Title: Experimental analysis of behavioral isolation among four sibling species of nemobiine crickets /
CITATION THUMBNAILS PAGE IMAGE ZOOMABLE
Full Citation
STANDARD VIEW MARC VIEW
Permanent Link: http://ufdc.ufl.edu/UF00098153/00001
 Material Information
Title: Experimental analysis of behavioral isolation among four sibling species of nemobiine crickets /
Physical Description: xii, 111 leaves : ill. ; 28 cm.
Language: English
Creator: Mays, David Lee, 1941-
Publication Date: 1975
Copyright Date: 1975
 Subjects
Subjects / Keywords: Crickets -- Behavior   ( lcsh )
Entomology and Nematology thesis Ph. D
Dissertations, Academic -- Entomology and Nematology -- UF
Genre: bibliography   ( marcgt )
non-fiction   ( marcgt )
 Notes
Thesis: Thesis (Ph. D.)--University of Florida, 1975.
Bibliography: Bibliography: leaves 102-103.
Statement of Responsibility: by David Lee Mays.
General Note: Typescript.
General Note: Vita.
 Record Information
Bibliographic ID: UF00098153
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: alephbibnum - 000569116
notis - ACZ5875
oclc - 02273916

Full Text












EXPERIMENTAL .ANALYSIS OF
FOUR SIBLING SPECIES


BEHAVIORAL ISO
OF NEMOBIINE


NATION AMONG
CRICKETS


David


Mays


A DISSERTATION
COUNCIL OF THE UNI
FULFILLMENT OF THE
DOCTO


PRE
VER
REQ
R 0


ENTED TO THE GRADUATE
ITY OF FLORIDA IN PARTIAL
IREMENTS FOR THE DEGREE OF
PHILOSOPHY













































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.,- L-O :


'lA


11hc( ], :


]B h -2 : 1


t:Si






















Among


no &; t:


c. m
C^h O Lltxfl--A


i1:ur -lI


singing


Orthoptera


th ouI: Jo)i CI


(1- .


of pet -instutli
--1< -*'-- _.-* , P- I -


Flo:da


somewhat


northward a


areL


. ,,' b .i -.


qrOund


rqi c c
'-J- './jft*'L*h


1iow


kniotn>


-s Pict.Oj mI.bius


ambi ti)..L-


rSc' r}


EBth


Exc


a- c


f iqht.e- L e -,,


possessing


only


forVwincf


case


f Co 1 V


th cCOe


1. j ra


arc


t re3 0: ^^c


c.:: r,


3]st


a "o-.:is al
aP'z o x a 1 -/Ji---- -< I


segOe9 t t


The


Ti~. i r. p
JL' L-^ t- S


, i- Tf-! li CE


dt K-


i1..: u? S !y'


clark


border


v w. q


tri xL,


St.-COated


appearance.


Be th


sexes f .,


have


prnomiii rant:


eves.


!whZ u


-3 --.-? ;- r-^


fr"ot


T.


their


WT 1k 11


head


S;-1d


between


sop-tenmr or


1964


mioorph


, T.


Sh' b.i


olK.og


ambit iouOS


frowp


noted


c- )or ation


d'f fe.en..


found


t hatit


b-ctwECC 1i


ecol oqical


Flori da


differences


population ns


s itua ti ons.


that


-^.t n


calli ] jf_


song


r'l J i


rate


drained

st.dy (

wJ ng st


bet-coen


co11ections


C. '-n


Walt cr

rake
T"7- 1 I ,
\ t '" "{. "


Sjrea :


, 1962)


a L-t


.n .-


in Soi_-L


a tcmpc:rature


one


indiv idual


appro :i.rnma tely


areas


vs.


cricket


weI 1 --


wirngstroke

maintained


(p/s)


tl ani


Otiw-rY $


over


range


^eii'P^4-~ C- /-LLL


tested,


s r t -


ti 9c


CWa-< ]


e1 .


no te-


T: MT' Pfi1 -p3r1j Or\


w i tL


pulses/ S second










Gun Club,


near


the municipal


airport.


"Slow"


ambitious,


with


a calling


song pulse


rate of


at 25C,


was


given


University


of Florida


Depar tmen t


of Entomology


library


specimftenl


recording


code


number


528.


"Fast"


ambitious,


with


a pulse


rate


25C


was


given


code


number


531.


Thes


types


crickets


could


be distinguished


the muni


cipal


airport.


Crickets


pro-


during


weedy


songs


were most


tur key-oak


areas


conunonl v


whereas


found


crickets


the disturbed


producing


songs were


found


surrounding


hammock.


In another


locality


(Goldhead Branch


State


Park,


Clay


Co.)


both


crickets


were


again


collected


in close


n r.
CtL 0 )


ociation.


Field


collected


adult


females


taken


laboratory


isolated


gallon


jars


provisioned


with


food


water


ap proxi mately


inches


autoclaved


sifted


sand


soon


p1odo uced


progeny


that


were


either


100%


100%


531.


Genera lly


females


were


more


black


white


sp- ,ckled


above


anid wb Lte


whi 3.o


were


markncd


softer


tones of


brown


with


J nLters.


nymphs


types


were


distinctively


consistently marked


colored


about


accuracy


separating


them.


nurb er


teeth


and


length


sound


producing


wing


file


varied


between


531.


These data


suggested


4rQ nnA




r.- To /Ir nT. cv. ri


Is\N r


-7 r- -i


Sf /""\ S-


tape


-t


allowing


_____










turkey-oak,


sandhill


area


3/4 mi.


west of Archer,


Florida, on 5 April 1967,


discovered


females


of Picto


nemobius that


were


dark brown with bright


straw


colored


markings


well. 1


the more typical dull 1-brown females of


Pictonemnobius 531.


Their differences


were


quite striking.


One individual female of each


color


type


was


brought to


the laboratory and each gave rise to progeny which were true


and bred true to their parental color type.


Laboratory tape


recordings of the resulting male progeny fell into two

categories corresponding with the two parental female color


types. The

in Florida,


dull-brown type,


known


(previously designated


song pulse rate of


47 p/s at 25C.


from

as 5


many other


31)


had a ca


The new, dark-b


brightly marked P. ambitious had a calling song


localities

11 ing

rown,

approxi -


mately


55 p/s at


5C.


Recordings


field-collected


ambitious


from


Archer locality confirmed the new male


song


type in the


field.


Numerous recordings


h"-d


already


been made under the


531 number for the two types,


the notation 531--1) for


the Dull brown


(47 p/s)


and 531-B for the Brightly marked


p/s)


Type


ambiti


31-B


osus


was


was


instituted.


found to breed true wherever


was


found and could be separated from 531-D and


in central











531-D,


making


this


character


less


dependable


terms


separating


species.


Type


531-B


appeared


to be distinct


from


531-D and


A fourth


anmbi


taosus


song


type.


was


discover


author


February


1969


along


a grassy


road


surrounding


pine


flatwoods,


.3 mi.


east


Clay


line on


Florida


road


The male


calling


song


attracted


attention


because


resembled


t of


Miogryllus


saussurci


(' Scuc1 r
" Li -V!.^^ -- ^ -


(whi ch


was


singing


from


burrows


e same


having


ti: me)


lasting


differed


more


than


from M.


secon11ds


saussui.cxi


A 531-D


male


was


located


same


site


calling


song


provided


a contrasting


reference


newil


song


type,


designated


as Pi ctoneitobius


525.


The


calling


son


pulse


rate


was


about


250C.


Both


ma I e s


females


were darkly


marked,


eas


from


other


three


types.


Early


tis


stud


four


song


type s


represent


biological


entities,


possessing


charac teristics


species


with regard


to reproductive


isolation


ecological


geog


raphical .


distribution


wi 11


refer


them


species


, 531-D,


531--B


f rom


this


point


will


present


proof


their


separateness


appeared


this


distinguishable

















SONG ANALYSIS


Equipment: Recording and Analysis

Tape recordings of male crickets were made


in the


Department of Entomology


noise


room


(LNR)


crickets


were


caged


individually in 4


in.


screen-


cages.


