
10 Ethogram of agonistic display of Corythalia aurata male from Gainesville, Alachua County, Florida 57
11 Ethogram of courtship of Habrocesturn bufoides Lake Oklawaha, Putnam County, Florida 59
12 Ethogram of courtship of Habrocesturn new species A male from Gainesville, Alachua County, Florida 61
13 Ethogram of courtship of Hellenes brunneiis male from Gulf Coast of Levy County, Florida 63
]k Ethogram of courtship of Hellenes coeoatus male from Jekyl Island, Glynn County, Georgia 65
15 Ethogram of courtship of Hellenes cf. coecatus male from Yuma Yuma County, Arizona 67
v i i Figure Page
16 Ethogram of courtship of Pellenes aalearatus male from the Muskoka District, Ontario, Canada 69
17 Ethogram of courtship of 'Pellenes viridipes male from the Muskika District, Ontario, Canada 71
18 Courting of Menemevus bivittatus 7^
19 Male of Menemevus bivittafAis mounting female 7^
20 Early courtship of Metaci/rba undata 78
21 Courtship pose of male Metacijrba undata 78
' 22. Early courtship of Ilentzia palmarum . . 82
23 Late stage in the courtship of Ilentzia palmarum 82
Ik Males of Eentzia palmarum in early agonistic display .... 85
25 Males of Hentzia palmarum in late agonistic display 85
26 Male of Hentzia grenada courting a female of H. palmarum . . 88
27 Male of Hentzia grenada mating with a female of H. palmarum. 88
28 Early courthsip of Corythalia aurata 93
29 Agonistic display of Corythalia aurata males 96
30 Fighting between males of Corythalia aurata. 96
31 Early courtship of Pellenes brunneus 101
32 Agonistic display of Pellenes tachypodus males 1 06
V i i i — —
KEY FOR SYMBOLS USED IM FIGURES 1-17
Direction of path (DR)
— Path direct
Path zigzag
Path an arc
I ^—I Movement forward
^ I— I Movement bacl I ( Stationary — Break Chel i cerae (CH) Chel i cerae in normal resting position ^ Chelicerae extended, fangs not exposed ^ Chelicerae extended, fangs exposed Palpi (PA) Palpi raised at angle ^/__ Right palpus raised ^^i^—. Left palpus raised Palpi lov/ered at angle —(— Palpi extended laterally "/i^ Palpi extended forward at angle Palpi extended directly forward Palpi moved up and down in unison fV^ Palpi moved up and down alternately Palpi moved in circular path, one clockwise, other counterclockwise I X First 1 egs (FL) First legs raised at angle —Ific Right f i rst " Left i rst First 1 egs II Fi rst 1 egs Fi rst 1 egs Fi rst legs Left first First 1 egs First 1 egs Fi rst 1 egs Fi rst 1 egs Fi rst legs First 1 egs Fi rst legs First 1 egs :Ui First ] egs Ta rs i and n 6 Ta rs i of fi Second legs (SL) — Second legs extended forward Third legs (TL) ''/v\1 Third legs raised in unison //VW Third legs lowered in unison /VXT Right third leg raised ^^^^ Left third leg raised AAV Right third leg lowered //^ Left third leg lowered Second and third pairs of legs (ST) Second and third legs nearly . parallel Second and third legs parallel and bent sharply at '^^^^^^ f emur-patel 1 a joints Prosoma (PR) Pro'soma raised ^ Prosoma moderately raised Prosoma lowered Prosoma tilted to right Prosoma tilted to left Opisthosoma (OP) ^3 Opisthosoma straight (normal posture in most species) Opisthosoma raised and tvjisted to right Opisthosoma raised and twisted to left Opisthosoma lowered Opisthosoma bobbed Mount i nc X i Abstract of Dissertation Presented to the Graduate Council of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy THE RELATIONSHIP OF EPIGAMIC DISPLAY TO THE SYSTEMATICS OF JUMPING SPIDERS (ARANEAE: SALTICIDAE) By David B. Richman March, 1977 Chairman: Jonathan Reiskind Major Department: Zoology A study was made of visual epigamic displays in representative species of North American jumping spiders, with a special emphasis on determining the value of using such displays in systematics. The displays of selected species representing Crane's behavioral groups were examined in detail. Several species were also selected because they presented special systematic problems. Other published descriptions of courtship and agonistic displays of salticids were also examined to provide a more complete picture of these types of behavior- In salticids. Crane's behavioral models for the salticids were found to be of little use In systematics, however the current study produced evidence that certain display characters can be used In systematic problems. Several presumed homologies In courtship behavior were described which largely support the most recent subfamillal arrangement based on morphology. Courtship was also found to be of use In some spec I es- 1 eve 1 problems, especially vjlth closely related sympatric species. Certain related sympatric species were found to exhibit courtship differences which presumably resulted X i I from selection for prezygotic reproductive isolation. Some ecologically separated species exhibited postzygotic genetic isolation. Agonistic displays were found to be of less use in systematics than courtship, because of the similar ways in which males engage in combat, even in distantly related genera. Correlations were found in most cases between tlie degree of sexual dimorphism and the complexity of epigamic displays. The orna- mentation of male salticids probably resulted from selection for prezygotic reproductive isolation and the recognition of the sexes. Sexual selection may also play a part in the evolution of such ornamen- tation. Female selection for males able to perform the proper sequence of the display and selection for males with greater persistence are probably the major elements in sexual selection. Some evidence was found for a contact sex pheromone in one species of saltlcid, MenemeTus bivittatus , during some peripheral experiments. The presence of such pheromones may indicate the utilization of non- visual cues in some species during visual courtship. This may also indicate a possible mechanism for the location of females during non- visual cohabitation courtship. X i i i CHAPTER I INTRODUCTI ON Salticidae (jumping spiders) is the largest of all spider families, with almost kOOO described species (Proszynski, 1971)- These spiders are diurnal vagabond hunters, which visually orient toward prey. They capture prey by jumping on it. Salticids have a characteristic f "squa red-of " prosoma and are easily identified by their enlarged anterior median eyes. The family is most diverse in the tropics, although a large fauna occurs in temperate areas. They exhibit a great variety of diversity in form and coloration. The males are often colorfully ornamented. This appears to be related to courtship (Peckham and Peckham, I889, I890). The systematic position of salticids with respect to other spiders is unclear. Most recently, Lehtinen (I967) placed the superfamily Salticoidea (including the Salticidae and Lyssomi n i dae) with the Zodariodea, Eresoidea and Thomisoidea in the branch Zodariides (p. 283, 288). He based this placement mostly on morphological similarities. This arrangement has not been universally accepted and the exact relationship of salticids to other spiders is still in doubt. Attempts to divide the family into subfamilies have resulted in a confusing abundance of systems, based primarily on morphology. These have ranged from the 70 groups of Simon (1901, 1903) and the 21 sub- families of Pet runkev i tch (1939) to the two subfamilies ( Lys soman i nae 1 2 and Salticinae) of Chickerlng {]3kS) . Pr5szynski (1976) disagreed with the placement of many of the genera in Pet runkev i tch ' s system. There remains a need for complete revision of salticid classification as pointed out by Proszynski (1971)- The Nearctic fauna of jumping spiders has not been examined in depth since Peckham and Peckham (1909). Several revisions of individ- ual genera (Barnes, 1955; Gertsch and Ivie, 1955; Galiano, 1965, 1966, 1969, 1972) and one "subfamily" (Barnes, 1958) have been published. Proszynski (1975) reviewed the Nearctic fauna in a short note and included some of this fauna in a larger work (1976). However, the lack of a modern systematic revision of North American salticids, as well as the lack of a recognized classification of the family as a whole, has made ecological and distributional studies very difficult. Disagreements in nomenclature can be found throughout the literature, even within the same publication. One possible hope for a natural systematic classification for the salticids lies in the utilization of behavior. As Platnick (1971) pointed out, courtship (defined here as i nt raspec i f i c male-female interactions, which precede and are preparatory to successful mating) is probably the most useful of behaviors for species level systematics because elements of it serve as isolating mechanisms. Aspects of courtship may also be more conservative than some other behaviors, and may show similarities between closely related groups. Crane (I9'(9b) proposed a behavioral classification (p. 173-175, 205-206) consisting of "runners", "intermediates", and "hoppers". Crane characterized each of these groups by their normal means of progression courtship and 3 agonistic display (defined here as i nt raspec i f i c male-male interactions, including those which are similar to courtship and those which involve fighting). Although this was not meant to be a natural systematic classification (as Crane points out, many displays may reflect parallelism among the subfamilies), Platnick (1971) assumed this, and stated: "That these groups do not coincide with the current systematic division of salticid species, casts more suspicion on the taxonomy than on Crane's analysis" (p. 43). Despite Platnicl<'s incorrect assumption, his conclusion that behavior should be considered as important a character as morphology Is basically sound. Braun (1963, 1964- Theridi idae) , Dondale (1964, I967, 1972- Thomisidae) and Vlijm and Dijkstra (I966- Lycosidae) have demonstrated the importance of courtship behavior in species level systematics in these families. Several important questions have arisen from this discussion. First, how well do Nearctic salticids fit Crane's behavioral cate- gories? Crane had well documented observations only from a limited number of Neotropical species. Only a similar study of the Nearctic temperate fauna could answer this question. Second, of how much use are visual epigamic displays (courtship and agonistic displays) in the classification of salticids? If these displays are useful, are they so at all levels or just at one level [e.g. species level)? These questions must be answered by comparison of a number of species, including both distantly related and sibling species. Third, what Is the relationship, if any, of visual epigamic displays to male ornamentation (sexual dimorphism)? An answer to this question might be found by examining the actions of males during courtship and . . 4 agonistic displays and seeing how closely the behavior correlates to the ornamentat i on Before such questions can be answered it is necessary to survey the literature and to make new observations. Table 1 is a list of references to salticid courtship and agonistic display, both in the literature and unpublished, and including the species examined in the current study. The subfamilies are arranged as Crane (l9^9b) presented' them in her Table II (p. 162-171), with the exception that the Lyssomaninae is placed first because it seems different enough from other jumping spiders to warrant more separation. Subfamilies which were not mentioned by Crane are placed where they probably would fit in her system. The Senses Involved in Epigamic Display Epigamic displays are widespread among the Metazoa. In salticid spiders such displays are primarily visual. 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ID C3 O in O o O O O o O o O O ai o O O o o t_) <_> <_> CJ o ro O o o C ro E _c o a) CO o Ln an cn 00 C s_ ro C_J -a E c cu JZ ro -C (= u JD JD _Q E r-~- in ro cn cn JZ cn d; -a- -d- u cn o cn cn cn 1- JD o D CO o Q. D- LA C cn ro ro ro _Q JD cn JO 3 cn -a c» CO CO cn cn CTi CTi c -cr -3- -3- ~a- ro cn cn cn cn cn CT\ C 0) o c c E c -a C in O ro ro 6 c 6 4-J E -C E 0) 0) 0) L_ 0) ro c N N 3 (U JZ .—^ +J 4-J O ^ o W c tz c "D c u o CS -c (U (D 3 o ro zc <^ E Wa o: cu ^ —' CO c 00 _j c « ro 0) 0 CU u c CL l--^i CO 4-1 c ro I/l CO Si L- ro i_ TO CO CO 1) i_ CJ c3 fx fJO ?H ^-^ E o <::! TO TO c M- LU 03 CY) Cj -4-i Sjj o O TO CO to o LU C« o tj CO TO TO UJ CO CL CO < UJ < CO s? in +^ ^ +i CO < CO in O -J TO ^'C II X O G o TO J3 0) _j n4-- UJ to Co Co Co Co CD Q. LU _J Q- D- (J in nn JZ c O ^ O >- cn (J cn ID TO 0) ID D_ Q. X) "D 1/1 c "O C ra C ID ra CL Q. E JZ JZ l/l t/1 4-J 4-" u 1_ ZI O CL 0 o CO - cn OD cn CO — QJ E c ID TO 0) JZ s_ o _^ o u u o D_ TO 13 cn cn C -iT 03 0^ E i_ C ra ID TO JZ E OJ C -C c o u TO TO D. D- q; <_) >• CU XJ TO C CD ^ J^ N O 4-1 — C sz c TO 0) TO :3 3: 1_ !_) 3 03 G 05 o 3 C JZ 4-1 o u 1/1 -a a. o 3 0) ca u X J3 II u 0) LU c ID Q. _I t/) 15 display to study. The components involved in visual displays include color, pattern, movement (timing and direction), form and size. Both color and pattern present some major problems to the observer of epigamic displays because it is difficult to determine the spectral colors perceived by organisms other than ourselves and to discover how color patterns might be altered by any differences in perception. Peckham and Peckham (I887, ]SSk) were the first to discuss the possibility of color vision in spiders, but their experiments viere not well controlled. Kaestner (19^9, 1950) and Crane (I9^9b) in- vestigated color vision in salticids, but had little idea of the actual spectral colors perceived by them. DeVoe (1972a, b, 1975); DeVoe and Zvargulis (1967); and DeVoe et al. (19^9) have shown that the retinal cells of at least some lycosid and salticid spiders possess dual sensitivities to color in the UV (360-370 nm) and in shorter visual wavelengths (510-532 nm). These are very similar to color sensitivities exhibited by insects and quite different from our own. Some attempts (Daumer, Eisner have been made 1958; et al. , 1969; Ghiradella et al., 1972) to photograph flowers and other natural objects in ultraviolet and thus gain some insight on what insects "see", but as yet no film has been developed which can show the dual sensitivity apparently present in most arthropods. Land (1969a) has suggested that some receptors in the salticid eye might be red sensitive, but DeVoe (1975) and Jackson (1976) have presented evidence contradicting this. Thus, it is difficult to evaluate the experiments on color vision made by earlier workers, up to and including Crane (19^9b), because they did not have this information available and were unable to properly analyze thei r data. 16 Although the role of color is extremely difficult to evaluate in salticid displays, movements, form and size are easier to study. Both Crane (1949b) and Drees (1952; found that the anterior median eyes could discriminate form. Crane found motion, form and size to be the most important factors in releasing epigamic display. Again, Crane's conclusion that color and pattern had little releasing value could possibly be the result of the inability of humans to perceive the same colors and patterns salticids do. Vie may tend to exaggerate the importance of motion, form and size because we can test them more easily in the laboratory. The structure and operation of salticid eyes have been examined in detail by several workers (Homann, 1928; Land, 1969a, b, 1971, 1972, the anterior median eyes 1974; Eakin and Brandenbu rge r , 1971)- Only are able to form images and they have great depth of field because of their tubular structure and cone-like retinae. To scan the visual field the retinae are moved by six muscles attached to each. Most of the evidence of the earlier workers (Peckham and Peckham, 1894; Homann, 1928; Bristowe, 1929; Kaston, 1936; Crane, 1949b) indicates the supremacy of visual cues in salticid courtship. It appears that the anterior median eyes and their keen visual abilities are by far the most important factors in salticid display. Chemorecept i on Pheromones (chemicals which serve as communicatory substances between members of the same species and which often function as location markers or stimuli for members of the opposite sex) have been found for 17 many animals (Birch, 197^)- So much literature on insect pheromones has accumulated in recent years (Jacobson, 1972; Shorey, 1973) that it is difficult to keep up with new developments in the field. A few arthropod sex pheromones are produced by males, but most are probably produced by females. The latter seems to be the most likely in spiders as females often are less mobile than males. The chemorecept i ve abilities of spiders and other arachnids are not well known and until fairly recently very little evidence existed for sex pheromones in spiders. Kaston (1936) in examining vagabond spiders was able to find evidence for such substances, only in a few of his tests with lycos ids. Long distance pheromones have been suggested as an explana- tion for mate finding in web-building spiders (Blanke, 1973; Enders, 1975). Crane (19^9b) suggested both long distance and contact chemorecept i on in salticids, and presented some interesting, but inconclusive arguments for the presence of such substances. Rovner (1968), Hegdekar and Dondale (I969), Richter (1971) and Dondale and Hegdekar (1973) have presented convincing evidence for contact pheromones in lycosid spiders, but Farley and Shear (1973) had ambiguous results. Contact pheromones in spiders may be associated with silk and thus an extract of the silk glands might be a possible starting point for the identification of such chemicals. Kaston (1936) and Crane (19'+9b) both indicated that some salticid males were attracted to locations where females had just been, but that actual courtship was initiated by sight alone. Hegdekar and Dondale (I969) indicated that even in lycosids the display was primarily visual, although males could be stimulated to display in some cases with only pheromone present. As yet no spider pheromone has been isolated and identified nor its source located. 