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HODGSON, Ronald Keith, 1946- SEX DETERMINATION IN THE AMERICAN HOUSE DUST MITE, DEKMATOPHAGOIDES FARINAE HUGHES, 1961 (ACARI: PYROGLYPHIDAE). The Ohio S ta te U niversity, Ph.D., 1975 Entomology

Xerox University Microfilms,Ann Arbor, Michigan 48106 SEX DETERMINATION IN THE AMERICAN HOUSE DUST MITE,

DERMAT0PHAGO3DES FARINAE HUGHES, 1961 (ACARI: FYROGLYPHIDAE)

DISSERTATION

Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate

School of The Ohio State University

By Ronald Keith Hodgson, B.S.

#*##*#

The Ohio S ta te U niversity

1975

Reading Committee: Approved by

G. W. Wharton Elton F. Paddock

Clayton B. Cook Adviser Department of Zoology ACKNOWLEDGMENTS

I gratefully acknowledge the guidance and patience of ny adviser, Prof, G. W. Wharton, throughout my graduate years. I thank Prof. Elton F. Paddock for his advice on the genetic aspects of this thesis and especially for the suggested approach. I owe a debt of gratitude to Mr. Fred S. Ruland and the Statistics

Laboratory, The Ohio State University, for their help in ordering and classifying the data and to Mrs. Sue Milne for typing the first draft and final manuscript. Thanks are also due to the Department of Zoology, The Ohio State University, and to the National

Institutes of Health for a training grant to the Acarology

Laboratory, The Ohio State University, for financial support.

Finally, I have to thank ny wife, Shu-ping, for her help with the statistics, but especially for her continuing support and my parents, Mark and Elizabeth Hodgson, who made i t possible for me to attend college in the first place. V2TA

February 25, 194-6 • • • Barn - Mt. Vernon, Ohio 1968 ...... B.Sc., The Ohio State University, Columbus, Ohio

1969-1970 ...... Research Assistant, Acarology Laboratory, The Ohio State University, Columbus, Ohio

1970-1971 ...... Graduate Teaching Assistant, Department of Zoology, The Ohio State University, Columbus, Ohio

1971-1972 ...... Graduate Teaching Associate, Department of Zoology, The Ohio State University, Columbus, Ohio 1972-1975 ...... Acarology Trainee, Acarology Laboratory, The Ohio State University, Columbus, Ohio

FIEUJS OF STUDY

Major Field: Zoology

Acarology. Professor George V. Wharton

Genetics. Professor Elton F. Paddock

i i i TABLE OF CONTENTS

Page

ACKNOWLEDGMENTS...... i i

VITA...... i i i

LIST OF TABLES...... v

LIST QF FIGURES...... v i

INTRODUCTION...... 1

MATERIALS AND METHODS...... 5

RESULTS...... 9

DISCUSSION...... 39

SUMMARY ...... 54-

APPENDIX ...... 55

LIST OF REFERENCES ...... 97

iv LIST OF TABLES

Table Page

1. Contingency table analysis of the and 0^ for 13 traits • 34

2. Two*tailed tests for skewness and kurtosis far the familial frequency distribution in the F? and 0, far 13 t r a i t s ...... 36

3. Contingency table analysis and estimated conditional [p(Rnw/Lnw) and P (Inw/Rnw)] and unconditional [P(Rnw) and P(Lnw)j probabilities of the right and left sides for those traits which occur on both sides of the body for the F^ and 0^, where R right, L left and nw non-wild phenotype e...... •••••••• 37

4 . Conclusions f a r each o f 13 t r a i t s when te s te d a g ain st any one of five appropriate hypotheses, where A accepted, R rejected and a blank means not applicable or insufficient d a ta ...... 4 0

5. Chi-square goodness of fit test far a ratio of 3 wildtype : 1 nnnr-wildtype for twelve families used to test the predictions of H3 using trait A8, presence or absence of small antero-dorsal shields associated with SCE and SCI se ta e ...... 44

6. Chi-square goodness of fit test for a ratio of 1 wildtype : 1 non-wildtype for 29 families used to test the predictions of H4 using trait A7, bursa*s position with respect to the anal slit in females ...... 46

7. Chi-square goodness of fit test for a ratio of 1 wildtype : 1 non-wildtype for 9 families used to test the predictions of H4 using trait All, additional small antero-dorsal shields in males ...... ••••••••• 47

8. Chi-square goodness of fit test for a ratio of 1 wildtype : 1 non-wildtype for all groups of females for trait A7, bursa*s position with respect to the anal slit in females • 46

v LIST OF FIGTRES

Figure Page

1* Karyotype of egg of D. farinae ...... 4

2. First eplmeres of male fused into a ! • • ...... 11

3. First epimeres of male fused into a V ...... 12

4« First epimeres of male not fused ...... 13

3. Protuberances on epigynial shield of female present • • • 14

6. Protuberances on epigynial shield of female absent and g a setae off epigynial shield of female .••••. 15

7* g a setae on epigynial shield of female ...... 1?

8. g a seta on cxlll shield of male ...... 18

9* Bursa to the left of anal slit in female •«•••••• 19

10. Bursa to the right of anal slit in female ...... 20

11. Small antero-dorsal shields absent ...... 21

12. Only SCE setae on small antero-dorsal shields • • • . • 22

13* Both SCE and SCI setae on small antero-dorsal shields f and small antero-dorsal shields unattached to propodosomal shield ...... 23

14* Small antero-dorsal shields attached to propodosomal shield and no additional antero-dorsal shields • • • • • 25

15* Additional antero-dorsal shields ...... 26

16. Normal s seta on tarsus I I I ...... 27

17* Stubby s seta on tarsus I I I ...... 28

18. Bifid s seta on tarsus III ...... 29

v i LIST OF FIGURES (c o n .)

Figure Page

19. Large spur on tarsus I I ...... 31

20. Medium spur on ta rs u s U...... 32

21. Small spur on tarsus II ...... 33

22. Male with first epimeres fused into a I emerging from tritonymphal exuvium ...... 33

v i i INTRODUCTION

Biological knowledge of the American house dust mite,

Dermatophagoides farinae Hughes, 1961, and the other pyroglyphids has increased dramatically in recent years (Fain and Lowry, 1974;

Keh, 1973; Brody [ed.^, 1971 j van Bronswijk and Sinha, 1970).

Information on sex determination in D. farinaemmm and the other pyroglyphids is lacking. In fact, only a few families of the Acari have been investigated in regard to sex determination: Argasidae and Ixodidae (Oliver, 1974; Sokolov, 1954)> Dermanyssidae (Oliver,

1965), Macrochelidae (Filipponi, 1964), Phytoseiidae (Wysoki, 1973),

Tetranychidae (Helle et a l., 1970) and Anoetidae (Hughes and Jackson,

1958). Three general reviews are available (Oliver, 1971, 1967 and

1964) as well as a thorough overview of arrhenotoky and variability in the Tetranychidae (Helle and Over me er, 1973)*

L ittle is known about sex determination in the suborder Acaridei to which D. farinae belongs. Sex determination among the Anoetidae is known best. Of the species examined all are either arrhenotokous and/or thelytokous (Oliver, 1971). No other investigated species of the Acaridei, except for the listrophcrid, Histiophorus numerous us

Dubinins, 1964, which can reproduce try amphoterotoky (the production of male and female offspring by parthenogenesis) (Oliver, 1971), has been found to be parthenogenetic. Four species more closely related 2 to the Pyroglyphidae (Glycyphagus domesticus (DeGeer, 1778),

Tvrophaeus nutrescentiae (Schrank, 1781), Rhizoglvphus echinopus

(Fumouze and Robin, 1868) and Acarus siro L., 1758) have been examined and are suspected to have chromosomal mechanisms of sex determination (Sokolov, 1954; Oliver, 1967). Only indirect information is available on sex determination in D. farinae and the other pyroglyphids (Furumizo, 1973; Larson et a l., 1969; Oshima and

Sugita, 1966; Oshima, 1964; Gridelet and LeBrun, 1973; Spieksma,

1967).

