THE COMPARATIVE BEHAVIOR OF

THREE SPECIES OF BLENNIOID FISHES

A Thesis

Presented to

the Graduate Faculty

California State College, Hayward

In Partial Fulfillment of the Requirements for the Degree

~~ster of Arts in Biology

by

David Gregory Lindquist

July 1971 ACKNOWLEDGEMENTS

I would first like to thank Dr. Boyd w. Walker, my former advisor at UCLA, for suggesting this group of fishes to study. I am indebted to Dr. John s.

Stephens, Jr., of Occidental College, for his sugges­ tion of the comparative problem and for his encourage­ ment and enthusiasm during the study. Dr. George s.

Losey, Jr., of the University of Hawaii, generously relinquished some of his unpublished methods for my use. Finally I wish to thank the members of my com­ mittee: Drs. James w. Nybakken, Jack T. Tomlinson and

Edgar Yarberry for critically reading the manuscript. TABLE OF CONTENTS

LIST OF 1rABI...:fSS • • • • • • • • • iii

LIST OF FIGtJRES • • • • iv

ABSfJ.'RACT • • • • • • • • • v

Il!l'l'RODUCTION • • • • • • • • • • • • l

I. GEl:lERAL PROBLEI;i • • • • • • • • • • l

~ II. REVIEI'J OF LITERATTJRE • • • • • " P-~ .. Systerratics • • • • 2

B. Behavior • • • • 7

.0-JETHGDS .t::-1:-JV MA'L'ER:W.LS • • • • • • • • • 9

a r. FISHES STUDIED • • • • • • ~

A~ Gulf California Fishes • • • • • • • 9

1. Ernblemaria hvpacanthus • • • • 9

2. Acanthernblernaria crockeri • • • 9

B. Honterey Bay Fishes • • • • 10

1. Neoclinus uninotatus • • • 10

c. Se]~Ual Dj,morphism • • • • 11

II. OBSER\.11\TION l·'ETHOD Al:ID EQUIPl·lliNT 13

p_ .. Laboratory Observations • • 13

B. Field Observations • • 16

i TABLE OF CONTEblTS (Continued)

III. COLLECTION AND TRANSPORT~TION OF SPECIMENS 17

GENERAL ETHOGRAM •• . . . 0 • • 0 0 21

COMFORT f-10VEMENTS • • • • • • • • ~ • 0 • 33 ELIMIN!\.TORY BEHAVIOR •• ...... 35

ENVIRONMENTAL ALTERATION'S . . .. . • • • • • 0 0 • 36 TERRITORIAL BEHAVIOR ••• . . 38

I. SPECIES DESCRIPTIONS • • • ...... 39

AGONISTIC BEHAVIOR . . . . . • Ill • • 45

I. SPECIES DESCRIPTIONS . . . Q • .. • 46 II. TEMPORAL ASSOCIATION OF AGONISTIC MOVEMENTS 56

III. MOTIVATIONAL ANALYSIS • • 0 • • 0 ...... 65

REPRODUCTIVE BEHAVIOR • • • • 77

COLORATION 0 • .. • • . . . 89

I. COLOR PATTERN DESCRIPTIONS 0 • • • 0 0 0 • • 89

II. COLOR PATTERN CHANGES 0 0 0 0 . . 99

DISCUSSION AND CONCLUSIONS • • 8 8 D 8 8 8 8 8 8 0 8 109

LITERATURE CITED • • • • • • • • • • .. • o • o • • • 114

ii LIST OF TABLES

Table !?age

Relationships between the species studied 14

2. The differences between the species' repertories in the general ethogram 28

3. Occurrence of agonistic motor patterns and their mean durations 55

4. Frequency with Which the nine different movements precede or follow each other in the agonistic behavior of Eroblemaria hypacenthus 59

5. Spearman's rank correlation coefficient for the agonistic movements of Emblemaria hypacanthus 60

6. Frequency with which the nine different movements precede or follow each other in the agonistic behavior of Acantheroble­ m

7. Spearman'o rank correlation ~oefficient for the agonistic movements of Acanthem­ bleP.taria crocker:i. 62

8. SWlil!lary of stn.t~.stically correlated move­ ment pairs for the chaenopsids 64

9. Fin Flicker subunit duration 80

iii LIST OF FIGURES

Figure Page

1. Circumorbital bones used to separate the three species studied 5

2. Study area for Neoclinus uninotatus 18

3. Agonistic behavior of Emblemaria hypacanthus 31

4. Flee~ttack diagrams for Ernblemaria hypacanthus, Acanthemblemaria crockeri and Neoclinus uninotatus 71

5. 1~ Emblemaria hypacanthus displaying the courtship inviting action~ Fin Flicker 81

6. Stimulus-response nature of the male invitation action 83

7. Early larva of Neoclinue uninotatus 87

B. Color patterns for Emblemaria hypacanth~ 90

9. Color patterns for ~canthemblemaria crocker! 93

10. Color patterns for Neoclinus uninotatus 96

11. Color pattern frequency in Emblemaria hypacanthus and Acanthemblemaria crockeri 104

• Color pattern frequency in Neoclinus uninotatus 107

iv ABSTRACT

Laboratory and field comparisons of the behavior, morphology, distribution, habitat, and ecology of the blennioid fishes Effiblemaria hypacanthus, Acanthemblemaria crockeri and Neoclinus uninotatus indicate the former two species of the family Chaenopsidae to be more specialized than N. uninotatus of the family Clinidae.

The comfort, eliminatory, and environmental alteration behavior of the three species is similar.

The agonistic behavior of each species is distinct.

The chaenopsids have elaborate threat codes and combat behavior. The agonistic behavior of the three species is apparently a function of the flee and attack motivation interaction. The action pattern sequence of pure attack or pure flee motivation differs from the sequence of mixed flee and attack motivations. The species differ in rela­ tive threshold levels of the action patterns, degree of ritualization, and in the relative importance of sign stimuli.

v Analysis of the preceding and following motor pat­

terns of the combat behavior of the two chaenopsids by the

Spearman rank correlation coefficient reveals correlated action pairs that fail to fulfill the fixed action pattern.

One explanation for this suggests that selection for a decrease in the monotony of sequences of agonistic action patterns is advan·tageous for occasions when novelty is necessary for arousal and attention; e.g. fishes that en­ gage in complex fights require that each animal continu­ ally and accurately assess the state of its opponent.

The general reproductive patterns for the three species does not differ. However, differences were ob­ served in the manner of the male invi·ting actions.

Intraspecific color pattern variations are apparently related to the behavior of the individual. Three basic color patterns for each species are demonstrated. The function of the color patterns appears to fall into three categori~s~ accentuation of postural displays, signaling the motivational state of the individual, and effecting reproductive isolation.

vi I~""TRODUCTION

I. GENERAL PROBLEM

The purpose of this study is to contribute to the

understanding of the behavior of three related blennioid

fishes. The comparative approach to animal behavior can assist in the assessment of relationships among related

species. Systematics too often evaluates the phylogenetic relationships among species on the basis of morphological characteristics •.

Comparison of the behavior within a closely related group of animals often reveals subtle differences in action patterns which may be analyzed in terms of their origins and adaptive value. Knowledge of these origins is in­ valuable for assessing the degree of inter- and intra­ specific convergence and divergence of action patterns and the value of these actions as taxonomic characters. These comparisons have proven valuable in assessing phylogenetic relationships (Hinde, 1970).

Furthermore, it is the author's intent to make the first step in contributing behavioral information that can 2.

later be used to elucidate the behavioral evolution of the fish families Clinidae and Chaenopsidae. Thus the system­ atics of the two groups might well be clarified with the additional use of behavioral information.

II. REVlEW OF LITERATURE

A. Systematics

The two fish families involved in the study are the

Clinidae and the Chaenopaidae. Neoclinus uninotatus, Hubbs

1953 (Clinidae) , Emblemaria hypacanthus (Jenkins and Ever­ mann 1888) (Chaenopaidae) and Acanthemblemaria crocker!

Beebe a:l".d Tee-Van 1938 (Chaenopsidae) are the species under consideration. Clark Hubbs (1952 and 1953) in his revision of the family Clinidae placed the subfamily

Chaenopsinae (which contained all three of the species above) under the family Blenniidae rather than under the

Clinidae. He based this placement on the extension of the lachrymal bone to a point below the middle of the eye and the absence of scales except in Neoclinua. Springer (1955) disagreed with HUbbs' 1953 classification. Springer be­ lieved the lachrymal extension to be a clinid character 3.

rather than a blenniid character. Thus, according ·to

Springer, the Chaenopsinae should go with the Clinidae.

Bohlke (1957) noted that the position of the Chaenopsinae may be about the same in either Hubbs' or Springer's classification since it is probable that the Blenniidae are derived from a clinid-like ancestor. But if the two families were to be maintained Bohlke preferred to follow

Springer. In addition, Bohlke supposed the position of

Neoclinus to be primitive to the other chaenopsin genera.

Stephens (1963) erected the family Chaenopsidae, com­ posed of 10 genera and 39 species. This assemblage is based on two bones in the circumorbital ring, a lack of scales, and the absence of a lateral line. Since Neoclinus has four circumorbitals (Fig. 1), is scaled, and has a well developed lateral line, Stephens prefers to include ~­ clinus within the Clinidae. In this scheme the chaenopsids are derived from an ancestry similar to Neoclinus. Springer

(1964) questioned Stephens' classification because of the scarcity of investigations as to the nature and number of the circumorbitals in clinids and blenniids. The major point Springer made is "whether the nuniber of circumorbi­ tals is a phyletic character, as Stephens believes, or a pointe'-! out

Stephans (3.970),

lateral line c~racteristics. Almost no variability

degree o£

lateral line reduction in chaenopsids was described in

pore counts. side ia a

present within too adapt.iv,;;; and 5.

Figure l

Circumorbital bones used to separate the three blenny species studies. A. NeoclimJs uninotatue, four circwnorbitals. B. Acanthemblema:ria c:rockeri, two circumorbitale and c. Emblemaria hypacanthus also two circumorbitals. (After

Stephens, 1963) 3rd

2nd

/

FUSION OF CiRCUMORarr.u.LS 3&4

LACHRYMAL 8 c 7.

presented in this paper Il'.ay be of use in assessing these

rela'tionships. Definitive '1-mrks on the behavior of the

many other species of the bro families will be needed to

form a firm basis for evolutionary considerations.

B. Behavior

The general behavior and natural history of the

chaenopsid fishes and those of the clinid genus Neoclinus

is almost unkn~wn. One of the first published works on

chaenopsid behavior was an anecdotal description of aquarium

observations on £Paenopsis ocellata (Robins, Vnillips and

Phillips, 1959). Wickler (1963) fiL-ned the ritual fighting

of Emblemaria pandio:nis and provided a very short written narrative. The specialization of organs having signal

function in :§.. :pandionis is treated elsewheJ;:e by Wicl~ler

(1967). Stephens, Hobson and Johnson (1966} presented

field observations for the reproductiv-e behavior of

Acanthemblemaria macrospi.lus. The literature contains

little on the habits of the fishes of t:lw genus Neoclinus.

Stephens, Q.t ll.J,, (1970) reported the habitat preference of Neoclinus stephenaae in their comparative ecological study of three California blennies. A manuscript concerning 8.

the comparative ecology of the three California clinids of the genus Neoclinus is now in preparation by Stephens, Os­ born, Hickman and this author.