18 MechanoreceptJ on In many araneids and other web spinning spiders, the sense of touch appears to be very important in courtship (Bristowe, 1958). This seems to be much less true of lycosids and salticids (Kaston, 1936; Crane, 19^9b). In fact touch seems to be important primarily during mounting in salticids and may be combined with chemorecept I on . Recently Jackson (1976) has demonstrated that at least some salticids may often mate without preliminary visual display if a male finds a subadult or adult female within her silk retreat. He has therefore defined two types of courtship for salticids. in type 1 (which will be referred to as "visual courtship" in this text) males go through the entire display before touching the female. Visual courtship does not occur in red light, which Jackson has shown to be undetected by Phidippus jofmsoni. Under red light males of P. joimsoni would only engage in type 2 courtship (which will be referred to as "cohabitation courtship"), sequestering subadult females until they matured and mating with adult females for longer periods of time than has been observed following visual display. Cohabitation courtship appears to consist primarily of tactile cues, although contact pheromones cannot be ruled out. Earlier workers, including Peckham and Peckham (I889), Nielsen (1932), McKeown (1936), Kaston (19^+8), Crane (I9^9b), Dewey (I962) and Forster and Forster (1973) reported cases of cohabitation, but did not elaborate on them. Jackson' (1976) showed that such behavior may be more widespread than previously thought and suggested tiiat it has distinct advantage for both sexes. 19 be used 1 al organs which appear to A few spiders have s t r i du ory of sound production in in courtship (Bishop, 1925). The examples Locket, 1926; Allard, 1936; Kaston, 1936; i (Bristowe and 1 ycos ds are Hallander, 1967; Rovner, 1967, 1975; Harrison, 1969; Buckle, 1972) with perhaps the most interesting, because these spiders often court visual displays and they are vagabond hunters like the salticids. it seems The hearing abilities of spiders are not well known, but likely that some hearing or vibration sensitive mechano'receptor may be present, especially in those species which make sounds during court- ship. Walcott (1963, 1969), Walcott and van der Kloot, 1959) demonstrated the existence of a vibration receptor in the theridiid Achaearmea tepidariorum and Rovner (1967) showed that females of Lyaosa rabida responded to playbacks of taped male courtship sounds by waving their first legs. in general, salticids seem to lack sound making structures. The one exception is that of the male of Phidivpus incevtus which produces a buzzing noise by striking of the palpi on the substrate during courtship (Edwards, unpublished observation). Hearing or vibration sense is probably of little importance in the epigamic displays of most salticids. . . . CHAPTER I I METHODS AND MATERIALS The behavioral observations necessitated the capture of fresh live specimens in the field and the maintenance of these specimens in the laboratory. The procedures were modified from those of Crane (I9^8b, 19^9b) Field Work Field sites where live material was collected are presented in Table 2. In addition, some of the data came from specimens collected in Pima, Santa Cruz and Yuma Counties in Arizona (several species of Okeefenokee Swamp, Charlton County {Metacijrba Pellenes-K\ chman , 1973), undata male) and Jekyll Island, Glynn County {Pellenes coecatus males and females), both in Georgia, and from the Muskoka District of Ontario, Canada {Pellenes calearatus and P. viridipes) At the Florida collecting sites specimens were collected by a variety of methods including beating shrubs and trees, sifting forest litter, sweeping through grass, herbs and shrubs with an insect net, peeling bark on dead trees and searching the ground and the surface of buildings. Of these methods, beating was the most successful for searching the most successful for Corijthalia auraba, Eentzia sp. , Menemerus bivittatus and often Pellenes sp., peeling bark the best for Metacyrba undaba and sweeping the best for Pellenes brunneus 20 e 21 Table 2. Collecting areas in Florida. Spec i es F i 1 mad Habi tat and Observed Local I on Al achua 1 Corijthalia. aurata. 1 a 1 1 eaf i tter Ga i nesv i 1 e and ong Archer Road (SR-24) Eabrocestvjn n. sp. A Marpissa sulcosa nea r Ga i nesv i 1 1 Metaphidippus sexmaculatus Neone I la vinnu. la Pellenes tac'hypodus p. trimaculatus Phlegra fasciata short grass Pellenes hrunneus bivittatus bu i 1 d i ngs Menemerus Plexippus paykulli .al pvarpera SR-2^*, near Archer old field Thiodina SR-325, 0.7 mi N. of understory in slash Phidippus pu Icherrimus SR-3A6, near Cross Creek pine flatwoods Thinodina sylvanna Corythalia aurata River Styx at 1 eaf litter Sarinda hen t si shrubs Lyssomanes viridis saw palmetto Synemosyna formica sulcosa SR-20, SE of Gainesville leaf and pine litter Marpissa along cypress pond Neon nelli near Pra i r i e Creek Sittious cursor Highlands Archbold Biological palmettoes Hentz^a grenada Research Station, Lake ^^^^ litter Habrocestum n . sp. Placid 22 Table 2 - continued Spec i es F i 1 med Habi tat and Observed Locat i on Levy Gulf coast at SR-2k grass Pe I lenes hvimneus Thiodina puerpera Way Key black mangroves Hentzia palmarim Mavpissa bina Seahorse Key black mangroves Hentzia palmarim Pelleries cf. viridipes SR-2k, near Alachua leaf 1 itter County 1 i ne Ma r i on grenada SK-kO, 7-10 mi E. of understory in sand Hentzia Lynne, Ocala National pine scrub H. palmariijri elegans Forest Tutelina low plants and ground Pellenes calcavatus in clearing n. sp. A 1 eaf 1 i tter in Pellenes cl ea r i ngs pikci Big Prai rie, Ocala tall g rass Mai'pisGa National Forest Monroe grenada pa 1 mettoes Hentzia P I necrest Pasco viridis Land '0 Lakes bushes Lyssornanes Putnam Hahrooestum bufoides Lake Oklawaha 1 eaf 1 itter rnvi ca pa 1 mettoes Synemosy na fo . 23 Care of Living Spiders Specimens brought to the laboratory from the field were l ni) the amount melanogaster) and cabbage looper larvae [Triohoplusia , and type of food appropriate to the size of the spider. Temperatures in the laboratory were relatively constant, approximately 2k°C. Lighting for the spiders was, for the most part, provided by over- head fluorescent tubes on a 12-hour light, 12-hour dark cycle, although the cycle varied somewhat during earlier work. Observat i ons Most displays and many of the experiments were recorded on super-8 color movie film. Two cameras were used during the course of the study; a Beaulieu ^(008 ZM II and a Bolex 280 macrozoom. In most cases epigamic displays were filmed under artificial light with a film speed of 18 fps (normal speed), but at least one film was made at twice this speed (1/2 s 1 ow mot i on) Both courtship and agonistic displays were filmed on scaled backgrounds, usually with metric scales. Most of the films were made while the spiders were inside a kO.'S x hO.5 x 10.5 cm Plexiglas chamber but because of the need for making horizontal views several films were made on a table top. Spiders of different (or the same) sexes were dropped down on the scaled paper (reused at first, but during later filming different paper . 2k was used for each species). The movie camera was focused on the pair and if courtship or agonistic display began the camera recorded it. Virgin females were, in general, more Tecept i ve to males than previously mated females, but with some species only the latter were available. Females were occasionally chilled to allow the male to go throught his complete display without interruption. This was not done as a rule unless virgin females could not be obtained. if not chilled, previously mated females would usually leave the area or attack the male. In several salticids (notably species of the genus Pellenes) matings of males even with virgin females were difficult to obtain in the labora- tory, in a few {e.g. species of Eentzia) matings with virgin females occurred readily. Agonistic displays were easily initiated in some species {Hentzia rjalmarim^ Sai-'inda hentzi and Corythalia auraba) but were difficult to initiate or possibly non-existent in others. At least two trials were usually attempted for both courtship and agonistic display. This was not possible in some cases, with difficult to obtain spec i es Only a few field observations on epigamic display were made, for the most part because of the difficulty of following the movements of such species as Hentzi pa'LmariMn in the thick growths of black mangroves brunneuG and other FeZZcn.^.? in grass and {Avicennia. nitida) , or Pellenes brush. Agonistic displays for both Corythalia auraba and Menemerus hivitiatus were observed in the wild. Also, for the purpose of examining the displays of Hentzia palmariMii on their normal substrate, specimens were filmed In the laboratory on stems and leaves of black mangrove. The reaction of males to their reflection in a small mirror was recorded, when possible, for comparison with Crane's observations and . 25 to obtain inforniation on possible agonistic displays when other males were not ava i 1 ab 1 e CHAPTER 1 1 I COMPARATIVE EPIGAMIC DISPLAYS This section consists of descriptions and analysis of the observed epigamic displays presented in outline in Table 3- The species are arranged in the subfamilies used by Petrunkevi tch (1928, 1939) and Crane (I9^(9b) so that a comparison can be made with Crane's Table II (Appendix A in this text). Whether the division of the family into these subfamilies reflect the natural phylogeny is debatable (see Chapter IV). The behavioral categories proposed by Crane {Ibid., 173" I (for 175) are noted for each species by the letters R (for "runner"), "intermediate") and H (for "hopper"). The sexual dimorphism present in each species is presented in the last column and can be directly compared with the courtship and agonistic behavior. A few selected species are examined in more detail in ethograms (Figures 1-17) and in the text. The ethograms represent individual courtship or agonistic displays which were thought to be most representative of the species under specific conditions. males In one case, that of Hentzia palmarum, ethograms of two different are presented for both courtship and agonistic display to show intra- specific variation. Because of the few repetitions of observations in many species, statistical analysis was not attempted. The types of move- ments themselves were found to be important .for separating related species. The numbers of observations and individuals involved for each species are recorded on Table 3. A direct comparison of important systematic court- ship differences in closely related species is presented in Table k. 26 S 27 Table 3- Courtship. »gonistlc display, mirror display and sexual dimorphism observed In North Anerlcan saltlclds. /igonistic Displays of SaUlcids Observed Hd'ie {Behavioral Croup of Crane) Mirror Display of Male Salticids Observed Observ. No. Observ. R " "runner" Agonistic Display (No. Males) No- Observ. e" Courtship Display . Kales) I • " i n I c rmcd i a t into Early and Late Parts) (Ho. Fif-hts) hales) ScAual Dirorphism and Late Parts) , Mount (ngs) (Divided Type of Display (NO- H • "hopper" (Divided into Early and enlarged chellcerae; for long 2(2)(0) Unknown No react ion 11'/ Kale with elongated Early nioven.ents : stationary front legs longer than in female periods, then approaches directly, repeat Later rnovcments: as in early Retinae of AHE : moved back and forth Palpi: held 90° to substrate (straight up) , Stat ionary First legs: extended laterally, larsl- _ metatarsi jerked alternately at i nte r va I Prosoma: raised off substrate Op i s IhosoTia : bent downward nAKT 1 SS INAE 1(2)(0) Similar to courtship 1(1) Sexes very similar: r,a les usually darker; Early moven>ents; direct toward female !(O(0) Like courtship females with central light Land on Later rtove^ients: zigzag op t sxhosofTta i ry Pa I p i : 1 owe red , si at ona First legs: raised Proson.a: low to Substrate Opi s I ho soma : ra i sed Similar to courtship 3(1) Sexes differ primarily in pattern: males Earl> movements: direct 2(1) (0) Unknown being darker, with dark band on op i s t hosOT.d Later niove'i^eni s : direct 1 i onary Pa I p i : o*vcred , stat First legs: raised Prosorra: low or occasionally slightly raised off substrate Opi s thosoma : ra i sed 6(I)C0) Unknown Unknown Sexes very siirilar >'. 8^lc:,3j (R) Early movenients: zigzag Later movements: zigzag Palpi : Stat ionary First legs: raised ProscwTia ; ra i sed Opisihosor^: raised, twisted to right or left No display seen in laboratory. 2(M(I) Simflar to courtship; male roves 3(2) Sexes differ primarily in pattern of dorsal /'i.t.c^r-ue iivit'.jtus (R) Early Piovenenis: direct, or at slight 12(3)(7) In field: back and forth in front of fii 1 r ror surface: males with dark central stripe ang le One display observed on Early novemenls: zigzag opisthcsomat females with wide light bano Later movenkcnis : direct Later if»ovements; fighting Palpi : Stat ionsary First legs: partly raised, occasionally Palpi : not observed Cbelirerae: extended during ^/'tghl I owef ed during stationary periods First legs: spaced widely at first, Proso.T.a : low extended and parallel to those Op I St hosof'.a : straight of cihier males during f/ight Prosorr,a: raised only slightly Opi s t nobor.a : s I r a I ght Table 3 " continued I Courtship Displays of Sattictds Observed Agonistic Displays of Saltictds Observed I Karic (Behavioral Group of Crane) I c I No. Observ. Mirror Display of Male Sa I t ds Observed R - "runner" No. Obscrv. " CoLirtship Display (No. Males) Agonistic Display (No. Males) No. Observ. I - 1 nl e rned i a te" (No. ftount ings) (Divided into Early and Late Parts) (No. Fights) Males H - "hc;per" (Divided into tariy and Late Parts) Type of Display (No. ) Sexua I D Imorphl sm Sexes very s i mi 1 ar /i tari^r:^ Ct^ilifcmica (R) Early rrovements: direct, jerky M3)(3) Later moven.ents: direct, jerky Palpi : Stat ionary First legs: raised, vibrated slightly Prosoma: low Op i s thosoma : s t ra i ght Similar to courtship 3(2) Sexes very similar Similar to M. califomica except: 2(l)(l) Unknov/n Palpi: extended laterally, during early courtship Stationary First legs: extended laterally during early courtship, raised to position like that of M. califomioa in late courtship, raised and lowered slightly during forward motion ProsoiT.a : ra I sed Op i 5 t hcsoma : Lcbb^d during mounting 2{2)(0) Unknown Kale with scale-covered eloncated chelicerae r^'->.3a c-^tleri (R?) Early fr«Dveiiien t s ; direct Later fnovements ; with some Indication of zigzag Palpi: extended laterally, stationary First legs: raised, vibrated prosofT.a : ra i sed Opi st hosonia : not observed Early movements: zigzag '.(M(2) Male with scale-covered elongated chelicerae 5. h.^tzi {R7) Early movefT'enis: zigzag 5(2)(3) Later novemenis: direct? Later moven.ents: fighting or retreat Palpi : extended lateral ly Pa I pi : extended laterally, Stat ionary Chelicerae: extended laterally, used to First legs : rai sed , waved In un t son Prosoma: raised bite with in fi ght First legs: raised, stationary Opi sthosDiT.a : bobbed up and down Prosoma : raised above substrate Dpi s I ho soma : lowered , twist ed to right or left observed 1(0(0) known Sexes very similar (R7) Early novements: not . Later moveinents : direct Pal p! : not o^se^ved First legs : rb i sed Prosor.a : ra i sed Op i s thosofna : tobbed up and down SiTT tCI KAE Unknown Sexes very similar Sc.K K.lli (17) Early movenients; zigzag 2(2)(0) Laier ntover.ent s : zigzag Palpi: raised and lowered In unison First I eg 5 : ra r sed , moved in un i son Prosoma: only slightly raised Opi stho^oi'.a : straight A : e t Table 3 " con! ir _ed Observed Courtship Displays of Saltlclds Ooserved Agonistic Displays of Saliiclds Ki->e (Behavior*! Sroup of Crane) Hlrror Display of Hale SaltldHs Observed No. Observ, No. Observ. Hales) Courtship Display (No. Hales) Agonistic Display (No. (No. Observ. I i r ijiTied i a t e" . i Sexual Dimorph 1 sm Parts) (No. hountlngs) (Divided into Early and Laie Parts) (No F ghi s) Type of Display (No. Males) H - "Mi;er" (Divided Into Early and Late S i T7 I C I NA£ Unknown Sexes very similar Early niovenients : who I c body vibrated 3(0(0) Late movtntents: jonped at female Palpi: stationary during vibration of body First legs: lowered prosoma: raised at least slightly 1 : t ra i Op i sthosofTia s ght Sexes very similar: male slightly darker Early movements: direct 5(2)(0) Unknown (17) than female Later rrovements: zigzag Palpi : not observed First legs: initially raised, then 1 owe red Prosoma: low or only slightly raised Gp I s Chosoma : straight differ prifr.arily in dorsal color pattern; l(l)(l)* Early rrovements: lateral l(2)(0) Similar to courtship; one jumped , 3(3) Sexes Early rove-iients : not well observed male generally darker than f er.ia le and with Later movements : not observed at re f 1 ect i on Later (noven.ents: not well observed dark or banded leg* Palp i ; i owe red , slat ionary Palpi : lowered, stationary : lowered? First legs: raised Flrsr legs ProscrfT.a : ra i sed Proso^na: raised Opi s t ho soma : I owe red? Opi St ho soma : lowered lar to courtship 2(3)(0) SirrJIar to courtship Dimorphism as in T. puerpera, from which this Early movenents: lateral or direct, ve 2(2)(0) Slmi Id) species differs in details of dorsal pattern slow, jerky Later novenienis: not observed Palpi: lowered, mostly stationary First legs: raised, then raised and lowered alternately and in unison, no apparent pailern Prosoma: raised : lowe red Op i £t hoson.a DENDhYPHAHT ISA£ [T'OvefTtents : zigzag I(2)(0) Kale with elongated and enlarged chelicerae, Early ruovemenis: zigzag 2(2)(0) Early t r fnovemen t s : f i ght i usually darker and with more contrasting Later movernents: zigzag? La e ng f Palpi : not observed markings than en^ !e; dorsal r^rkings Palpi: lowered, stationary Chelicerae: used in fighting (Edwards, d I f ferent First legs: raised, jerked bi I shed) Proson-.a: low to slightly raised First ! egs : ra i sed Opi sthoso-a: lowered Proson.a : ra i sed Opi St hosonia : 1 o^er ed Hiovenients: zigzag 9(5)(1) Male with elongated chelicerae; body more Early movements: often zigzag, soroetlmes 25(2)(l8) Eari> tV'.:-:- jre^^i^ (l) + r.cven.t:ni : rocking niol ion of both 2 fights n.a 1 straight Lalfcr s elongate than fenale; clypeus of e while rr.aies into Synchrony, fight- with ma ) first legs dark; dorsal Later movements, as in early falls and r^rkings differ en ing or retreat results ."^j Pa I pi : ex.1 eniied 1 at era 1 1 y Palpi: e-^ienaed laterally, stationary ^. First legs: extended laterally, raised, Chelicerae: extended laterally, fangs '-i' lowered alterndidy and in unison extended, used to bile severa 1 t imes extended * First legs: raised slightly, . Proson.3 ; ra 1 sed \. laterally, nearly parallel to rival Op f slhoso~.a : twisted to right or left, during fight 5 1 ra i gr.tened at intervals i Prnsrii. a : r a sed i • -, r I nt. 0;- I ' . I w 1 I r-rl I <- 55 COttftshlp DtsyfftyiS of SsJtfelcts Ofcser*«d Agonistic DfspJays of Saif.eids Ob,rr,ed R ". tinners" No, Observ, • Ho. - : cdlate" Observ. Mirror Display I Courtship Display (No. Hales) of Hale Saltlcids Observed (Divided into Early and Late Parts) (No. Mountings) (Divided Into Earty and Late Parts) (Ho. Fights) Hq. Observ. Type of Display (No. Hales) Sexual Dimorphlsrt! As ip ^r^nacia except front legs Chases rivals away (Peckham and raised llflfcnown and lowered more often, usually Peckham, I889) Halo with fringe of with each forward movement. white hairs on first legs- first legs n»re elor.g:,ie. dorsal markinos B. pa:-,-.