Caged


singers


were


located by using an


ear-


phone monitor,


and recorded with


a dynamic microphone


Scotch 202 magnetic recording tape at 15


inches/sec.


on an


Ampex models


tape


recorder.


The


tape


record er


was


located


an adjoining laboratory and


was s


activated


remote controls from within the LNR.


Some


tape


recorded


sounds


were


analyzed with a Kay


Electric


Company


audio-


spectrograph


(Sona-Graph)


but calibration


procedures


were


leng thy and production


was


slow.


More


often I


used


oscilloscope with


"stora


sc reen.


" Several


different


complete chirps could be displayed


one


time on the


osciilosco


using a


sweep


rate of


sec/cm.


From


tota 1


30 pulses the pu


rat


wa s


determined by the


formula


^- 30


I .-3


sided










where D


-= mm required for


30 pulse display and


the sweep rate


(.05 sec/cm)


Calling song pulse rates


perature


are


the individual cricket


a function of the tern-


therefore can he


corrected to some temperature standard


comparative


purposes given the


slope


of the pulse-rate-vs.-temperature


plot,


Pulse rate


corrections


25C


were made with the


form la


p/s at 25.00C


(25. 0


- T)


2.232


+ p/s


where T


= temperature


time of recording


2.232


slope


of the pulse-rate-vs.-temperature plot


(Walker


1962)


Description of Cal ing Songs


The calling songs of Pictonemobius,


like the calling


songs


of other Gryllidae,


are p


reduced


itary adult


males and attract sexually


responsive


females.


songs


have been shown to be


species


speci fic with


respect


to the


females that are attracted


(Walker


1957


, Alexander


1967


Hill


et al.


1972,


Ulagaraj


1974).


On numerous


occasions


or more of the four Picto-


nemobius


species


have


been heard calling in the


same


area.


In such


areas


individuals with intermediate


sonCs


have not










pulses


of sound


("chirps")


as opposed


trilling


crickets


which


produce


long


sec.


seFries


of pulses


("thrills")


Each


pulse


created


a single


clos ie C


upheld


mesothoracic


wings,


during which


file


(ventral.,


right


wing)


in contact


with


scraper


(dorsal,


left


wing)


The


ses


are


discrete


bursts


sound


havLrng


fundamenLal


oscillations


5,000-7,000


range.


These


fundamental


oscillations


correspond


with


strikes


file


teeth


scraper.


Species


long


chirp


, appro.x.yimte..
\^- ^1~ J YU- n- .tEIN7-


sec.


in duration,


with


a pulse


ra te


at 2


(n=21)


municipal


airport,


Gai,-esville,


Florida.


0O -.il graphic


traces


of a 528


calling


song


are


in i.1


A.


10. O/


chirp


makes


pulse


rate


sound


even,


slowe r


listener,


compared


with


Species


531-D


a chirp


interm-d i iatr


length,


to 0


sec.,


with


a pulse


rate of


47.0


(n=9)


from


the municipal


airtportj
T) -


Gaime Jvi


l e,


Flor ida -


A complete


cbhirp,


pulses


, an d


spaced


chirps


are


shown


Fig.


Species


531-B


short


chir


! .. fl


S -e .


length,


with


a pulse


rate


54.9


(1n-21)


from


west


of Archer,


Florida


Some


individual


ctlisi


stently


yn\ ry ,-r A -i a


- ---_ - : -


h an


1, less


I 1


long


regul.arly


^ ,^ ^


I


*


fw J h ^ -


II


'I












































































































.. Calling


song
r- ^ r


S
- - -


4 sp
* 1


>ec


of Pi


- a .


ctonemobius


^-


..lla


% rl


-_-r-


f 'i


"*


A AII










Species


long


chirp,


long


longer


than


528,


usually


sec.


long,


fastest


Pictonemobius


pulse!


rate,


59.9


250C


(n=8)


.3 mi.


east


Clay


line


on SRI 6.


trend:


of decreasin


g pulse


rate


with


increase


chirp


length


from


31-B


makes


song


quite


unusual


to hear.


Adding


distinctness


a slow build-up


intensity


early


part


chirp


(Fig.


ID) .


PictoneInohius


song


individual


variation


exists


the number


of chirps


unit


time,


intensity


changes within


chirp,


the maximum


intensity


chirp.


These


differences


rem ra in


consistent


throughout


sample of


individual' s


calling


song.


Several


laboratory-


reared


pulses


531-D


within


were


a chirp


found


recorded


to have


shortly


irregularly


after


spaced


their


maturation


adult


stage.


field-collected


laboratory-reared


ec_, in e- 11 -


more


than


one


week


were


found


to be


irregul ar


this


mcinner. t


geographic


distribution


corn


pi nation


cal in n


song


pulse


rates


presented


later


this


section


will


serve


define


four


species


of Pi c tonemobius.


Description


of Cour


ship Songs


acoustical


s iqrials


associated


with mating


are











prin


cipal


courtship


song-


(with


or without


spermnatophore


characteristic cal


a long


seri s


soft


chirps


(often


irregularly


spaced)


in which


f irsi


pulses


pulse


are


rate,


produced


followed


at almost


exactly


resumption


calling


of the


typical


song


raLte,


increase


intensity


(Fig.


When


a male


loses


contact


with


female


court-


he often


produces


loud


extended


pulsed


chirp


Fig.


usually


beginning with


court tship-type


wide-spaced


pulses.


Prior


"backing


under"


both


"brief


spermatophoreless


mounting"


" lengthy


mountinigt


[sperma tophore ]


produced,


transfer"


entirely


(Mays


calling


1971)


a long


SOn l


trill


rate


(Fig.


Courtship chJ rps


are


than


calling


song


chirps.


Courtship


interruption


chirps


backing


under


trills


531-D


531--B


are


than


their


call in


song


chirps.


untrained


listener


would


likely


find


it difficult


to d.sLinquish


courtship


chirps of


four


species


easy


to di sti anguish


courtship


chirps


from


calling


chirps.


Geographic Dstribu tion


Crickets


orinciiallv


belonging


in Florida


genus


. ,,<.. . . .


Pictonemobi


immed i 5te1 v


occur


.n1 *n


5-10


hiuchh


Singer


shorter


, --\-i t
a mJ












































I(f *
r- T .Q
C f-H 0)
.0 Mf A


a C










Cleburne Co., Alabama


I have been unsuccessful in locating


Pictonemobius


west


the Florida panhandle, and north


southern Alabama and central Georgia.


Collections and laboratory tape


record nrgs


of more than


700 Pictonemobius from


over


1.00 Florida localities fall into


the four


categories


of Pictoremobius


discovered


in the


Alachua


area.


The identity of field-collected specimens


was determined by


analysis


laboratory recordings of male


calling songs,


habitat specificity,


color


patterns.


areas


distribution overlap do not


indicate intimate sympathy.


Localities in which


necessarily


such


sympathy


was


evidence


are


listed


(Table 1)


and illustrated


(Figs.


& 6).


The ana


lysis


of calling songs from several


areas


sympathy defined the


species


numerous times.


Marler


(1962)


states "Judgments of degrees


species


speci ficity


are


relative and


arise


from


compari sons


between


signals


under


consideration


other


signals


wh ich


are


likely


to be


present at the


same


t ine a nd place"


After


several


areas of sympatry


were


understood,


widely separated popula-


tions


each of the four kinds could be related


over


a wide


geographic


area


including most


Florida.


A detailed


analysis of allopatric variation in


Pi Ctonemob i us


is not


I _


u
























































































































v r


I 0'




























































































Ob C


'a'






















0


U,
CC
o
rto
in
(Q
JiJ
to t
tv M
(N r


i0 r(


.4- +- 4-


I 0%


0 (T


r- in


tn W


CT f- *
CJ .Q "**


'- .




C3 0 -

+ac
so OO
r^ kW

m-oo

+NCU






dUO4
#1 0 0


O C .C
orm >.t,






U r ld













Notes to Table 3


Alachua Co.


Municipal
R.20E., 'iT


airpo t,


. 9S.


Monteocha
Fairbanks


"Gun


Gaines vis Ie


Club"


10 rm.