The sex determining mechanisms known to occur in the Acari are arrhenotoky, simple sex (XX:XY and XX:XO) and multiple sex

(X i X2.^2^2:4 i 22^) chromosome mechanisms, thelytoky and amphoterotoky

(O liv er, 1971 and 1967). Monogeny, th e production by some fem ales in a population of only male or female offspring, is reported for two species of Trombiculidae (Holman, 1961; Neal and Barnett, 1961).

A population with a monogenic mechanism has at least three sex genotypes: males, females that produce only male offspring and females that produce only female offspring. In addition, some populations exhibiting monogeny also have females which produce both male and female offspring rather than unisexual broods. In this investigation of sex determination in D. farinae all of the above mechanisms w ill be considered.

Sex determining mechanisms in the Acari have been investigated largely by cytogenetic analyses, except for breeding experiments conducted on some parthenogenetic farms. Inadequate resolution has been a barrier in the cytological examination of D. farinae (Figure 1), Therefore, the present investigation of sex determination in D. farinae will consist only of breeding experiments based on two assumptions, other than, of course, that sex determination in

D. farinae is genetic. First, it is assumed that hypotheses developed prior to the experiment using models of simple Mendelian inheritance, simple sex-linked inheritance and the sex ratio can discrim inate among monogeny, arrhenotoky, sex chromosome mechanisms, thelytoky and amphoterotoky. The term ‘’simple11 used in regard to

Mendelian and sex-linked inheritance refers in this investigation to a single locus with two alleles, the wildtype allele always being dominant to the non-wildtype allele. Second, it i3 assumed that some data collected on some variable traits of D. farinae. selected after the breeding experiment, with the help of phase contrast microscopy, and not necessarily known to be inherited, will prove to be inherited and amenable to testing the predictions of the proposed hypotheses. Figure 1. Karyotype of egg of D. farinae MATERIALS AMD METHODS

Two sources of D. farinae. one from Columbus, Ohio, and the other from Washington, D.C., were reared in the laboratory as by

Larson et al. (1969) except that the medium consisted of human hair and baker*s yeast. The amount of medium for isolations and crosses was small enough so that individual mites could be seen.

Crosses were conducted in the following manner. Active tritonymphs were collected from cultures of the Columbus source.

Each was isolated and cultured as above. Each was periodically examined to see when the adult stage was present. Every time an adult male and an adult female were observed, the last seen was put into a jar with the first seen. Afber thirty days both of the adults were removed and mounted on a slide, tfeually, none of their offspring were adults at that time. If, when any of the mated pairs were mounted some of their offspring were also adults, then they too were mounted. The mated pair and their adult offspring are indistinguishable and thus the parent (s) and offspring encountered on these occasions could not be labelled with distinguishable identi fication numbers, but proved useful when the observed phenotype for a trait was the same for the parent (s) and the offspring. For the next thirty days the jar was periodically examined and whenever an active tritonymph was seen it was removed and isolated. As soon as it emerged a3 an adult it was mated either to one of its sibs or to an adult from the Washington source, depending only on uhich was available at the time. The Washington source adults were obtained in the same manner as the in itial Columbus source adults. The mite pairs of these two types of mating, brother-sister and outcross, were cultured for thirty days. At the end of thirty days the mated pair was removed and mounted on a slide. Over the next thirty days as the offspring became adults they were removed and individually mounted on a slide.

Thus there were three classes of matings: (1) an initial mating

o f 66 p a irs of v irg in a d u lts of th e Columbus so u rce, 38 of which

produced offspring, (2) 129 brother-sister matings of some of the

offspring of mating (1), 88 of which produced offspring, and (3)

137 outcross matings of some of the offspring, 83 males and 54-

females, of mating (1) with virgin adults of the Washington source,

80 of which produced offspring. This gives five groups of mites:

(l) the 132 in itial Columbus mated adults, P, (2) the 395 offspring

of the initial Columbus mating, (3) the 137 Washington source

adults, 0, (4-) the 1013 offspring of the brother-sister matings, F^,

and (5) the 976 offspring of the outcross matings, 0^.

Qualitative and quantitative traits were selected for three

surveys in order to pick a group of traits that showed the most

promise of being variable genetic traits. The three surveys were

as follows: (l) a survey of 30 individuals randomly chosen from

group P, (2) a survey of 37 individuals randomly taken from a culture

of the Washington source, and (3) a survey of one P mating and all of their descendants. What were thought to be qualitative traits were 7

given numerical values for the observable phenotypes. Both the

classifying of the qualitative traits and the measuring of the quantitative traits were carried out under phase contrast microscopy with the aid of a drawing tube for the quantitative traits. The appropriate numerical codes for the 13 traits selected from the

surveys were then recorded for each of the 2521 experimental animals whose pedigrees could be traced. The first four digits of the

specimen*s identification number designates its family. The data

gathered were then used to test the predictions of the hypotheses

prepared for discriminating between the possible mechanisms for sex

determination.

Ordering and classifying of the data was accomplished by a

packaged program (Nie et al. , 1970) and programs written by

Mr. Fred S. Huland. Tests for skewness and kurtosis, g-j_ and g£>

of the fam ilial frequency distribution of observed ratios and the

contingency table analyses (variance tests for homogeneity of the

binomial distribution) were conducted using programs written by my

wife, Dr. Shu-ping Hodgson. All Chi square goodness of fit tests and

conditional and unconditional probabilities were carried out by me

on a Hewlett-Packard 9100B Calculator. S tatistical hypotheses were

tested using both one per cent and five per cent levels of

significance, except for the two-tailed gj^ and g2 tests where the

2% and 10 % levels were used. The specimens used in this investigation, including those vith non-distinguishable identification numbers, have been deposited in the collection of the Acarology Laboratory, The Ohio State University,

Columbus, Ohio 43210. RESULTS

The survey of 30 randomly selected P individuals (18 females and 12 males) entailed the examination of 108 qualitative traits and the measurement of 22 quantitative traits. Eighty-four of the qualitative traits and six of the quantitative were dropped for the

survey of 37 randomly selected individuals (25 females and 12 males) taken from a culture of the Washington, D.C., source. The 84 quali tative traits were dropped either because they were considered not to vary sufficiently or because the variation observed would have to be subjectively evaluated. The six quantitative traits were dropped because they were correlated with the same measurements on the other

side of the body. The same 24 qualitative and 16 quantitative traits

used in the survey of the Washington culture were also used in the

survey of one P pair (1500) and all of their descendants (28 females

and 1 4 males) except that an additional eight qualitative traits were

examined. All of the quantitative traits were afterwards dropped

because they demonstrated little variability or were correlated

with general body size and thus promised little assistance in

testing the available hypotheses. The last 19 qualitative traits

dropped were eliminated by combining traits on both sides of the

body which had been previously treated as two traits and by

discarding those traits considered to be continuously variable or 10 easily distorted by the process of slide preparation and were therefore difficult to objectively codify. The 13 traits eventually selected and the possible phenotypes, along with their absolute frequency in the and 0^, follow. The right and left sides for traits A8, A9, A12 and A13, although considered as a single trait when testing the hypotheses* were treated as separate traits during the evaluation and recording of the phenotypes in order to minimize the number of phenotypic classes and provide additional support, by the coincidence of the two sides, far the conclusions drawn concerning their possible inheritance. The wild phenotype is given first for each trait except Al, where the first is the wildtype in females and the second is the wildtype in males.