Fishes have been the object of extensive behavioral research. The majority of this work has been carried out under laboratory conditions since fishes are difficult to observe in the field. Most of these studies are anecdotal.

Few quanti·tative analyses of fish behavior have been published. Nelson's (1964) work on glandulocaudine fishes represents quantitative excellence in ethological analyses.

Problem solving involved the use of probabilistic and cybernetic methods relatively new to ethology, I'Jiepkema

(1961) was the first, however, to quantitatively analyze the temporal patterning of a fish's behavior. The entire behavior pattern of a species was analyzed in such a way that the variables obaervad were ordered according to sets of common causal factors. Losey (1968) presented an ex­ tensive comparative study of the behavior of a closely related group of marine bottom fishes. Be studied the

Pacific members of the genus Hypsoblennius in the field as well as in the laJJoratory. 9.

t1ETHCIDS AND MATERIALS

I. FISHES STUDIED

A. Gulf of California Fishes

1. Ewblemaria hyPacanthus

svstematics and Distribution - The genus EmblenBria

(Jordan and Gilbert 1883) may be separated frorr, ·l:he other

chaenopsid genera by the sexually dimorphic spinous dorsal which is high and sail-like in mature males (Fig. 5) • !]L. hypacanthus is distinguished from the other four Pacific

members of the genus by its relative lack of specialized

characters {Stephens, 1963), Emblemaria hypacanthus is

the nuu~rically dominant rrember of the genus in the Gulf

of California. This species occurs throughout the Gulf

of California from La Paz to Puerto Penasco and is endemic

to the region. !>'ly specimans were collected at Puerto

Penasco, Sonora and Puerto Refugio, Isla Angel de la

Guardia, ~Exico.

2. Acanthernblemar ia crocker i

Systen~tics and Distribution - The genus Acanthemble- 10.

maria is separated from the rest of the chaenopsid genera by the presence of spines on the parietal bones and teeth in a doUble rm~ on the palatines. Acanthemblemaria crock­ sri is distinguished from the other members of the genus by the elongated sharp cranial spines, a single dark br~~n spot covering the cheek in front of the operculum end a more striking sexually dimorphic color pattern (Fig. 8).

A. crockeri is also endemic to the Gulf of California.

1'1Y specimens '\'Jere taken at Bahia de Los Angeles end Bahia de San Francisquito, Baja California~ Puerto Refugio,

Isla Angel de La Guardia and Guaymus, Sonora, r~xico.

B. Monterey Bay Fishes

1. Neoclinus uniotetus

systew~tics and Distribution - N. uninotatus can be distinguished from tl1e chaenopsids by its larger adult size (90-210 rr~i and the presence of scales and a distinct lateral line. li· uninotatus may be distinguished from the other members of the genus by the jaw seldom extending as far posterior as the origin of the dorsal, several ocular cirri with one exceptionally large anterior one over the eye, and a single ocellus on the anterior portion of the 12.

Acanthemblemaria crockeri shows distinct sexual

dimorpnism in its color pattern and to a lesser extent in

spinous development of the skull. The color patterns are

shown in Figure 9 (page 94).

Neoclinus uninotatus shows distinct sexual dimorphism

in orbital cirri development and proportional body measure­

ments. My data indicate male cirri length to average 12.1

mm and branching of the cirri to average 22.3 branches for

8 males ranging 109 to 140 mm standard length. Four fe­ males ranging 120 to 143 mm had an average cirri length of

6.3 mm and average branching of 9. These data are cor­

roborated by Stephens and Hickman (pers. comm.) , who re­

port: "Males have a mean cirri length of 1. 7 orbits, never having cirri shorter than one orbit. Females have a mean

of 0.6 orbits, their cirri never exceeding one orbit. The male cirrus is stouter and branches soms,~hat differently,

the male branches being palmate distally, the female's branches occurring someWhat more proximal."

Clark Hubbs (1953) reports five proportional body measurements expressed in per cent of standard length which separate the sexes. The males have statistically

significant greater measurements of the upper jaw length, 13.

body depth, first anal soft-ray length and fifth dorsal

soft-ray length. The distance from the anal origin to

the pelvic insertion is statistically greater in females

than in males.

I have found the priF~ry ocular cirrus length to be

the best character is separating the sexes in the field.

Hubbs' proportional body maasuremants are not practical to

use in the field and often give confused results with

preserved specimens.

II. OBSERVATION METHODS AND EQUIPliJENT

A. Laboratory Observations

The investigation of each species began with groups of

fishes in laboratory aquaria. Tank I, 0.5 x 0.3 x 0,3 m.,

used for observing the chaenopsid species, was constructed

of glass on all four sides. The bottom of the tank was

covered with a 3.5 crm. layer of mixed aquarium gravel and broken shell. Two subsand filters were placed under the

gravel mixture. The refugee for the fish were scallop

shells containing many empty bivalve burrows, col~~r beads of the coral, Porites californica, serpulid worm 14.

TABLE 1

Relationships between the species studied

Emblemaria Acanthemblew~ria Neoclinus hypacanthus crockeri uninotatus

Family Chaenopsidae Chaenopsidae Clinidae (after Stephens)

Number of 2 2 4 circumorbitals

Lateral line and scales absent absent present

Size range adults 25-55 ll1lll 25-5 0 lll!ll 90-210 rnm (both sexes)

Faunal range Gulf of Gulf of La Jolla to California california Monterey Bay, California

Habitat Intertidal Porites Subtidal sandstone californica sand-trash reef pools coral basins and subtidal algal mats

Sexual High dimor­ low dimorph­ moderate dimorphism phiSID7 males ism: color dimorphism, have high patterns males have sail-like distinct larger dorsal fin cirri 15.

tubes and sections of opaque plastic tubing. These ref- uges were scattered over the bottom of t~~ aquarium. The

0 temperature of tank I was kept at 15 Celsius.

Tank II, 1.0 x 0.4 x 0.3 m., used for observing

Neoclinus uninotatus, was constructed of fiberglass, with one glass face. The bottom was covered with gravel and filters as above. Refuges were empty barnacle tests, a pint bottle and five blade polyvinyl coring tubes of 3.5 and 5 em. diameters. The temperature of tank II was kept at 10.5 ~ 0.5° Celsius.

Still photographs were obtained with a 35-mm Nikon camera and an electronic flash. Motion pictures were taken with a super 8-mm Nikon, at film speeds varying from

18 to 24 frames per second, using a flood lamp. Notes and tape recorded Observations were used for essentially all data collection.

Observation periods ranged from 1 to 4 hours. At least 50 hours of observations were obtained for each species.

All fish were fed an abundance of cut frozen brine shrimp, live brine shrimp and pieces of clam meat. Food was added to the tanks with a minimum of collateral cues. 16.

B. Field Observations

Behavioral observations were extremely difficult to make under natural conditions using SCUBA in the field.

The fish do not engage in their normal undisturbed activity

14hen the diver is visible but spend most of their time observing the diver. This is especially true when the observer must get close to his subject When visibility is poor. Also the diver may be in constant motion due to surge. However, valuable information concerning duration of territory maintenance, territory size and population densities can be gained through the use of underwater transects and grid systems. Six 30 meter transects ~~re made by myself and three other divers of the Moss Landing

~~rine Laboratories Makrele cruises to the Gulf of Calif­ ornia during the spring of 1969. We attempted to count all macroscopic organisms located within one meter of the transect line. The following transect locations included the chaenopsid Acanthemblemaria crocker!, Bahia de San

Francisquito, Puerto Refugio and Bahia de Los Angeles.

A grid system was employed at the commercial wharf number 2, Monterey, California, (Fig. 2) to observe a population of Neoclinus uninotatus. Three 5-meter square l7o

quadrats were constructed of eight 20 em concrete blocks with eyebolts set into cement added to the centers of the blocks. Rope was attached to the eyebolts and run between the blocks to complete the squares" The grid was located in 10 to 12 meters of water approximately 30 meters east of the Wharf. A varying number of glass containers {e.g. bottles, jars, jugs) were placed within each grid square and thirty-five N. uninotatus were subsequently released into the grid area.

Fifteen fish were tagged by means of modified

Peterson discs made of glass beads beat beaded on nylon monofilament line (Stephens !!:!:, g, 1970) • However, these tags were lost within a few weeks after release of the fishes into the field because of abrasion between the refuge and tag. Individuals could often be recognized on the basis of location, coloration, size and sex. The glass containers were also tagged when occupied by a fish.

III. COLLECTION AND TRANSPORTATION Of! SPECIMENS

Specimens were collected using SCUBA and quinaldine, an anesthetic. The quinaldine, dissolved in isopropanol

(lilO dilution), was delivered, a few cubic centimeters at 18.

Figure 2

Study area for Neoclinua uninotatus at the commercial Wharf number 2, Monterey, California.

The upper expanded view shows the grid quadrats in relation to tr.e north end. of the Wharf. See text for further details. Monterey 8'""'''a; 20.

a time, to the water surrounding the fish by means of a one pint polyethylene squirt bottle. Gibson (1967) and

Stephens et al, (1970), have described the use of quin­ aldine in behavioral and ecological studies. The mode of action of the drug is considered to be analogous to that of barbiturates (Muench, 1958).

The Gulf of California species were packed in plastic bags with sea water and an oxygen atmosphere, then placed in insulated containers. These containers were trans­ ported via auto, to the laboratories for further studies.

The fish remained in these containers as long as three days, and sustained little mortality. 21.

GENERAL ETHOGRAM

The following description serve to acquaint the reader with the generalized patterns of movements and displays seen in the three species studied. The organi­ zation underlying the code of comple'c signals and inter­ specific differences will be dealt with separately.

First it is necessary to define several broad types of behavior in terms of their function. The categories used by Losey (1968} will be follu~ed here. These broad types of behavior will be related to the specific movements where possible.

Comfort Movements: behavior which serves to make the

fish comfortable and to groom the body.

Ingestive Behavior: movements involved in the ingestion

of food material.

Eliminatory Behavior: movements related to the elimina­

tion of metabolic wastes.

Maintenance Behavior: movements directed to the altera­

tions of the refuge such as digging, and removal of

objects from the refuge with the mouth. 22.

Agonistic Behavio:q any non-selcual inter-individual

behavior.

1. Submissive: subordinate behavior involving a

low possibility of attacking.

2. Aggressive: dominant behavior involving a

high possibility of attacking.

Sexual Behaviorr any inter-individual behavior that in­

creases the possibility that eggs will be laid and

fert.ilized.

1. Courtship: behavior between male and female

not involved in actual egg laying and fertili­

zation.

2. Mating: behavior involving actual deposition

and fertilization of the eggs.

Parental Behavior; movements associated with the care and

maintenance of the eggs by the male in his refuge.

'l"he movements observed for the three species are as follows:

Locomotion:

1. ~ - Lateral sinusoidal flexures of the body

which may be accompanied by pectoral fin move­

ments. This type of swimming would be classified 23,

as moderately anguilliform (Brader, 1926}.

2. Hop - Short ju.Ttlps over the substrate, propelled

by pectoral and pelvic fins which may be accom­

panied by caudal fin flexures.