- Early movements: zigzag 3;(3)(i) Later movef^cn ts : 2 i gzag Sexes djrfe.r P^lpi: extended laterally, raised In color pattern: male with more distinct *arkings and wit* and lowered i« ajnl.5,xjn *fhne moving v*;lte s»rip« on clypeus and First legs: Towered, extended iateratJy chelicerae at wide angle at first, narrowing as rr^le approaches female. Jerked at i nte rva 1 Prosoma: low or only s 1 i ght !,y raised Opi s inosoma : straight Early movements: direct ]Z(t|:fQ) Unknown Later movements; direct Sexes very Palpl: moved with np p^^tte*^^ simi lar First tegs: raised, moved with no pattern Prosoma: low Opis t hosoma : straight Early niovements: zigzag, with some arc- 3(2){0) LlivfcmawTT I ike (not ion Unknown Sexes very Later movements; as In early similar: males darker in ,olor Palpi : Stat ionary First legs: raised, crossed, jerked at i nteryal Prosoma: low Oplsth&s©ma: srr.aFght Ffci; ^.Slif*. U ) Early movements: zigzag 1(1) (O) Early movements: direct or circling K27)(2) Later mov^m^gnts: direct BftfcFiewn. Later movepieiflitst fighting Hale w,th black and Palpi : statitjfiary PalpT: white fringes on first StalTonary, extended soncwhat legs; First legs: raised, jerked opisthosomal markings more distinct- 1 era 1 a t ly palpl with Proscwna : ra ! sed contrasting white First )e.gs: raised ca. 30* and extended and black ar«s Op is thesooia : ra i 5 1 ght latcally ^5" during early st^gp, extended laterally at rSyht angles during f t qht Prosoma: raised, especially jtjst error to and during fight Opisthosoma: lowered a-^d twisted opposite to dlreclion of movement ) t t p T^b Ic 3 - con r i nu^d Agonistic Displays of Saltlclds Observed Cojrtshlp Displays of Saltlclds Observed I K^r'r (Behavior*? Group of Crane) Mirror Display of Male Salticids Observed No. Observ. No. Observ R - "runner"' Agonistic Display (No. rales) No. Observ. • "lntcrr»edlate" Courtship Display (No. Kales) I SeKua 1 D i morphl sm (Divided into Early and Late Parts) (No. F(ghts) Type of Display (No. Males) H - "hoppers" (DI /\6cfS into Early and Late Parts) (No. Mount Ings) DEN?RVPHAHTIKAE Unknown Male with black clypeus moven»cnts: zigzag or spiral MOO Unknown Zisvac-:^e ^.af fr'n~ei (l) Early Later moveir^ents: as In early Palpi: not observed First legs; raised, crossed, raised and lo*'ered later during lateral movements ProsoTia: low Op i 5 1 hosona : s t ra i gh 3(1) Sexes differ in color: males with grey scales 'i(l)(0) Raised first legs, moved back and Similar to courtship; some luteli-rc clec Prosoma : ra r sed Op i s tho'^-tvna : straight PELLENtNAE Seres differ in color patlerpi: males with nore and »»(2)(0) Raised 3rd pair of legs 2(M(0) nove rrien t s : body move d back .^:l^^^:rc^;u- rufcidee (H Early contrastingly r.arVcd palpi; rr^arklngs riorc shifting position over forth by distinct than those of female legs, sonie zigzag motion, stationary for long periods of tlfJW Later movements: as in early Palpi: raised and lowered alternately or in unison First legs: lowered Prosor\a ra i sed Op ' s thosof^.a : s t ra i ght (O) Soo>e zigzag motion, one male junped 2(2)(0) Unknown Sexes very similar; males with somewhat more Early -riovements : zigzag or arc, body 2(2) E. p-uU- (H) distinct pattern tilted toward direction of Fnovement at another Later ry^vements: zigzag or arc Palpi: lowered, stationary, occasionally raised and lowered slightly In unison or while body tilted, one lowered and one raised First legs: lo-ered during earlv courtship; sh raised during later court I Prosorr^: lowered early, raised later Op i s thobcwr.a : s t ra i ght Note: fcnale actively participates in display, usually causing mnle to rnove In arc or circle around her 1(2)(0) Unknown Sexes differ In color pattern: nales with stationary periods. I0(2)(0) No d 1 spl ay seen B. n. sp. A(H) Early movements: bright red cepha'Ic area and with more rapid arc-zlgrag run, follo^'cd by distinct dorsal pattern with jump at end of sideways run Later TnovcTients : as in early? Palpi: raised and lowered alternately or in un I son First legs: lowered Prosoma: slightly raised Op i s t hosoma : s t ra i gh s , 32 Table 3 " contli^ued Courtship Displays of Saltlclds Observed Agonistic Displays of Saltidds Observed hA-- (8e^avior^l Group of Crane) Mirror Display of Male Saltlclds Observed R - "runner" No. Observ. No. Observ. " I (No. Nsles) Agonistic Display (Ho. I I n tf^d a te" Courtship Display Males) No. Obsrrv. I Into Early end H • "horpers" (Divided Into Early and Late Parts) (No. Mount ngs) (Divided Late Farts) (No. Fights) Type of Dl splay (No. r.ales) Sexua I D i norph I sm fELLlMINAE F^llenee agilie (h) Early movements: zlg?a9 3(2)(0) No display seen 2(2)(0) No reaction (1) Hale with long fringes Later movements: direct on first legs; dor>.al color pattern differs from that of female Palpi : Stat ionary First legs: bowed downward, then raised Prosoma : slightly raised Op I s thosoma ; bobbed up and down P. arizanr.nEie (H) Early movements: zigzag n(7)(l) Unknown Unknown Dicnorphism Later movements: direct essentially as in P. agilie, from which Palpi: lowered, son>e movement In onlson this species differs in details of dorsal pattern First legs: bowed downward, then extended laterally and later forward, touching female Prosoma: slightly raised to low Opi sthosoTia : ral sed s. I 1 ght ly , bobbed none. At Early move">cnts: zigzag 32(3)(0 Like courtship, or least B(6)(0) Similar to courtship; male moves Hale vith t-JO Later movements: stationary for long one male Jumped at rival prominent spatulate spines on pro- back and forth in front of mirror, lateral surface periods of time, may arc around of each first leg; third legs often climbs onto surface modified with female projection on each patella and knobby distal end Palpi: moved back and forth, raised and to femora; latter with dark spot and stripes; dorsal lowered In unison early, later color pattern differs from that of extended forward female, male darker First legs; raised, tarsi jerked at 1 nte rva 1 Third legs: raised and lowered Proso^a: Ictv Optsthosoma: straight, bobbed slightly Early moverT>ent5: direct 3(2) (O) Like courtship, or none 5(5)(0) Unknown Males with greenish first legs (Oniar lo) Later movements: direct Including 2 "ith white fringe; and spatulate spines; third Palpi; touch substrate alternately during F lor ida legs modified with spike on each patella; part of sequence with first legs bent spec imens dorsal color pattern downward, otherwise stationary differs from that of female First legs; raised at start of sequence bends legs downward at t i b i a- me t a t arsus joint and vibrates them slowly upward, legs suddenly straightened, then vibrated slowly downward and quickly raised ca- 6 times-'sequence repeated Third legs: raised, lo%./ered at intervals Prosoma: low, lowered further during lower- ing of f i rs t 1 egs Op i s thosoma : 5 tra i ght (H) Ear ly movements : somewhat zigzag, dls- l(I)(0) Unknown Hale with spatulate orgari i zed spines on first legs; stripes on Later rovements: as in early clypeus; dorsal pattern differs from that of female, male darker Palpi : lowered raised, First leas; then raised and lowered in ur i son Prosoma; lowered Op i s t hos oma : straight e e 33 Tab^e 3 " cont inucd Salticids Observed Agonistic Displays of SaUlcids Observed Courtship Displays of Observed HaiT>e (Behavioral Croup of Crane) Mirror Display of Hale SaUlcids No. Otserv. No. Observ. R "runner" No. Observ, (No. Males) Agonistic Display {t^o. Males) - "inle'r^ed i a t Courtship Display I e" (io. Hales) Sexual Di»norphIsm (No. Mountings) (Divided into Early and Late Parts) ^''o- fights) Type of Display H - "hopper" (Divided into Early and Late Parts) Male with long greenish fringes and spatulste PEILEHINAE spines on first legs; third legs modified with Like courtship, or none 5(2) (0) Unknown FelleKpe c^fcct:^ (H) As In P. ti'unneUG . Al most identical. 7(2)(0) projection on each patella; clypeus red; v/hile spots on cheliccrae; dorsal color pattern I differs from that of female, male darker. Unknown Dimorphism similar to that of P. ccecatntt^ 17(n)(I) Unknown but r;oIe with different shaped projections on P. cf. c^catut (H) Courtship similar to P. brunneue ^ patellae; no white spots on chelicerae; except that the palpi are lowered third red cn clypeus more extensive and rK)5tly stationary i Unknown Kale covered with iridescent scales; clypeus P. KarUni (M) Early movements; zigzag 10(3)(l) Later mover>ents: zigzag, in both early iridescent pearl; femora of first two pairs of and later stages makes short, explosive legs flattened or concave anterioraMy and "jurnps" at end of lateral movements iridescent; dorsal color pattern nore distinct Palpi: lo^vercd, rrmstty stationary or raised than i n f ema 1 and towered slightly In unison First legs' raised, occasionally raised and lowered In unison Second legs; displayed by bending backwards, exposing Iridescent femora Proso'^.a : low Op i s t Hoscna : straight zigzag 5(3)(0) Unknown Hale with long black fringes and iridescent F i.-lie-^f.is (H) Early moveT^enls; Un known Later movements: not observed blue metatarsi on laterally flatt ened first with reddish or golden Palpi ; not observed legs; prosoma often First legs: raised. lowered alternately cephalic area; body covered with iridescent prosoma: not recorded metallic scales; much darker in color than Opisthosoma: raised f ema 1 8('»)(0) Early movements: direct toward-rival 3(2)(0) t^c'-jrcd-s (H) Tarly mover>ents: direct Sexes differ in dorsal color pattern: males Later mover-ents: one rr.^le pushes other Later novements: direct with more distinct pattern, usually darker back in relatively direct path Palpi: lowered, stationary Palpi: stationary First leas: raised, waved In unison Chellcerae; not extended at intervals First legs: raised Proson-^ : raised slightly Proso(r.a: raised only s'lghtly Opislhosona: bobbed up and down Oplsthoso^a: bobbed at first, then held S t ra i Q^^t II n known 5. tire:7iB (H) Early movements: zigzag Hal e with 1 ongcr front legs t ban f ema lei tars I later r*over^nts' zigzag of first legs black; palpi black and white; Palpi: fT»oved back and forth in unison, dorsal pattern of male differs from that of Iheri alternately, then held fo'ward feTiale; nalc darker First legs: raised and bowed, tarsi jerked Prosoma: low t Op i s t hosoma : straight LIVe courtship 6(M(o) zigzag 12(M(0) that r. u i-lTuUt'jF (H) Early novements: Unknown Sexes similar except male has different Later moven^cnts: zigzag cephalic markings (white stripes in eye region) Palpi: extended, moved alternately or In than female (dark bands on clypeus); r^ale color un i son pattern more distinct Fi rst legs: raised prosoma: low Opi St hosoma: st ralght , Table 3 - continued Courtship (Behavioral Croup of Crane) DIsploys of Saltlclds Observed AoonUtlc Dlsplnys of S«ltlcld% Observed ft "runner" No. Observ. Mirror Display of Hale Saltlclds Observed I - Obierv. "Intermediate" Courtship Display (No. Hales)r Agonistic Display H "hopper" (hq. Kales) (Divided into Early and Late Parts) (No. Mount' ngs) No. Observ. (Divided into Early and lat^Parts) (Ho. r.ghis) Type of Display (No. (Ules) Sexua t Dimorphi SfTt ^. viridifee (H) Early nwvements: direct 3(2)(0) Like Courtship Later movements: M2)(0) direct Male with greenish first legs with long white Palpi: touch substrate r-early in unison fringes and spstulate spines; third legs as first legs are lowered otherwl se modified with triangular patellae, r.alc dorsal s tat iooary color pattern differs fro.n that of fe-^le; First legs: raised, then slowly lowered dorsa 1 color pattern very sir>ilar to that while being vibrated back and forth and of P, c.2Zj:Li\itue bend I ng downward and i nward at the tibla-metatarsus joint, suddenly jerked upward, sequence repeated Third legs: raised, lowered at intervals, not correlated to other irovements, may be alternate or In unison Prosoma: low Oplsthosoma: St ralght , bobbed at Intervals F. cf. viT-idipee (H) Early movfrr^ents: not well observed l(l)(0) UnlcnoHn Later novc^nts: not observed Un known Dintorphism siriilar to that of ?, -.'iriJi-eB Palpi : lowered , s tat iooary but third legs of ii.ale not nodified First legs: raised Prosoma: low i Opi s thosoma : s t raight r. n. sp. A (H) Early movements: direct 2(2) (O) Male Jumps at other male, no real display Later movements: direct, 1(2)(0) almost motionless Male with long golden-yellow fringes on first for long periods legs; clypeus bright red; dorsal color pallern Palpi : not observed more distinct than that of fe-.ale First legs: raised Prosoma: low 1 Opi s thosoma : s tra I ght riLCcrc faeciata (17) Early movements: zigzag l{t)(0) Dnwn Later movements: direct Unknown Sexes similar: njle with cephalic area brighter Palpi : not observed red than that of feir.ale; clypeus turquoise Prosoma: not recorded, probably low b1 ue Op is thosoma: straight hlexippi>;ae \ ^r-rythalia cui'ata (H) Early movements: usually direct Il(5)(l) Early movements: zlazaq Later TOvenents: zigzag La ter fnov^rifnt s : direct, fray result in Sexes similar: male with white scales Palpi: rotated, one clockwise, other on cl>pei-s. fight Ing chelicerae; palpi counter-clockwise more contrastingly rrjrked than' Palpi: t~>Dve€ up and down In un'son; those of female First legs: lowered, raised slightly during c lockwi sc- counter- c lockwi 5.C mot Ion mot i on observed once P rosoma : ra i sed Chellcerae: extended during fighting Op i 5 thosoma : s t ra r ght First legs: lowered, aligned with second and third pairs to form "fan" Prosoma: raised Opisthosoxa: straight or lowered d " i Tat I e 3 con inuf Courtship Displays of Saltldds Observed Agonistic Displays of Salilcids Observed Harrv fBrhavioral Group of Crane) Mirror Display of Male Saltictds Observed It " "runner" No. Observ. No. Observ. (No. Males) Agonistic Display (Ho. Males! Mo. I « "intermediate" Courtship Display Observ. Early end . I (Divided into Late (*;o. H - *'hoDper" (Divided Into Early and Late Parts) (No Mount ngs) Farts) Fights) Type of 01 splay (No. Hales) Sexual Dimorphism PLEIIPPINAE No display seen (Crane, Unknown Flc^ifpus par/ F i rs t legs : ra 1 sed female P rosoma : ra 1 sed Qplsthosoma: straight KAGOWINAE first legs mover-en t s : zigzag 5(l)(0) Males wave at one another UnkrKJwn (H) Early "Norrr^l" phase male similar to female, movements: direct (Peckham and Peckham, I889) Later except for having yellow hairs on palpi mostly stationary Palpi; and stripes and spots on femora of legs; First legs: raised, moved back and black phase male differs from f e^-^a 1 e in forth having black body and three tufts at>ovc : ra i sed Prosoma an I er ior eyes Opi sthosoma : s t ra i ght Mated with fei-^le T. Sulvann. 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O 72 The Epigamic D isplay of "Runners" Crane (I9^9b, p. 173~17^) characterized "runners" as being those salticid species in which "progress is a scurrying run" and that in these chemotaxis and distance chemorecept ion are important, there are no intermale displays or fighting, the prosoma is low and the spines few. She used as examples Ashtabula, Menemerus , Semoyhia, Lyssomanes and possibly Hijatia {Marpissa in part). For the current study Menernerus bivittatus and the closely related Metacyrha undata were used as major representatives of this behavioral class. Other species which may be classified as "runners" include Lyssomanes viridis, Marpissa bina., M. pikei, M. suloosa and possibly Neonella vinrula^ Savinda cutleri, S. hentzi and Synemosyna formica. Courtship in Menernerus bivittatus The courtship of Menemerus bivittatus is very simple. Crane (I9^9b) gave the following sketch: "Carapace low, 1st legs raised slightly, stretched an ter i or 1 ra 1 ; 2nd legs extended forward; abdomen level ate 1 y (p. 162)." My observations revealed essentially the same display (Figures 1 and 18-19)- Males almost never deviated from a direct approach to the female. The motion of the male was very jerky, with forward motion interupted by 6-1^ pauses. The first legs were usually raised slightly higher when the male was at a distance of approximately k cm from the female and this was repeated three more times in one case. Some males on the other hand never even raised their first legs. V/ith the exception of the forward extensions of the second pair, the rest of the legs were not utilized except in moving forward. The palpi were never observed in motion. The first legs were finally extended further forward, touching Figure 18. Courting of Menernerus hivittatus . The male is on the right. Specimens are approximately 8 mm long. Figure 19- Male of Menernerus hivittatus mounting female. i 75 never moved or dis- the female, just before mounting. The abdomen was the female prosoma, played in any way. Mounting was accomplished over with few other preliminaries. Menemerus hivittatus Agon i Stic Display in pay no attention Crane {Ibid., p. 163) states that "usually males and that males to each other," that there are no distinctive motions occasionally court their mirror images. This description seemed to be However, accurate, based on the few observations made in the laboratory. two males of this species a chance observation was made in the field of were observed on involved in a distinct agonistic display. The two males silk retreat in a the side of a building. The smaller of the two had a male approached space between a beam and the outside wall. The larger first legs from below, displaying in a side-stepping manner with the widely spaced. The smaller male, still inside the retreat, oriented millimeters towards the movements of the larger male and advanced a few male forward several times, only to retreat again. Finally, the smaller extended the emerged and engaged in fighting with the larger male. Both of the first legs laterally so that they were nearly parallel to those After other male. They appeared to push each other back and forth. the wall grappling for one to two minutes and moving over the surface of from each other for several centimeters, during which time they broke away ran under the beam. at intervals, the smaller male finally broke away and retreat for a The larger male occupied the entrance of the smaller male's exemplifies short time, before being disturbed. The foregoing description negative evidence the difficulties involved in the interpretation of agonistic obtained in the laboratory. Bhattacharya (1936) reported display in this species observed in India. 