Al Sex: P2 °1 fem ale 542 535 male 471 441

A2 First epimeres of male (Figures 2 ,3 and 4): f 2 0 l fused into a Y 465 435 fused into a V 4 0 not fused 2 4

A3 Protuberances on the epigynial shield of female (Figures 5 and 6)5 F 0 2 °1 absent 444 4 6 3 present 98 70 Figure 2. First epimeres of male fused in to a 7 Figure 3. First epimeres of male fused in to a V Figure 4. First epimeres of male not fused Figure 5. Protuberances on epigynial shield of female present Figure 6. Protuberances on epigynial shield of female absent and g a setae off epigynial shield of female 16

Placement of g a setae of female (Figures 6 and 7): F2 °1 both off epigynial shield 306 359 both on epigynial shield 45 34 right off and left on 41 51 left off and right on 42 56

A5 Placement of g a setae of male (Figure 8):

both on cxIII shield 390 4 0 4 both off cxIII shield 1 0 right off and left on 1 1 left off and right on 1 2

A6 g a setae: F2 °1 both present 830 914 both absent 36 7 right absent and left present 71 24 left absent and right present 61 24

A7 Bursa*s position in female (Figures 9 and 10):

to the left of anal slit 292 281 to the right of anal slit 246 24 9

A8 Presence or absence of small antero-dorsal shields associated with SCE and SCI setae (Figures 11, 12 and 13): F2 °1 p resen t r ig h t sid e 798 754 l e f t sid e 809 765 r ig h t sid e absent 204 209 l e f t sid e 191 201 Figure 7* g a setae on epigynial shield o f fem ale 18

Figure 8. g a seta on cxIII shield of male 19

Figure 9. Bursa to the left of anal slit in fem ale Figure 10* Bursa to the right of anal slit in female esS S

Figure 11. Small antero-dorsal shields absent Figure 12. Only SCE setae on small anfcero-dorsal shields Figure 13. Both SCE and SCI setae on small antero-dorsal shields , and small antero-dorsal shields unattached to propodosomal shield A9 SCE and SCI setae: F2 °1 r ig h t sid e 968 both present 1004 l e f t sid e 1003 971 0 both absent r ig h t sid e 0 l e f t sid e 0 0

r ig h t sid e 0 SCI absent 5 l e f t sid e 7 0

right side 0 2 SCE absent left side 0 0

A10 Small antero-dorsal shields*, vhen present, relationship to propodosomal shield in male (Figures 13 and 14): F2 °1 attached 356 345 unattached 70 67

All Additional small antero-dorsal shields in male (Figures 14 and 15): F2 °1 absent 237 157 present 225 276

III (Figures 16, 17 and 18): F2 °1 r ig h t sid e 943 normal 915 l e f t sid e 951 913 r ig h t sid e stubby- 39 34 l e f t sid e 40 35

r ig h t sid e 22 b if id 19 l e f t sid e 14 21 Figure 14. Small antero-dorsal shields attached to propodosomal shield and no additional antero-dorsal shields Figure 15. Additional antero-dorsal shields

i Figure 16. Normal s seta on tarsus III Figure 17, Stubby s seta on tarsus III 29

Figure 18, Bifid s seta on tarsus III 30

A13 Size of spur on tar3U3 II (Figures 19, 20 and 21)

right side 484 medium l e f t sid e 497

right side 209 86 sm all l e f t sid e 230 87

right side 233 38 3 la rg e l e f t sid e 230 374

The recorded phenotype for each tra it on each of the 2521 specimens is presented in the Appendix. Those specimens with non- distinguishable identification numbers are not listed.

To ascertain if the 13 traits are non-randomly distributed, a contingency table analysis of the and 0^ for each trait and analyses for skewness and kurtosis of the familial frequency distri bution of each trait were conducted. If a trait is inherited one would expect that the offspring from the brother-sister matings would be at least as variable if not mare variable far any trait than the offspring of the outeross matings, due to the segregation of alleles in the F2. Statistically, each of the traits is equally or more variable in the F 2 than in the 0^ (Table 1). I therefore suggest that all of the traits might be inherited, in particular traits A3, A6, A9 and All. The coincidence of the left and right variances of each of the four traits A8, A9, A12 and A13 provides addi tional support for the conclusions drawn concerning these traits.

In addition to a contingency table analysis to detect possible segregation of alleles, tests for skewness and kurtosis of the familial frequency distribution of each trait were conducted. Unless a quali tative trait is distributed at random and thus not inherited or is always Figure 19. Large spur on tarsus II Figure 20. Medium spur on tarsus II Figure 21, Small spur on tarsus II T able 1

Contingency table analysis of the Fg and 0^ for 13 traits (•significant at 5% and ••significant at 1%)

T ra it F2 Variance 0^ Variance X 2

A1 252.030 2 a . 713 0.344 A2 5.906 3.959 0.275

A3 80.275 60.792 4.990*

A4 90.239 101.238 0.189

A5 2.964 2.990 0.002

A6 139.689 51.915 60.892**

A7 133.502 132.014 0.169

r ig h t 162.425 163.647 0.518 l e f t 154.519 159.172 0.897

r ig h t 5.020 2.033 1.175 l e f t 6.921 0.000 6.754**

A10 59.193 56.807 0.004

A ll 115.915 100.216 20.162**

r ig h t 57.354 50.142 0.296 l e f t 51.069 52.771 0.155

r ig h t 244.051 238.176 3.822 l e f t 244.764 239.156 0.342 35 inherited in a testcross (backeross) manner, then one would not expect a normal distribution. The results of such analyses for each trait are presented in Table 2. All of the traits, except Al, A7,

A13 and possibly All, are not normally distributed and thus are probably inherited, but not in a testcross manner. Traits A3, A6 and A9 are particularly noteworthy since both the contingency table analysis and the tests for skewness and kurtosis support the contention that they are inherited. Traits Al, A7, A13 and possibly

All are either randomly distributed or inherited in a testcross manner. The coincidence of the and gg values of the right and left sides of each of the trait3 A8, A9, A12 and A13 provides added support for the conclusions drawn for these traits (Table 2).

An additional contingency table analysis was conducted on those traits which occur on both the right and left sides of the body in order to ascertain if both sides are independent of or dependent upon each other. Trait A5 for both the P2 and 0^ and trait A9 for the 0^ were not analyzed due to insufficient data in some classes.

The null hypothesis of independence of the sides for each tra it is rejected at the 1% level of significance (Table 3)* In addition, the estimates of the conditional and unconditional probabilities far both sides of each trait are presented far comparison. If both sides are independent of each other for a trait, the conditional probability is expected to be equal to the unconditional probability. It is obvious that the estimated conditional probability is much larger than the unconditional probability in each case. This then supports the contention that both sides for Table 2

Two-tailed tests for skewness and kurtosis for the familial frequency distribution in the F 2 and 0^ for 13 traits (^significant at 10% and ^^significant at 2%)

sample V ariable 81 S2 sample S i g2 s iz e n s iz e n

Al 0.2495 3.2360 88 0.1642 3.1657 80

A2 6.1003** 42.7145** 81 6.8275** 51.6239** 71

A3 1.5600** 5.5182** 81 2.7866** 11.5428** 74

A4 0.7110 2.3323** 76 0.9434** 3.3639 71

A5 5.6570** 35.2597** 78 5.6946** 35.1744** 68

A6 1.1690** 4.4131* 88 3.2763** 14.5312** 78

A7 0.4173* 2.7485 81 0.1731 2.5121 74 r ig h t 1.552** 88 A8 4.930** 1.604** 5.391** 79 l e f t 1.734** 5.611** 88 1.607** 5.256** 79 r ig h t 8.1959** 72.0542** 88 38.826** A9 6.1235** 79 l e f t 5.8651** 40.1246** 88 _____ a ___- a 79