3. Forewalk and Backwal~ - This movement is accom­

plished by walking movements of the pelvic and

pectoral fins over the substratum. Backwalking

may associated with Submissive behavior.

Comfort:

4, ~ - The mouth is opened maximally then closed.

5. Coug~ - The mouth is opened sub~~ximally, the

operculi are pushed outward, and the branchiostegi

dot'lnward, while the opercu1ar membrane closes the

branchial chamber. The mouth remains open and the

operculi and branchiostegi are quickly adducted,

forcing water and detrital material in the bran­

chial cavity out of the mouth.

6. Chafe - The fish rolls over on its side from the

swimming position and chafes the flank against the

substratum by a quick caudal and pectoral fin beat.

This is mainly a comfort moverrent but may take

place as a displacement activity during courtship. 24.

Ingestive:

7. Feed -The fish first visually fixes the food

item then quickly swims out of the refuge to

take it into the mouth in a snapping motion.

The fish then returns to the refuge and usually

backs in caudal end first. The food item may

also be taken while the fish is in the refuge

or on the substratum. The piece of food may be

tested by taking it into the mouth and then

spitting.

Maintenance:

8. Spit - The fish picks any debris from the sub­

stratum in its mouth and coughs, expelling the

debris.

9. Stationary Swim - Attenuated Swimming motions of

the trunk directed toward the substratum. This

movement serves to dig into the substratum and

for the clearing of debris. (Ha intenance)

Submissive:

10. Flee - The fish Swims quickly away from the

opponent. The dorsal fin may be depressed and

color changes may take place (Fig. 3B; page 27}. 26.

18. Push - The mouth is opened maximally and

pressed against the opponent. The fish then

begins to Swim against the opponent.

19. - The fish &.~ims quickly after the

opponent. The unpaired fins may be erected.

20. Head Butt - The mouth is opened maximally and

directed in a ramming fashion toward the

opponent.

21. Mouth Lock - The fish interlock jaws. This

position may be held for 2 to 3 seconds.

22. Head Arch - The front of the head is pointed

downward while the predorsal fin area is arched

up. This movement occurs within the refuge and

serves to make the fish appear larger.

(Aggressive)

Sexual:

23. Fin Fliclcer - Rapid spreading and folding of

the unpaired fins coupled with out and in move­

mente from the refuge. The frequency is quite

stable at approximately 1 per second. The

amplitude does not vary. (Courtship} 27.

24. Tail Beat - The fish directs an incomplete

Swimming motion of the trunk and caudal fin

towards the side on which another fish lies.

(Courtship)

Parental:

25, Pectoral Fan -The fish beats one of its

pectoral fins or both, alternately. The rate

varies from 0.6 to 1 per second. The amplitude

is highly variable. A few to all the pectoral

rays may be involved. The beginning of a beat

begins with the uppermost rays moving forward

followed in succession by the next lower rays.

The return beat follows the same pattern except

in reverse direction. The resultant feathering

action moves water in a posterior direction.

Ambivalent:

26. Head Quiver - Extremely high frequency and low

amplitude movements of the bead. (Ambivalent

or Displacement Activity) 28.

TABLE 2

The differences between the species' repertories in the general ethogram categories:

0 does not occur

-1- does occur

* does occur, but slightly altered in form from

the general ethogram description

? the situation in which it may occur was not seen

Erriblemaria Acanthemblemaria Neoclinus hypacanthus crockeri uninotatus

Locomotion:

1. Swim +

2. Hop + ?

3. Forewalk, + Backwalk

Comfort:

4. Yawn +

5. Cough +

6. Chafe + ? ?

Ingestive:

7. Feed + + +

Maintenance:

8. Spit + ... 29.

'rABLE 2 (contd.) l'laintenance E. ~- !· (Continued) hypacanthus crockeri .uninotatus

9. Stationary + + + Swim

SUbmissive:

10. Flee t 'I" t

11. Fin Fold 1" 'I' + Agressive:

12. Head Up ..! .J ?

13. Fin Erect + 'I" +

14. Branchiostegi + .. + Lower

15, Gape + + + 16. Mouthfight + * *

17. Bite .j. + + 18. Push * * * 19. Chase + ... +

20. Head Butt 0 + 0

21. Mouth Lock + + +

22. Head Arch 0 0 +

Sexual:

23. Fin Flicker + 0 0

24. Tail Beat 0 0 '1- 30.

TABLE 2 (contd.)

~- A. N. hypacanthus crockeri uninotatus

Parental:

25. Pectoral t Fan:

Ambivalent:

26. Head 0 0 Quiver

.._, 31.

Figure 3

Agonistic behavior of Ernblemar1a hypacant.hus.

A. Two males displaying Mouthfighting acti;d.ty.

B. A male refuge holder displaying Fin Erect

and Gape postures to a Fleeing male.

Enlargement: 3X. A

8 33.

COMFORT .MOVEMEm"'TS

I-lci>ougal (1933) proposed the term "comfort movements."

La , Kortlandt (1940) introduced comfort movements as

those actions that apparently serve to remo-ve discomfort

in the animal, e.g., scratching and preening in a cormo­

rant. Baerends and Baerends-Van Roon (1950) included in

this category stretching movements, which had previously

been termed "preparatory movements" :by Heinroth (1910).

All species studied were nearly identical in respect

to their comfort moVIaments. These movements may be sepa­

rated into two groups, stretching movements (Yawning, No. 4)

and those related to removing irritants or detritus from

the body (Coughing, No. 5, Chafing, No. 6).

Yawning is primarily a stretching movement of the jaw

musculature. Yawning is most frequent when the fish is

otherwise unoccupied e.g., the fish has been inside its

refuge for a prolonged period. This movement is common to

many fishes e.g., the stickleback (.Morris, 1958), and

Cichlidae (Baerends and Baerends-Van Roan, 1950), and is most similar to that described for the blenny genus 34.

Hypsoblennius (Losey, 1968).

Coughing apparently serves to clear the oral-bran­ chial chamber of foreign material. This action is espe­ cially common in silt-laden water, during feeding, and after handling when mucus collects in the oral cav:lty.

This movement is common to many fishes. Barl~J (1961) has described it in Badis badis and Losey (1968) in

Hypsoblennius.

Chafing apparently aids in the removal of irritating particles from the body (Baerends and Baerends-Van Roon,

1950). This action was commonly associated with fishes infected by the ciliate protozoan, Cryptocarion irritans.

This movement also occurs occasionally as a displacement action in Emblemaria hypacanthus. Losey (1968) frequently found species of Hypsoblennius to use this action as dis­ placement activity.

In summary, Comfort movements generally occur in fishes during periods of relative inactivity. The excep­ tion occurs when the movements appear as displacement actions in cases of conflicting or ambivalent motivations. 35.

ELIMINATORY BEHAVIOR

Generally no s_pecial eli:minatory behavior was ob­ served. This is co:m:mon in most fishes in which the feces usually emerge continually, particularly during strong flexures of the body (Losey, 1968). W"ickler (1957), how­ ever, observed a special defecation movement "/hereby the fish curves the body and beats a pE:Ctoral fin down tl1e concave side excreting a fecal ball.

Emblemaria hypacanthus exhibits both the unspecial­ ized and a specialized behavior. A string of fecal material may be passively excreted when the fish is out­ side the refuge. However, inside the refuge the pelvic fins are used to sweep out the fecal pellets depc.sited there. This behavior is in keeping with the extensive tube cleaning activities of the fish. 36.

E~v lRONI1E:NTAL ALTERATIONS

Alteration of the environment such as moving foreign

objects, digging and ventilation is a common occurrence in

many animals. The blennies studied also exhibited these

actions in the form of Spit, No. 8; Pectoral Pa~No. 25

and Stationary Swim, No. 9.

Individuals may dig shallow holes under rocks or may

clear material frOitl inside of an existing hole by Station­

ary Swimming, or Spitting. Stationary Swimming produces

sweeping currents along the substratum that carry away

lighter particles. Spitting is used to carry particles of

sediment or debris from the digging site. After a fish has

occupied a territory for two to three weeJ~s, it 'Vlill re­

sist the addition of foreign objects to tbe immediate

vicinity of the refuge. This was witnessed for Acanthem­ blemaria crockeri in the field. After I placed a few

small pieces of algae over the entrance to the refuge the

fish would carry them away with the mouth.

Ventilatory activity in the form of pectoral fanning

serves to increase the flow of water around the fish. 37.

This convection may prevent lowering of the oxygen tension in the refuge. These respiratory currents are usually associated with the fanning of the eggs by the male. 30 ..

The territory is "!:hat area surrounding the central

refuge and is defended against intraspecific intru;Ition.

Both sexes of the blennies studied hold territories, ex­

cept during sexual behavior when males become strongly

territorial and females wander throughout the available

habitat. Generally, the larger the individual, the larger

the territory. Larger dominant fishes attempted to defend

more than one refuge within their territory.

The area defended by a refuge holder is a function of

the sex, size and previous encounter with the intruder. A

female or smaller male may pass through the territory un­

noticed while an evenly matched fish elicits an aggressive

display from the owner, particularly when the intruder is

an individual that has been involved in previous disputes with the owner.

The kinds of territorial habitats, spatial distribu­

tions and densities of fishes are dealt with in the

accounts of individual species. 39.

I. SPECIES DESCRIPTIONS

A. Emblemar:l.a hypa.canthus

l, Habitat

Two different habitats were found for !· hypacanthus.

The habitat of Puerto Penasco was a low, flat sandstone reef containing many srrall shallow tide pools. The fishes were commonly occupying empty mollusk holes in the sand­ stone. Other fishes common to the pools were the blenny

Hypsoblennius gentilus, the goby, Gobiosoma chiquita, the clinid Paraclinus sini and juveniles of Apogon retrosella

{Apogonidae) and Abudefduf troschelli (Pomacentridae). At

Puerto Refugio the fish were found among subtidal mats of brozoan encrusted mats of Sargassum.

Densities of !· hypacanthus were estimated at Norse

Beach, Puerto Penasco, Sonora, M

2. Behavior

The territorial behavior of E. hypacanthus does not 4-0.

depart from the generalized descriptions. The territory size ranges from 5 to 20 em. radius in ·the aquaria. (The smaller terri·tories may be an artifact of crowding and confinement in the smaller tank I) •

The fishes '.'/ere provided with pholad tulles, serpulid worm tubes and 1 em. translucent plastic tubes. Indivi­ duals of both sexes took up residence in the pholad ·tubes.

The plastic tubes and '.·mrm tubes were occupied in a tran­ sitory manner and never used as permanent refuge.

B. Acanthemblemaria_ crocker! l. Habitat

A. croqkeri showed a marked preference for the aban­ doned fan 'iJorm holes and crevices in the green coral

Porites californica. However, this fish is not unique as two gobies (Elacatinus puncticulatum and Lythrypnus dalli) also occur on the coral. It is not uncommon to see the newly settled post larvae inhabiting the siphon holes of a low encrusting brown sponge or in mollusc holes. How­ ever, as these individuals grow larger they must find larger holes or be eliminated. i\. crockeri cryptically mimics the same general coloration and the shape of the 41.

~iving f3n worm in the "I·Jithdrawn position.