(T J.


north


(NE.


Walker


of Sec. 24,
collections)


of Gainesville,


east


Gainesville


sever


inc.


39th


Ave.


15th


Mill Ihopp


Archer
Plan L


Di vi


block


Industry


of F.


campus


Alachua-Levy


1 i]_ c


Sr. 24


mi.


west


of Archer,


scou t


SR.24


cast


S2nOA0


from


jct..


w ith


US. 301


Baker Co.


.1 mi.


south
west


GeorTij. a


scate


line


SR 121


from


Bay Co.


Panama


City


Leac"h,


city


limits,


east


Highland


Clay Co.


Goldhead


Starke


Branch


ea st of
Country


State


Cl] a
Club


Park


line on


Walker


SR 16


collections)


vicinity


Collier Co.


sou-th


jct..


jct .


-1 r- I


i 4


I ___


vicini .











Flagler Co.


South


of Marineland


Franklin Co.


Carrabelle,


nr.


cemetery,


north


side


town


Gilchrjst Co.


Jenny


Springs


Hernondo Co.


south


Co.
Archbol d


Croom west


biolocrical


1-75


Station


Venus


Jefferson Co.


.5 mi.


north


on SR.


LaPayette Co.


north


Dixie


line,


S357


Lake Co.


LTesburg


city


limits ,


south


US.27


Leon Co.


Ta 11


Ti;ues


Research


Stai.tion/,


north


Lake


lamoni a


Levy Co.


west


of Otter


Creek,


then


4.5 mi.


nor th


north Bronson,


flatwoods


1.5 mi.


north


Ma r i on-Levy


line


on US.41


Manatee


Springs


"18.












Mar ion Co.


.7 mi.


north of


Sparr


Ocala,


city


limits


& S336


US.27


& 441


at Orangeblossom Hills


S484 at Dunellon


jet.


S200


S 484;


east


jct.


S200


S484


Pa Im Beach Co.


Jupiter


PuLnam Co.
-- -- -_- _


easi-


Me 1 rose


S iinole Co.


Sanford, nr.
northwest of


25th St.


"F or t-


Grand view,


1/2 mi.


Reed"


Sumter Co.


SumL to -Lake


line


SR.50


Tavlor Co.


Blue Sp-ings 1 mi.


north


of Cedar


Island


Forest


Capi tal


Statc


Park


south!


of Perry


Wackulla Co.


1/2 mr.


west


of jct.


SR.365 on


SR. 98


on "old"












Brooks Co.


Quitman


Decatur Co.


West


of Climax


_ _





















35.7


41.0


36.7


'. 38.3

38,3 1

35.9


37.5'

37.2k

38,5


35.9


33.1


S, .**' -


Fig. 3.
T-Tnn/ nr-^i ni-1nA-


Distribution o
1 nn~ml 4a1 4-,, ntw^ -


Pictonemobius


n -S. f a-


.L. -- I-----------


528
__ .1 -I


in Florida.


















47.3


46.5


43.7


45.8


46.7


46,3


47.2


46.4


48,7


e1.0***




























52.8


" 55,4


55,4-


54,9'

46.6"


53.3


51.2


, *0*


Fig. 5.


a.


distribution o


f Pictonemobius


531-B
*


in Florida.
a


52.9


J






















59.9


60.4


a 50,6


, .'.* '*"
















ECOLOGICAL DISTRIBUTION


General


Habitat


Crickets


genus


Picton


"ius


are


found


from


dry


sand dunes and


central


ridge


areas


margins


swamps


tidal


salt marshes.


They


are


rarely


found


in mature,


dense hammocks with


a closed


canopy where


fire


been


ex-


eluded


years


or more.


Most of


Florida


is maintained


fire


sub-climax


with


additional


removal.


of pine


from


the central


sand


ridge


area,


clearing


agriculture,


creation


of pasture


ran


ge grazing


land


road


dredge


fill.


these


disturbed


areas


found


largest


populations


of Pictonemobius.


Alexander


and


Thomas


(1959


their


studies. s


three


sibling


soecir es


the Allonrn.ob


.US


fasciatus


group


found


that


three


were


nmijxed


with


indication


of interbrcedi ng


areas


Vw;her


artif


icia


ci earina


taken


place.


Similar


occurrences


with


Picto-


nemrnobi. us


are


discussed


later


in this


chapter.


The


effect


burni


size


of Pictonemobiu s


poDulations


was


evident


in comipar


unburr


annually


biannually


burn ed


plots


Ta ll


Ti. be .rs


Resear ch


Station


(TTRS)


rnA c' t


T -.


-i


11 C, I 1-- I r -. .-. I


- f E .


T r f..-


*m /, -' --i


__


,-^lr^ J -


i










A plot


whi ch


fire excluded


past


years


not contain


Pic Lonemobius.


They


ware


scarce


from


mar-


gins


absent


from within


"woodyard


hammock k


which


appears


lightning


to have


fires


been


and


und isturb ed


a record


except


logging


localized


northeast


Gum Pond


area


1945.


A 26 January


1913


account


from


Beadel

tall p


Diary


ine,


records


live oak,


the "w

holly,


Joodyard"


magnolia,


being


etc.


"thick


" (Betty


wood,

Ashler,


Historian,


TTRS, Personal]


Communication, 10


March


1973) .


Habitat descriptions


four


song


types


Pictonemobius


follow the


classification


Laessle


(1942)


The


531--D


Habitat


Species


531-D


is most


COL'lmo1n


in xeric


hammocks


mesic and hydric

thereby opening


hammocks

the canopy


that

and


are


burned


thinning


or disturbed,


understory.


Many


of Florida's


ruderal


commruni ties


such


as old


fields


fire


lanes,


roadside


ditches,


sand


lC)mounds,


fall


into


this


category making


531-D


most


commonly


encountered


Pictonemobius.


Species


531-D occurs


ii ammock


communities


over most


of mid


-central


Florida


even


into


"the


sandhills"


longleaf-pine


turkey-oak


(Pinus


palustris


Quercus


laevis


association,


predominantly


beneath


live-oak


virginiana)


They


appear


in xeric


hammock


sandhill


lawns


. .











(Vitis


Virgini a


crceperr


Parthenoci.ssus


quinquefolia),


saw palmetto


(Serenoa


J__-s1r -_,' r


persirunon


Diospyros


virginiana),


are


prcsenti


an areas


where


large


populations


531-D are


encountered.


In Alachua


surrounding


counties,


gum


they


(Liquidambar


persimmon,


hackbe


abounded


under


styracifl uaa

rry (Celtis


several


species


hawthorn


of oaks,


(Crategus


nississippiensi.s)


and


sweet


sp.)


wax myrtle


(Cerothamnus


ceriferus)


growing


solitarily


small


clusters


in pasture


s and


along


g assy


roads


ides.


They


are


also common


in partially


shaded


lawns


with


soime


leaf


litter


in and


around


clumps
c].u_,4-


fallen


Spanish


moss


(Tillandsia


usneoides)


the well-drained


portions


of weedy-grassy


roadsides


grown-over


road


heaps.


typical


531-D


hammock


habitat,


kept


open


grazing


cattle,


shown


Fig.


The 531-B Habitat


Species


531-B


is associated


with


"scrub


" i.e.


sand-pine


Pinus


clausa-Qutrcus


spp.


association


longleaf-pine


turkey-oak


associa


tion


wi. th


some


tran-


sition


into


]onglea*-pine


bh uejack-oak


(Pinus


palustris-


Quercus


cinerca)


association.


In most


tnese.


situ:.t tons


rosemary


ericoides)


, sEnd-pine


(Pinus clausa)
I---


"vPr TV


.-4A A -1l- '' J A IUA ^.* > f- ..J.- \ V- ^- <' / t -J A .t *- -


nrhrr r


nlnnl


(Geobalanus


oblancifolius.


r ITm mon


(Ceratiola


are r


sp.),








27














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years


ago.


East


west


coast


531-B


occur on


well-drained


dunes,


shell


mounds,


and


dry


sandy


hills


that.


contain


live-oak


Quercus


virginiana


var. .


geminata)


sandhill


plant


associations.