A10 1.844** 6.080** 79 1.221** 3.654* 69 A ll 0.0460 2.1797* 80 0.4470* 2.5742 70 right 3.0218** A12 11.9247** 88 1.6151** 4.7589** 79 left 2.8501** 10.3800** 88 1.3839** 3.4801 79

right 0.2801 2.7298 88 0.0461 2.5307 78 A13 l e f t 0.2998 2.6613 88 0.1962 2.6209 79 a and g2 do not exist 37

Table 3

Contingency table analysis and estimated conditional [P (Rnw/Lnw) and P(Lnw/Rnw)J and unconditional [P(Rnw) and P(Lnw)j probabilities of the right and left sides for those traits which occur on both sides of the body for the Fg and °1> where R = right, L -left and nw = non-wild phenotype

(••significant at 1% le v e l)

T ra it G eneration X 2 P (Rnw/Lnw) P(Rnw) P(LnwAnw) P(Lnw)

M F2 69.726** 0.523 0.200 0.517 0.198 A4 33.582** 0.400 0.180 0.378 0.170

A6 F2 78.184** 0.371 0.107 0.336 0.097 A6 0.226 0.032 0.226 0.032 °1 38.845** A8 F2 886.153** 0.984 0.203 0.931 0.192 AS 0.928 °1 851.964** 0.975 0.217 0.209 A9 F2 26.586** 0.143 0.005 0.200 0.007 A12 654.176** 0.870 0.061 0.770 F2 0.054 A12 °1 496.869** 0.714 0.055 0.755 0.058

A13 F2 476.551** 0.819 0.449 0.861 0.472 A13 570.109** 0.896 0.877 °1 0.493 0.493 each trait are dependent upon one another. Therefore, if inherited, the tvo sides far each trait are probably inherited as a single t r a i t .

Considering that none of the evidence is overwhelmingly against the contention that each trait is inherited, all of the traits are used to test the predictions of the appropriate hypotheses. DISCISSION

Twenty-nine hypotheses were developed to differentiate among thelytokous, arrhenotokous, amphoterotokous, monogenic and sex

chromosome mechanisms for sex determination. Of the 29 hypotheses tested using the data on 13 traits, only five provided pertinent

information on the mechanism of sex determination in D. farinae.

Sex determination in J). farinae was ascertained to be most

likely a sex chromosome mechanism with the male as the probable

heterogametic sex. The first two hypotheses based on the expected

sex ratio and its statistical properties reveal that parthenogenesis

is absent or extremely rare in D. farinae. The second hypothesis

also eliminates monogeny as a viable candidate mechanism. The

third hypothesis detects a probable autosomal tra it, thus dispelling

the contention of parthenogenesis as a probable mechanism and

suggesting a sex chromosome mechanism. The last two hypotheses

detect a probable sex-linked trait and the heterogamety of the male.

Each of the hypotheses and the conclusion regarding each trait is

presented in Table A, where A = accepted, R = rejected and a blank

means the hypothesis was not applicable or there was insufficient

data available for testing.

Each of the five useful hypotheses will be presented and the

conclusions drawn concerning the possible mechanism w ill be discussed.

39 40

Table 4

Conclusions for each of 13 traits when tested against any one of five appropriate hypotheses, where A - accepted, R® rejected and a blank means not applicable or insufficient data

Trait hypothesis

HI H2 H3 H4 H5 Al R R A

A3 A

M A

A6 A

A7 R A8 R

A10 A

A ll R R

A12

A13 A The vildtype designation was always assigned to the phenotypic state for each tra it, except sex (Al), which occurred most frequently and the wild phenotype was always assumed to be dominant to the non-wild phenotype. The hypotheses dealing with genetic transmission w ill be concerned with the following crosses.

Sex^linked Autosomal

AA x A aa x a AA x AA aa x aa 4' 4^ AA A aa a AA aa

AA x a Aa x a AA x aa Aa x aa * lAa:lA !Aa:laa:lA:la Aa lA aslaa

Aa x A aa x A AA x Aa Aa x Aa 4' * 4' !AA:lAa:lA:la lA aila lAA:lAa lAA:2Aa:lai

HI: If offspring are produced by a female without previous contact with a male then the sex determining mechanism is most likely not by amphimixis.

HI is rejected because neither I, Furumizo (1973) or Larson et al. (1969) have ever observed a female oviposit that had not had previous contact with a male. This is also true for the related species D. pteronyssinus (Trouessart, 1897). Amphoterotoky (the production of males and females by parthenogenesis), thelytoky (the production of females by parthenogenesis) and arrhenotoky (the production of males by parthenogenesis) are not, therefore, likely 42

candidates as mechanisms for sex determination in D. farinae. or in

D. pteronvssinus. It has been reported, however, that copulation

is a necessary prerequisite to oviposition in the following arrhenotokous mite3S Dermanyssus gallinae (DeGeer, 1778) (Oliver,

1965), several anoetids (Hughes and Jackson, 1958) and practically

all of the Phytoseiidae (Vfysoki, 1973). Thus, if D. farinae does reproduce by parthenogenesis it would appear that copulation is at

least a prerequisite to oviposition.

H2: If the frequency distribution of the familial sex ratios

is not normally distributed then the sex determining mechanism is

most likely not by sex chromosomes.

H2 is rejected because the frequency distributions of the F2

and 0^ familial sex ratios (Al) are not significantly different

from a normal distribution based on the results of tests for

skewness, g^, and kurtosis, g2 (Table 2). The frequency distribu

tion of the familial sex ratio for a monogenic population would be

bi- and possibly tri-modal; therefore, monogeny is a non-viable

candidate mechanism for sex determination in D. farinae.

Parthenogenetic populations often have a skewed frequency distribi>*

tion of their familial sex ratios and, therefore, any parthenogenetic

sex determining mechanism is even more unlikely to be a candidate

mechanism for D. farinae.

H3: If a trait that appears in both sexes, and where the

parents are both wildtype, is expected in a phenotypic ratio

significantly different from 3 wildtype : 1 non-wildtype in all

fam ilies, when both phenotypes are present among both sexes in the offspring and after all families that produced offspring of only-

one phenotype were eliminated from the analysis, then the trait is probably not autosomal.

H3 is rejected for the trait A8, presence or absence of the

small antero-dorsal shields associated with the SCE and SCI setae, because the ratio of the tvo phenotypes is not significantly

different from a ratio of 3 wildtype : 1 non-wildtype far the total

of, =0.152, and for each of eleven out of the twelve families

that were testable for this trait (0202, 1108, 1109, 2110, 2612,

2804, 4205, 4401, 4505, 4900, 5000 and 5110) (Table 5). A

heterogeneity test, )(^ - 14-198 with eleven degrees of freedom,

permitted pooling of the data. It bears repeating that for a trait

that occurs in both sexes no such families should produce offspring

in which both phenotypes are present in both sexes if the trait is

sex-linked. Such is not the case for an autosomal trait. Some of

the above families have both wild and noi>-wild phenotypes present in

both sexes (0202 w ith 5w?:4nw^:2w^:2nwtf*; 2110 w ith 9w£:4nw?:9w*':lnw*;

2612 with lOwfsSnw-P^w^slnw^ and 4900 with 3w?:6nw&7w<*':lnw<3') and

thus trait A8 appears to be a simple Mendelian trait. The

demonstration of a single autosomal tra it renders the parthenogenetic

mechanisms untenable and suggests that sex is probably determined by

a sex chromosome mechanism.