76 Courtship in Metacyvha undata only slightly more complex than The courtship of Metacyvha undata is and Peckham (I889) recorded the that of Menemevus hivittatus . Peckham summarized it in her Table II courtship of M. undata and Crane (19^+96) and 2nd legs raised slightly, stretched as follows: "Carapace high; 1st the few observations available laterally; abdomen pendent (p. I62)." From while sketchy, is accurate for (all based on one male), this description, stages of courtship (Figures 2 and the most part, at least for initial raise its second pair of 20-21). The male was not, however, observed to Figure 17). A male approxi- legs as shown by Peckham and Peckham (1889, first legs half way, then finally mately 7.5 cm from a female raised his lowered his first legs and nearly 90° to the substrate. The male then forward he extended his first repeated the display. As the male moved second legs of the male could legs laterally. When viewed from above the legs (Figure 20) The first . be seen projecting forward anterior to the and the palpi were spread body of the male was raised off the substrate male was, as in Menemevus widely (Figure 21). The forward motion of the also raised his first hivittatus, direct. During the approach the male stopping and pausing for as much legs slightly with each forward motion, reached approximately h cm from as k seconds in between. When the male upward once and then extended them the female he raised his first legs first legs and the male forward. The female responded by raising her after touching the female retreated. The male moved forward again and, process of mounting over her with his outstretched first legs, began the taken by all salticids prosoma. The male assumed the mating position and during the mounting the (Kaston, 19^48, Figure 2010). Just before The male is Figure 20. Early courtship of Metacyrba undata. on the right. Each Figure 21. Courtship pose of male Metacyrba unda.ta. square is equal to 1 mm. • 79 male moved his first legs in a bacic and forth see-savj or vibration approxi- mately 16 times (rate 5/second). No agonistic displays were observed in this species, although two males reacted to mirror images by courtship- like movements. Observations of a similar species, Metaoyvba californica, indicate that this species has a very simple courtship, perhaps less complex than, but similar, to that of M. undata (Table 3) The Epigamic Display of "Intermediates" Crane (l9^9b, p. 17^) characterizes her "intermediate" behavior class as those salticids which alternate between hopping and running in normal progression and in which "distance chemorecept i on" is important, courtship is little differentiated from agonistic display, male-male fighting is frequent, the prosoma is of medium height and the spines are "moderate." She includes Vhiale, Sassacus, Paraphidippus [Eris) , Salticus and possibly Gertschia, Hentzia and Phidippus In this group, llcntnia, according to Crane, was just in between "runners" and "intermediates." I have used Uent2.ia palmarmi, H. grenada and, to a lesser extent, H. mitvata as examples of this behavioral class. Other "intermediates" on Table 3 include Eris marginata. Metaphidippus galathea, M. manni, M. sexrnaculatus , M. iritis, Phidippus pulcherrimus Sassacus papenhoei and possibly Neon nelli, Tutelina elegans, Phlegra fasciata, Thiodina puerpera and T. syLvana. Courtship in Hentzia pa Imarim The courtship of Hentzia palmarum was found to be relatively complex, although in a few cases the male approached a female and mated with her after very little visual display. Early and late stages in the courtship 80 are shown in Figures 22-23 and variations in the courtship are shown in two ethograms (Figures 3-^)- At the start of most of the displays observed the male was facing the female at a distance of 3-4 cm. Usually any movement of the female initiated a display by the male. He would then spread his first legs widely apart (approximately 100°) and then advance toward the female, usually with the opisthosoma twisted to the right or left and raised approximately 30°. Most males proceeded in a zigzag path toward the female, with pauses about once a second, during which the opisthosoma was straightened and then twisted again. A few males (Figure k) switched the opisthosoma from the right to the left or left to right during the display, but most l opisthosoma was twisted. The magnitude of the zigzags varied from a nearly straight path to sideways motions of as much as 1 cm. When the male reached within 1 cm of the female he usually straightened his abdomen and extended his first legs forward, toward the female (Figure 23). During the final stage of the courtship the male alternately raised and lowered his first legs two or three times, in addition to raising them slightly at the start of each forward movement. Occasionally only one would be raised and lov;ered. If the female was receptive (which she usually was, if virgin--of 12 virgin females, 10 accepted males) the male moved forward, alternately touching one first leg to the female and then the other in a see-saw manner approximately five times. He then mounted and twisted the female's opisthosoma around so that the was able to insert one of his palps into one side of the epigynum, later repeating with the other palpus on the othr side. Males often shifted from one side to the other several times, until the female started moving and Figure 22. Early courtship of Hentzia palmarum. Male is on the left. Specimens are approximatly 6 mm long. Figure 23. Late stage in the courtship of Hentzia palmavim. 83 apparently disengaged herself. Males removed from a female after mount- ing usually resumed copulation without repeating courtship. Agonistic Display in Uentzia patmarmi The agonistic display of Hentzia palmarum (Figures 5"6 and 2^-25) X'jas quite distinct from that of courtship in that the males exhibited an exaggerated side-to-side rocking motion, often in synchrony with each other, with the fangs extended and the body raised higher off the substrate. In a typical encounter the males faced each other and one initiated the display, with the other following. The males approached each other in rocking, zigzag to nearly straight, forward motions. They usually fell into synchrony when within 1 cm of each other. The raised and extended first legs touched the substrate with each dip to the right or left. The males usually extended their fangs at this point. V/hen they were nearly in contact the opisthosoma was finally straightened, the chelicerae were extended as widely as possible and at contact one male's first legs were nearly parallel to those of the other male (Figure 25). The males, if one had not retreated during the preliminary sparring, fought, grappling with their chelicerae. Usually the larger of the two males drove the smaller away, but if the males were of the same size the contest was not decided as quickly. One combat between roughly equal sized males lasted approximately 5 seconds, whereas another between a smaHer and a larger male lasted less than 1 second. Once a smaller male apparently injured a larger one by biting him on one of the first legs. Observations of males placed together on stems and leaves of the black mangrove in the labora- tory confirmed that such displays take place easily in natural settings, Figure 2^*. Males of Hentzia palmarum in early agonistic display. Specimens are approximately 6 mm long. Figure 25. Males of Hentzia palmarum in late agonistic display. 85 1 86 and that fighting is often the result of the display. The only similarities to courtship were that the first legs were spread widely and the opistho- soma was raised and twisted to the right or left. In mirror displays the males reacted exactly as they would with another live male, attacking the image and trying to bite it. Courtship in Eentzia grenada Not counting hybridization attempts, the observations were made of curtship exhibited by two males from Highlands County, Florida. The early observations were made of a male from Marion County courting, and mating, with females of H. palmarum from Levy County (Figures 26-27). The courtship of H. grenada (Figure 7) was very similar to that of //. palmarum and virgin females of the latter species would always accept males of the former. In one case a female of //. grenada accepted a male of palmarum. Viable eggs were produced in at least two cases, but the immatures never reached adulthood (Appendix C) . The close relation- ship of H. grenada and H. palmarum and their often close proximity in nature could lead to hybridization in the wild on occasion. 1 have examined several hundred //. palmarum and approximately one dozen H. grenada specimens of both sexes without finding such an intergrade. Agonistic Display in Hentzia grenada The agonistic diplay of Hentzia grenada was indistinguishable from that of H. palmarum. No mirror displays were attempted with this species, however, several encounters between males of H. palmarum and //. grenada were staged. One male of H. grenada easily defeated males of H. palmarim of his own size. He mated with a total of four H. palmarum females. Figure 26. Male of Eentzia grenada courting a female of H. palmax'um. Specimens are approximately 6 mm long. Figure 27- Male of Hentzia gvemada mating with a female of //. palrnavum. 88 ^ 89 Finally he was defeated by a larger //. palmarwn male. Another male of H. grenada also was quite successful in defeating an palmarwn male of similar size. Males of H. grenada of a given size have slightly more elongate chelicerae than males of //. palmarum of the same size and this may given them some advantage in such staged fights. Probably the two species seldom meet in nature. Their ecological isolation (Appendix C) probably insures their genetic Isolation. Courtship In Hentzia mitrata. One pair from Florida was observed, along with a very short segment of courtship by a male from Ontario. The male from Florida also courted a mated female of H. palmarwn for a very short time. Peckham and Peckham (1889) described the courtship as follows: "The body is somewhat raised; the first legs are held at a right angle to the cepha 1 othorax , and the abdomen is twisted to one side, and, as he dances before the female, Is changed now to the right, now to the left " (p. 50). This description agrees with the observations made during the current study. The male from Florida was observed during four successive courtships of a chilled female (already mated) and the courtship on all occasions was quite similar (Figure 8). The male began facing the female at a distance of several centimeters. Usually a slight movement of the female started the male's display. The male spread his first legs widely (approximately 100°) and advanced toward the female In a more or less straight path. The opistho- soma was twisted to the right or the left. The male stopped and paused for short intervals (none longer than 1/3 second once the forv^ard motion had started), raised and lowered his first legs and sometimes straightened his abdomen. Unlike the other species of Henizia, the H. mitrata male 90 also would in some cases straigten his abdomen while in motion (not seen in courtship recorded in Figure 8). As the male approached the female he narrowed the angle of his first legs, similar to the actions of both H. palmavum and H. grenada, but did not finally straighten the abdomen until he started to mount. He moved his first legs up and down in unison, or sometimes alternately at intervals as he moved forward. Unlike the other two species he raised his first legs sharply even during the early stages of the courtship. The male did not appear to move his first legs as much during mounting as did //. palmarum males. Hentzia rnitrata, unlike the other two species, has vertical, short chelicerae and white fringes of hair on the first pair of legs. The latter may be related to the more frequent high up and down movements of the first legs during early courtship (see Table k) . Peckham and Peckham (l889, p- 50) observed that males usually jumped on females at distances of "one or two inches" before mating. Such behavior was not observed in the few examples seen. No agonistic displays were staged because of a lack of enough males. The Epigamic Display of ''Hoppers" Crane {]3h3b, p. 173~17^) characterized "hoppers" as being those salticids which normally progress by hopping and in which chemical stimul are unimportant, courtship and agonistic display are distinct, no male- male fighting occurs, the prosoma is high and the spines "moderate to numerous." She included such genera as Corythalia, Plexippus^ Mago, and possibly Maevia, Eustiromastix and Philaeus . I have used Corgthalia auratU:, Habvocestwn bufaides , H. n. sp. A, Pellenes brunncus, P. coecatus P. cf. coecatuS:, P. calcavatus and P. viridipes as major examples of this . 91 group. Other "hoppers" observed include Habrocestum pulex, Fellenes agitis, P. arizonensis, P. alypeatus, P. hallani, P. hivsutus, P. tachypodus, P. tarsalis, P. tvimacutatus P. cf. viridipes, P. n. sp. A, Plexippus paykulli and possibly Maevia inolemens. Courtship in Corythalia aurata Courtship in Corythalia aurata (Figures 9 and 28) is similar to that of C. fulgipedia (Crane, ]3k8a, p. 28) in that the male's first legs are kept down on the substrate and the palpi are rotated in a clocl The path of the male showed considerable variation, being zigzag to nearly direct. The male moved forward in jerky motions, with his body somewhat elevated off substrate op i s ra t . Females the and the thosoma st I gh usually responded to the approach of a male by raising the first legs and elevating the prosoma (Figure 28) (also seen in Habrocestum pulex and some Pellenes) All females observed either exiiibited this behavior or ran from the male. At this point, with the female approximately 1 cm away, the male usually zigzaged back and forth, with his palpi in motion, his first three pairs of legs extended laterally and nearly parallel to each other, and his body elevated. The male then attempted to mount, raising his first legs up and raised the prosoma higher. One male finally managed to mount over the female's elevated prosoma, and successfully mated, inserting his embol i a 1 ternate y into the female's epigynum. 1 Figure 28. Early courtship of Corythalia auvata. The i.(,.le (larger than the female in this case) is on the left. Specimens are approximately 5 mm long. 93 3k Corythalia aurata, while similar to C. fulgipedia in its courtship, differs in morphology. Males of most species of Corythalia are often quite different in appearance from the female. However, the males of C. aurata have no fringes of hair on their legs and they seem to differ from females of the same species only in having numerous white scales on the cylpeus and chelicerae and slightly more contrastingly marked palpi- Males of C. fulgipedia have the first three pairs of legs fringed and compressed. The other South American species males have fringes on various legs. Agonistic Display in Corythalia aurata The agonistic display of Corythalia aurata (Figures 10 and 29-30) differed from courtship in that the opisthosoma was often bent downward, the palpi were moved up and down, and the prosoma was raised off the substrate to a greater degree. The displays occasionally ended in fight- ing, but generally were terminated with the retreat of one of the males. Individuals sometimes initially started to court, but quickly shifted to agonistic display. Males began the agonistic display with the prosoma elevated, the opisthosoma either straight or bent downward, and the first three pairs of legs nearly parallel and touching the substrate. Either both males approached one another or one initiated the approach. As in courtship the palpi were in motion most of the time, the movements being mostly up and down, rather than in circles. The approach of the males was jerky, with very short pauses (Figure lO) after forward runs. One male was observed to raise his third pair of legs off the substrate while he was under attack. Before contact was finally made in the encounters observed, one or both of the males raised his first legs. Usually one Figure 29- Agonistic display of Covythalia aurata males. Specimens are approximately 5 mm long. Figure 30. Fighting between males of Covythalia aurata. 96 ] I 97 male raised his first legs and tine other follov-jed, but it was not dependent on which individual initiated the display. If one of the males had not been driven off at this point, contact was made and the first legs of both males were lowered and held parallel to those of the opponent (similar to the pose of Menemerus or Hentzia) . They fought pushing'each other back and forth and seeming to bite with their chelicerae (Figure 30). One of the males eventually broke away and ran. No fighting took place in one confrontation between males observed in nature. The smaller of the two males retreated after displaying for only a few seconds. Males of Covythalia often differ markedly in size. It is doubtful that fights take place between smaller and larger males as the smaller usually retreats before that stage is reached. A total of 30 interactions between five marked males were observed to determine if the same males succeeded in driving away their opponents. In all cases the larger males defeated smaller males. Males of nearly the same size as their rival fought until I one eventually defeated the other. Repeated fights between such males always resulted in the same male "winning." These encounters were observed over three days and are not included in the encounters used for analysis of the display (Table 3). in Courtship Habroaestim hufaides and B. n. sp . A Habrocestwri hufaides and a very similar undescribed species were collected in different types of leaf litter and these species present a very interesting systematic problem. The courtships are different in several respects (Figures 11-12). Individuals of both species are very small in size (2-3 mm) and both have zigzag courtships. In the court- ship of H. bufoides the males make sideways runs of a much shorter . 