Two 15 m. transects at Bahia de San Francisquito and one at Bahia de Los Angeles revealed densities of 1 to 8 individuaJ.s, all of which were clumped around or on

Porit~~ californica.

2. Behavior

The territorial behavior does not depart from that of the general description (page 38). The sizes of the terri­ tories in the aquaria were generally the sarr~ as those for

E. hypacanthus. Five :'1· crockeri divided up a single 10 em. diameter piece of columnar coral into territories rang­ ing from 4 to 7 em. in radius. However, a 20 em. diameter rock scallop was divided among seven fish resulting in territories of 4 to 15 em. A. crockeri preferred aban­ doned mollusc holes, worm tubes and crevices fvund in

Porites and the rock sc~llop Hinnites multirugosa to artificial translucent plastic refuges.

c. Neoclinus uninotatus

1. Habitat

In l4onterey Bay Neoclinus uninotatus is most commonly associated with subtidal sand and trash areas such as 42.

found at Monterey municipal wharf number 2. N. uninotatus prefers to inhabit the empty containers (e.g. bottles, jars and cans) which have been discarded and lay abun­ dantly on the sandy bottom next to the wharf. I have also seen one individual inhabiting an empty pholad burrow on the subtidal shale in the vicinity of the 1'1onterey sewer outfall which is located at a 1,000 meter distance east of the wharf. Another single individual was taken under the Highway 1 bridge located at the Elkhorn Slough at Moss

Landing. ~~les are more sedentary and territorial than females. Females meander over the sand bottom more often than males.

2. Behavior

· 1'1. uninotatus, a larger bler>..ny, guarded la:~:ger terri­ tories ranging from 10 to 40 em. radius in lab aquarium

II. Refuges were supplied in the form of barnacle shells, bottles, and black plastic tubes. No pxeference was indi­ cated for any particular refuge. Some individuals exca­ vated caves underneath rocks.

Territory size, as estimated from the spacing of individuals, appeared to be larger and quite variable in th8 dominant sand and trash habitat at the Monterey harbor. 43.

Three permanent 25 square meter grid quadrats (Fig. 2) were observed 2 to 4 times per month for a period of 7 months. Ample refuges v1ere supplied in the form of glass bot·!:les. The shortest distance observed between two in­ dividuals within a refuge was 25 to 60 em. More commonly distances ranged from 2 to 3 meters.

The population density in the grid area ~ncreased during the period from Narch 16 to June 3, 1971. The numbers rose from a lov1 of 6 to a high of 21 individuals vY!th!n the combined 75 square meter area. Apparently in­ dividuals were immigrating into the \~harf area.

The increase in nu:rJ:>er of N. uninotatus appears to be greatest during the late spring and summer months. ry:·his seasonal appearance of the fish cot1ld be linked to the abundance of dying, post spawned squid in the l'ionterey area. I have observ<:d the squid fishing hoats to discard portions of their catch into the water surrounding ·the wf>arf when pumping their bilges. Gut analysis of eleven

~· un!notatus during May and June, 1971 revealed decapod crustacea, Onuphid polychaetes and squid tentacles.

Fields (1965) reports the peak squid catch for l'1onterey to be during the months of April to July for the years 1931 44.

to 1960. This period corresponds to the sEasor.a l irmni­ gration of N. uninotatus. 45.

AGONISTIC BEHAVIOR

~e fight behavior of fishes has most often been

dealt with in an anecdotal manner. This simply involves

an account of the various displays or fixed action patterns.

For example, Robins, Phillips and Phillips (1959) have

described the threat and attack behavior for an Atlantic

chaenopsid Chaenopsis ocellata. This type of study re­

veals little about the quantitative patterning of the

individual movements or acts that make up the so-called

"fixed action patterns."

In theory then, if a fixed action pattern consists of

several components (acts), say A,B,C, and D, these should

occur in some predictable sequence (Hinde, 1970). It is

generally recognized that the fixed action pattern is a

graded response (Barlow, 1969). This gradation is ex­

pressed as the intensity of the action which is directly

related to the stimulus for expression (Morris, 1957).

How predictable are the sequences of actions and what

are the underlying causal factors responsible for the expression of various threshold levels? This section will attempt to answer these questions in terms of the agonistic behavior of the blennies studied.

The agonistic behavior of the three blennies centers mainly around the defense of the territory. Agonistic encounters occur when an individual infringes upon the territorial area o£ another. The encounters usually start with a threat display such as Fin Erection and

Branchiostegi Lower and never proceed beyond the initial threat stage. Encounters are terminated by separation of the antagonists and by the assumption of a submissive posture, such as Fin Folding, by one of them. The motor patterns below signify agonistic behavior and are arranged in order of increasing aggressive \~lence in the display hierarchy (Barlow, 1962). The adjectives, aggressive and submissive, refer to the immediate behavior of the fish as evidenced by its overt behavior in terms of motor patterns.

I. SPECIES DESCRIPTIONS

A. Eniblemarie hypacanthus

There are no obvious sexual differences in the motor patterns of the agonistic behavior. However, males demon- 47.

strated a greater amount of agonism and differed in respect to sexual dimorphism (see p.ll). The patterns are listed below in order of increasing intensity (Barlow, 1961).

Nurobers in parentheses refer to the general etbogram.

1. Flee (10 ): submissive (Fig. 3B)

2. Fin Folding (11): submissive

3, Backwalk (3): submissive. This movement

rarely occurs. Backwalk progresses quickly

into Flee, making distinction between the

two difficult.

4. Headup (12): Submissive and aggressive

5, Fin Erection {13): aggressive. The form of

this movement is highly ritualized, The high

sail-like dorsal fin of the male accentuates

the act.

6. Branchiostegi Lower (14): aggressive

7. Gape (15): aggressi'~ (Fig. 3B)

8. Mouthfight (16): strongly aggressive. This

differs from the general ethogram in that the

opened mouth is often directed at the opercular

spot of the opponent. {Fig. 3A)

9. Push (18): strongly aggressive. This movement 48.

and mouthfighting form a continuum and the

two are difficult to separate. The push is

directed at the opercular spot of the opponent

as in Mouthfighting.

10. Bite (17): strongly aggressive

11. Chase (19): stro~g1y aggressive

Nonterritorial agonism - These encounters never pro­ gressed pest the initial threat stages. Head Up and Fin

Erection are the first steps in the threat hierarchy.

Fin Erection is followed by Branchiostegi Lowering and

Gaping. The displays do not progress further as the sub­ ordinate individual will Flee without returning the display.

The subordinate fish is always the smaller of the two. The intensity or valence of the motor patterns expressed .is a direct function of the stimulus situation e.g., posture, size, proximity, coloration, and any previously determined dominance of the opponent.

Intraterritorial agonism - The valence of the ago­ nistic acta 'l'lithin the territory also depend upon the stimulus situation and previous encounters. The proximity of the refuge is moat important. The territory may be defended from within the refuge. A Gaping display from 49.

the entrance usually suffices to drive away a smaller or subordinate individual.

The following account presents an idealized encounter that develops into the highest intensity motor patterns.

All of these acts are rarely observed during a single en­ counter. The hypothetical fishes are equal in size and are referred to as 1, the defender, and 2 the intruder.

Fish 1 assumes the Head Up display as 2 approaches the territory. As 2 approaches within 5 em. 1 protrudes from the refuge and assumes the Fin Erect posture. Two responds by also assuming the Fin Erect posture. As the two fish come into lateral juxtaposition, the threat dis­ play increases to the next higher intensity. Both l and 2 simultaneously assu~e the Branchiostegi Lowering and

Gaping displays. Two aims its Gape at the operculum of 1 and Pushes 1 out of the refuge. One darts at 2 with an open mouth and 2 backs up momentarily. They Mouthfight for 3 to 4 seconds interrupted by periods of 5 to 10 seconds when both fish resume the lateral displays. One

Bites down on the lm~r jaw of 2 momentarily. ~1o slips out of the Mouthlock and both diaplay Gaping. Two's in­ tensity of display decreases as 2 moves slowly away, and suddenly Flees. 50.

B. Acanthemblemaria crockeri

No differences in agonistic motor patterns were ob­ served between the sexes of ~. crockeri. Males were more aggresnive then females and differed in the sexually dimorphic color pattern (Fig. 8). As most of my observa­ tions were wade dltring spring and summer, it is possible that sexual motivation (e.g., hormones) may have affected the fish, even though it was not always evidenced by court­ ship. The actions that differed from ;§_. hyPacan·thus are listed below. The numbering has the same reference as in the previous section.

1. F~ad Up (12); mild aggression. The mouth may

be subma~dmally opened at low intensities of

aggression.

2. Fin Erect (13): mild aggression. This movement

is much less ritualized than that seen for E.

hypacanthus and the dorsal fin is not highly

developed but is low and e'\ren in shape.

3. Mouthfight (16)1 strongly aggressive. The jaws

are opened maximally to an angle of 90° or less

and directed at the gaping jaws of the opponent.

But instead of a pushing at the opponent's mouth 51.

or opercular spot, the head is moved laterally

from side to side. This effects a raking move­

ment over the opponent's head.

4. Head Butt (20): This action is described in the

general ethogram and was observed only for A.

crockeri.

Intense fighting was observed, filmed, and will be dealt with in detail. A. crockeri differs from E. ~­

canthus and Neoclinus uninotatus in the manner in which

fights develop. A challenging fish circles around the

opponent's refuge displaying the Fin Erected, Branchio­

stegi Lowered and mouth Gaped. The challenger attempts

to Push the opponent out of his refuge by using a swift

Head Butt aimed at the operculum. Intense Mouthfighting

commences and may last 3 to 4 minutes. During this time

the intruder attempts to take over the refuge by entering and defending it. On one occasion the refuge was ex­

changed 3 times with the owner ultimately winning out.

Nonterritorial agonism - These encounters consist of

a dominant fish approaching a subordinate, who often

Backwalks and/or Flees. If the approached fish does not

Flee, the aggressor responds with a low intensity Gaping 52.

display.

Intraterritorial ?sonism - A subordinate fish, not displaying, does not elicit aggressiveness unless he ap­ proaches to within 4 em. of the refuge. An evenly matched intruder causes the owner to assume l~N intensity Head UP and Branchiostegi Lowering and Fin Erection. The intruder responds by returning the lateral display and attempts to push the owner from the refuge. This leads to intense

Mouthfighting, Pushing and Biting. The intruder is driven off by a series of Pushes and Bites. The owner then re­ enters his refuge.

t-!ales were almost always successful in defending their territories for periods up to 30 days in aquaria. Males without territories were subordinate and rarely aggressive.

Females are dominant only when attempting to defend a territory. Females rarely hold territories for more than a few days and are observed to roam about more often than not.

c. Neoclinus uninotatus

This clinid species shows a lesser degree of agonis­ tic activity than the chaenopsids. Displays rarely pro­ ceeded to actual contact. When contact was made it was 53.

brief and never advanced to intense fighting as seen in

the chaenopsids. No sexual differences existed for the

intensity of aggressiveness or the motor patterns.

The following motor p3tterns T.1ere not observed to be associated wit1l agonistic behavior• Backwal'k, HeaCl Up and

Branchiostegi r,owering. The actions that were notably different from the chaenopsids are (numbering as before):

1. Head Arch {25): aggreAsive. The front of the

head is pointed d01.>1n While the predorsal fin

area is arched up. This movement occurs within

the refuge and serves to make the fish appear

larger.