H4: If a qualitative trait expressed in one sex only is

expected in a phenotypic ratio significantly different from 1 wildtype

1 non-wildtype in all families after all families that produced

offspring of only one phenotype were eliminated from the analysis, AA

Table 5

Chi-square goodness of fit test for a ratio of 3 wildtype : 1 non-wildtype for twelve families used to test the predictions of H3 using trait AS, presence or absence of small antero-dorsal shields associated with SCE and SCI setae (^significant at 5% le v e l) fM X family wildtype non-wildtype t o t a l

0202 7 7 14 4.667*

1108 16 6 22 0.061

1109 28 6 34 0.980

2110 18 5 23 0.130

2612 13 6 19 0.439

2804 14 5 19 0.018

4205 7 5 12 1.778

4401 21 5 26 0.462

4505 36 6 42 2.571

4900 10 7 17 2.373

5000 17 6 23 0.014

5 1 1 0 9 5 14- 0.857

2 196 69 265 14.350 45 then the trait is most likely not sex-linked.

H4 is rejected for the following traits: A7, bursa*s position

in females, X2 =1.114* based on 249 wildtype and 226 non-wildtype

individuals (Table 6) and All, additional small antero-dorsal

shields in males, X2 =0.031, based on 66 wildtype and 64 non-wildtype

individuals (Table 7). Heterogeneity tests, X ^ =13.964 with 28 degrees of freedom (d.f.) for A7 and X2 - 9.601 with eight d.f. for

All, permitted pooling of the data. It was also found for trait A7, however, that the phenotypic ratio for all 1 3 3 3 fem ales was not

significantly different from a ratio of 1 wildtype : 1 non-wildtype,

X2 =3.572 (Table 8). A heterogeneity test, X ^ = 5.013 with four

d .f., permitted the pooling of the P, F1, F2» 0 and 0^ females. It

will also be recalled that A7 is normally distributed (Table 2) and

thus it appears that the bursa*s position with respect to the anal

slit in females is not sex-linked but a random event, i.e ., occurring

on both sides with equal frequency and without a discernable pattern.

This was not found for trait All, additional small antero-dorsal

shields in males and thus it is apparently a sex-linked trait.

The demonstration of the probable sex-linkage of at least one

trait and the autosomal nature of at least one other trait suggests

that sex determination in D. farinae is probably by means of a sex

chromosome mechanism. It is not possible in the present investigation

t o d if f e r e n tia te between sim ple and m u ltip le sex chromosome

mechanisms. Such a task would require an extensive linkage a n aly sis

and/or a karyotype analysis. 4 6

T a b le 6

Chi-square goodness of fit test for a ratio of 1 wildtype : 1 non-wildtype for 29 families used to test the predictions of H4 using trait A7, bursa*s position with respect to the anal slit in females fam ily w ildtype non-wildtype t o t a l X2 0707 8 13 21 1.190 0709 6 6 12 0.000 0902 7 7 H 0.000 0904 13 11 24 0.167 1 1 0 8 6 7 13 0.077 1 1 0 9 9 1 2 2 1 0.429 1503 13 5 18 3.556 1 7 0 1 1 0 6 1 6 1.000 2 1 1 0 8 5 13 0.692 2 1 1 1 8 5 1 3 0.692 2 6 1 2 7 6 13 0.077 2705 6 7 13 0.077 3908 5 5 10 0.000 4004 6 5 11 0.091 4202 6 5 11 0.091 4401 7 7 H 0.000 4502 11 11 22 0.000 4504 11 7 18 0.889 4505 13 14 27 0.037 4 9 0 1 8 7 1 5 0.067 5 1 0 2 1 0 8 1 8 0.222 5 1 0 3 9 9 1 8 0.000 5 1 0 4 11 1 9 30 2.133 5 1 0 5 5 6 1 1 0.091 5 1 0 8 10 7 1 7 0.529 5 1 1 1 9 8 1 7 0.059 5 1 1 2 13 6 1 9 2.579 5502 7 7 H 0.000 6 0 0 1 7 5 1 2 0.333

U 249 226 475 15.078 47

T a b le 7

Chi-square goodness of fit test for a ratio of 1 wildtype : 1 noit-wildtype for 9 families used to test the predictions of H4 using trait All, additional small antero-dorsal shields in males (^significant at 5% le v e l) family wildtype non-wildtype t o t a l X 2 0204 6 5 11 0.091

0701 6 5 11 0.091

0904 6 9 15 0.600

1 1 0 9 7 6 1 3 0.077

1 7 0 1 1 4 7 2 1 2.333 2 1 0 0 1 1 5 1 6 2.250

2 1 1 0 5 5 1 0 0.000

4201 5 7 12 0.333

5 1 0 4 6 15 2 1 3.857*

£ 66 64 130 9.632 4 8

Table 8

Chi-square goodness of fit test for a ratio of 1 wildtype : 1 non-wildtype for all groups of females for trait A7, bursa's position with respect to the anal slit in females (^significant at 5% le v e l) group w ildtype non-wildtype to ta l X 2 P 21 15 36 1,000

Fi 67 83 150 1.707 292 246 P2 538 3.933* 0 AO 39 79 0.013

281 249 530 1.932

£ 701 632 1 3 3 3 8.585 4 9

H5: If a sex-linked trait is expressed in only males and all

brother-sister mating pedigrees consisting of a non-wildtype P

grandfather and F^ father have only non-wildtype male offspring in

th e F2 then the heterogametic sex is most likely not the male.

The hypothetical pedigree of particular concern is as follows*

where nAH is dominant to Ma", cf^male and ?= fem ales

P ?Aa x ^ a

F-^ ?Aa x<^a

F2 1? Aa: l? a a :1p"A : 1-^a

H5 is rejected for the trait All, additional small antero-

dorsal shields in males. Four brother-sister mating pedigrees

consisting of non-wildtype P grandfather and F^ father had both

wildtype and non-wildtype male offspring in the F 2 (3904 with 4w and

7nw, 3906 with 4w and 2nw, 3908 with 3w and 7nw and 3 9 1 1 with lw and

2nw). Thus, males, rather than females, appear to be the hetero

gametic sex (XX:XI, XXsXO, X2X2X2X2 1X^X2?) in D. farinae. As mentioned previously, a chromosomal mechanism with male heterogamety

is known in the Acaridei. More specifically, T. putrescentiae.

A. siro and G. domesticus are known to be XX:X0 (Oliver, 1967). It

should also be noted that all of the pedigrees for the traits A8,

presence or absence of the small antero-dorsal shields associated

with the SCE and SCI setae and A ll, additional small antero-dorsal

shields in males, have been examined and are consistent with the models of inheritance proposed for each. 50

A simple chromosomal mechanism of sex determination with male heterogamety in D. farinae is expected to produce a primary sex ratio of 0.5* where sex ratio is equal to the number of males divided by the number of males plus the number of females. The sex ratios observed in this investigation are as follows:

= 0.535 based on 212 males and 184 females* X 2 s 1.980j

F£ = 0,465 based on 471 males and 542 females* X ^ = 4*976*j and

0^ - 0.452 based on 441 males and 535 females, X ^ - 9.05**. The

F2 sex ratio is significantly different from 0.5 at the 5% level and the 0^ sex ratio is significantly different at the 1% level. It should be noted that the F 2 and 0^ sex ratios are not significantly different from each other which is what one would expect if sex is determined by sex chromosomes. Sex ratios observed in natural populations are significantly different from 0.5 too for both

D. fa rin a e and D. pteronyssinus (Oshima* 1964; G rid elet and LeBrun,

1973)* Such observations do not present serious difficulties far the proposed mechanism since none of the sex ratios observed are primary sex ratios, i.e., at fertilization. All of the sex ratios that are significantly different from 0.5 favor the females and can be attributed to a differential mortality of the sexes after fertilization. It has, in fact, been demonstrated that females of both D. farinae and D. pteronyssinus live approximately twice as long

as males in laboratory experiments (Furumizo, 1973; Oshima and

Sugita, 1966; Spieksma, 1967).