The more


northern


sandhills


are


more


heavily


forested


less


likely


to have


sandpine,


rosemary,


and


cricket


531-B.


531-B


habitat


a xeric


, open


habitat,


occupying


highest and


"scrubby


best drained


1latwoods"


chapmanii)


asso


soils


(Quercus


var.


which


entire


state.


gemninata


occurs


myrtifolia,

n places


I


where


general


there


level


is a slight


of the


rise


flatwoods"


one or


(Laessle


feet ab


1942)


ove


usually


sufficiently


open


or well


drained


531-B.


Species


-D and


531-B


were


plentiful


an area


where


hammock


woods


adjoin


a turkey-oak


sandhill, 3/4


west


Archer,


528 with


north


Florida


Figs.


531-B occurred

Bronson, Levy


S A


site


Co.,


Florida


similar


distribution


5 mi. west

(Figs. 10


The


Habitat


Specie


s 528


does


covers

occurs


much

in


same geographic


the more disturbed,


area

weedy


recently

spottily


burned


alonic


sections


field


of a site.


martin.


- ... -----


Commonly


a road margqi


occurs


the center


citation


isolates


- Q.


r


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z


S. ,










males


(Fig.


The


opposite


situation,


where


531-B


surrounded by


occurred


center


of a large


weedy


field


(Fig.


number s


have


been


found


recently


cultivated


fields


weedy,


grassy


pastures,


and


in young pine

now border, 1


plantations?


ongleaf-pine


which

and t


were


formerly part


urkey-oak


woods


and


containing


531-B


as shown


in Fig


northern


associated


Florida


with disturbed


p-in


counties

e forests


528 appears

that contai


closely

n some


turkey-oak.


Here


they


often


border


531-D


areas


with


predominating


underbrush


higher


fire


sandhills


or mechanical m


that are


cans.


kept


clear


A detailed


study


of a 528


, 531-D


habitat


presented


later


this


paper.


Habi tat


Species


paulustris


COcurs


association,


slash-pine


and


flatwoods


black-pine


Pinus


fetterbush


flatwoods


(Pi nus


seroti na


- Desmothamnus)


association.


These


assoc


river


ceriferus)


Ilex


atiorns


flood


wir


coriace .)


often


plains

e grass


and

(A


blackberry


border

contain

ristida


(Rubu


bayheads


or extend


wax myrtle

spiciformis)

s betulifolil


outward


(Cerothamnus

, gall-berry


sundew


(Drosera


capil laris


usually with


-A


Spec


packed


dark


sites

colored


have


rich


soils


pine needles,


mossy


_11_ _










herbage


often


grow


these depressions.


Alachua


sites


have


include


been


northeast


opened by


Gainesvil e' s


occas onal


wet


burning


flatwoods


clearing.


that


Simi-


sites


of south


occur


Georgia.


northward


The.


into


river


Okeefenokee


flood


Swamp


plains


region


ground


situations


where


further


Florida


occurs.


northward


panhandle


possible


throughout


are


that


pine


additional


species


flatwoods


areas


occurs


of Georgia


since


this


habitat


fragmen ted


in North


Florida


into


east


west


coast


areas


separated


central


sand


ridge.


528,


53J -D


Habitat, a Case_ Study


temrporal


spatial.


distribution


531-D


was


intensively


examined


.n an


area


near


the Gainesville


municipal


airport


of Sec.


R20E,


in August


1971)


study


area.


was


selected


because


contained


both


531-D


over


an ecologically


diverse


area.


Components


xeric,


mros


, andc


hydric


hamrmock


communities


plus


long leaf-pine


turkey


longJleaf--pine


and blue-


jack-oak


communities,


both.


disturbed


undisturbed,


occur


together


this


study


arac


The distribution


principal


types


vegetation


are


shoun


in Fig.


woods


area


was


a mixture of


turkey


'-oak


longleaf-pine


with


bluejack-oak,


water-oak,


wild


cherry


one0


zone.


y--a ,

























































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* af r tin.
<3 .- **. *.r..

S
i
*e
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C


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S**


- S
*
S
*5
- -


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*
* S
5 0
*~*S I
S
S.. *~g.. *5
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.S
*

S..
99 -*


.

*
*
S


** 2



-* -


a-o
0- Q

0 Cn
-I 0


I



I
I
"l



t*

'^3
8S


<1o


,w


***,.:










sumac


is overgrown


with


grape


(Vitis)


and


virginia


creeper


(Parthenicissus


quinque fol i.a)


area


where


tree


s and


brush had


been


close


ground


level


appeared


to have


been


periodically


treated


this


a U.S.


manner


Government


Jame s


Lloyd.


at least


(1955-1960)


The most


last


aerial


recent


years


photo


judging


loaned


clearing


was


from


to me by


about


months


beforehand


and resulted


in about


are


ground.


remainder


water-oak


was


brush,


littered


palmetto


with


fronds,


dead,


turkey-oak


and numerouS


heaps


bottles,


paper,


cans,


and


other


debris.


Daytime


high


temperatures


in the


shade


ft.


above


the ground


were


23rd


2C on


24th,


, 21 and


about


22 August


270C


5th.


Niqhttime


low temperature


s were


20-22C.


The midday


low


relative


humidity


was


-30%


first


days,


but


creased


50-60%


remaining


days.


Methods


irregular


plot


approximately


one


acre


was


mapped


with


respect


the major


physical


features and


vegetation


(Fig.


12 A)


Hourly


surveys


were


made


ong


a predetermined


route


(Figs.


August


over much


p.m.


25 August


period


1971.


from 1:00


Sampling


p.m.


continued


* * S a


-1 *


, Smilax,


I


A










1624


locations


individual


singers


(528


531-D)


were


recorded


on separate


hourly maps.


S ong Per i od i c ity


Song production


was diurnal


bimodal.


number


calling males


increased


during


early portion


dropped


to about


0.2X


(528)


0.5Xv


(531-D)


the morning peak


y midday.


nurmbe r


of calling males


increased


agaa n


afternoon


(Fig.


dark


numbers


of calling males


of both


31-D


had


again


decreased.


This


bimodal


peak


number


calling


males


agrees


closely with


isolated


males


plot


under


calling


conditions


song


produced


constant


tempera-


ture


controlled


photoperiod


laboratory


(Fig.


Similar


singing


occurs


on sunny


days


(Figs.


A-C) ,


on cl


ou.cy


(Fig


D-F)


under


uniform


diurnal


light


intensity


laboratory


(Fig.


sug-


gestingc


internally


controlled


diurnal


cycle


song


production.


Alexander


(1960)


states


"Light


intensity


seems


to be


mo s t


import an t


single


factor


determining the


exactU


time on


each


day when


different


species


begin


song "


42)


This


seems


appropriate


nocturnal


crepuscular


singing


species,


does not


offer


suggestion


to how


Pictonemobius


spp.


"anticipate"


1 S


S *-


& .. LI 4. -.4 4---'- A-- --- ~ -1---- L


bimodality


universally


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Spatial Distribution


Calling males


of Pictonemobius


were


heard


in both


open


areas


wooded


areas -


Male


s of


in open


areas


decreased


their


calling


the approach


of midday more


than


compared


with


those


wooded


areas


which


decreased


by only


(Figs.


& E)


There


was


indication


any movement


individual


to or


males


from


woods


females


were


this


observed


time,


though


moving


just


after


rain,


on moist


ground.


Calling males of Pictonemobius


the woods,

areas. Th-

(Figs. 15


gers


following


cloudy


with


only


ie distribution


& C)


n cloudy

except


days


the open


The


(Fig.


approximate'


days,


on:


(Figs.


531-D


heard,


singers

d the d

5 D, E,


the morning


15 F)


there


appeared


were

locate


sunn


istribut


wi


after


to be mo


areas.


like] ihood


number


highest


occurs

are low

heard a


of hearing


calling


proportion


at night


yet


night


when

the

7 in


courtshi p


songs


courtship

relative n


relative


sampling


songs


given


songs


umbers


umbers of

periods)


tim


to ca

of ca


courts


approx


usually in

d in open

y days,

ion of

th the

oon on the

re singers



is independent

e. For 528,

lling songs

lling songs

hip songs

inmates





















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- -


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L0


OS .