98 maximum length (0.25-0.50 cm) than those made by //. n. sp. A (1-2 cm). Males of H. bufoides tilted their bodies to the right and left, depending on the direction of the run. Males of the undescribed species did not tilt their bodies, but did raise and lower one or both of their palpi at the end of each run. Hahroaestum n. sp. A males started the court- ship with very wide arcs (approximately 2 cm) and rapidly contracted these to 0.25 cm. This display took only 2 seconds, the male then paused for 1-2 seconds, and repeated the zigzag display. Habrocestim bufoides males did not vary their sideways run much in the few observed court- ships. Their distinctive tilting motion readily separated them from H. n. sp. A (Table 4). Uabvocestum bufoides is at least somewhat ecologically isolated from the undescribed species, being found primarily in mesic and flatwoods litter. Eabrocestum n. sp. was found almost totally in turkey oak and hickory litter. In at least one instance in a mixed forest they were found within a few meters of each other and in some cases may occur together. Little in the way of agonistic display was observed, although one male of the undescribed species raised his second pair of legs during one encounter with another male of the same spec i es Courtship in Pellenes brunneus, P. ooeoatus and P. cf. coeoatus Another interesting problem in systematics is posed by members of the coeoatus complex of Pellenes. Three members of this complex (which also includes P. bovealis , P. captiosus and P. mexicmms) were observed. enes Pell bvw^neus and P. ooecatus , while they have overlapping ranges, have not been found together. They live in similar habitats and may be mutually exclusive. Pellenes cf. ooecatus, which may be an undescribed 99 is a species, western counterpart of P. coecatwi . It is found in grass and alfalfa throughout southern Arizona and California, and in Mexico. The courtship of Pellenes brunneus (Figures 13 and 31) usually began when the male was approximately 3 cm from the female. The male initially raised his first legs and spread his palpi as widely as possible laterally. He usually moved in a zigzag during early courtship, switching later to an arc around the female. During early courtship the palpi were raised and lowered at a rate of approximately 3/second for 5-10 seconds, while still being kept widely separated. The male moved forward toward the female while raising and lowering his first legs and pausing several times until he was approximately 0.5 cm from the female. The male then assumed a characteristic pose of members of the Pellenes coecatus species group, with the first legs held high, the palpi 1 owered, extended forward and held parallel and stationary, and the body low to the substrate. The third legs were pressed to the sides of the op i sthosonia. During this pose the male often remained stationary for 15-30 mi'nutes, especially if the female was not turned toward him. If the female turned toward the male, he raised and lowered his first tarsi very rapidly several times and this activity was often preceded or followed by the raising of the third legs, which were also moved backward and forward. The third legs were usually raised one at a time and the male ended the sequence by touching both third patellae together over his dorsal surface and then lowering them to their original positions. Only one mounting was observed and this was with a chilled female. In this case the male continued his display for several minutes, creeping slowly forward until he was able to touch the female. The male then leaned forward and touched the female with one of his first legs several times. This activity was repeated alternately Figure 31- Early courtship of Pellenes hvunneus . The male is on the left. Specimens are approximately 5 mm long. 101 1 I 102 with the right and left legs. The third legs were moved again several times, the last time just before mounting. The male then mounted and attempted to mate as in other salticids. Virgin females of P. brunneus were difficult to obtain and all females seemed to be quite resistant to male advances. This seemed to be true with most other species of Pellenes and it is possible that they required conditions which were not present in the laboratory before they would mate. Because of the close relationship between Pellenes hrunneus and P. coecatus and because of an earl ier study of what was thought to be P. coecatus in Arizona (Richman, 1973) it was thought useful to obtain specimens of P. coecatus from the eastern United States and record their courtship display. Fortunately three males and two females of the eastern form were collected by D. and W. Maddison near Jekyll Island, Georgia and the courtship was filmed. The display (Figure 15) was very similar to that of P. brunneus, within variation shown by that species, but differed from the Arizona specimens (Figure 15 and Table k) in that the palpi of the eastern form (P. coecatus) were extended laterally and moved up and down during early courtship, whereas the western form (P. cf. coecatus) males kept their palpi lowered and relatively stationary. The eastern form males differed from those of the western form also in coloration, the chelicerae having white patches, the red area of the cylpeus not extending past the anterior lateral eyes and the dorsal pattern also being somewhat different. W. Maddison (personal communication) has pointed out some differences in the palpal morphology of the males and there are also some differences in the morphology of the third leg, ' those of the western form resembling P. brunneus more than P. coecatus. 103 Pellenes brimneus , while agreeing vvith P. coecatus in courtship behavior, differs from both P. coecatus and P. cf. coecatus in having a dark brown clypeus, partially covered with white scales or hairs; in lacking heavy fringes on the first legs and in having a different shaped projection of the patellae of the third legs. It is probable that at least P. brunneus and p. coecatus are sibling species and that P. cf. coecatus is a closely related, possible sibling, but still distinct species. No specific agonistic display has been observed in these species, although encounter- ing males often raised their first legs. Courtship in Pellenes calcaratus and P. viridipes Like the P. coecatus species group, P. viridipes and the related species P. calcaratus seem to form a natural closely related species group. The courtships of these two species were similar in that the first legs of the males vibrated downward (at least during most of the courtship), the third legs were raised and lowered, while being moved back and forth, and long stationary periods were spent in front of the female. In the courtship of P. calcaratus (Figure 16) the first legs were bent at the tibia-metatarsus joint and vibrated slowly upward at the start, followed by a sudden straightening of the legs and then a series of approximately 8 slow lowerings of the rapidly vibrating first legs. At the end of each lowering the first legs were suddenly raised again. The palpi were moved only during the early raising of the vibrating and bent first legs. The palpi were raised and lowered alternately at a rate of 1.25/second for k seconds. In the courtship of P. viridipes (Figure 17) the male's first legs were bent at the tibia-metatarsus joint, both downward and inward. The legs were never raised while so bent, but they 1 ok were not lowered at a steady rate. Instead they were at times almost stationary in height while the tarsi were rotated in a small circle. The first legs were finally lowered the rest of the way and suddenly straightened and raised. This sequence was repeated. During the initial movements of the first legs the palpi were moved up and down rapidly nearly in unison at a rate of approximately 2.7/second for 1.5 seconds. The third legs of both species were moved up and down and back and forth at intervals, apparently with little pattern. The movements of the first legs and the palpi (see also Table k) serve to separate these remarkably similar species (the dorsal patterns of the males are nearly identical). A female of P. calcaratus was used to obtain courtship from P. vividipes males because no P. viridipes females were available. It is unlikely that the male varies the basic movements of the courtship, even in such an artificial situation. The courtship observed for P. viridipes matches a published account by Peckham and Peckham (I89O). No distinctive agonistic display was observed. Agonistic Display in other Pellenes After observations on P. agilis, P. brunneus, P. calcaratus^ P. coecatuSj P. tachypodus, P. trimaculatus , P. viridipes and P. n. sp. A, I have only seen a definite agonistic display in P. tachjpodus. In most species the males may court, ignore or jump at each other (prey capture behavior), but P. tachypodus males have been observed on three occasions to go through a ritual, similar to courtship in the movements, in which one male pushes another male backwards (Figure 32). This shoving match resulted in one male eventually breaking away and retreating. Figure 32. Agonistic display of Pellenes tachypodus males. Specimens are approximately 5 mm long. 106 107 Sumn ' a ry The courtship behavior of k8 species of North American salticids was presented on Table 3- Agonistic and mirror displays were also shown, where possible, and sexual dimorphism for all species was presented. Ethograms for selected species were presented in Figures 1-17. Observations on species of Crane's "runner" behavioral class indicated that while these species did seem to have relatively simple courtship and usually no agonistic displays, at least one species {Menemerus bivittatus) exhibited agonistic display and fighting. Observations on species of Crane's "intermediate" behavioral class indicated that while these had more complex courtship rituals and often agonistic displays similar to courtship, at least some [Hentzia. palmarwn and U. grenada) exhibited agonistic display that is distinct from courtship in many ways. Observations on species of Crane's "hopper" behavioral class indicated that while these generally had the most complex courtship displays, a possible lack of combat in agonistic behavior and the highest degree of sexual dimorphism, at least one species [Corythalia aurata) engaged in fighting during agonistic displays and exhibited a low degree of sexual dimorphism. Several sympatric spec i es pa i rs [llahrocestwn hufoides and H. n. sp. A, and PeVlenes calcaratus and P. viridipes) were found to be easily separable by their courtship behaviors. . 108 6. Some ecologically isolated species (e.f/.j Hentzia palmarvjn and H. grenada) were found to lack differences in courtship and to. be able to hybridize. These species were found to be postzygot i cal ly Isolated. 7. Two closely related parapatric species {Pellenes brunneus and P. coecatus) were found to have indistinguishable courtships, but these are probably isolated by competitive exclusion. 8. Two closely related allopatric species [Pellenes coecatus and P. cf. coecatMs) were found to h ave similar, but distitiQuishable courtsh i ps CHAPTER IV EPIGAMIC DISPLAY IN JUMPING SPIDERS AND ITS SYSTEMATIC IMPLICATIONS Earlier in this work several questions were asked about Crane's behavioral groups, the uses of epigamic displays in systematics and the relationship of such displays to ornamentation. Crane's behavioral groups have been found to be much too simple to be useful in system- atics. Her classification of salticids as "runners", "intermediates" and "hoppers" utilized characters which probably reflect convergent behavioral adaptations by different phylogenetic lines. Also, at least several of the species used in the current work had behavioral charac- teristics which did not coincide with their behavioral group as defined by Crane. Courtship behavior is useful in some systematic problems, especially in distinguishing closely related sympatric species. The possible use of such displays in higher level systematics within the subfamily is more problematical, but some probable homologies at the generic level can be seen. Agonistic display is much less useful In systematics than courtship because some species may lack this behavior and distantly related genera may exhibit similar displays. Sexual dimorphism played a role in all the observed epigamic displays and a positive correlation v;as found to exist between the degree of male ornamentation and the complexity of courtship display. Crane's Behavioral Groups and the Systematic s of Jumping Spiders Crane's behavioral groupings (19^9b, p. 173-175) were not meant to be a systematic classification, but a series of evolutionary grades 109 110 through which different phylogenetic lines passed. This was misunder- stood by some workers [e.g. Platnick, 1970 and behavior was suggested as a replacement for morphology in discovering phylogenetic relation- ships. Unfortunately some of the genera placed by Crane in her behavioral groups do not possess some of the characteristics of the assigned group. Crane had to draw on the literature for reference to other than South American species and much of the published descriptions of epigamic displays are incomplete. Very few species of jumping spiders had been adequately analyzed. Crane thus had no indication that at least one species of Covythalia {C. aurata) , unlike her Venezuelan species, did engage in fighting during agonistic display. Also, while she reported that the agonistic display of Eentzia mitrata (based on Peckham and Peckham, I889) was the same as courtship, observations of such displays In Hentzia patmarim and //. grenada have Indicated some distinct differences between courtship and agonistic display. Crane's placement of Hentzia between "runners" and "intermediates" was based in part on the presumption that there were no such differences between the displays. In her own observations, Crane apparently missed the agonistic display of Menemerus bivittatij.s , possibly because she may have seen only laboratory encounters. Menemerus , as a "runner" is not supposed to have such a display. The complexity of visual displays, which made it difficult for Crane to analyze them, is probably a direct result of the consid- erable speciation and consequent selection for prezygotic isolating mechanisms exhibited by this family. Many similarities In behavior and morphology within the salticids may have resulted from either 1 1 1 parallel or convergent evolution. Thus analysis of these characters for homologies used in systematic work must be done with caution. Morphological classifications of the salticids have not been successful because many have been based on a few relatively simple characters [e.g. ret roma rg i na 1 cheliceral teeth or proportions) which may be derived in unrelated forms by convergence. Genitalia, although secondarily reduced in this family, are perhaps the most promising morphological characters for systematic use and Proszyiiski (1976) has proposed a classification based on these. Courtship Behavior and Phylogenetic Relationships Crane's behavioral groupings should now be discarded in favor of groupings based on comp 1 ex homologous courtship behaviors between genera. The latter can, with some caution, be used as additive evidence to sub- familial arrangements based on genitalia or other morphological charac- teristics and either support or detract from the probability that these arrangements are phy 1 ogenet i 1 ca 1 y correct. In Table 5 a total of Ih genera are presented as placed in various subfamilies by Pet run kev i tch (1928) and Proszynski (1976). Differences i ' between Pet runkev tch and Crane (I9'f9b) are also noted. Pet runkev i tch s system was based on external morphology, excluding genitalia. Proszynski 's system was, on the other hand, based totally on genitalia. The last column shows a possible placement of genera based on presumed courtship homologies and some morphological evidence. In many cases courtship homologies are not found throughout the subfamily, but seem to relate groups of genera through similar behavior in certain included species. — — ' — — — —. — - — 112 4—' c O OJ . E > cu 0) cu 0) 0) 0) CD CD CD CD u c _c o« CD C C C C ro O 0) 0) (U CU 0! c CQ (H 0) QJ CD CD CD cu 4-1 4-1 4-1 4-1 Q_ c (H CB C C C C CD CU a) C C c C OJ D- C C C CD CD CD CD CD CD 0) l/l — c , , , ._ C c JZ x: JZ _IZ . ro -C TO l/l l/l , , CD . CL Cl CL Cl .— .— CQ Ul E l/l — 1_ CU 0 >- >~ >- >- • — -t-l O .— L L. 1_ CD tn s_ s_ i_ i_ L- in (3- 3 D E CD 4-1 4-1 ~a "D XI T3 ID D l/l 1_ 1- , , , L- c 4-1 4-J c C c C O o >- CD CD CD CU > > cu (U cu CU D Q. o _j s: s; 2; 00 l/l CO Q Q Q Q •— < < < U ID M- O (U 1) CU CU TO CD CD CD CD I. 