In addition to the probable sex-linkage of sex (Al) and additional small antero-dorsal shields in males (All) as well as the 51 autosomal nature of the presence or absence of the small antero- dorsal shields associated with the SCE and SCI setae (A8), some knowledge ms gained on the possible inheritance of the other traits* The traits, bursa*s position (A7) and size of spur on tarsus II (A13)» were both normally distributed. A13 is probably

quantitative in nature while A7 appears to be greatly influenced by

norr-genetic factors, as mentioned in connection with H4.

The other eight traits [farm of the first epimeres in males

(A2), protuberances on the epigynial shield of females (A3),

position of g a setae in females (M) > position of g a setae in

males (A5), presence or absence of g a setae (A6), presence or

absence of SCE and SCI setae (A9), relationship of the small antero-

dorsal shields to the propodosomal shield in males (A10) and the

form of the s seta on tarsus III (A12)J were not normally

distributed and are, therefore, probably greatly influenced by

genetic factors. The analyses, however, did not suggest a pattern

of inheritance consistent with a simple Mendelian or a simple sex-

linked trait.

Two of the latter traits are of especial interest. Trait A2,

form of epimeres in males, has been discussed by Fain and

van Bronswijk (1973) in the following manner, ttIt is to be noted that

a slight tendency to produce heteromcrphism in the males exists also

in Dermatophagoldes farinae Hughes. In that species there are males

with epimera I completely separated and others with these epimera

fused into a V or a Y. As the specimens with fused epimera I were

more sclerotized than those with separate epimera we have surmized 52

(Fain* 1967) that this fusion was in relation with the degree of sclerotization and thus with the age of the specimens. We think now that this fusion is merely the expression of a tendency to heteromorphism. . The data presented verify Fain and van Bronswijk’s conclusion in regard to this trait, i.e., it is not likely a function of age, but rather appears to be influenced b7 genetic factors. During the present investigation additional evidence was obtained to support the contention of an early expression and therefore probably an hereditary nature for this tra it. A male with epimeres fused into a I is pictured in Figure 22 emerging from the tritonymphal exuvium.

The second particularly noteworthy tra it was the form of the s seta on tarsus III (A12). It produced the most unusual phenotype, i.e ., a seta that often appeared stubby or bifid and although scored for both males and females it appeared most frequently in females.

There is no apparent difference between the normal seta of males and females and thus it is unexpected that it does not vary similarly and in approximately the same frequency in males. A similar condition, trifid s setae on tarsi III and IV, has been used as a diagnostic trait in another species of the Pyroglyphidae,

Sturnophagoides kivuana (Fain, 1971)• Thus it is possible that alleles far this trait are more common in species other than £• farinae. Figure 22* Male with first epimeres fused into a Y emerging from tritonymphal exuvium SUMMARY

In summary, 13 qualitative traits were used to test the predictions of five hypotheses that discriminated among thelytoky, arrhenotoky, amphoterotoky, monogeny, a sex chromosome mechanism with the female as the heterogametic sex and a sex chromosome mechanism with the male as the heterogametic sex. It was concluded that sex determination in D. farinae is most likely by means of a sex chromosome mechanism with the male as the probable heterogametic sex .

54 APPENDIX

This appendix provides the raw data from which the analyses were made. Each of the specimens possesses a distinguishable identification number. Mites with the same numbers in the first four columns and a two in column nine belong to a single F 2 fam ily.

Mites with the same numbers in the first four columns and a five in column nine belong to a single 0^ family. The parents of a sp e c ific F2 family have the same numbers in the first four columns as the F2, but a one in column nine. The 0 parents of a specific

0^ family have the same numbers in the first four columns as the 0^, but a four in column nine. A one in column eight designates an individual from the Columbus source (this individual is also part of an F^) and a two in column eight designates an individual from the

Washington, D.C., source. The P parents of a specific F^ will have the same numbers in the first two columns as the F^, but zeros in columns eig h t and n in e. Column te n (sex) d istin g u ish e s between th e two individuals in each F mating and each F^ brother-sister mating.

The WGM above columns eight and nine represents generation. The date each specimen was mounted is presented in columns 1 0 -1 5 * The rest of the columns present the coded value far each phenotype for each trait far each specimen. The following is a list of the traits, the column, in parentheses, in which it occurs (four traits, AS, A9,

55 A12 and A 1 3 , occupy two columns, the first column representing the right side and the second the left side) and the numerical code for

each phenotype for each tra it for each specimen. A code value of

one always represents the wild phenotype and is presented first,

except for tra it A1 where the first code number given represents

the wild phenotype in females and the second code number given represents the wild phenotype in males.

Al Sex (16): female - 1 and male = 0

A2 First epimeres of male (17): fused into a I = 1, fused into a

V = 2 and not fused = 0

A3 Protuberances on the epigynial shield of female (18): absent

= 1 and present = 0

A4 Placement of g a setae in female (19): both off epigynial

shield - 1, both on epigynial shield = 0, right off and left on

- 2 and left off and right on = 3

A5 Placement of g a setae in male (20): both on cxIII shield = 1,

both off cxIII shield = 0, right off and left on = 2 and left

off and right on - 3

A6 g a setae (21): both present = 1, both absent - 0, right

absent and left present = 2 and left absent and right present

-- 3

A7 Bursa*s position in female (22): to the left of anal slit = 1

and to the right of anal slit - 0

AS Presence or absence of small antero-dorsal shields associated

with SCE and SCI setae (23* 24): present 1 and absent = 0 A9 SCE and SCI setae (25, 26): both present =1, both absent