-- a-


o o 0o 00ooi


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'C

I c.
1ii
1_


a l:-.o
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*
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may have gone

slow pulsed c


field


unnoticed


ourtship


than


since


shorter


song


courtship


song


more-

of 5


[than

easily


cal

re


31-D.


Pulse Rate Overlap in Callina Songs


The

from one

calling


four sp

another

songs.


initially


detects t

involved,

singers h


ecies

in t

When


of Pictonemobius


he

one


difficult

differences


short


ieard.


species-specific


between

tected.


open

It


Once

song


sunny


this


to differences


prompted my


rates be

should o

example,

males.

made mea


var


twee

ccur


the


field

first


are


basis


arrives


identify the


between


time


m


at a new sit


singers


songs


can


listener


differences


shady

e in


in microhabitat


postulating

n species s


the existence


singing


heterospecific


,


attraction


find


isurements


iety


crickets


of cool


such


identify


"tuned


variation


sites


can


individual

temperature


together.

attraction


531-D

overla


on clear winter


ambient

located.


temperatures

Recordings


all

in"


in pul

often


pulse


thi


overlapping


If

could]


females


days


such


ov'


occur


to warm


pulse

in ho


at ground


rate


level


temperatures


f-

wa

a


42



ling]

cognized







ishable

ale

e it may

soon one

ecies

he

the

se rate

be de-

rates,

at

g pulse

erlap

--for

28 singing

occurs, I

finding

here

t various


sites


difference


g


distingu


O<


t


_


___



















































































































































































































































































































H Hm











Pulse


overlap


was


iinves ticated


three different


sites


Pictonemobl us


species.


were


known


to occur


sympatrically.


Calling


songs


C. l--r- J-


ind] ividuals


slowestL


pul se


rate


s ec-' C 1-


slowest


individuals


fastest


corded


pulse


to establish


rate


whether


were


pulse


sought


rate overlap


out


exis ts


un der


these


conditions


area


approximately


square


yards.


studv


si te


was


orti


transitional.


habitat


airport,


used


son.g


Gainesville.


iodic


A cool,


study


clear


wJ th


te'm


municipal


peratures


15--200C


shade


30-350C


under


leaves


full


sun-


light


was


selected,


Songs


individuals


531-D


were


recorded


over


1-hour per


iod -


Whenev er


possible


specimens


were


captured


andi


location


ecC


h singer


noted.


Du r :i ng


survey


period


temp ecratre. Te


ground


funll


shadee


went


from


17.0


while


temperature


under


a leaf


ful 1


sunliqhtt


remained


.00C,


results


are


g i.ven!


in Table


A simi.


2-hour


study


was


conduct ted


a site


nta r


Croom,


Hernando Co.,


Florida,.


with


531-B


(n=9)


(n=30


that


were


found


together


in a disturb


partial


logged-over ,


longleaf-pine


area.


irl- *I --1 I


*


re-


wee(oy


race


Sere


species


1 -


trke'-uc.k


I.


m i^/^


_ I


-1


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study


period.


The


temperature


under


a leaf


full


sunlight


11:45


a.m


was


41.60C but


dropped


p.m.


shadows


from nearby


trees


shaded


results


are given


Table


A similar


3-hour


was


conducted


with


531-D


(n=16)


531-B


(n=21)


at a


site


west


of Archer,


Florida,


in an


ecotone


between


longleaf-pine


turkey-oak


sandhill


live-oak hammock


(Table


the Archer


site one


case


pulse


rate


overlap


occurred


between


specimens


identified by


chirp


length


531-B and


531-D.


This


531-D male


was


singing


open


a pul


rate of


66.6


which


was


faster


than


four


slowest


531-Bs


that hour.


There


is no correlation


between


pulse


rate and


degree


shading


where


song


was


heard:


however,


mos t


specimens


531-D


were


in more


shaded


situations


than


those


531-B.


assuming


temperature


linear


(Walker


relationship


1962)


between


using


pulse


the mean


rate


pulse


rate of


same


conspecific


locality


laboratory recorded


a reference,


individuals


temperature


from


of each


going


mal e


was


approximated.


The


results


indicate


that


calling males


are


areas


where


temperatures


are


25-35C


even


though


a wider


range


temperatures


are


available


r I I - 1 1 i - I-


invest nation







47


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(Table


may


actually


less


than


given.


Furthermore,


ecies


with


fastest


pulse


rate


appeared


to be


singing


the highest


temperature


at each


three


localities


studied.


This


would


tend


accentuate


calling


song


differences


that already


exist between


song


types


at each


three


study


sites.


primary


conclusion


from


this


study


that


calling


song pulse


rate


overlap


areas


sympathy


extremely


rare.


Coldhardiness


Study


life


stages


of Pictonemobius


have


been


found


during


seasons


the year,


during winter months


at drier


times


reasons


early


early


juveniles


juveniles


appear


become


scarce.


scarce


Possible


include


lack


female


wintering)


oviposition,


behavioral


delayed


adaptation


development


(i.e.


(egg


burrowing)


over-


or the


eggs


juveniles


being


unable


to survive


periods


severe


cold


or drought.


Laboratory


vs.


Field


531-D Coldhardiness


Test


Pictonemobius


spp.


breed


continuously


, all


stages


being


present


throughout


year,


even


in areas


of northern


Florida


where


weather


service winter


temperatures


below


-8C


occur


yearly.


Pictonemob ius


531-D


survives


winters


-I r 0 Xl IV fl rN r fl


fl Tk \ ,.-^ 4--


4- ,-v1nV,- 4 -% t.. r *


+-1- 0 F'I -1 l0CT


--* Y


~UI1










colonies


same


species,


which


life


stages


were


best adapted


any,


following


study


was


conducted.


From both


laboratory-rear


ed colonies


recent


field


col-


elections


(April),


individuals


five


classes--small


juveniles,


(large)


juvenile


males,


(lar


juvenile


females,


adult males,


and adult


females--were


placed


in rearing


jars


put


in an environmental


chamber


to be


subjected


series


successively


colder


night


temperatures,


approxi-


mately


10 hours


each


night,


until


crickets


had


died.


Each midday,


at approximately


20C


, mortality


counts


were


made.


In general


laboratory-reared


531-D


suffered


more


mortality


beginning


-40C


than


field-collected


individuals


(Table


appears


that


small


juveniles were


least


tolerant


temperatures.


This


experiment


was


in part


repeated


with


10 male


female


nymphs


from


same


field


laboratory


sources


days


later.


single


10-hour


cold


period


-80C was


given


first


night


to eliminate


effects


starvation


dessication


which


were


suspected


to have developed


previous


experiment.


test


crickets


were dead


the day


following


treatment


suggesting


that


a single


-hour


exposure


to -8C was


suf-


ficient


to kill


531-D collected


field.


Under


natural













































































)041
4m>0 -











528,


May


531-D,


1971,


531-B,


varying


were


numbers of


investigated.


field--collected


In early


individuals


each


species were


divided


into adult


male,


adult


female,


juvenile


groups,


placed


into


roaring


ars


containing


sand


that was moistened


on one


side


, and


a small


piece


of paper


toweling.


jars


were


placed


the environ-


mental


chamber,


which


was


to provide


midday


high


temperatures of 7 0C


followed by


successively


lower


night


time


temperatures.


successive


nights


temperature


declined


was


held


four


hours at


-80C.


Census ing


took


place during


following midday warm-up


periods.


The


stepwise decline


was


give


greatest


chance


acclimation


occur


as well


to provide more


than


one


period


where


temperature


might


enough


cause


mortality.


temperature


towel-sand


interface,


where most


crickets


congregated,


was


monitored


during


most


cooling-down


and


warming-uo


periods


showed


a 1-


to 2-hour


reaching


same


temperature


ambient


the environmental


chamber.


A pretest


census


census


following


first


cold


night


(+30C


were


identical;


no crickets


died.