2. Gape (15): aggressive. ~. l1ninotatus is capable

of a trernendotls Gape1 the ja\'>~S 3re opened maxi­

mally to 180°.

3. Head Quiver (26): ambivalent. This movement

occurs during the entry of an intruder into the

territory. The Head Quiver takes place within

the refl!ge.

4. Mouthfight (16): strongly aggressive. The two

opponents face each other, Gape simultaneously

and press the jaws together one against the other& 54.

5. Push (18)' strongly aggressive. With the mouths

.r:n:·essed tightly together, the opponents commence

sinusoidal swimming movements toward ·!:he other.

Nonterritorial agonism - Fresh specimens from the field often Gape, Mouthfight and Push in the transportation container. These intense encounters last 3 to 5 seconds.

After the fish are introduced to the aquaria and establish territories, nonterritor:!.al encounters end with only a mild display of Fin Erection or a low intensity Gape.

Intraterritorial agonism- High intensity Gaping, l"iouthfighting and Biting can be continually elicted from a territory owner by forcefully introducing another con­ specific or sometimes by a mirror. 'l'he intruding fish, if smaller, was always met by the owner extending from the refuge Gaping maximally, a sufficient display to cause the intruder to Flee. If the intruder is larger, the owner may stay within the refuge and Gape submaximally, Head

Quiver and/or Head Arch. A larger individual often evicts a smaller one from his refuge by Gaping wEximally at the entrance of the refuge. If this is not successful the larger fish will Bite down on the lower jaw of the owner and jerk him from the refuge. These encounters last up to 10 seconds. 55.

TABLE 3

Occurrence of agonistic motor patterns and their mean durations (in seconds ± standard deviation, as determined from cine analysis where available); order as in general ethogram (page 19).

0 - does not occur

+? - probably does occur but observations insufficient

+ - does occur, in the form of the general ethogram * • does occur, but in slightly altered form from general ethogram

? - the situation in which it might occur was not seen

Emblemaria Acanthemblemaria Neoclinus hypacanthus crocker! uninotatus

1. Backwalk (3) + (0.45fO.OB) + (0.93f0.58} 2. Flee (10) + + (0.93;!0.58) + 3. Fin Fold (11) + (1,9±1.4) + +

4. Headup (12) + (6.9±11.0) + (0.58;!0.41) 5, Fin Erection (13) + (8.6±9.5) + (11.9±5.8) + 6. Branchiostegi + (5,7±5.1) + (1.5±0.48) Lower (14) 7. Gape (15) + (2.7;!:2.3) + (l.03;!(J.68) 8. Mouthfight (16) + * (0,47;!0.22.) 9. Bite (17) + (.47;!:0.25) + (.53;!:0.23) + 10. Push (18) * (.30;!:0.21) * (.91;!(J.41) * 11. Chase (19) + ( .54;!:0.39) + + 12. Head Butt (20) 0 + 0 13, Head Arch (22) 0 0 + 14. Head Quiver (26) 0 0 + 56.

II. TEJHl?OR!cL P.SSOCIATI0~1 OF T:tl.E AGONISTIC HOVEk,iENTS

It is suggested, in the previous section, that groups of movements are commonly associated. For example, the lateral display of the chaenopsids, ~. hypacanthus and

A. crocker!, is composed of three motor patterns: Fin

Erect, Branchiostegi Lower and Gape. Each of these move­ ments represents an increasing intensity of agonism.

Thus the lateral display fits the definition of a fixed action pattern given on page 45. The problem of the pre­ dictability of the patterning of the individual agonistic movements will now be considered.

One can arrange the movements that commonly precede or follow each other, i.e., show a high temporal associa­ tion (Baerends et al., 1955). If the movements compose certain fixed action patterns, then correlated pairs of motor patterns should be expected. On the other hand, if the movements described hav~ no common causal factors, one would expect them to precede or to follow each other randomly. The randomness of the sequence of these move­ ments can be tested.

For this analysis, over 550 feet of movie film was used for a frame by frame recording of the various ago- 57.

nistic movemen·ts. This represents 50 to 60 hou.rs of aquarium observai;ion time. Every separate performance has been recorded and time units computed from the film speed and frame count (Table 3). No distinct.ion has been made bet\~een low and high intensities of the same movement.

'l'he filmed agonistic movements and post•1res of 7 individuals (5 males and 2 females) of ErrJ:>lemaria hypacan­ thus included in·teractions bet'I-Jeen males, females and male-female pairs (not courtship). The films of Acanthem­ blemaria crockeri included 4 males. I counted the number of times each movement preceded or followed itself or each of the other movements. For nine movements the results of this counting have been summarized in Table 4 for ~. hypacanthus and in Table 6 for ?:_. crockeri. Here in each cell the top-most figure indicates the number of times a certain muvement preceded (reading the column), or followed

(reading the row) itself and each of the other movements.

The second fig·ure in each cell indicates the number expected for each cell in case of random sequence; it has been ob­ tained by dividing the product of the marginal totals for each cell by the sum total.

To get comparable values of the differences between 58.

the numbers obtained and expected, for each cell the former was divided by the latter. These ratios, the third value in each cell of Tables 4 and 6, indicate the relative frequency of each of the nine movements that precede or follow each other. The ratios can now be ranked according to decreasing size; in this way two rank numbers (one for preceding and one for following movements) are found for each cell (Siegel, 1956).

The follo1ving is a list of abbreviations used in the tables of this section. FE = Fin Erect, BL =

Branchiostegi Lm~er, G =Gape, Bt -=Biting, HF =Mouth­ fight, HU ~ Head Up, B"l = Backwalk, HB = Head Butt,

P = Push, FF = dorsal Fin Folded, Chs ~ Chase, and

Fl = Flee. 59.

TABLE 4

Frequency with which the nine different movements precede or follow each other in the agonistic behavior of Emblemerie hxpacanthus (see page 57 for explanation of cells)

Preceding component

FE BL G Bt r!F HU BW FF Chs Marginal Totals FE llO 39 102 19 11 19 20 0 0 310 39.9 30.2 30.0 11.6 13.0 17.8 9 ,l, 1.6 1. 0 2.8 1.3 3.4 1.64 0.85 1.06 2.13 0 0

BL 52 55 85 10 10 18 0 0 0 230 29.6 22.4 22.2 8.6 9.7 13.2 7.0 1.2 o.a 1. 75 2.45 3.8 1.16 1.04 1.36 0 0 0

G 90 98 70 21 29 60 20 0 0 388 50.0 37.8 37.5 14.5 16.3 22.3 11.7 1.5 1.3 1.8 2.6 1.86 1.45 l. 78 2.7 1.7 0 0

Bt 21 29 9 11 10 20 0 0 0 100 ... 12.9 9.7 9.7 3.7 4.2 5.75 3.0 0.5 0.3 ., 1.63 3.0 0.93 2.97 2.4 3.5 0 0 0 c" 0 0. MF 21 0 42 ll 10 0 0 13 9 106 8 0 13.7 10.3 10.2 4.0 4.5 6.1 3.2 0.5 0.4 u 1.54 0 4.0 2.75 2.2 0 0 26 22.5 s::"" .... HU 31 22 13 0 12 22 0 0 0 100 5 12.9 9.7 9.7 3.7 4.2 5.75 3.0 0.5 0.3 ...... 0 2.4 2.26 1.32 0 2.85 3.8 0 0 0 "" nw 24 19 41 29 32 9 31 0 0 185 23.8 18.0 17.9 6.9 7.8 10.6 5.6 0.9 0.6 1.0 1.06 2.3 4.2 4.1 0.85 5.5 0 0

FF 0 0 0 0 0 B 11 0 0 19 18.4 1.9 1.8 0.7 0,8 1.1 0.6 0.09 0.06 0 0 0 0 0 7.3 17.8 0 0 Cha 0 0 0 0 12 0 0 9 0 21 2.7 2.0 2.0 0.8 0.9 1.2 0.6 0.1 0.06 0 0 0 0 13.2 0 0 90 0 Marginal Totals 349 264 262 101 114 156 82 14 9 2710 Sum Total 60.

TABLE 5

A. Spearman's rank correlation coefficient for preceding movements of Emblemaria hypacanthua

FE BL G Bt MF HU BW FF Cha

FE 1 ~ 0.535 0.110 -0.265 0.140 0.583 0.213 -0.185

BL 1 0.880 0.658 -0.119 0.370 0.170 0.177 0.040

G 1 0.677 0.175 0.713 0.291 0.513 -0.035 Bt 1 -0.351 0.723 0,189 0.330 0.131 MF 1 -0.360 -0.360 -0.092 0.621

HU 1 0.608 0.330 0.275 BW 1 0.025 0.081 FF 1 0.318 Chs 1

B. Spearman's rank correlation coefficient for following movements of Emblemaria hypacanthus

FE BL G Bt MF HU BW FF Chs

FE 1 0.640 0.550 0.140 -0.260 0.221 -0.060 0.040 0.247

BL 1 0.150 0.341 0.008 0.473 -0.077 0.190 0.176 G 1 0.516 0.205 -0.351 0.382 0.060 0.091 Bt 1 0.220 -0.230 0.320 0.221 0.565 MF 1 -0.440 -0.359 0.501 0.200 HU 1 0.307 0.092 0.070 BW 1 0.283 0.283 FF 1 0,663 Chs

NOTE: Underlined values are significant to the 0.05 level see text for abbreviations 61.

TABLE 6

Frequency with which the nine different movements precede or foll~w each other in the agonistic behavior of Acanthemblemarie crocker! (see page 57 for explanation of cells) Preceding component G BL IUl MF Bt p FE Fl HU Marginal Totals G 69 57 60 44 9 21 9 24 3 296 36.5 12.1 36 26.7 6.3 10.6 7.8 8.8 1.44 1.9 4.7 1.67 1.65 1.43 1.98 1.15 2.7 2.08

BL 51 15 6 15 0 3 0 24 3 112 14.3 4.57 13.6 10.1 2.4 4 2.9 3.3 0.54 3.56 3.J 0.44 1.48 0 0.75 0 7.J 5.5

HB 68 6 59 54 15 12 4 9 3 230 29.3 9.34 28.02 20.7 4.9 8.23 6.01 6.84 1.1 2.3 0.64 2.1 2.6 3.06 1.46 0.67 1.32 2.7 ...c '" MF 56 0 42 54 10 9 45 8 0 224 "'0 28.5 9.09 27.3 20.17 4.8 8.0 5.9 6.7 1.1 1.96 0 1.54 2.68 2.1 1.1 7.6 1.2 0 u~ i'J' Bt 7 3 15 27 0 4 0 0 0 56 .... 7.13 2.3 6.8 5.0 1.2 2.0 1.5 1.7 0.27 ~ 2.2 5.3 0 2.0 0 0 0 ...... 0.98 1.3 0 !>< p 6 0 75 0 3 14 7 0 0 105 13.4 4.7 12.8 9,5 2.2 3.8 2. 7 3.1 0.5 0.45 0 5.65 0 1.36 3. 7 2.6 0 0