- 0, SCI absent = 2 and SCE absent - 3

A10 Small antero-dorsal shields1, when present, relationship to

propodosomal shield in male (27): attached = 1 and

unattached * 0

All Additional small antero-dorsal shields in male (28): absent

= 1 and present - 0

A12 Form of s seta of tarsus III (29, 30): normal = 1, stubby

= 0 and bifid - 2

A13 Size of spur on tarsus II (31, 32): medium =1, small = 0

and large - 2 mite g date trait 0100000 00 021174 0 0 9 9 1 1 9 11 1 1 1 11 11 0100000 00 021174 1 9 1 1 9 1 1 11 1 9 9 11 00 0101000 01 040174 0 1 9 9 1 1 9 11 1 1 0 11 11 0101000 Ol 040174 1 9 0 3 9 1 1 11 1 9 9 11 11 0102000 01 040174 0 1 9 9 9 2 9 11 1 1 0 11 11 0102000 01 040174 1 9 1 1 9 1 0 11 1 9 9 11 11 0102001 02 041874 1 9 1 1 9 1 0 00 1 9 9 11 11 0103000 01 041174 0 1 9 9 1 1 9 11 1 1 0 11 11 0103000 01 041174 1 9 1 0 9 1 1 11 1 9 9 11 22 0104000 24 042274 0 1 9 9 1 1 9 11 1 1 0 11 11 0105001 02 061174 0 1 9 9 1 1 9 11 1 1 1 11 11 0105002 02 052074 0 1 9 9 9 9 9 11 1 1 0 11 21 0105003 02 052074 1 9 0 9 9 0 0 11 1 9 9 11 22 0105004 02 052074 1 9 1 1 9 1 0 11 1 9 9 11 12 0105005 02 050274 1 9 1 9 9 3 0 11 1 9 9 11 10 0105006 02 050274 1 9 1 3 9 1 0 00 1 9 9 11 22 0105007 02 050274 0 1 9 9 9 3 9 11 1 1 0 11 11 0105008 02 050274 0 1 9 9 1 1 9 11 1 1 0 11 11 0106000 14 042474 0 1 9 9 1 1 9 11 1 0 0 11 11 0106000 24 042474 1 9 1 1 9 1 1 11 1 9 9 11 21 0106001 05 061474 1 9 1 1 9 1 1 11 1 9 9 11 11 0200000 00 021174 0 0 9 9 1 1 9 11 1 1 1 11 00 0200000 oo 021174 1 9 1 1 9 1 1 11 1 9 9 19 11 0201000 01 040174 0 1 9 9 1 1 9 11 1 9 0 11 11 0201000 01 040174 1 9 1 1 9 1 O 11 1 9 V 11 22 0201001 02 042974 0 1 9 9 1 1 9 11 1 1 1 11 11 0201002 02 041074 1 9 1 2 9 1 1 00 1 9 9 11 22 0201003 02 051474 0 1 «* 9 1 1 9 11 1 1 0 11 11 0201004 02 051474 0 1 9 9 1 1 9 11 1 1 1 11 11 0201005 02 051374 1 9 1 3 9 1 1 11 1 9 9 11 22 0201006 02 051374 0 1 9 9 1 1 9 11 1 1 1 11 11 0202000 24 042274 0 1 9 9 1 1 9 11 1 1 0 11 11 0202000 14 042274 1 9 1 O 9 1 O 11 1 9 9 11 22 0202001 05 052474 o 1 9 9 1 1 9 01 1 9 1 11 11 0202002 05 052474 1 9 1 1 9 1 0 11 1 9 9 10 22 0202003 05 052474 0 0 9 9 1 1 9 00 1 9 1 11 00 0202004 05 052474 1 9 1 1 9 1 1 11 1 9 9 11 22 0202005 05 052474 0 1 9 9 1 1 9 11 1 1 0 11 11 0202006 05 052474 1 9 1 1 9 1 1 11 1 9 9 11 22 0202007 05 053174 0 1 9 9 1 1 9 11 1 0 0 11 11 0202008 05 053174 0 1 9 9 9 1 9 00 1 9 1 11 11 0202009 05 053174 1 9 1 1 9 1 0 00 1 9 9 11 22 0202010 05 050374 1 9 1 1 9 1 1 00 1 9 9 11 22 0202011 05 050374 1 9 1 2 9 1 0 11 1 9 9 11 22 0202012 05 050374 1 9 1 3 9 1 O 11 1 9 9 11 22 0202013 05 050374 1 9 1 0 9 1 0 00 1 9 9 11 22 0202014 05 050374 1 9 1 1 9 1 1 00 1 9 9 11 11 0203000 14 042574 0 1 9 9 9 2 9 11 1 0 0 11 11 0203000 24 042574 1 9 1 1 9 1 0 00 1 9 9 11 11 0203001 05 052474 1 9 1 1 9 1 0 11 1 9 9 11 22 0203002 05 052474 1 9 1 1 9 1 O 00 1 9 9 11 22 0203003 05 051074 1 9 1 1 9 1 1 00 1 9 9 11 22 0204000 14 042574 0 1 9 9 1 1 9 11 1 1 1 11 11 0204000 24 042574 1 9 1 1 9 1 1 00 1 9 9 11 11 0204001 05 061474 1 9 1 1 9 1 1 11 1 9 9 11 22 0204002 05 061474 0 1 9 9 1 1 9 11 1 1 0 11 11 0204003 05 061474 o 1 9 9 1 1 9 11 1 1 0 11 11 0204004 05 061474 0 1 9 9 1 1 9 11 1 1 0 11 11 0204005 05 061474 1 9 1 1 9 1 1 11 1 9 9 11 22 0204006 05 061474 1 9 1 3 9 1 0 11 1 9 9 11 22 0204007 05 061474 1 9 1 3 9 1 0 11 1 9 9 11 22 0204008 05 061474 0 1 9 9 1 1 9 11 1 1 1 11 11 0204009 05 061474 1 9 1 1 9 1 0 11 1 9 9 11 22 0204010 05 052474 0 1 9 9 1 1 9 11 1 1 1 11 11 0204011 05 052474 0 1 9 9 1 1 9 11 1 1 1 11 11 0204C12 05 052474 1 9 1 1 9 1 1 11 1 9 9 11 22 59 roite g date tra it 0204013 0 5 052474 0 1 9 9 1 9 11 1 1 0 11 11 0204014 05 052474 1 9 1 1 9 1 11 1 9 9 11 22 0204015 05 052474 0 1 9 9 9 9 11 1 1 0 11 12 0204016 05 052474 0 1 9 9 1 9 11 1 1 1 11 12 0204017 05 052474 1 9 1 2 9 1 11 1 9 9 11 22 0204018 05 05 2474 1 Q 1 3 9 1 11 1 9 9 11 22 0204019 05 052474 1 9 1 1 9 1 11 1 9 9 11 22 0204020 05 051074 0 1 9 9 1 9 11 1 1 0 11 11 0204021 05 051074 0 1 9 9 1 9 11 1 1 1 11 11 0205000 14 042574 0 1 9 9 1 9 11 1 1 0 11 11 0205000 24 042574 1 9 1 1 9 1 01 1 9 9 11 00 0205001 05 06 1474 0 1 9 9 1 9 11 1 1 0 11 11 0205002 05 061474 0 1 9 9 1 9 11 1 1 0 11 11 0205003 05 061474 1 9 1 1 9 1 00 1 9 9 11 22 0205004 05 061474 1 9 1 1 9 1 11 1 9 9 11 22 0205005 05 061474 1 9 1 1 9 1 11 1 9 9 11 22 0205006 05 052474 1 9 1 1 9 1 11 1 9 9 11 22 0205007 05 052474 1 9 1 2 9 0 00 1 9 9 11 22 0205008 05 052474 1 9 1 1 9 0 00 1 9 9 11 22 0205009 05 051074 1 9 1 2 9 1 11 1 9 9 11 22 0205010 05 051074 0 1 9 9 9 9 11 1 1 0 11 19 0205011 05 051074 1 9 1 2 9 1 11 1 9 9 19 22 0300000 00 021174 0 0 9 9 1 9 11 1 1 1 11 00 0300000 00 021174 1 9 1 3 9 0 11 1 9 9 11 00 0301000 01 040174 o 1 9 9 1 9 11 1 1 1 11 01 0301000 01 040174 1 9 1 9 9 1 00 1 9 9 11 01 0302000 01 041174 0 1 9 9 1 9 11 1 1 0 11 99 0302000 01 041174 1 9 1 1 9 0 11 1 9 9 11 00 0303000 24 042274 0 1 9 9 1 9 11 1 0 1 11 11 0303000 14 042274 1 9 1 1 9 1 GO 1 9 9 11 99 0304000 14 042274 0 1 9 9 1 9 11 1 1 0 11 00 0304000 24 042274 1 9 1 1 9 1 00 1 9 9 11 11 0305000 01 042674 0 1 9 9 1 9 11 1 1 1 11 00 0305000 01 042674 1 9 1 1 9 0 11 1 9 9 11 99 0306000 01 042674 1 9 1 1 9 1 11 1 9 9 11 11 0306001 02 061174 0 1 9 9 1 9 11 1 0 0 11 00 0306002 02 061174 1 9 1 1 9 1 00 1 9 9 11 11 0306003 02 061174 1 9 1 3 9 1 00 1 9 9 11 00 0306004 02 061174 0 1 9 9 1 9 11 1 0 0 11 10 0306005 0? 