The


results


successively


colder


ni.ahts


on adult


males,







54








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levels


one


after


adult


exposure


female


to -30C.


remained


Only


after


one


juvenile


an exposure


-80C.


additional


treatment


-80C


killed


remaining


crickets.


During


these


studies


laboratory


rearing


Pictonemobius


continued


normally.


Crickets


controls


these experiments


were


not


always


available,


but


surplus


crickets


that


were


collected


reared,


lived


weeks


under


laboratory


conditions.
















PHONOTAXTS


TO CALLING


SONGS


Introducti on


Species


specific


"calling


songs"


"pair


formation


songs"


are


produ


by male


Pictonemobius


(and most


other


crickets

females


female


as well) t

for mating.


toward


0o attract sexually

Phonotaxis is th


sound.


responsive

directed


strength


conspecific


movement


phonotaxi s


that


female


shows


toward


source


of a male


calling


song


or test


signal


difficult


to measure.


development


a consistent


reliable method


of quantifying


female


phonotaxis


to precede


attempts


evaluate


specifi-


city


of phonot


actic


response.


deve lopmen t


such


method


a maj


part


this


section.


assay


531-B,


virgin


their


female


response


Pictonemob i.us


to a wide


528,


range of


531-D,


natural


synthetic


sounds


further


defined


those


Pictonemobius


species.


Methods


Remote Event


Recording


EIqui. ment


Recordings


song


cycles


movement


crickets










Alton


Electronics


following


Company


transducers


were


Sensitive


used


Relay

trigger


(SR).

the


The

relay.


Micro


phone transducers:


A n um ber r


inexpensive


crystal


microphone


, such


The ro


Test


TRM-1A


"tape


recorder


microphone,


or dynamic microphones


such


Astatic


Corp.


model


level


DN-HZ


such


studiomike


a cricket


were


used.


singing


increase


caused


relay


sound


trigger.


Cadmium disulphide photoelectric cell:


This


photocell


was


used


to record


exac


t inme


that


LNR lights


were


switched


on or off.


Phonocartriidge:


A "sensitive


platform"


was


constructed


from nylon me


strung


tightly


across one end


inch hoop


and held


tight


small


spring


diameter


metal


pulling


disc,


downward


(Fig.


agci nst


The


a central


needle of


inch


an Astatic


type

disc


414-2


phonocartridge


Vibrations


applied


rested


lightly


anywhere on


this


platform


central

m were


transmitted


phonocartridge


resulted


triggering


relay-event recorder


system.


relay


could


acti-


vated


slight


stroking


a human


hair.


Whenever


crickets


walked


onto


or moved


slightly


while


upon


plat-


form,


relay


was


activated


several


times.


preserve


con


Lac ts


to eliminate


unnecessary


movement t


s of


event


recorder,


capacitors


I













































































































































































































































r










closed


longer


than


pulse


interval,


thereby


remaining


closed


for the entire


duration


of a chirp.


Further


capaci-


tance was often


applied


cause


relay


contacts


remain


closed during


chirp


intervals,


causing


recorder


remain


"on"


position


until.


singing


had


stopped.


Depending


transducer


signal,


various


resistors


with


values


100,000


range


were


applied


across


relay


input


to locate


relay


sensitivity


controls


some-


where


near


their mid


range.


Both


capacitors


and


resistors


effect


of preventing


spurious


noise


from


being


event on


ER chart


averaging


discontinuous


signa


into


continuous


positive


responses.


Photo-


cell required

output whereas


a 100

most


resistor


microphones


input

were


and

used


with


76,000


across


input


output


respectively,


Synthesis of Artificial Test Signals


Cricket

single


songs ge

frequency


nerally


between


ave

3,00


their

0 and


energy

8,000


concentrated


and


consequently


sounds


they


produce


are


often musical


"bell


like"


human


listener.


This


single


frequency


result


rapid


wing vibration


developed


wing


file


teeth


are


driven


over


scraper


during


a wing


-S










7,000


This


tone was


switched


a General


Radio

trill


type

with


1396--A T

an on-of


'one--burst


f ratio


Generator

1:2. Th


(TG)


to create


e AO output


was


monitored


a Monsanto model


100B


Counter


Timer which


gave


visual


readout


the AO


frequency


intervals.


A sine wave generator


used


(SWG)


vary


was


pulse


used


timing


rate.


inout


signal


thus


produced had


on-off


swi thing


noise


with


a harsh


on-off


contrast.


This


type


of signal


was


used


once


test


females


resp


onise


synthetic


signals


Further


refinements


involved


replacing


TG with


Alton


Electronics


Company


Field Effect


Transistor


Gate


Signal


Switch


(FET)


which


switched


7,000


AO signal


on and


off.


An externally


appi


drive


signal


produced


an Alton


switching


Electronics


rate.


Company


timer


rise-time


determined


fall-off


the FET


each


pulse


wa s


modified bv


second


drive


apple


ication


input


FET.


a .04


This


capacitor


equipment


across


made


possible


to shape


beginning


each


pulse


(Fig.


An example


an artificial


signal


illustrated


in Fi


D along with


a natural


pulse


(Fig.


19 C)


TG was


used


to chop


this


signal


into


chirps


various


pulses


long.


output


from


19).


numb






























END AND BEGINNING OF .5 MSEC/DIV
"SQUARE CUTr PULSE,
SYNTHETIC SIGNAL. 7000 CPS.


PAIR OF NATURAL PULSES.


5 MSEC/DIV


BEGINNING AND END OF .5 MSEC/DIV
"ROUND CUT' PULSE,
SYNTHETIC SIGNAL. 7000 CPS.


PAIR OF SYNTHETIC PULSES.


5 MSEC/DIV


7000 CP














COUNTER


(Mon


TIMER


santo model


TONE BURST
GENERATOR


(Gener
type


TIMING


Radio


1396-A)


INPUT


TIMER


(Alton


Electronics


Company


TAPE


RECORDER


(Ampex model


351)


100B)


1-IN
68f
/ -OUT

DRIVE
/I


WIDE


RANGE


(Hewlett


OSCILLATOR


Packard


-7


201C)


.004uf


47Th


OSCILLOSCOPE


(Tektronix


EXTERNAL.


type


564)


TRIGGER


AUDIO MONITOR


(Alton


Electronics


Company)


SPEAKER


(Sphericon


model


T-202)


Fig.
cun+^-i-hnF r


Schematic


equipment


used


in production


nrv r-lro- nal


r(* > II r











TG was displayed


the Tektronix where


final


adjustments


were made


upon


amplitude of


the AO output,


pulse


rate,


pulse


interval,


and


pulse


shape.


finished


signal


was


switched


input


the Ampex


and recorded


tape


ips.


A section


this


tape


was


then


removed and made


into


model


loop


T-202


for testing


(Sphericon)


females


or Realistic


phonotaxis.


40-2338


Sphericon


(Realistic)


exponential


horn


tweeters


(8n)


were


used


to broadcast


signals


(Fig.


Equipment


for Tests


of Phonota


xis


-foot-diameter,


sand-bottom arena


with a


6-inch-high


screen


side


lightweight nylon-mesh


was


test


enclosure


in which


female-responses


were


studied


(Fig.


21) .


Sitting .atop a


6-foot


step


ladder,


experimenter


looked


downward


arena,


recorded


data,


and by


remote


con-


trols


started


stopped


broadcast of


sound


arena


speaker.


arena


center were


two or


three


water vials with exposed


moist wicks


and a


small


dish


crushed. dogfood.


arena


experiments


took


place


low noise


room


(LNR)


under


16L:8D photoperiod.


All


experiments


were at


to 50%


The Ampex


was


used


to record


and


play


back


tapes


20).


































































































source


Fig. :
Ra ta


Test


arena


containing


-U


sens


* U


itive


- a r n n 4.. a a 9L 9 m.L-n I a


platforms


- .3 m- -


* 1


<


I










output


the Ampex was


tested,


and


sound


level


adjusted


sound


laboratory.