FE 28 4 21 19 5 22 0 5 3 107 13.6 4.3 13.0 9.6 2.3 3.8 2.8 3.2 0.5 2.05 0.93 1.61 1.98 2.17 5.!J 0 1.56 6.0

F1 17 9 25 18 11 4 0 0 0 86 10.9 3.5 10.5 7.7 1.8 3.0 2.2 2.5 0.4 1.55 2.57 2.37 2,34 6.1 1.3 0 0 0

HU 16 10 0 3 0 0 0 4 0 33 4.2 1.3 4.0 2.97 0.7 1.1 0.86 0.98 0.15 3.8 7.7 0 1.05 0 0 0 4,08 0

Marginal Totals 317 101 303 224 53 89 65 74 12 2487

Sum Total 62;;

TABLE 7

A, Spearman' s rank correlation coefficient fo:c preceding wovements of Acanthemblemaria crockeri

G BL HB Ml' Bt p FE Fl HU G 1 0.796 -0.360 -0.900-- 0.022 -0.440 0.084 -0.120 0,633 BL 1 -0.050 -0.560 -0.018 -0.640 0.160 -0.390 0,665

Rll 1 0.190 -0.069 -0.010 0.766 0.243 -0.320 MF 1 0.066 0.322 -0.300 0.118 -0.123 Bt 1 0.252 0.015 0.444 0.261 p 1 0.067 0.392 -0.500

FE 1 0,016 -0.320 Fl 1 0.225 l!U 1

B, Spearman's rank correlation coefficient for following movements of Acanthemblemaria crockeri

G BL Hll Ml' Bt I! FE F1 HU

G 1 0.438 -0.950 -0.180 -0.128 -0,400 -.o. :no 0,850 0.540 llL 1 -0.450 -0.32.1 0.219 -0.321 -0.425 0.570 0.170 HB 1 0.150 0.385 0.632 0.367 -0.820 -0.200

Ml' 1 0,350 0.100 -0.275 -0.399 -0.020 Bt 1 0.190 0,267 -0.300 0.250 p 1 0.200 -0.440 0,284 FE 1 -0.119 -0.037 Fl 1 0.634

HU 1

NOTE: The underlined values are significant to the 0,05 level. See text for abbreviations. 63.

F~om the ~ankings, Spearman's ~ank co~~elation co­ efficient has been calculated for preceding (Tables 5A and

7A) and following~ables 5B and 7B). The significance of

these values can be tested (Siegel, 1956-210) and in

Tables 5 and 7 the unde~scored values are significantly correlated at 0.05 level. Table 8 presents the correlated movement pai~s, both positive and negative, for Emblemaria hypacanthus and Acanthemblemaria crocke~i.

It is evident each pair of the 10 movement pairs for

Emblemaria hypacanthus and of the 7 for Acanthemblemeria

crocker! have statistically significant common causal fac­

tors. Four of the movement pairs for A. crocker! do not demonstrate any statistically significant common causal factors. The common causal factors for the correlated move­ ments are not clearly a result of the fixed action pattern.

The lateral display consists of Fin Erection, Branchiostegi

Lawer and Gaping, in that order. For E. hypacanthus the

co~relation of Fin Erection followed by Branchiostegi Lower only is fulfilled and the reverse is also significant. The other chaenopsid, A. crocker!, shows Branchiostegi Lowering

to be significantly followed by Gaping. Suffice it to say for now that the agonistic fixed action pattern of lateral 64,

TABLE 8

Summary of statistically correlated movement pairs for the cbaenopsids,

E. hypacanthus A. crockeri

correlation ••• positive positive negative

BL - FE BL G MF G

G BL HU G 1' BL

FE - BL HIJ llt G ll1J Bt BL FE - HB HB - Fl Bt G G - Fl HU - G HB p HU - Bt Fl - HU Chs - MF llW - HU FF Cbs 65.

display may not be as fixed as it appeared to be in the general observations.

III. MOTIVATIONAL ANALYSIS

In order to understand the relationships among the pairs of correlated movements, it might be best to consider their underlying causal relationships. A review of the ethological concepts regarding causation is given by Hinde

(1970: section 2). Motivation can be defined as the ten­ dency to perform an action, or set of actions (Losey, 1968:

50}. According to Hinde (1970) the performance is deter­ mined by:

1. The balance of the combined motivational

states which may facilitate or inhibit

one another~

2. The absolute threshold level of the action

in terms of the stimulus intensity necessary

for its expression;

3. The relative threshold level of the action

in terms of the conflict between two actions

where the one with the highest threshold will

be expressed. 66.

Many studies indicate that the motivation of ago­ nistic behavior is the resultant of conflicting motiva­ tions (Hinde, 1970; Morris, 1958r Losey, 1968). In this analysis sexual motivation is assumed to be low only for the purpose of simplicity. Thus only two fundamental tendencies remain: attack (A) and flee (F) (Tinbergen,

1940). Since these two tendencies are incompatable, a fighting animal finds itself in a state of conflict. The exact nature of the conflict state at any one moment will be determined by: (1) the absolute level of the fleeing tendency1 (2) the absolute level of the attacking tendencyr

(3) the relationships between these two levels, e.g., a high A/F ratio is indicated by the probability that more, aggressive activities will foll~J. Furthermore, the hypo­ thetical threshold levels (see 2 above) for the various agonistic actions can be presented as a function of the flee and attack motivations (Morris, 1958).

The motivational analysis presented here attempts to follow Losey (1968), Morris (1958) and Tinbergen (1952).

This method describes the observed behavior in terms of stimulus intensity and is particularly useful in compara­ tive outlines. Figure 4A (page 72 ) (after Losey, 1968) 67.

may be used for purposes of discussion and interpretation.

The intensity of the agonistic actions increases with the distance from the origin. Each action is represented by a range of A/F intensities included in the shaded areas.

Mouthfighting is a more intense action than Head Up threat, and Action B is more intense than Action A (see Fig. 4A).

The actual expression of the agonistic action is deter­ mined by the relative threshold level as discussed in 3 above (page 65). These relative threshold levels are subjectively determined from the data given in Tables 4 and 6, from notes and from tape recordings of agonistic activity patterns. The more intense actions are hier­ archically superior. ONhen the threshold for Mouthfight­ ing is breached, this action would be expressed in place of the hierarchically inferior Head Up threat in Fig. 4A.

Action B would be expressed in favor of Action A). The probability that the individual will actually attack or fight is positively correlated with the intensity of the actions as represented by their A/F ratios.

As an example, in the simplified behavior pattern of

Figure 4A the sequence of behavior resulting from an in­ crease in pure flee stimulation is Backwalk, Fin Fold 68.

and Flee. Similarly, an increase in pure attack stimula­ tion leads sequentially to Head Up, Mouthfight and Chase.

Hmvever, you will note in Figure 4C that as the stimula­ tion becomes a mixture of both flee and attack moti,Tations, the sequence is different from the pure attack stimulation.

For example, the pure attack sequence for Acanthemblemaria crocker! (Fig. 4C) reads Head Up, Fin Erect, Branchiostegi

Latl, Chase, Gape, .1>1outhfight, Bite, and l?ush. As the stimulation becomes mixed the sequence changes to Head Up,

Fin Erect, Branchiostegi Low, Mouthfight, Gape, Bite, and l?ush. The slopes of the threshold lines of the motor patterns are also subjectively determined from the amount of attack and flee stimulus presented by the opponent.

For example, the movement Chase has a greater amount of attack stimulus resulting in a slope towards the attack axis.

The most important implication of the A/F hYPothesis is that at equal intensities of the F and A motivations, they exhibit a mutual inhibition (Tinbergen, 1952).

Tinbergen points out that when the tendencies to attack and to flee are more or less in balance, each prevents the other from expressing itself fully, so that the animal can 69.

neither attack its enemy, nor flee from it. In such a situation an animal performs one of two basic types of activity: ambivalent actions, or displacement activities.

The range of equal attack and flee intensities (Actions A and B, Fig. 4A) has been found to correspond to ambivalent behavior and displacement behavior respectively (Morris,

19581 Moynihan, 1953). The ambivalent behavior has a l~Ner threshold.

The A/F motivational lev-els in the blennies studies are determined by: the stimulus situation presented by the opponent, refuge proximity, and to a lesser degree any immediately preceding agonistic encounters. This includes such factors as the posture, distance, size, relative dominance, and perhaps coloration (see pages 89 to 107) of the opponent.

Several of the agonistic actions are never, or only rarely seen in the absence of an aggressive response by the opponent. This aggressive response may be in the form of a sudden move toward the individual, an attempt to Bite, or more commonly, an aggressive posture. The points La and Lf represent the point of maximum motivational levels that can be attained in the absence of this aggressive response (Fig. 4A). points, l_ and 1..,, represent the "' ,_ threshold line, low which, it is supposed, the animal will insufficiently stimulated to shoN any overt ago- nistic responses.

Species differences in the motivation of agonistic behavior, both qualitative and quantitative, are discussed separately below. For simplicity, shaded areas rep- resenting the range of A/F intensities have been replaced by single lines Which de:mark the threshold va (Figs.

,c,andD).

A. Eniblemaria hvpacanthus

The hypothetical A/F diagram that portrays the be- havior observed i~ E. hypacanthus, is presented in Fig.

4B. The factors delimit the appro~irnate placerrant of each action are summarised below" A more explicit discussion would require a detailed account of many in- dividual encounters. Suffice it to say that many lesser factors also entered into the construction of the diagram.

l. Actions were not observed where attack and

flee motivations are low; i.~., most non-

territorial encounters when the oppor.ent

is not displaying. 71.

Figure 4

A/F diagrams for Effiblemaria hypacanthus, i'\cantheffiblemaria crockeri and Neoclinus uninotatus.

The threshold levels for the agonistic actions are presented as a function of flee and attack motivations. A is a hypothetical diagram for discussion. The shaded areas represent zones of A/F ratios for the action pattern. See text.

Abbreviated is: roth-fight - mouthfight.

(after Losey, 1968) 7? '~.

A 0 0 ,...., ? La A- 73.

2. Backwa lking appears over a range of lrrw and

medium A/F intensity encounters. It might

be considered 1~1 intensity Fleeing.

3. Fleeing is seen in a wide range of encounter

intensities. It may be followed by aggressive

action if the fish remains at some distance

from the opponent.

4. Fin Folding appears in subordinate fish when

the opponent is much larger or is displaying

aggressively. It usually precedes Fleeing in

these situations. A wide range of intensities

of this action is seen. The higher intensities

are associated with higher A/F ratios.

5. Eead Up occurs during low and high intensity

encounters as well as conflict situations.

The higher intensities are accompanied by:

a. Fin Erection, which is the next

higher intensity state.

b. Branchiostegi Lowering

6. Chasing is seen during aggressive behavior when

the stimulus situation is provided by a Fleeing

fish. This suggests a reduction in Flee motiva- 74.

tion.

7. Gaping is seen at high A/F ratios and leads into:

a. Mouthfighting, Which consists of

an advancing ano retreating com­

ponent, suggesting that the A/F

ratio may oscillate near unity with

the performance of the action. The

retreat portion diminishes at higher

A/F ratios, and the fish come into

juxtaposition, a necessary p"!:erequi­

site for:

8. Push. This action also contains an advancing

and retreating component while occurring at a

slightly higher intensi t;r.