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CO rO CO CO CO CO CO ;^ S ^) o O CO o H C CO O \o ca OJ o t Menemevus in the Ael ur i 1 1 i nae . From published accounts of the court- u1 ship ^u-'tuvillus (Bristowe, 1929, 19^1, 1958) I am also somewhat reluctant to use this subfamily. However, the genera Menemevus and Metacyrba are probably closely related and they do not seem to belong to the Marpissinae. Several different courtship behavioral characteristics were presumably homologous and may be useful in establishing higher relation- ships. Crane touched on this in her diagram (1949b, Figure 2) of a hypothetical phylogeny by using such terms as "poser", "foreleg waver", "eye-color shifters", "abdomen twister", "foreleg lifter", etc. However, she did not expand this idea in her discussion of phylogeny (p. 205-206). The following characteristics of courtship seem to link species and genera into distinct groupings. Posture appears to be the most conserv- ative, and thus the most useful systematic character, with movement less conservative and less useful. Prosoma kept elevated, opisthosoma pendent and long periods spent in posing The species of Lyssomanes [L. bradyspilis and L. viridis) exhibit these characteristics. They seem to warrant at least a separate sub- family, if not family. I doubt that the movements of the anterior median eye retinae (these are Crane's "eye-color shifters") in courtship 115 are unique to Lijssornanes . Such shifts have been seen in other salticids (Bristowe, 19^1; Land, 1969b) especially in relation to prey capture. In Lyssoinanes these may be simply scanning movements by the retinae of the male, which are accelerated because of the heightened stimulus of the female presence. The unusually transparent prosoma of Lysomnanes makes such retinal movements more noticeable than they would be in other salticids. Opisthosoma raised or raised and twisted, some species exhibiting a zig - zag motion with prosoma elevated The species of the genus Marpissa tend to exhibit courtships in which the prosoma is raised nearly perpendicular to the substrate. One species, M. snlcosa, differs from the others in twisting the op i s t hosoma to the right or left and exhibiting a zigzag motion with the prosoma elevated. It is the latter motion and elevation of the prosoma which seems to link Marpissa to Maevia. The genera are also similar in several morpho- logical points, such as in having four pairs of spines on the ventral first tibiae. These two genera are placed in the Marpissinae on these bases. The subfamily is more restricted in Proszynski's classification, including only Marpissa. In later stages of courtship first legs raised, opisthosoma strai ght and path to female direct; prosoma usu ally low The genera Menemerus , Metacyrba and Phlegva seem to be related to the similarities of their very simple courtships. David Hill (personal communication) has found the opisthosoma scales of the first two genera to be nearly identical and unlike those of Marpissa. There is some . . y , 116 doubt as to whether these should be placed in the Ael u r i 1 1 i nae , but they do not seem to belong to the Marpissinae as thought by Pet runkev i tch (1928) and Barnes (1958). Fhlcgra has been placed in the Pelleninae (Petrunkevi tch) but does not seem to belong there based both on court- ship and gen i ta 1 i a First legs raised, opisthosoma bobbed, and palpi extended lat e r a 1 1 The members of Sarinda seem different enough in their courtship that they should be separated from the Synagelinae, rather than being placed with the genera from the latter into the Synemosyn i nae , as Proszynski has done. The raising of the first legs is also seen in Synemosyna. First legs lowered, opisthosom a raised vertically or moved. from side to s i de The members of the genera Gertschia, Synageles, Semovina and Feckhamia (see Crane's Table II, reproduced as Appendix A) have a completely different courtship approach, and their genital morphology seems also to separate them, from the My rama rach i nae First legs raised, courtship a slow zigzag There is little argument on the placement of Sittioiis and Feon in the Sitticinae, based on morphological and behavioral similarities. First legs raised and prosoma elevated or first legs and pros oma 1 owe red ~ often with a typical side-stepping movement and often hav i ng op i sthosoma twisted to the right or lef t The genera in the Dend ryphant i nae are in some ways, difficult to characterize. While this subfamily is well characterized morphologically, 117 behaviorally it is pol y thet i c--no one courtship characteristic being common to all. Yet, most of the examined genera are related to one another by one or more behavioral traits. For example, opisthosomal twisting is common in some genera [Hentzia, Zygoballus and some Pliidippus) and one of these, Phidippus is , linked to Eris, Thiodina and Tutelina by its characteristic side-stepping hesitating movement, with first legs raised and prosoma elevated. Metaphidippus and Sassaaus , which are related to Phidippus and Eris by the morphology of the genitalia, have species [Metaphidippus vitis and Sassaous papenhoei) which exhibit very similar courtship postures, with the first legs raised and crossed and the prosoma lowered. Both species jerk the first legs up and down at the start of their sideways runs and both have zigzag or spiral paths to the females. Zygoballus has both a similar morphology and similar epigamic displays to the dend i ryphant nes . In morphology it resembles Evis and in courtship and agonistic display it resembles Hentzia. Thiodina is unusual in many ways, but it has similarities in genitalia and behavior to several of the dend i ryphant nes . Many of the genera exhibit mosaics of derived and ancestral characteristics. First legs lowered, body moved in a rapid zigzag or vibrated and palpi often utilized in display The genera examined in the current study [Covythalia, Habrocestum and Ueonella) seem to be related by their courtship. This relationship is especially evident between some species of Covythalia and Hahvocestum. Proszynski (1976) has placed all three genera into the Euophrydinae and has presented convincing evidence of the relationship of these with 118 each other and with EuopJiPys, based on genital morphology. However, the courtship of Comjthalia auvata and the three species of IJabpoaostwn differ from EuopJirijs in that the male of Euopiwjs poses with the first legs raised nearly vertically (Bristowe, 1929, 19^1, 1958). Again, the presence of similarities in courtship between certain species in different genera, which appear to be related morphologically, is taken further evidence of their phylogenetic relationship. Genera difficult to c ha racter i ze Salticus does not belong to the Marpissinae, as thought by Petrunkevi tch (1928) and Crane (1949b). Its zigzag courtship (Bristowe, 1929, 19^1, 1958) is in some v^ays similar to Eentzia, but the abdomen is not raised or twisted. It seems to belong to its own subfamily, as suggested by Proszyriski (1976), although it is morphologically related to at least one other genus, Bvedayici. If SciZ'b'Lcus had been found to be related to Marpissa, the subfamilial name would become the Salticinae rather than the i Marpissinae (Pet runkev tch , 1939). Pellenes , while morphologically distinct, is such a diverse genus with respect to courtship that it is difficult to characterize. It seems best to retain the Pelleninae for this genus, while noting the difficulties involved. Plexippus has a rather simple courtship. It is certainly not related to the rapidly moving Cori/thalia as has been suggested. The subfamily Plexippinae is retained for this genus mainly because it does not seem to fit elsewhere. . 119 Possible phylogeny of some salticid subfamilies Some morphological and behavioral similarities between subfamilies suggests a basis for phylogeny. The Marpissinae seems to retain more "primitive" traits, such as four pairs of ventral tibial spines on the first legs and relatively complex male genitalia. Some courtship behaviors, such as the twisting of the opisthosoma in a Marpissa suloosa- like ancestor, may have been the origin for similar behavior in the Dendryphantinae and Synagelinae. The latter seem to be related to the Dendryphant inae by the genital morphology as well. The ant mimics pose a difficult problem, but differences in courtship, ant mimicking behavior and morphology seem to separate the Synagelinae from the Myrmarachn inae. It is doubtful that ant mimicry arose only once in the family, as Proszynski (1976) would seem to have indicated by combining the two subfamilies into the Synemosyn i nae . The phylogenetic lines leading to the other subfamilies are, at least for the present time, difficult to un rave 1 Courtship Behavior and Species-Level Systematic s The details of courtship can be used as a basis for distinguishing two species when the possible two species are closely related and sympatric (Table k) . Examples of closely related sympatric species which can be easily separated by their courtships include Pellenes calcaratus and P. viridipes , and Hahvocestwn bufoideo and H. n. sp. A. Both species pairs are similar morphologically, although they can be separated by some details of the male ornamentation. Pellenes calcavatiis males have a different pattern of first legs and palpal . 120 movements than do males of P. viridipes during courtship (see Figures 16-17)- Habroaestmi bufoides males exhibit shorter, more constant zigzags than males of H. n. sp. A and also differ in tilting the body to the right or left and in the pattern of palpal movement (Figures 11-12). Closely related allopatric, parapatrlc or ecologically isolated species may not have developed the premating behavioral isolating mechanisms that permit easy separation on the basis of behavior. No distinct differences have been found between the courtships of Pellenes P. brimneus and coeaatus^ Hentzia palmarim and //. grenada or Thiodina puerpera and T . sylvana. In the first case the species seem to be para- patrlc (possibly due to competitive exclusion) and in the last two cases the species are ecologically isolated. In at least one pair of allopatric species [Pellenes coecatus and P. cf. coeoatus) some differences have evolved in the courtship behavior, although the court- ships are similar enough to show a close relationship. Some differences in the morphology of the male palpus (W. Maddison, personal correspondence) indicate that these are probably separate species, and not just geographic varjants. In those cases where both morphology and courtship fail to separate two ecologically distinct species, it may be necessary to hybridize them to see if postzygotic isolation occurs (see section on sexual selection and Appendix C) Sexual Dimorphism and Epi garni c Display From the species which I have studied in Florida and Arizona and from published accounts, it was generally found that those with the most 121 complex courtship ritual [e.g., Pe'llenes) had the highest degree of sexual dimorphism. This dimorphism was primarily based on ornamentation of the male, but. in some cases {Ex'is^ Hentsiaj Savinda, Zygoballus) males had enlarged or elongate chelicerae which were used in male-male fighting. The least sexual dimorphism was exhibited by species of Metacyrha. These have relatively simple courtships. An even simpler courtship was exhibited by Menemevus bivittatus , which showed some unexplained pattern differences between males and females (Figures I8-I9). These differences were not involved in the courtship display. There was some evidence for contact sex pheromones in both Metacyrha and (Appendix B) Menemerns , which might aid males in finding adult or subadult females within their silk retreats. This and the simplicity of the displays suggests that cohabitation may be common in these species and therefore morphological sexual dimorphism is not necessary. Sexual dimorphism was found to be generally more pronounced in the other species examined. One exception was the so-called "hopper", Covythalia aurata, which did not show the complex sexual dimorphism of Crane's South American species of the same genus but did have some epigamic ornamentation. The courtship of Corythalia aurata was quite similar to that of some of Crane's species. Agonistic display was well developed in C. aurata and, unlike Crane's species, males often fought, using their chelicerae at close range (Figure 30). Smaller males usually retreated before actually fighting and this type of behavior may have led to the totally ritualized agonistic displays found in the South American species. 122 The most extreme examples of sexual dimorphism are exhibited by species of Pellenes . Males of such species as P. brunnend , P. coecatus, P. cf. aoecatus, P. aalaaratus and P. viruKpss have highly modified third legs. Males of many species have fringes of setae on the first legs and some have spatulate spines. In most species the male and female exhibit different patterns and coloration. The modifications of the male legs are always displayed in courtship. Most species do not seem to have agonistic displays, contrary to Crane's hypothesis, but the difficulties involved in interpreting negative laboratory evidence have not permitted firm conclusions. From Table 3 it can be seen that the genus is very complex and difficult to characterize on behavior alone. Pellenes is one of the largest genera, with at least 60 species in the United States and Canada and with 11 species in Florida alone (Appendix D) . There are fewer species in the tropics and probably none are found in Venezuela where Crane did her work. It seems that courtship may have been the major factor in the evolution of much of the sexual dimorphism observed in salticid spiders. This has been primarily the result of selection for prezygotic isolating mechanisms, selection for differences between the sexes functioning for the identification of sex, or a combination of these factors (Mayr, 1972). Sexual selection may also be involved, but selection pressures are probably so intertwined that separating them into discrete units may not be possible. in salticids the sequence of the courtship movements and persistence of the male are behaviors probably controlled in sexual selection by the female, as has been found in Drosophila (Maynard Smith, 1956 and others). 123 Sexual Selection and Sexual Dimorph i sm Sexual selection is now recognized as natural selection, but Darwin (1859, 1871) made a distinction between natural selection (selection by competition for resources due to the overproduction of progeny in variable populations) and sexual selection (selection by female choice and male competition). He even suggested some form of rudimentary aesthetic sense in females which allowed them to pick the most ornate male to fertilize their eggs. Thornhill (1976) reviewed the subject and retained the term for female choice (intersexual selection) and male competition for mates (intrasexual selection) where the selection can be demonstrated quantitatively. This does not necessitate the presence of any aesthetic sense in the female. Female choice has been demonstrated in a number of organisms, but often the selectional process which led to such choice has been misunderstood. Mayr (1972) points out that there are other possible interpretations of the existence of sexual dimorphism without the necessity of invoking sexual selection. The advantage obtained in having ornamentation may be, as Mayr states, that males are more readily recognized as the correct sex and species. If male orna- mentation proceeds to the point where predator and parasite pressures overcome the advantage in obtaining a mate, these majes and their genetic line ai'e eliminated. Females have a higher investment in reproduction than males and it is thus to the female's advantage for her to resist until the male has demonstrated his vigor by performing the species-specific courtship correctly and persisting in his attempts to mate with her (Williams, I966, 1975; Trivers, 1972). Examples of this are found in species of Pellenes, Habrocestum and Cory thalia. In ]2k some species of Pellenes males may court for as much as 30 minutes before being able to mount. Females of Babrocestwv pulex and Corythalia auvata raise their first legs and elevate the prosoma during courtship. Success- ful males have to climb over the elevated prosoma, while forcing the female's first legs down. On the other hand, females of some species (those of Metacyvba^ Menernerus and Hentzia) will, if virgin, mate with almost any male, even if the courtship display is not fully performed. If females of a species are very aggressive it is advantageous to the male to avoid direct contact until after he has "gauged" the initial reaction of the female, rather than running directly to her and in turn being killed and eaten. Historically males which did not behave in this manner were selected against. This might explain the usually zigzag or spiral approach in many species (Table 3) and does not require some mystical female appreciation of the dance. The presence or absence of specific ornamentation and courtship movements may be criteria utilized by females for rejecting or accepting the male's attentions prior to mounting. Males of Pellenes often court females of different species (as also noted by Crane for several other genera, 1949b, p. 210), but the display usually ended with the female either leaving or jumping at the male. Between some similar species, which are geographically or ecologically isolated in nature, such barriers are not necessary to isolate and may not exist. Males of Heiitzia palmaiVMn and fJ . grenada readily courted females of the other species and virgin females of either species vjould mate with them. Several masses of fertile eggs resulted, but the young never matured (Appendix C). A large male of Thiodina puerpera successfully courted a virgin female of . 125 T. sylvana, but she laid only infertile eggs. Finally, G. B. Edwards (personal communication) successfully hybridized Phidippus regius with P. otiosus (reciprocal), obtaining an generation in which males were sterile and had a courtship display intermediate between the two species. All of these examples are of species which are ecologically separated i n the w i 1 d Agonistic displays are less common, at least in the laboratory, than courtship displays among salticids. Male competition for mates is probably the basis for such displays. It is rare in nature for more than one male to find a female at the same time, except in those species which usually have a very high population density (such as in Hentzia or Cory thalia) or perhaps utilize pheromones (such as in Menemerus) . Corythalia aurata males seem to recognize superior size and smaller males often retreated from larger males after only a short display. If many species practice cohabitation courtship, as in the case of Evis marginata (Peckham and Peckham, I889) and Phidipvus ,johnsoni (Jackson, 1976) and drive away other males, fighting and agonistic display would be at a selective advantage. Males of Corytlialia aurata and Thiodina sylvana have been found within or near the retreats of adult or subadult females of the same species in the wild during the current study. Also, in the one observed agonistic display of Menemerus hivit-tatus , one of the males was inside a silk retreat initially. Such circumstantial evidence may indi'cate that males guard their own retreats, or the retreats of females with which they are cohab i tat i ng . Further study of this problem is necessary before any conclusion can be made. 126 Summary Crane's behavioral groupings are mucii too simple to be used for classification in the Salticidae and should now be discarded. Presumed behavioral homologies of complex courtship behaviors can, with some care, be used as a way to arrange related genera into subfamilies and to serve as supporting evidence for classifications based on morphology. Closely related and sympatric species can, in several cases, be separated by courtship behavior. Some closely related allopatric, parapatric or ecolo-g ical ly separated species are not so easily separated because they may not have evolved premating behavioral isolating mechanisms. Hybridi- zation experiments may demonstrate postzygotic isolation in closely related species not separable by courtship or morphology. The evolution of male ornamentation Is probably related directly to the advantage derived from correct species and sex identification by the female. Sexual selection (choice by the female of certain males within her species) may play a lesser role in the development of male ornamentation. Possible criteria used by the female in sexual selection include the proper sequence of the male's courtship and the persistence of the male. The adaptive radiation of the salticids has left the systematlst with a complex pattern of parallel and convergent evolution, and there is little fossil evidence with which to interpret it. The evolutionary pattern within the family can only be worked out through indirect evidence, including the use of courtship behavior and morphology. APPENDl CES APPENDIX A COMPARATIVE DISPLAY DATA IN SEVEN SUBFAMILIES SALTICID SPIDERS (Crane, 19'49b, Table II, p. 162-171) . 