Experimental Designs


Controlled


experiments


demonstrating


female


response


to natural


for Oecanthus


synthetic

by Walker


male calling

(1957), for


songs


have been


Ephippiger


spp.


reported


Busnel


(1963)


, Teleogryllus


Hill


(1972)


Scapteriscus


by Ulagaraj


(1974)


It appeared


that


Pictonemobius


would


lend


itself


quite well


oassay


virgin


female


response


since


they


are


plentiful,


small,


wingless,


diurnal.


factors


controlled


several


equipment


test


pilot


that


will


investigations


were


into


necessary


be described


various


a reliable,


in detail,


some


these


findings will


given


preface


larger


and more


inclusive


tests


discussed


later.


Earl


response


y experiments


periodism,


were made with a


one of


single


speaker


female


"square


test


locations


response


cut"


speaker,


to sound


synthetic


located


surrounding


level,


trills


randomly

arena


(Fig.


with


"dummy"


speaker placed


opposite


con-


trol.


arena


Two wire


that


arcs


when


were


viewed


placed


from above


mesh


an arc with a


inch


radius


extended


outward


from


eac


speaker,


demarking










period


9 minutes.


Following


this


speaker positions


were


changed


a different


test


sound


was


presented


same crickets

females were


after

tested


another

each ti


m


9-minute

e. If t


silent


wo or more


period.


females


were within


either


zone of


attraction another


test


speaker


position


was


randomly


chosen before


test was


begun.


sound


period


was


divided


into


three,


3-minute


periods.


number of


females


each


zone of


attraction


test


control)


each


3-minute


period was


recorded.


Control,

tended t


or dummy,


.o respond


speaker

visually


counts were made


speaker


in case


or remain


females


beneath


wire


arc.


Later


experiments


female


response


"round


cut"


trills


chirps


was


evaluated by


using


three


test


speakers,


addition


of ER counts of


cricket activity.


The


random selection


speaker


locations


used


earlier


experiments


sometimes


resulted


in a


test


starting when


several


crickets were


concentrated


in an


adjoining


speaker


location.


To eliminate


this,


a sequential


presentation


test


control


signals


was


developed


which


three


speakers were


spaced


equidistant


the outer margin


arena


(Fig.


21).


A control


(natural


calling


song


instead


of the


previously


used


silent


control)


test broadcasts











change


the next


tape


loop


speaker


position,


and


adjust


signal


intensity.


It also


allowed


enough


silent


time


before


next


test


presentation


the experimenter


to return


room and


crickets


resume


normal


movements


around


the arena


after


the experiment


was


readied,


After


each


test


signal


presentation,


3-minute


silent


period


preceded


each


control


tape


loop


presentation.


Each


6-minute


control


broadcast had


effect


of clearing the


area


around


next


test


speaker


prevy


Ous


test


speaker


attracting


test


subjects


to one


location,


served


test


a before


presentation


could


after


control


be compared.


signal,


The


to which


6-minute


each


test


control


periods


allowed


enough


time


record


res


ponse


present


a number


different


signals


same day


without


loss


sensitivity


and


responsiveness


part


crickets.


The


sequence was


follows:


6-minute


control


3-minute


signal,


silence.


6-minute


This


silence,


sequence


6-minute


was


test


repeated


signal,


eac


different


test


signal,


and


ended


with


6-minute


control


broadcast.


this


manner


it was


possible


to present


series


order,


seven


different


to a group of


virgin


test


signals,


female


in a randomly


crickets


less


chosen


than


3 hours.


third


consecutive










early tests


phonotaxis,


age of virgin


females was


the only


standard


used


select


test


subjects.


Because


some


females


showed


a lack


responsiveness,


a new


method


of selection


virgin


females


was


developed


females


used


later


experiments


were


selected


only


they


began


a mating


sequence


(courtship


"spermatophore


formation"


stage


(Mays


1971)


with


a male


their


own


song


type


24-48


hours


prior


testing.


They were


selected


taking


stridulating male,


placing


him in


jar with


several


virgin


females,


observing


courtship


then


re-


moving


responding


female


soon


the male


formed


spermatoph


ore.


Differential


Response


Taped


Calling


Songs


Species


531-D virgin


females


were


tested


response


tape


recordings


their


own


(conspecific)


each


other


s (heterospecific)


calling


songs


three


reasons.


First,


such


test


would


determine whether virgin


females


would


res


pond


tape-recorded


songs of


their


con-


specific males.


Second,


it would


demonstrate


response


attracted


was


species


to each


specific


other'


s songs


or whether


Third,


they


could be


it would


give


some


indication


suitability


test


terms


signal


presentation,


sound


pressure


level,


response










females was


tested


three


replicates of


each


species.


numbers


that


responded by


entering


zone


attraction


test


speaker


are


shown


Fig.


There was


Such


indication


sharp discrimination


attraction


females


the dummy


favor


speaker.


calling


songs of


their


conspecific males


demonstrates one way that


531-D avoid


interbreeding.


The


test


signals


were


528-100,


35-1


s at


26.0C


-78,


44-8


25-80C.

Sound Level


test


to evaluate


more-intense-than-normal


response


male


calling


of virgin


song was


females


necessary


before


an appropriate


sound


level


future LNR arena


tests


could be


selected


The effect


a more


intense


signal


phonotactic


behavior


virgin


females


was


evaluated


compared


with a


tape


recorded


song


similar


that


produced


singing males


5-inch


distance.


A General


Radio Company


type


1551


B sound


level


meter


(SLM)


(scale A at


inches


from the


sneaker)


was


used


adjust a


tape


loop


copy


calling


song


levels


intensity,


75 db.


speed


female


response


sound


was


greater


during


db signal


broadcast


than


during










attraction.


test


signal


was


effective


in demon-


strating


a rapid


positive


response,


but


responding


females


moved


aro


much


more


rapidly


than


ever


observed


in response


living


males


(Fig.


23).


The distribution


sound


level.


arena


was


measured with


SLM.


The


zone of


attraction


close


approximated


area


to which


75 db extends


Fig.


Broadcast


sequent


sound


levels of


experiments


because


75 db were


adequate


chosen


response


sub-


occurred


a relatively


short


period


time


at sound


levels


approxi-


mating natural


Response


ones.


Perlodism


Daily


chan


female


response


were


suspected


view of


response


daily


ten,


rhythm of male


3-week-old,


song


virgin,


production.


laboratory-reared


females,


a pair


9-minute


broadcasts of


taped


calling


sured


song


four times


separated by


daily


minutes of


three


silence


consecutive days.


was mea-


Once


daily the


broadcasting


schedule


was


offset


two


hours


resulting


broadcasts


every


2-hour


period


(composite)


24-hour


day.


this


study


percent


those


entering


zone


attraction


during


initial


minute


of broadcast


24).























SONG


531-D


SONG


100-


FNOTESTED-

NO .TESTED-


531-D


531-D


Fig. 22.
Pictonemobius


spons
and


e to tape recorded
531-D females.


calling


songs


S* *.
* ** *
*g *
* "...
* a
.me:..*
*. *.
* *. C
*/ -*.
* *g *
**
genS
* S*
- **-
,,d *<.
.'.* .* i

! a *
* *. .
e* *


** I
49-<


Teste


_ _L T E S_ I__T Y


C


. rn II 1 II / t*4tflflC nfl S


F fl nt -


......


* n ^rW ^1 f- ^/I I


/ 2\


I


9 _
























































1 foot


Fig.


Distribution of sound level in test


arena.




















* t
* *


0


. ....



^~~~~~-


-4 ^
SN


4.1

0
O
-4-

0
4 -)


'-4


-r4 C


O3
-"-4
WH Cn
CC)











average


response


broadcast


dark


period


(1.25)


was


similar


that


the day


(1.31)


average


first


response was


average


2.11


response


compared


.88.


with


silent


a second


period


third


response


day


averaged


.05.


In general


crickets


responded


poorly


during


last


days


test.


One


cricket


died


on 10


February


the others


spent much


their


time


center


arena


around


the water vials


food


dish.


These data


demonstrate daily periodism


female


responsiveness


re-


corded male


calling


songs,


overall


level


response


was


so low that


such


periodism,


exists


could


have


been


missed.


results


tions


pointed


physiological


need


readiness


further


virgin


considera-


females.