Intense behavior is rarely seen in the absence of an aggressive display by the opponent. The more aggressive the display, the greater the augmentative value tLosey,

1968). High augmentation is correlated With high motiva­ tion. The augmentative value of the aggressive actions inc~eases from left to right in Figure 4B. It is intui­ tive that these aggressive displays ai.J

B. Acanthemblemaria crockeri

The hypothetical A/F diagram that depic·ts the be­ havior seen in !!.· crockeri is presented in Figure 4C.

The relationships between the threshold values and the

A/F motivations are qualitatively much the same as in

:§.. hypacanthus. The points that differ quantitatively are as follows:

l. Head Up is much less frequent and is seen in

more intense situations. It is still exhibited

by individuals in conflict situations, such as

a territory owner who sees an approaching in­

truder. This indicates that the threshold is

higher in A. crocken, while the A/F ratio is

similar.

2. Fin Erection and Branchiostegi Lowering occur

less of·ten and are much less ritualized. 'l'his

suggests a higher threshold for each of the

actions.

3. Mouthfighting and Pushing are seen more often

and occur over a wider range of A/F intensities.

The former implies 10\~er threshold values.

4. Biting occurs less frequently which suggests a

higher threshold value. 76.

c. Neoclinus uninotatus

T'ne hypothetical A/F diagram that describes the a9o­ nistic behavior observed in N. uninotatus is presented in

Figure 4D. The relationships between the threshold values and the A/F motivations are markedly different in quanti­ tative as well as qualitative respects. Due to the sparsity of quantifying evidence Figure 4D must be considered as mm:e provisional than those preceding.

1. Head Quiver is rare. It is observed only in

strong conflict situations, and may be followed

by intense aggressive or submissive actions.

2. Gaping occurs frequently over a wide range of

A/F intensities and is extremely ritualized.

The latter suggests a lower threshold.

3. Biting is often seen in situations which lack

augrrentation by the opponent. This suggests

a lower threshold value.

4. Push only occurs when two individuals have

their mouths Gaped and pressed together.

This indicates a high augmentative value and

hence a higher thrashold for expression. 77.

REPRODUCTIVE BEHAVIOR

The general reproductive pattern in the three blennies is for the female to enter the male's refuge,

deposit her eggs, and leave. The male remains with the

eggs and guards them until they hatch. The larvae then become planktonic before settling out as benthic pre­

juveniles.

The males eJthibit various courtship actions, or dis­

plays, while the females generally maintain a submissive

posture. Mating proceeds similarly for the two chaenopsid blennies, but there are marked differences in the court­

ship actions of the male. Complete mating was not observed

for Neoclinus uninotatus.

A. Species Descriptions

1. Emblemaria hypacanthus

I have observed male courtship activity throughout the

observation pariod from February to August. However, mating and spawning only occurred during April and June.

The actions that are included in the reproductive behavior of E. hypacanthus are listed below: 78.

(1) Chafe (6): males and females. This action does

not differ from that given in the general

ethogram.

(2) Fin Erect (12): males only. The movement is

not as intense as it is during agonistic be­

havior. The coloration of the body and dorsal

fin differs from the agonistic color pattern

(see pages 89 to 92 ) •

(3) Gape (14): males only. Again, neither the

intensity nor the duration of this movement

are as great as the agonistic response.

(4) Fin Flicker (22): males only (Figure 5). Of

the species studied this movement is character­

istic only of E. hypacanthus. The rapid spread­

ing and folding of the unpaired fins is coupled

with and out (spread} and in (fold) movement of

the body in the tube. Cine analysis reveals

the out and in motion to cover only 3 to 4 mm

and is barely perceptible in some individuals.

The pelvic fins are held out and away from the

body and the pectorals may be fanned for

stability. See Table 9 for duration of folded

and erect positions, and bout length. (5) Pectoral Fan (24): males only. Does not differ

from general ethogram. This action serves to

move water past the eggs in the refuge.

Cine analysis reveals a finer description of the Fin

Flicker action pattern. Four Fin Flicker bouts of a single male were filmed on different occasions. The move­ mant appears in two distinct subunitSf the up position of the dorsal fin and the down position. The mean duration in seconds of these subunits is presented below for each of the four separate sequences. The mean bout and interval durations, and bout per interval ratio are also included.

An idealized courtship sequence of a fully ripened

(determined from overt behavior or appearance} male and female is described below.

The female approaches the territory of the male with her dorsal fin about half erected. The male extends two­ thirds of his body from the tube and simultaneously assumes the Fin Erect and Gape postures. As the female approaches the side of the male's refuge, the male begins the Fin

Flicker movement. The numbers of Fin Flicker bouts varies from 4 to 9. The female then chafes 2 to 3 times in succession. As the female begins to enter the refuge, 80.

TABLE 9

Fin Flicker Subunit Duration

Sequence Duration (seconds)

Fin Up Fin Down N

#1 0.375 0.413 8

#2 0.57 0.80 5

4f3 0.435 0.645 5

#4 0.475 0.66 6

mean bout duration - 3.5

mean interval - 2.7

bout/interval ratio- 1.32 8lo

Figure 5

Male Emblemaria hypacanthus displaying the courtship inviting actions; Fin Flicker.

Taken from actual photographs. Enlargement: X4. 82 .. 83.

FEMALE STIMULUS MALE RESPONSE

A. E. hypacanthus

Nearby Extends from tube

In territory and approach Fin Erect

Displacement and chafe Fin Flicker

Enter Back into refuge

B, ?.'!_. crockeri

Nearby Extends from tube

In territory and approach Branchiostegi Lower Enter Out & in movement

Back into refuge

Figure 6. Stimulus response nature of the male invitation actions. The responses that may be caused by certain female stimuli in terms of actions are presented, with an indication of the degree of overlap, as in Barlow {1962). 84.

tail first, the male may leave the tube temporarily but he immediately re-enters. This is especially common when the size of the refuge is such that it is difficult for the female to enter. They remain in the refuge for one to three minutes. After the first female departs, a second and then a third enter in the same manner. ?ne awaiting female positions herself out>3ide the refuge until the mating female departs, at which time she enters. See

Figure 10 for stimulus-response chain.

The number of eggs deposited by the female were not counted. Stephens' (pars. comm.) fecundity counts of ripe females in his collection reveal a range of 92 to 126 eggs per female with a size of 0.7 to o.a mm.

2. Acanthemblemaria ~ckeri

~. crockeri spawned in the laboratory during the middle of June. I hnve observed newly settled post larvae during June and JUly at Guaymus, Sonora, Mexico. This may suggest a seasonal spawning during long photoperiod. Their reproductive behavior is similar to that of E. hyuacanthus except for the deletion of the Fin Flickering movement and the addition of Branchiostegi Lowering.

The courtship movement of the male again consists of a series of out and in movements from the tube refuge.

The dorsal fin is held in the ersct position and the radii branchiostegi are l~dered throughout the out and in move­ ments. One series may contain 6 to 8 complete actions.

These bouts are repeated 3 to 5 times v>~ith a variable interval between. A cine analysis th:l.s action was not available. The mating pair reme.:l.ned within the refuge for

3 to 6 minutes.

The males again guard the eggs until they hatch.

Stephens' (per a. comm,) fecundity studies shC'i'J 74-116 eggs per ripe female with an egg size 0.8 to 1.0 mm.

3. Neoclinus uninotatus

Mating was not observed. Males were observed guard­ ing eggs :from January through September with peak periods during the summer months. Initial stages of courtship were observed and both sexes appeared to use Tail Beat as a sexually inviting action. The motor pattern of Tail

Beat does not differ from the general ethogram.

I observed a mated pair within the same refuge

(bottle) in the ld. The fereele appeared upside do;.Jn and deposited her eggs on the upper surface and sides of the inside of the refuge. One refuge contained over 86.

15,000 eggs by volumetric estimation (Lagler, 1956: 107).

This was evidently the result of multiple spawnings as there were at least 2 to 3 distinct developmental stages.

A few larvae hatched from eggs that 11Jere brought to the laboratories from the field. They were kept alive four days and subsequently preserved, mounted, stained and photographed. The larvae is illustrated in Figure 7. 87.

Figure 7

Early larvae of Neoclinus uninotatus.

Drawn from a preserved specimen. 88.

5.5 mm 25 X 89.

COLORATION

The blennies studied exhibit: a wide range of color patterns that may be shown by any one individua:J., depandent on the sex some cases. They have ability to change

their color pattern within a matter of smconds. B~~~r, the me les of Embl.emar i:a bypacanthus and

Acanthemblemaria crockeri acquired the dark courting colora­ tion over a period of days.

A. Species Descriptions

1. Emblemar!!, h;[P

The predominant color patterns exhibited by~. ~­ canthus are presented in Figure a. Pattern A il!! shown for the female of the ~peci~s (note the sexual dimorphism).

This pattern more common to females end consists of

:tight brown mottling over portions of t.h.e heed and belly region but predominant over the doraal surface and extend­ ing into portions of the dorsal fin.

Color pattern B is indicative of rneles. Most apparent are the brO\vn vertical bar!ll covering middle portion of 90.

Figure 8

Color patterns for Emblemaria hypacanthus

A. Female (37.0 mm. in standard length)

depicting cryptic mottled pattern.

B. ~Ele (39.5 mm. in standard length) depict­

ing agonistic pattern. c. Same male as above depicting courtship

pattern.

Tne drawings were made from a series of

color photographs. 91., 94.

A

8

c Color pattern B is exhibited only by males. This

relatively nondescript pattern consists of an overall

yellow to red color with no strong pigmentation on the body. The anterior dorsal fin membranes beb1een spines

1 ·through 5 are heavily pigmented in red. The ventral

one-·third of the anal fin is tipped in black.

Color pattern C is also only observed for males of

the spacies. The anterior half to ~~o-th!rds of the fish

is a dark red to brown overall. The anterior half of the spinous dorsal is a dark red brown. An intermediate pattern beb..,een B and c exists and is designated B/C.

This pattern is the same as B except the body has a greater amount of the red brawn pigment present.

3. Neoclinus uninotatus

Pattern A is indicative of either sex of N. uninotatus

(Fig. 10). ~ne body colorations may range from grey-white to brawn-blacl{. Color pattern A is strongly mottled com­ pared to the other s.pacies already described. This pat­ tern is formed by darkening the saddles on the dorsal sur­ face. Orange to pink colors cover the anal fin and small parts of the dorsal fin. The fish illustrated in pattern

A is a female. Notice the sexually dimorphic shorter 96.

Figure 10

Color patterns for Neoclinus uninotatus a. Female (153 mm.) depicting the cryptic

mottled pattern b. Fa le ( 164 mm.) depicting the agonistic

pattern c. Male (as above) depicting courtship pattern

see text for full descriptions 97.

B

c 98,

ocular cirri. The single dorsal ocellus is dark blue and ringed in yellow in all three patterns depicted.

Pattern B is exhibited by a male of the species but is common to both sexes. This pattern is distinguished by the dark longitudinal stripe that runs the length of the body. Again orange to pink colors cover the anal and parts of the dorsal fin. The head is also heavily pig­ mented a brown to black color. A triangle is left un­ pigmented to the posterior and ventral to the eye.

Pattern c is pres~~d to be the courtship coloration of either sex. Here it is represented in a male. The body is strongly mottled in brown and red. The dorsal fin contains five red bars, the anteriormost being largest. The anal fin is solid red and tipped in b1ack.