129 Name Display References Courtship Posture (Stage 1) Marpissinae M arpissa undata Peckham, '89, p. 43; fig. Carapace high. (De Geer) {"Marptusa familiaris") 1st & 2nd legs raised slightly, (U.S.A.) stretched laterally. Abdomen pendent. Marpissa riimpfi Bristowe, '29, p. 330 352.. Carapace high. '41, fig. 1st legs raised vertically, parallel. Scop. Bristowe p. 480 ; (England) Abdomen raised. Mencmerus bhnitatus Crane Carapace low. (Dufour) (Unpublished observations) 1st legs raised slightly, stretched (Venezuela) antero-laterally; 2nd legs ex- tended forward. Abdomen level. forward of Hyctia 7iivoyi Luc. Berland, '27, p. IB; fig. 1st legs stretched @ Z 45°; raised & lowered. Bristowe, '29, p. 329 ; fig. (cf. '41, Abdomen often bent sideways, Bristowe, p. 484 ; fig. (Europe) Dendryphantinae) low. Hyctia pikei Peckham Kaston, '48, p. 455; -fig. Carapace (U.S.A.) 1st legs raised vertically @ Z of 45°. 2nd legs extended forward. Abdomen raised. scenicus. Saltimis cingulatus Pani;. Bristowe, '29, p. 332. Similar to S. (England) raised, stretched Salticiis ttceyiicus (Linn.) Peckham, '89, p. 39. Ist legs slightly {" Epihle.mum scenicum") laterally. Gerhardt, '21, p. 131. Cheliccrae opened. {"Epihle.mum scenicinn") Palps stretched laterally. Monterosso, '24, p. 1. Bristowe, '29, p. 332-. Bristowe, '41, p. 480, 499; fig. (Europe & U.S.A.) Synagelikae parallel. Syjiageles vejiaior Luc. Bristowe '41, p. 485. 1st legs stretciicd forward, (England) Abdomen wriggled or waved from side to side, sometimes raised a little. 90° Seniorina bracJiyclielyjie Crane, '49, p. 37. 1st legs stretched forward, @ slightly raised. Crane (Venezuela) Z , Abdomen raised; tends to remain liigh with increasing excitement. GeriRchia noxio:ia (Hentz) Kaston, '48, p. 451; fig. Ist legs braced laterally. (U.S.A.) Abdomen raised vertically &: swung from side to side. Peckhaniia picata (Hentz) Peckham, '89, p. 43. Carapace high. '90, 1st legs on ground, bent, convex sur- Peckham, p. 121 ; fig. (U.S.A.) face forT.vard. Abdomen raised vertically. .Spider sways from side to side. 1 30 Principal Apparent Locomotion & Remarks Threat Posture & Fighting Morphological Adaptations to Display Marpissinae (cent.) Slight enlargement basal half 9 vibrates palps. of 1st & 2nd legs. Jaws opened; pressed against 1st legs elongated, enlarged, those of opponent. darkened. enlarged, Runs; jumps only on prey or Uusually $$ pay no attention to 1st legs elongated, to cross gaps. each other; no distinctive mo- darkened. 1st legs little used in walking. tions; rarely courtship to mir- 9 vibrates pale palps. ror image in closed vial. 1st legs elongated, enlarged, darkened. 1st legs elonga-ted, enlarged, 1st legs not used in walking, darkened. extended forsvard & raised Abdomen boldly striped. to clear ground. Chelicerac elongated, en- larged, darkened. 1st legs slightly elongated. Jerky walk; hops when pur- Like courtship, but jaws opened Chelicorae elongated, en- sued. (Crane obs. U.S.A.). wider and palps not .stretched larged, darkened. slightly elongated. to sides. 1st legs (No mirror display. Crane obs. U.S.A.) Synagkunab (cont.) 1st legs enlarged. enlarged, Runs; jumps only on prey or Apparently no inter-male dis- 1st legs elongated, to cross gaps. play. No mirror display. darkened. Abdomen dark with 2 dorsal 1st legs stretched to front, white spots. parallel; do not touch ground. 9 vibrates palps. 1st legs elongated, enlarged,' Runs slowly, irregularly, like darkened. ant. dorsal (Emerton, 1909). Abdomen , dark with white bands. 1st legs elongated, enlarged, darkened. Abdomen dark with dorsal white bands. — , 131 Name Display References Coui-tship Posture (Stage I) Lyssomaninae Lyssojnanes bradyspilus Crane, '49, p. 34; see also Carapace high, occasionally bobbed. braced Crane Text-fig. 8, this paper. No leg.s raised; 1st 3 prs. (Venezuela) forward. Palps sometimes tap ground. Abdomen pendent, twitched. Prolonged posing in this position, as Rate of retinal motions within AME are accelerated. Dendryfuantinae Ashtabula fur cilLata Crane, '49, p. 41; fig. Carapace high. Crane (A'enezuela) 1st legs stretched laterally, waved up & down. Palps jerk; later arc quiet. Abdomen turned to side. Hentzia mitrata (Hcntz) Peckham, '89, p. 49; fig. Carapace high. {"Icius mitratus") 1st legs stretched laterally, held at (U.S.A.) right ^ to body. Abdomen twisted to side. Jcius elegaiis (Hentz) Peckham, 89, p. 46. Ist legs waved "in way that reminds ("Dendryphantex' elegans' one of a wind-mill". Later, re- (U.S.A.) volves on tip-toe. SasKacus ocellaius Crane Crane, '49, p. 45. Carapace scarcely elevated. (Venezuela) 1st legs stretched up & out at right /£ to each otlier. Cheliccrae closed. Palps vibrated occasionally. Abdomen sometimes twisted slightly to side, held motionless. Sassacus fUivicinctus Crane, '49, p. 41. Carapace moderately high. Crane (Venezuela) 1st legs stretched up at wide i. to each other. Chelicerac stretched sideways, but closed. Palps stretched sideways. Abdomen trailed inconspicuously from side to side. Later: Carapace & abdomen slowly rocked from side to side. M ctaphrdippus protarvus Peckham, '89, p. 45; fig. Carapace low. (Walck.) and/or ("Dendryphantes capitaUis") 1st legs stretched forward, close to M. galalhca (Walck.) (U.S.A.) ground, slightly curved with tips turned up. Palps given circular movement. Later, lies on side, legs still extend- ed. Paraphidippusmarginatus Peckham, '89, p. 50; fig. 1st legs raised, curved toward each (Walck.) ("Philaeus militaris") other, tips nearly meeting. (U.S.A.) Palps moved up and down. 132 Principal Apparent Threat Posture & Fighting Morphological ' Locomotion & Remarks Adaptations to Display Lyssomaninab (cont.) Threat display rarely induced; Distal portion of black retina Runs in short spurts. Jumps posture as in courtship, but no contrasts with green sur- only in final pounce on prey. bobbing, no abdominal twitch- roundings. Palps pendent, pat ground ing, no acceleration of retinal Chelicerae elongated but not during pauses. motions. specially displayed. 1st legs take part in locomo- Fighting & mirror display ab- tion. sent. 9 sags to side when watching display; muscular activity in eyes increase as in $. Dendryphantinae (cont.) Apparently no inter-rnalc dis- 1st legs elongated, enlarged, Runs; jumps only on prey or play. No mirror display. darkened, fringed. to cross gaps. Abdomen with lateral white 1st legs unused in progress, stripe bounded by irides- held forward, they & palps cence. tap ground; waved in air in pauses. 9 vibrates palps. Same as courtship. 1st legs elongated, fringed. 9 "indifTerent"; sometimes at- Chases rivals away. Abdomen with lateral white tacks stripe bounded by darker. Carapace with frontal tufts. 9 irritable, prone to attack; 1st legs elongated; tibia later rai.'^es abdomen or fringed, black-spotted. turns it sideways. Both sexes covered with iri- descent scales. Stage I: Similar to courtship, 1st legs elongated, enlarged, Runs; jumps only to catch except chelicerae partly un- darkened, with white scales. prey & cross gaps. sheathed, & no distinct ab- Chelicerae elongate, enlarged 1st legs held forward, scarce- dominal twisting. darkened. ly touch ground; these & Stage II: Actual fighting, 1st Clypeus with white band. palps wave in air & tap legs raised vertically; chel- Abdomen with large, sub- ground, during pauses. icerae wide open; palps ex- distal, white-barred black 9 once did weak mutual dis- tended laterally. Clinching spots. play. frequent, with occasional in- jury; no deaths seen. Stage I: Like courtship except 1st legs elongated, enlarged, Runs; jumps occasionally no rocking. darkened. when pursued, as well as to Stage II : ( Rare) . Chelicerae un- Chelicerae elongate, en- catch prey & cross gaps. sheathed, extended forward; larged, darkened. Palpates ground & waves 1st 1st logs raised vertically; Clypeus with yellow band. legs less than S. ocellatus. clinching; no injuries seen. Occasional mirror display. Distinct from court.ship. 1st legs 1st legs elongated, enlarged. Runs, but liops freely when raised. Sparring, chasing & White markings on dark- pursued, as well as to catch clinching without injury. palps & clypeus. prey to cross gaps. Palps often jerk. 1st legs not waved. Take some part in walking. (Crane obs. U.S.A.) Distinct from courtship. 1st legs elongated, enlarged. 3 3 guard immature 99 until 1st legs raised. Chelicerae elongated. molt. Palps vibrated. Abdomen dragged to either side. Chasing and clinching without injury. ) 133 Narnc- Display References Courtship Posture (Stage 1) Dendryphantinae (cont.) Phidippus audax (Ilentz) Peckham, '89, p. 45. Carapace moderately high. ("Pliidippiis morsitans" 1st legs raised in 2 jerks, stretched Kaston, '36, p. 120; fig. obliquely out at Z of 45° to (U.S.A.) gi-ound ; later waved alternately. Palps raised and lowered rapidly when close to 9. PhidippiLS clams Kaston, '36, p. 118; fig. Carapace moderately high. Keyserling (U.S.A.) 1st legs stretched out, tlic lemur obliquely up, other segments paralleling ground. Palps sometimes widely spread. Abdomen now & then moved from side to side. Phidipims purpuratus Kaston, '36, p. 121; fig. Carapace high. Keyserling (U.S.A.) 1st legs stretched out, held higher than in clarus & audax; waved. Palps held wide apart, parallel with each other, occasionally drummed on ground. Abdomen dropped. P}iidippus whitrnanii Peckham, '89, p. 44, Carapace high. Peckham ( "Pliidippus rufiis") 1st legs stretched forward k up, (U.S.A.) crossed at tips. Palp.s held wide apart, parallel with 2nd legs. Abdomen dropped. Sways during advance. Hyllinae Evarcha falcata Bl. Bristowe, '29, p. 333. Some waving of 1st legs & twitching Bristowe, '41, p. 480. of abdomen, but tends to leap on (England) 9 with almost no display. Homann, '28, p. 249; fig. ("E. hlanchwrdi") (Germany) Pliilaeii^ chrysops Poda Borland, '14, p. 116. Carapace high. Thomas, '29, p. 267. 1st legs raised & waved up & down. Bonnet, '33, p. 139; fig. Palps vibrated. (France) Abdomen slightly elevated. Courtship sometimes almost absent. Leaps on 9 suddenly. Pliialc flxnnviea Crane Crane, '49, p. 48. Carapace high. (Venezuela) 1st legs raised at 45° / to each other & ground. Palps vibrated irregularly. Abdomen lowered. Carapace rocked from side to side. Later, sinks lov.', 1st legs stretched to front. Principal Apparent Threat Posture & Fighting Morphological Loconaotion & Remarks Adaptations to Display Dendryphantinae (cent.) Like courtship until opponents 1st legs elongated, enlarged, Both sexes savage, prone to close; then cholicerae &.fangs with white fringes and cat mate. opened wide; some 9 fighting. scales. Palps with white scales. As in P. audax. 1st legs elongated, enlarged, lightly fringed. .Palps with white band. 1st legs elongated, enlarged, heavily fringed. Clypeus with white fringe. 1st legs elongated, enlarged, Hopping well developed. heavily fringed. Hyujnae (cont.) 1st legs elongated, enlarged, darkened ; palps pale, white- haired. 1st legs elongated, darkened Excellent visual acuity; hops with light tufts & scales. skilfully when pursued. Berland & Thomas saw fre- quent leg-waving when spiders were alone in field and in clean boxes; Bonnet saw waving only before 5$. Differs from courtship in quiet 1st legs elongated, darkened. Basically a runner although palps & absence of low stage. Clypeus & palps with bufly jumps readily over gaps; At close quarters, chelicorac yellow bands against black. 1st legs little used in walk- open, 1st legs spread widely, ing, habitually waved dur- often touch opponent's; no in- ing pauses; true also of $9 juries. & young. Displays freely to mirror. 9 raises ist legs & vibrates palps during display. . . 135 Name Display References Courtship Posture (Stape I) Plexivpinae (unpublished ohs.) Carapace very hiph. Plexippus paykullii Aud. Crane ; PI. I, fig. 1, this paper. Ist leps stretched forward, up & out, (A''cnczueia) at varying wide ^, higher with excitement; no waving. Palps quiet. Abdomen pendent. Prolonged posing- in display posi- tion. Eustiromasiix sp. Crane (unpublished obs.) Carapace high. (Venezuela) 1st legs raised antero-laterally @ /_ of 45° to ground; sometimes jerked, & raised higher. Palps vibrate. Abdomeji horizontal. Some posing in display position. Saitis barbipes Sim. Berland, '23, p. 206. 3rd legs wave. Berland, '27, p. 15; fig. (France) CorytJi.alia xanUiopa Crane, '48.1, p. 35; figs, (also, Carapace moderately high. Crane see footnote, this paper, p. Palps hanging, motionles.s. 183). Abdomen slightly below horizontal. (Venezuela) Stage la: Side-to-side rocking, all feet on ground. Stage lb: 1st legs stretched for- ward, straight, parallel (a) I q( 45" to ground. No posing in display position. Coryth.alia clmlcea Crane Crane, '48.1, p. 21; figs. Carapace moderately high. (Venezuela) Palps hanging, motionless. Abdomen about horizontal. 3rd legs stretched out, waved up & down in unison above the hori- zontal. No posing in display position. Corythalia fulgipedia Crane, '48.1, p. 28; figs. Carapace progressively lowered. Crane (Venezuela) Palps jerked in unison. Abdomen about horizontal 1st, 2nd, & 3rd legs stretched far out to sides, the 3rd legs being slight- ly intermittently raised & vi- brated up & doATO in unison with body. No posing in display position. 136 Principal Apparent Threat Posture & Fighting Morpholog-ical Locomotion & Remarks Adaptations to Display Plexippinae (cont.) No intor-malc displays seen. 1st legs elongated, enlarged, Hops often even during ordi- Mirror response not tested. darkened. nary progress. White clypeu.^ band against 9 completely passive, black. Scarcely developed; rarely, brief 1st legs slightly longer, thick- Hops often even during ordi- threat similar to early court- er, blacker than others. nary progress. ship but abdomen lowered & 1st legs take active part in palps quiet. walking. Occasional brief mirror display, Palps quiet. sometimes ending in attack. § extremely savage both be- fore & after mating; often kills $. Same as courtship. 3rd legs elongated, com- Performs display motions pressed, fringed. even when alone in clean boxes. Completely distinct from court- Palps & clypeus with yellow Ordinary progress consists ship. bands. largely of hopping. Carapace very high. 2nd, 3rd & 4th legs fringed, 1st legs take active part in Palps motionless, flexed, their compressed, with iridescent walking. yellow scales continuing band patches; 3rd & 4th legs Palps quiet. of clypeus. elongated. 9 not aggressive; when watch- Abdomen pendent. ing display, sits high with 2nd, 3rd & 4th leg.s off ground, braced legs until receptive, raised successively higher. then crouches. Prolonged posing. No fighting. Displays freely to mirror. Distinct from courtship, although 1st, 2nd & 3rd legs fringed, Locomotion as in xajithopa. same legs used. compressed, with iridescent 9 sometimes aggressive, may Differs in waving 3rd legs below patches; 3rd legs elongated. do some form of reciprocal horizontal, often in an arch; display. prolonged posing. Palps flexed & stiff. No fighting. Displays freely to mirror. Distinct from courtship, although 1st, 2nd & 3rd legs fringed, Locomotion as in xanthopa. similar in very first phase, & compressed, with iridescent 2 more aggressive than most same legs used throughout. patches strongly developed; salticids, although not seen in legs Whereas courtship carapace 3rd elongated. to injure $ ; when stimu- sinks lower, in threat it rises Palps with white patches. lated perfonns rough, re- higher. At peak, all legs are ciprocal display. drawn close in & stretch up, 2nd tarsi leave ground & 3rd legs are raised slightly; body rocks from side to side; X'dsing wth 3rd legs arched frequent. Palps jerked at beginning, later flexed & stiff. No fighting. Displays freely to mirror. . 137 Name Display References Courtship Posture ( Stage I) Magoninae Mncvia vittata (Hentz) Peckham, '89, p. 54; fig. Displays of the 2 forms differ. ("Astia vittata") Gray form: Carapace high ; 1st legs (Wisconsin, U.S.A.) raised, waved, palps stretched Painter, '13, p. 625. sideways; abdomen down; later, (Connecticut, U.S.A.) whole body low, 1st & 2nd legs forward, tips touching; palps forward. (Peckham, Wise). Low stage precedes high stage. (Painter, Conn.) Black, tufted form: Carapace very high; 1st legs raised & waved, or held high in pose. (Peckham & Painter). Mago dentichelis Crane Crane, '49, p. 48; fig. Carapace slightly raised. (Venezuela) 1st legs stretched sideways & slightly forward; waved alter- nately up and down or held in pose. 2nd tarsi touch ground, far out & slightly forward. Abdomen sometimes vibrated. Baling depressiis Walk. Bristowe, '31, p. 1409; fig. All legs on ground, drawn in, sways Bristowe, '41, p. 484; fig. body from side to side. (England) 138 Principal Apparent ThreJit Posture & Fighting Morphological Locomotion & Remarks Adaptations to Display Magoninae (cont.) Wave 1st leg-s at each other but Tufted form is black anteri- Peckham & Painter disagree "quarrels . . harmless" orly, has cephalic tufts. on selection value of di- (Peckham). morphism Dijplaya to mirror. gg irritable; fight each other. Stage I: Similar to courtship. 1st & 2nd legs enlarged, 5 other Venezuelan magoni- Stag-e II: 1st legs waved higher, blackened. nids use all legs actively in aLmoat meet overhead; when walking, hop freely during lowered, 1st & 2nd tarsi rub ordinary locomotion. together. Rarely, chelicerae All include rubbing of 1st & opened and knocked repeatedly 2nd tarsi in display. against opponent's; no inju- 9 usually extends 1st legs dur- ries. ing courtship. Displays to mirror. 