Greater


response.


response


Rather


been


than


noted


repeating


other


this


test


tests


female


s-ubsequent


testing


virgin


female


Pictonemobius


was


the morning


hours


corresponding


first


daily


peak


in male


song


activity


noted


in both


field


and


labor


atory


Results


Analysis of Female Response

Square-cut trill s, 2 species


When


531-D


laboratory


reared


virai n


fPma 1 r


a ictI i n-m i i rmrc


f rnrn nr


I


1











the pulse rate was appropriate.


y varying


audio


oscillator


from


range


nr.


31-D


range


nr.


45 p/


experimenter


could


attract


one


direction


switching


speaker


pulse


rate)


attract


531-D


opposite


direction.


No differences


could


shown


cut,


in response


synthetic continuous


531-D


trills of


virgin


females


appropriate


square-


pulse


rate


their


tape-recorded


natural


calling


songs


(Fig.


Round-cut trills, 4 species tests.


Virgin


females


528,


531-D


531-B


that


were


exposed,


one


species


time,


a series


seven


synthetic


round-cut


trills


each


apart


range of


their


expected response


(30.0


67.5


p/s).


Tape


loops


of natural


calling


songs,


38.1,


47.2


3.8,


p/s,


res


pectively,


were


used


controls


throughout.


Sixteen


different


synthetic


trills


ranging


from


30.0


67.5


p/s,


were


used.


Portions of


three


such


trills,


, 32.5,


are


shown


(Fig.


Synthetic


trills


were


used


because


chirp


length


varied


between


four


song


types


would


likely


introduce


additional


variable.


Each


series


seven


test


signals


were


presented


once


each


day


between


8:00


12:00


a.m.


EST


to a group of


crickets


one


species.


randomly


chosen


sequence of


presentation


began


at a different broad-


27)















531-D


FEMALES


FEMALES


100-


-4


* ,-S
*:'
** >
f4 f S 44
* .45


*, .
**.
*
* **
So
*. *

* *
**
9 ** .
*
* a *
*
"* **
. 4-
C
*


* *
* S S
* a

-,


TESTED


BROADCAST

SIGNAL*


*The re


was


no difference between the broadcast periods and


the silent periods at the
of the arena).


"dummy"


speaker


(opposite


side



































SYNTHETIC TRILLS (TOP TO .10


BOTTOM)


30.0,


32.5 AND


SEC/DIV


NATURAL NEONEMOBIUS


SEC/DIV


aL.iaiA a A


*.1 ^^I


-.,,.\


-- =


-


A i I I ii, |


~IICLI










response


to each


test


broadcast


period was


evalu-


ated


following


four ways


data


compiled


each method


separately.


results


show


four


peaks


female


resp


onse


each of


four methods


corresponding


calling


song pulse


rates of


each


four


species.


Each of


four methods


analysis


was


attempt


measure


same


test


response


in different


ways.


Event recorder


counts.


As crickets moved


area


front


of a broadcast


speaker,


they walked


onto


sensitive


platform


(Fig.


triggered


phono-


cartridge-sensitive


"clicks"


relay


the event


system.


recorder pen


number


scribed


events


on a chart


each


6-minute broadcast


period


was


tallied.


Unavoidable


individual


ferences


between


eacn


sensitive


platform


system with respect


responsiveness


given


cricket


movement


were


averaged


having


each


test


signal


broad-


cast


once at


each


platform during


three


test


periods


each


species.


This


tally was


compared


with


average


number


of events


broadcast


the controls.


control


This wa


preceding


signal


following


expressed


an attempt


6-minute


a percent


to reflect


intensity


attraction


time


each broadcast


test


signal


to eliminate


effect


any


changes


in responsiveness










test


trill


pulse


rates


were


range


of +


appropriate


male


calling


song


(Fig.


Number


responding.


Any


female


which


turned


toward


direction


broadcast


sound


and moved more


than


body


lengths


toward


broadcast


speaker with


short


quick


steps


characteristic of


phonotaxis


was


given


a posit


response.


The number


responding was


expressed


a percent


the average number


responding


during


preceding


following


control


periods


(Fig.


thereb


y eliminating


effect


of different


proportions


females


being


ready to


respond


rate


produced by


conspec


ific


males


Number


moving.


Movement during


each


6-minute


experimental


period


whether


silent,


test


or control


was


quantified by


totaling


the number


individuals moving


during t

2-minute


he first-

segments


2-minute,


index of


second


2-minute


third


movement during each test


or control


period


maximum--10

subtracting


during


index was


was


cricke


preceding


arrived

ts during


total m

silent


to demonstrate


place during the


various


at by

q there


movementt


period.

change


test broad


taking the total mov

e, 2-minute periods)

30 maximum) that occ

The purpose of this

in activity that too

asts. The controls


ement



urred



k


29).


















fiaaa


U
r
-rnnnrnsrnrnnnfltfl
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greatest


appropr


activity


iate calling


occurred


t^ ci-Jt^L
r]an-
ew


four


song


spac e "


females


tested


. 31)


Weighted response i n d ex.


Ti- .


vre.asure


a ttmp nts


score


individual


femal


to how


strong lyi


they


respond


to a gi', en broadcast


siane 1


was


done


include


comparison


with


the event


recorder


counts. -


index


was


evaluated


total


number


cricket Ls


"zone


attraction"


(less


those


already


present.


during


pre-


ceding


plus

Some


6-minute


silent


3-4-minute


females


climbed


period)


count


0-2-minute


plus


screen


period


4-6-miniute


direct .y


front


count.

the


speaker


were given


sum of


these


previously


index


outlined


values.


at tractiveness


shown


Fig.


528,


531-D,


531-B,


525.


plot


this


index agrees


closely with


event


r ecor i


counts


and


describes


similar peaks


response


, orientation,


activity


already


described


(Fig.


32).


preceding plots


female


response:


showed


a wide


range of


peak


response


each


song


type


tasted.


This


lack


sharp discrimination


under


theE 2


test


conditions


may


have


resulted


from usina


fema les


with


low threshold


response


in a


"single


choice"


. e.


response


vs.













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similarly


deprived


virgin


females


of Teleogryllus


responded


two choice


about


cases


situations,


i.e.


to nonconsoecific


no response


songs.


. response


conspecific


sound


vs.


response


to nonconspecific


sound,


Hill


et al.


(197


noted


that


females


discriminated nearly


100%


time.


Artificial


trills


stimulated


females


to greater


response


than


control


calling


song,


suggesting


that


continuous


trills


present


above-normal


stimulus,


which


have


also


effect


lowering pulse


rate dis-


crimination.


Chirp


Length


effect


changing


chirp


length


while


maintaining


a constant


pulse


rate


chirp-to-interval


ratio,


was


evaluated


using


virgin


females.


Species


was


chosen


because


it had


longest chirp of


Pictonemobius


seemed


the most


likely


species


to be affected by


shortened


chirps.


test)


Round-cut


trills


were chopped by TG


used


into


chirps


previous


, 64,


pulses


long with


intervals


chirp


length,


that


each


test


signal


presented


same


number


pulses


minute


increase


28)


Plots


response


response


chirps


(Fig.


approached


show


typical





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It appears


that


a chirp of


some


length


important,


short


4-pulse-on,


2-pulse-time-off


chirp


received


some


response.


When


the number


pulses


per minute


kept


same,


short


pulse-number


chirps


are


close


together


and may


elicit


response


which


reflects


pulse


rate effect


rather


than


a chirp


rate


effect


(Fig.


The calling


song


of Neonemobius


cubensis


(Saussure


537-16,


which


a nearly


continuous


trill


similar


that


525,


was


added because


cubensis


occurs with


some


situations.


The N.


cubensis


test


signal


(Fig.


57.1


s received


lowest response of


despite


nearly


continuous


trill


close


pulse


rate


being within


range


expected


response


noted


pulse


rate experiment


(Figs.


33)














































128/64 64/32 32/16 16/8 8/4 4/2 537-16


COPY


128/64 64/32 32/16 16/8 8/4 4/2 537-16


PULSES ON


PULSES OFF


COPY


COPY


/




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