Since mating was not observed, this pattern must be considered more provisional than the previous. •

II. COLOR PATTERN CHANGES

The coloration of fishes often changes with the affective state of the individual. Many of these changes are correlated with changes in behavior and probably serve as visual signals, or social releasers (Barlow, 1963).

The measurement of the reaction of an individual to a color change, or to different color patterns (e.g., model presentation), is necessary to demonstrate the visual signal values of a color pattern.

The correlation between color pattern occurrence and behavior was investigated quantitatively. The methods of

Losey (1968) are followed. The laboratory notes and cine­ matographic data were used. These data were tabulated as to sex, overt behavior in terms of actions, and coloration.

The overt behavior was divided into 6 artificially separated categories. The categories represent major changes in the motivational state of the fish. The choices were based on the motivational analyses of ago­ nistic behavior. Categories 1 through 4 exclude court­ ship situations.

1. Submission: subordination. 100.

2. Mild Aggression: indicated in all species by

movement toward another individual, without

any agonistic display.

3. Aggression: a relatively high A/F motivation

ratio, but not including the higher absolute

attack motivations that can not be attained

in the absence of an aggressive response by

the opponent1 threat.

4. Strong Aggression: a very high A/F motivation

ratio, beyond the attack motivation level of

3 (above), fighting.

5. Courtship: any act increasing the probability

of mating.

6. Exploration: movement around the habitat with­

out social interaction.

The color patterns were listed in the approximate order of decreasing crypticism (rare patterns were omitted).

The frequency distributions of the color patterns were calculated for each of the behavioral categories as cumula­ tive percentages.

The cumulative representation requires that the preceding color pattern frequency be subtracted in order 101.

to arrive at the absolute percentage value of the color pattern for that behavioral category. The cumulative method is more desirable than plotting the absolute values because the cumulative curves rarely erose over one another and are therefore easier to visualize as a group.

Furthermore, the family of cumulative frequency curves that describes the occurrence of a particular coloration in all behavioral categories may be presented in one figure (Fig. 11) eo that differences can be esti­ mated. Families of curves for different species can be easily compared.

The differences between the curves for different behavioral categories are used to indicate the possible signa 1 value of a coloration. For example, the efficiency

of a signal is determined by the uniqueness of its fre­ quency curve. The concept of signal value here refers to the degree that it serves to differentiate between behavioral categories, and thus serves to communicate these to other individuals.

A. Species Descriptions

1. Emblemaria hypacanthus

The motor patterns used in the analysis of color 102.

patterns are as follows:

1. Submission: Flee, Fin Folding, Backwalk.

2. Mild Aggression: as indicated above.

3. Aggression: Fin Erect, Branchiostegi Lower, Gape.

4. Strong Aggression: Mouthfight, Push, Bite, Chase.

5. Courtship: Fin Flicker.

6. Exploration: as indicated above.

The families of cumulative frequency curves (Fig. 11) represent the occurrence of the different color patterns.

Color pattern A, the fewale mottled or cryptic condition, occurs most often in the female courtship category. But this pattern also occurs during submission and exploration and to a lesser degree during mild aggression. The transi­ tion from the cryptic pattern A to the intergrade A/B to the non-cryptic B pattern occurs with a rise in the in­ tensity of male aggression. Pattern B is particularly indicative of strong male aggression as is pattern c for male courtship. The difference in the shepe and position of the curves for male courtship and aggression indicates the possible signal value of the colorations. In summary,

·the coloration of E. hy:pacanthus could serve as an e££:1- cient signal to differentiate between male courtship, m~lfj 103.,

aggression and "female courtship-submission."

2. Acanthemblemaria crockeri

The actions used to indicate the behavioral categories in A. crockeri were the same as those for E. hypacanthus, with the following exceptions:

1. Courtship: Out and in darting movements from

the refuge.

Again, families of frequency distribution curves

(Fig. 11) describe the occurrence of the coloration fea­ tures. Pattern A occurs only in females and is indicative also of female courtship. The male patterns also change to the darker state (B/C and C) with an increase in aggressiveness. But the aggressive coloration in A. crockeri is not distinct form the courtship coloration as it is for E. hypacanthus. This increaees the possible efficiency of a signal of increasing aggressiveness, but does not differentiate courtship from aggression.

3. Neoclinus uninotatus

The motor patterns that were used to indicate the behavioral categories in H• uninotatus were the same as those for ~· hypacanthus, with the following exceptions: 104.

Figure 11

Color pa·l:tern frequencies in Emblemaria ~canthus

and Acanthemblemaria

Shown below are the number of observations:

Behavioral cateqo~y E. hypacanthus A. crockeri

exploratirm 21 17

submission 16 13

mild aggression 15 12

aggression 11 9

strong aggression 14 B

male courtship 13 5

female courtship 4 2 105.

100

!

50 ....

0 -0 w > f-- < _J ::J e) 2 0 I ::J B (.) A NB c

>- (.) Acanthemblem . z 100l 0·· ...... aria crockeri w fem a 1·······e cour\sli ...... IP /~·..:.:~.:_: .. :..::_: ::.: ..:: .. :: .. :.: ~:..: :.:Q:.:..:.:..: ~~'-''-'-""~~· ::J I ~--:::=;·-, 0 I __....~ _. w I / --- ! I _,n@ij)r 7 / / / .... . & ,#s::b~/• / ...

~Ii:y/ / // .:.... / , . ../ . /1''# #./' . / :' / . ;;:/' / ////// i1I .'4- 1 Z ~ 0~;·;~_/A __ /------~--it~~------~~~~ & .. _ ------B c COLOR PATTERN ~C lOGe

(1) Aggression: Gape, Head Arch, Pin Erect.

(2) Strong Aggression: Push, Bite, Chase.

(3) Courtship: Tail Beat.

In N. uninotatus also, families of frequency dis­ tribution curves (Fig. 12) describe the occurrence of color patterns. A rise in aggressive activity is accom­ panied by a shift from the mottled cryptic pattern A to the striped pattern B.

The courtship colorations are different from those of the preceding species. Both sexes of N. uninotatus develop the bright red markings on the dorsal fin, the body and anal fin.

Color pattern A was seen predominantly in exploring or subordinate individuals. 107.

Figure 12

Color pattern frequencies in l~eoclim;s uninotatus shown below are the number of observations:

Behavioral categorv

exploration 27

submission 19

mild aggression 16

aggression 9

strong aggression 7

courtship 3 lOB.

Neoclinus un·lnotatus ~~ 1hmissjoo . ----

..

5 ..

>­ t.) 2 LU :::l 0 LU- It

c

COLOR PATTERN 109,

DISCUSSION AND CONCLUSIONS

The results of the analysis of the temporal pattern­

ing of agonistic behavior suggest that the order of the

action patterns are not as fixed as theory or subjectivity predicts them to be. Barl0\'1 (1968) argues for factors

that might favor variability over stereotypy of action patterns. To support his argument, Bar 1011 uses !·1orris'

(1957) example of "typical intensity." This model re'­

sembles a simple dose-response curve. The "intensity"

of the response is directly proportional to the strength

of the causal factors. Consequently the intensity or

completeness of the behavior reflects the motivation of the sender; the system is said to be rich in information.

Havlever, there immediately arises the possibility of ambiguity because of the gradations between the various degrees of expression of the behavior. This is just the

situation we might expect in fishes that engage in complex

fights (e.g. ?1_. crockeri and E. hypacanthus), requiring that each animal continually and accurately assess the state of its opponent. Barlow also points out that reduction in monotony may be selected for occasionally 110.

when novelty is necessary for arousal and attention

(Berlyne, 1960}. One way in reducing monotony, Barlow suggests, is ·to vary the sequencing of the motor patterns.

The numerous flow diagrams given in the literature (Neil,

1964) indicates the variability in sequencing is much greater tl1an one would have predicted from the original notions of chains of releasers and instincts (Tinbergen,

1951). In conclusion Barlow suggests replacing the term fixed action pattern with modal action pattern (M.A.P.).

Barlow submits, "This conveys the essential features of the phenomenon: there is a spatiotemporal pattern of coordinated movement, and the pattern clusters about some mode, making the behavior recognizable."

Description of agonistic behavior, in terms of moti­ vation, summarizes the evidence and facilitates inter­ specific comparison and inferences as to the possible evolution of behavior. The threshold levels of the actions presented in the A/F diagrams are of course only approxi­ mated.

Primitive agonistic behavior, "all or nothing" attack or flee, is rarely observed in higher animals. Each spe­ cies possesses a threat code that usually precedes all-out 111.

attack. Threat behavior alleviates the damage done to

individuals without decreasing the survival value of aggression. It arises out of a conflict between the motiva­

tions of Flee and Attaclc, and in its most primitive forms

it may consist of the simple superposition of motor pat­

terns simultaneously activated by both (Lorenz, 1964).

The possession of a territory seems more important

in determining the motivations 1 sta,te in Neoclinus unino­ tatus than in the chaenopsid species studied. I have ob­ served a population increase at the ~1onterey wharf of year class 2 and 3 individualo (as determined from otoliths).

This indicates that these year classes are particularly mobile. Hickman (pers. comm.) reports a very similar situation for ~· uninotatus at the Redondo King Harbor.

~nus the actual importance of territory v~ning, in deter­ mining the motivational state of the fish, could serve to reduce aggression among migrant individuals.

The function of the color patterns appear to fall into three categories: accentuation of postural displays, signaling the motivational state of the individual, and effecting reproductive isolation. The darkening of the body and fins and the reduction of crypticism accentuate 112.

the postures of the body and increase the apparent size and background contrast. This serves to increase the stimulus value of ·the displays in intimidating rivals and in attracting a mate. The changes that accompany increas­ ing aggre:;;siveness and courtship in chaenopsid males fall into this category. The over a 11 pallor of the female

Emblemaria hypacanthus courtship coloration might also re­ duce the apparent size.

Certain features of the coloration and sexual dimor­ phism appear to be signal organs. For example, the male

Emblemaria hypacanthus changes from the barred pattern dur­ ing aggression to the solid dark brown color during court­ s11ip. This signals a change in motivation due to presence of a ripe female. The high dorsal sail-fin of the male is accentuated by blue and orange colors during courtship.

This signal organ is used in rapid spreading and folding manner to attract the female. Sexual dimorphism is impor­

·tant when considering courtship behavior because as females come to look more and more distinct from males, males tend initially to respond less aggressively and instead exhibit stronger sexual responses (Johnsgard, 1968: 125).

Neoclinus uninotatus must be considered the least 113.

specialized studied. The species is relatively unspecialized ecologically since it can periodically in­ habit the artificially created sand and trash basins associated with harbors. ~· uninotatus shows extremely little morphological specializations in comparison to the chaenopsids. Sexua 1 dimorphism is generally lacking and the sexes of ~T. uninotatus are much more difficult to sep­ arate. Behaviorally, N. uninotatus shews few specializa- tions except for the l1ighly ritualized and tl:1e distinct aggressive color pattern. The threat code is simple and effective since actual combat was rarely observed. 114.

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Barlow, G.W. 1961. Social behavior of the desert pupfish, Cyprinodon rnacularius, in the field and the aquarium. Amer. Midl. Nat, 65: 339-359

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