1st legs enlarged, partly 9 vibrates light palps. blackened. APPENDIX B A CONTACT SEX PHEROMONE IN MENEMERUS BIVITTATUS Crane (l9^9b) hypothesized that there might exist some chemical substances which were utilized by salticid spiders in courtship. These substances, or pheromones (chemical's which are utilized for communication within a species, often in relation to reproduction) have been found in many diverse organisms. The recent discovery by bioassay techniques of contact sex pheromones in some lycosids (Rovner, I968; Hegdekar and Dondale, 1969; Richter, 1971; Dondale and Hegdel interest in spider pheromones, although none has been isolated to date. The main reason for examining pheromones in the present study was to determine the importance of such chemicals in displays and the extent to which they might cause difficulties with the interpretation of visual displays {e.g., distraction of male or atypical courtship). No evidence of such a problem was found, but at least one species [Menemerus bivittatus) was found to have a contact sex pheromone which may allow the male to locate the female in the absence of visual feedback. Males of Menemerus bivittatus (and also Uentzia palniarum and Pellenes brunneus) were placed into petri dishes containing two half circles of filter paper (a modification of techniques used by Hegdekar and Dondale, 1969)- One of these halves had been in contact with a virgin female for 2k hours and the other half was clean (control). Each male was given three minutes In the dish and each test was filmed 139 1^0 at 2 fps. Tfte last two minutes of each test was used For analysis so that the male would be as little disturbed as possible. The males of M. bivittatus spent 73% of the time with the female-contacted paper (n = 10, SD = + 1.8?;, males = 2) and also exhibited distinct behavior while in contact with the paper. They were especially active around areas where females had been known to spend some time. They drummed with their palps on the substrate, circled and returned to the same spot. Specimens of Metacyrba undata were observed to exhibit similar behavior, but this species was not tested. The other species tested produced ambiguous and inconclusive results. The pheromone of Menemerus bivittaiMS is probably produced on the silk dragline of the females and so the silk glands would seem to be a good possibility for extract bioassay. This is the first report of a contact sex pheromone in salticids. APPENDIX C THE GENUS EENTZIA IN FLORIDA The genus Bentzia was last revised by Peckhani and Peckham (1909)- They listed three species for North America, H. gvenada, U. mitrata and //. palmax'wri. All three species are found in Florida. During the course of the present study specimens of these species were collected. Of these, H. mitrata was the least abundant, being collected occasionally on shrubs and herbs. This species is known from Alachua, Dade, Levy, Palm Beach (Banks, ]30ka) and Pasco Counties. Hentzia grenada is widespread, but seldom abundant except in the south. It seems to live almost exclusively palms, espec i 1 the saw palmetto . on a 1 y {Seronoa repens) Records include Alachua, Dade (Banks, ]30ha), Duval, Highlands, Marion, Monroe and Putnam Counties, and as far north as Charlton County, Georgia. It was the dominant species at the Archbold Biological Research Station, Highlands County, and at Pinecrest, Monroe County. Hentzia palmarim is by far the most abundant species in the north and ranges across most of the state. It is known from Alachua, Baker, Charlotte (Banks, 1 904a), Citrus, Dade, Duval, Highlands, Lake, Levy, Marion, Monroe, Orange, Palm Beach (Banks, 1 90^a) and Putnam Counties. This species was very abundant on black mangrove {Aviaennia nitida) at Way Key, Levy County during 1975- As many as ^0 specimens could be collected in one half hour by beating the trees, llentzia palmarum was also abundant on under- story shrubs in the sand pine scrub of the Ocala National Forest, Marion County and on willow [Salix carotiniana) at Lake Kanapaha, Alachua 142 County. Adults and immatures have been found in just about all months of the year around VJay Key. Egg sacs were produced in captivity from March to October. This species seems to be one of the most versatile of the salticids in regard to life history and habitat. The close relationship between H. gvenada and //. palmanm has been demonstrated by at least six matings between males of //. gvenada and females of //. palmarum and one mating between a male of H. palmavum and a female of H. gvenada. In three of the former matings, the female laid eggs, and two batches hatched. One of these broods was preserved and another was raised. Of 18 eggs laid by one female August 30-31, 13 shed the chorion on September 10. These were removed from their dish on September 28 and fed 1st instar cabbage loopers. All but one died between October 5 and October 28. One lingered on until April 1-2, but never matured. R. Skinner (personal communication), using techniques he has successfully utilized to raise //. palmavum from the egg, was unable to raise another hybrid brood. The data, as incomplete as it is, would seem to indicate that //. gvenada and //. palmarum were at one time isolated from each other. Hentzia gvenada was found in northern South America and probably the West Indies and palmavum on the mainland of North America. When the Florida peninsula rose during the Pleistocene, the two species had routes to Florida open to them. Eentzia palmavum invaded f rom the north, and finally reached the tip of the peninsula. Hentzia gvenada, probably arrived by balooning or by accidental transport (possibly even in historic times), moved up the peninsula from the south, reaching into Georgia. The species had evolved somewhat differently in relation . 143 to preferred habitat and H. palmarum seems more versatile in this, being found on many shrubs and trees, both along the coast and inland However, although easily recognized by their color patterns and pro- portions of their bodies, their courtship and agonistic displays are vertually identical. As yet, no hybrids have been found in the wild. A Key to the Species of Ilentzia found in Florida Ma 1 es la. Cheltcerae short, vertical; first legs white, with a fringe of white hairs /7_ rnitrata lb. Chelicerae elongate, horizontal; first legs dark, without fringe of wh i te ha i rs 2 2a. Dorsal white stripe on prosoma widening toward and reaching front eyes; first legs banded white and brown; body elongate //. ,jrcnada 2b. Dorsal white stripe on prosoma narrowing toward, and never reaching, front eyes; first legs usually solid brown; body less elongate than //. grenada //. palmarum Fema 1 es la. Abdomen with distinct pattern of dark brown spots • //. rnitrata lb. Abdominal pattern obscure 2. 2a. Abdominal markings consisting of obscure dark stripe, often broken into elongate chevrons; body elongate. ... H. qrenada 2b. Abdominal markings mostly spots and chevrons (never a stripe), usually quite obscure; body less elongate than in //. grenada ' palma.rum^ APPENDIX D A LIST OF THE SPECIES OF PELLENES FROM FLORIDA Like Bentzia, Pellenes [including Habronattus (Lowrie and Gertsch, has 1955)] not been examined closely since 1909 (Peckham and Peckham). Following is an annotated list of the species known to occur in Florida. Records are from my own collection and that of G. B. Edwards, and from the Division of Plant Industry. Pellenes [Pellenes) longimanus Emerton. Males from Lee and Alachua Counties in the Wallace-DPI collection Pellenes {Habronattus) agilis (Banks). Males and females of this colorful species were collected at Seahorse Key and on the mainland near the Gulf Coast in Levy County, during July and August on leaf litter along the beach and on the ground. Pellenes (H.) brunneus Peckham and Peckham. This is perhaps the most widespread of the species of Pellenes in Florida. Records indclude Alachua, Baker, Date, Charlotte (Banks, 190^a), Collier, Levy and Monroe Counties. This species is often taken by sweeping grass, especially low grass. It is very abundant along roadsides near Way Key in late summer and fall. Pellenes [H.) calcaratus Banks. Records of this, species include Charlotte (Banks, 1904b), Marion, Palm Beach (Banks, igo^b), and Polk Counties. A number of specimens were taken in spring and summer from a clearcut area in the sand pine scrub in the Ocala National Forest, Marion County. These spiders live primarily on low grass and on the ground. 1^5 Pc-il'Lenes (//.) coecatus (Hentz). Records include a male from Tall Timbers, Leon County and an immature from Alachua County. Several males and females were collected during March in Glynn County, Georgia by D. and W. Maddison. The habits of this species are very similar to those of P. brunneus. Pelleries {H.) decorus (Blackwall). Several males and one female were collected by sweeping along S-325, Alachua County^ in July and another male was collected near a cypress pond in Alachua County in April. All the specimens were associated with low wet places. Petlenes (ff. ) elegans Peckham and Peckham. Males and females have been taken in Levy, Alachua and Marion Counties. These have been on leaf litter in turkey oak forests and in clearcut areas in sand pine scrub. Pellenes {H.) tachypodus Chamberlin and ivie. Males and females have been collected in Alachua and Marion Counties on leaf litter, especially that of turkey oak, usually in the spring. Pellenes {H.) trimaculatus Bryant. Described from Sebastian, Indian Pxiver County, Florida, this species has been found in Alachua County primarily in turkey oak leaf litter. Usually taken in the spring. Pellenes cf. viridipes (Hentz). This species is almost identical to P. viridipes, except that males lack any modifications on their third legs. Records include Alachua, Hamilton, Levy and Putnam Counties, usually in leaf litter. Banks (1904a) had P. viridipes from Dade and Palm Beach Counties. . 146 Pe'Llenes n. sp. Males and temales of this striking species were collected in scrub oak leaf litter and on the ground around Central Tower in the Ocala National Forest, Marion County during the spring. The males have a red clypeus and yellow fringes on their front legs. They are not related to P. coeoatus . . APPENDIX E CHARACTERISTIC SALTICIDS IN FLORIDA HABITATS During 1973-1976 salticids were collected in a number of habitats and niicrohabi tats around the state of Florida. As little seems to have been published about the relationship between salticids and natural habitats in Florida, I have listed the habitats (including a few artificial ones) and the characteristic salticids associated with them. Many of the natural habitats listed are based on those described by Laessle {]3h2) 1. Sand-pine scrub [Pinus olausa- Querous spp. association). a) Understory [Quercus-Lyonia-Ceratiola-Ilex] . Species associated with this part of the understory included LyssonianGs viriclis, Hentzia pa'Lmaviim, Phidippus putnconi, Tiitelina elegans and Thiodina sylvana. b) Understory {Seronoa-Sabal) . Two species which were apparently associated with this habitat were Hentzia grenada and Paramaevia michelsoni c) Leaf litter. A few species have been collected on leaf litter within sand-pine scrub. These include Habvocestum n. sp. A, Pellenes tachypodus P. n. , sp. A and Phlegi^x fasciata. Pellenes n. sp. A was most commonly collected in small cleari-ngs. d) Clear-cut areas. Clear-cuts are found scattered throughout the sand-pine scrub. The characteristic salticids of these areas 1^7 148 included Pellenes oalearatus, P. elcgans,, Phidippus oavdinalis, P. apacheanus and Zygoballus sp. 2. Sandhills or old high pine {Pinus palustris-Queraus 'Laevis association). Much of the longleaf pine forests have been cut. These are now represented mostly by pure stands of Q-uereus laevis or a mix of Q. laevis, Ceratiola eviooides and occasionally Quercus virginiana and Q. ciyierea. a) On turkey oak {Quercus laevis) bark. One specimen of Maevia intermedia, a dark phase male, was collected on the bark of a young turkey oak. b) Understory (var ious, herbs and shrubs- usually very sparse). A few specimens of Metaphidippus galathea, Phidippus putnami and Zygoballus sp. were collected on understory plants, mostly by sweeping. c) Leaf litter. Species which were characteristic of turkey oak leaf litter included Corythalia aurata, Habrooestwv n. sp. A, Parainaevia miclielsoni, Pellenes tachypodus, P. trimaculatus , P. cf. vivid-ipes, Phlegra fasciata and Sitticus cursor (rare). 3- Xeric hammock and in live oak {Quevous virginiayia) on the edges of meslc hammocks. a) On live oaks. Two species, Metacyrba undaba and Phidippus otiosus were commonly collected on live oaks. Phidippus regius was occasionally found (especially during winter) under loose bark and Ballus youngii was collected once under the outermost layer of bark. . . . 1^9 b) Understory (various shrubs). Hentzia niitrata vjas collected occasionally on shrubs under live oaks. c) Leaf litter. Three species, Corythalia aurata, HabrocesfAAm n. sp. A and Fellenes cf. viridipes were collected in live oak 1 eaf 1 i tter Disturbed xeric communities. Areas of disturbed woodland characterized by turkey oak {Quercus laevis) , laurel oak {Q. laurifolia) and hickory {Carya sp.). a) Under bark, on trees and logs. Species collected especially during the winter include Phidippus cca'dinalis, P. otiosus and p. regius. Metacyrba taeniola and M. undata have also been found in this m i crohab i tat b) Leaf litter and. in grass hammocks. Several species were found in this mixed microhabitat including Habrooestum n. sp. A, Fellenes elegans, P. trimaculatus and P. cf. oiridipes. Mesic hammock [Magnolia grandiflora-Ilex opaaa association). True mesic hammocks are rare and most have been disturbed. The best examples can be seen in San Felasco Hammock (Alachua County) and Highlands Hammock (Highlands County). Most of the wooded areas with a distinct canopy are characterized by only a few salticid spec i es a) On trees and understory within the hammock. Two species were found on trees and shrubs, Lyssomaiies viridis and Thiodina sylvana, the former most often on Magnolia. b) Leaf litter. Species which were found in mesic leaf litter include Corythalia aurata, Habrocestim bufoides and Sarinda hentzi. . 150 6. Hydric hammock- cypress ponds nnd swamps {Toxodium distichum- Pinus eVliotti association). a) On tree trunks and under bark. One species, Metacyrba imdata has been collected on both cypress and pine trunks. b) Pine litter. This litter microhabitat contained one of the most diverse of the salticid litter faunas. Species found here included Corythalia aurata, C. delicabula, Marpissa sulcosa, Metaphidippus castaneus, Neon nelli, Neonella vinnula, Pellenes decorus (accidental), Sarinda hentzi, Sitticus cursor and Zygoba I lus bettini 7. Slash pine flatwoods {'Pinus elliottii association). Much of this consists of disturbed flatwoods, with some longleaf pine {Pinus palustris) and several hardwoods, such as QMercus nigra and Q. laurifolia. a) Under bark and on dead branches of trees. Several species were collected under bark, or by beating dead branches of trees. These included Admestina tibialis (immature), Ballus sp. (probably undescribed) and Metaayrba imdata. b) Understory shrubs and herbs. Several species were collected by sweeping through understory plants. These included Metaphidippus galathea, Phidippus pulcherrimus and Zygohallus sp. c) Saw palmetto understory [Serenoa rep ens) . Species taken by beating saw pal met toes included Maevia inalemens^ Peokhamia americana, Synemosyna formica and Thiodina sylvana. d) Pine and leaf litter. Species collected in flatwoods litter included Corythalia aurata, C. delicatula, Habroccstum bufaides. . 151 Maevia ( inolemens immatures) , Marpissa siilcosa. Neon nelli and Sarinda hentzi. Roadsides and grassy areas. Along many Flroida side roads various species of salticids were swept from grass and associated herbs. These included Marpissa pikei, Metaphidippus galabhea, Paramaevia hohhsi, Pellenes brunneus , P. decorus, Phidippus audax, P. clavus, Thiodina puevpera, 7yaohallus bettini and Z. sexpunctatuG Grassy areas near ponds. Specimens of Eris marginata and, occasional Marpissa pikei were collected by sweeping tall grasses. Oak litter on sea beaches. Two species have been collected on live oak {Quercus virginiana) litter just above high tide on Seahorse Key, Levy County. These were Ilabrocestimi n. sp. A and Pellenes agilis. The latter species was also collected on the ground near the coast on the mainland in Levy County. Mangroves {Avicennia-Phisophora-Lagunaularia associations) . Mangrove communities seemed to be dominated by Hentzia palmarum although Marpissa hina was occasionally found on black mangroves (Aviaennia nitida) and one specimen of Phidippus regius was collected, also on a black mangrove. Saw palmetto [Ssrenoa repens association). Open stands of saw palmetto were found to have several characteristic salticids includ- ing both Maevia inclemens and Paramaevia michelsoni (never found together), as well as Peckhamia americana., Phidippus regius and Synemosyna. formica. Spanish moss [Tillandsia usneoides) . Only one species, Metaphidippus tillandsiae, was collected on Spanish moss. . 152 14. Floating vjater plants {EichornLa crassipes association). Mavpissa bina was occasionally collected oh these plants. A few specimens of Zygoballus nervosns were also collected on floating plants, mostly Eichornia. 15. Old fields and prairies. 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Spiders of southern Africa. Books of Africa, Cape Town. 200 p. BIOGRAPHICAL SKETCH David B. Richman was born November 6, 19^2, in Dunkirk, New York. His parents took him to Arizona in 1948 and he grew up in and around the city of Yuma on the Colorado River. He attended Arizona Western College and received his A. A. in I968. While at Arizona Western College he was elected to Phi Theta Kappa and Who's Who in American Junior Colleges. In September, I968, he entered the University of Arizona in Tucson, from which he received the B.S. in 1970, and the M.S. in 1973, both with majors in zoology. While at the University of Arizona he was elected to Beta Beta Beta, Phi Kappa Phi and Sigma Xi. During the years in Arizona he made numerous trips into the deserts and mountains of the southwestern United States and northwestern Mexico. In October, 1972, he traveled through Baja, California, from Santa Rosalia to La Paz via Sonora and across the Gulf of California by ferry. He was admitted to the Graduate School of the University of Florida in September, 1973, with a major in zoology and a minor in entomology. He has published articles on jumping spiders, brown spiders, tiger beetles and butterflies. 162 . 1 certify that 1 have read this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is fuT adequate, in scope and quality, as a dissertation for the degree of Doctor of Philosophy. 4~ ^^---__JLo^athan Reisl I certify that I have read this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is fully adequate, in scope and quality, as a disseration for the degree of Doctor of Philosophy. ohn F. Anderson 'Associate Professor of Zoology 1 certify that 1 have read this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is fully adequate, in scope and quality, as a dissertation for the degree of Doctor of Philosophy. CI ilfford "jojinson Professor of Z*oology I certii'y that I have read this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is fully adequate, in scope and quality, as a dissertation for the degree of Doctor of Phil osophy ;3. hi. OJUeit D. H. Habeck Professor of Entomology 1 certiPy that I have read this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is fully adequate, in scope and quality, as a dissertation for the degree of Doctor of Philosophy. 'Professor of Entomology This dissertation was submitted to the Graduate Faculty of the Department of Zoology in the College of Arts and Sciences and to the Graduate Council, and was accepted as partial fulfillment of the requirements for the degree of Doctor of Philosophy. March, 1977 Dean, Graduate School