061174 I 9 1 1 9 1 11 1 9 9 11 22 0306006 02 061174 0 1 9 9 1 9 00 1 0 1 11 01 0306007 02 061174 1 9 1 3 9 O 10 1 9 9 11 11 0306008 02 052074 1 9 1 2 9 1 11 1 9 9 11 10 0306009 02 052074 0 1 9 9 1 9 11 1 0 1 11 00 0306010 02 052074 1 9 1 2 9 1 11 1 9 9 21 00 0306011 02 052074 0 1 9 9 1 9 11 1 1 1 11 10 0306012 02 052074 0 1 9 9 1 9 11 1 1 1 11 11 0306013 02 052074 0 1 9 9 1 9 11 1 0 1 11 11 0306014 02 052074 0 1 9 9 1 9 11 1 1 1 11 99 0306015 02 050274 1 9 1 2 9 O 11 1 9 9 19 11 0306016 02 050274 0 1 9 9 1 9 11 1 0 0 11 09 0306017 02 050274 1 9 1 1 9 0 11 1 9 9 11 11 0306018 02 050274 0 1 9 9 1 9 11 1 1 0 11 11 0307000 24 042574 0 1 9 9 1 9 11 1 1 1 11 00 0307000 14 042574 1 9 1 1 9 0 11 1 9 9 11 11 0307001 05 061474 1 9 1 1 9 1 11 1 9 9 11 22 0307002 05 061474 0 1 9 9 1 9 11 1 9 0 11 00 0307003 05 061474 1 9 1 1 9 1 00 1 9 9 11 00 0307004 05 052474 0 1 9 9 1 9 11 1 0 1 11 11 0307005 05 052474 1 9 1 1 9 0 11 1 9 9 20 21 0307006 05 052474 1 9 1 1 9 I 11 1 9 9 22 22 0307007 05 052474 1 9 1 2 9 1 00 1 9 9 11 22 0307008 05 052474 0 1 9 9 1 9 11 1 1 0 11 22 0307009 05 051074 0 1 9 9 1 9 11 1 1 1 11 99 0307010 05 051074 0 1 9 9 1 9 11 1 1 0 11 22 0307011 05 051074 1 9 1 1 9 0 11 1 9 9 22 11 60 mite g date tra it 0307012 05 051074 0 1 9 9 1 1 9 11 11 1 1 11 21 0307013 05 05 1074 1 9 1 3 9 1 1 11 11 9 9 11 11 0307014 05 051074 0 1 9 9 1 1 9 11 11 0 1 11 11 0307015 05 051074 1 9 1 1 9 1 0 11 11 9 9 11 11 0307016 05 051074 1 9 1 1 9 1 1 11 11 9 9 02 11 0307017 05 051074 0 1 9 9 1 1 9 11 11 1 O 11 00 0307018 05 051074 0 1 9 9 1 1 9 11 11 1 0 11 22 0308000 24 050974 0 2 9 9 1 1 9 11 11 0 1 11 00 0308000 14 050974 1 9 1 1 9 1 0 11 11 9 9 11 22 0308001 05 052274 1 9 9 9 9 1 1 00 11 9 9 11 11 0308002 05 052274 0 1 9 9 1 1 9 11 11 1 1 11 11 0308003 05 052274 1 9 1 1 9 1 1 01 11 9 9 11 22 0308004 05 052274 1 9 1 1 9 1 1 11 11 9 9 11 21 0308005 05 052274 0 1 9 9 1 1 9 11 11 1 0 11 11 0308006 05 052274 1 9 1 9 9 2 0 11 11 9 9 11 11 0500000 00 021174 0 0 9 9 9 3 9 11 11 0 1 11 00 0500000 00 021174 1 9 1 0 9 1 1 00 11 9 9 11 11 0501000 14 042274 0 1 9 9 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92 mite g date trait 5105021 0 2 052074 0 1 9 9 1 1 9 11 1 1 1 1 1 0 1 5105022 0 2 050274 0 1 9 9 1 1 9 11 1 1 1 1 1 11 5105023 0 2 050274 0 1 9 9 1 1 9 11 1 0 1 11 11 5105024 0 2 050274 O 1 9 9 1 1 9 11 1 1 1 11 11 5105025 0 2 050274 0 1 9 9 1 1 9 11 1 1 0 11 11 5105026 0 2 050274 1 9 1 1 9 1 0 0 0 1 9 9 11 2 2 5105027 0 2 050274 0 1 9 9 9 1 9 GO 1 9 1 11 11 5106000 0 1 050874 0 1 9 9 1 1 9 11 1 1 0 11 09 5106000 0 1 050874 1 9 0 1 9 1 1 0 0 1 9 9 90 11 5106001 0 2 052174 o 1 9 9 9 3 9 11 1 1 1 11 11 5106002 0 2 052174 0 1 9 9 1 1 9 11 1 0 0 11 09 5107000 0 1 050874 0 1 9 9 1 1 9 11 1 0 1 11 11 5107000 0 1 050874 1 9 1 1 9 1 0 0 0 1 9 9 11 2 2 5107001 0 2 061674 o 1 9 9 1 1 9 11 1 1 0 11 1 1 5107002 0 2 061674 1 9 1 1 9 1 1 00 1 9 9 11 0 0 5107003 0 2 061674 0 1 9 9 1 1 9 11 1 1 0 11 0 1 5107004 0 2 061674 1 9 1 1 9 1 1 0 0 1 9 9 2 2 11 5107005 0 2 061674 1 9 1 9 9 3 1 11 1 9 9 11 2 2 5107006 0 2 061674 1 9 1 1 9 1 1 11 1 9 9 11 0 1 5107007 0 2 06 ±674 0 1 9 9 1 1 9 11 1 1 0 11 11 5107008 0 2 061674 0 1 9 9 1 1 9 11 1 1 1 11 0 0 5107009 0 2 061674 1 9 1 1 9 1 0 0 0 1 9 9 11 2 2 5107010 0 2 061674 o 1 9 9 1 1 9 11 1 1 1 11 0 0 5107011 0 2 061674 0 1 9 9 I 1 9 11 1 1 1 11 11 5107012 0 2 061674 i 9 1 9 9 3 1 11 1 9 9 11 0 0 5107013 0 2 061674 0 1 9 9 1 1 9 1 1 1 9 1 1 1 0 0 5107014 0 2 061674 0 I 9 9 1 1 9 11 1 1 1 11 11 5107015 0 2 052974 1 9 0 1 9 1 1 0 0 1 9 9 11 2 2 5107016 0 2 052974 0 1 9 9 9 9 9 o o 1 9 1 11 0 0 5107017 0 2 052974 0 1 9 9 1 1 9 11 1 0 1 11 1 0 5107016 0 2 052974 0 x 9 9 1 1 9 11 1 1 1 11 11 5107019 0 2 052974 1 9 1 1 9 1 1 0 0 1 9 9 11 11 5107020 0 2 052174 1 9 0 1 9 I 1 0 0 1 9 9 11 2 2 5107021 0 2 052174 o 1 9 9 1 1 9 11 1 1 O 11 11 5107022 0 2 052174 0 X 9 9 9 9 9 0 0 1 O 1 11 0 0 5107023 0 2 052174 0 1 9 9 1 1 9 11 1 1 1 11 11 X 9 1 1 1 1 5107024 0 2 052174 0 9 9 11 1 X 11 19 5107025 0 2 052174 0 1 9 9 9 2 9 1 1 1 X 1 1 1 11 5108001 0 2 061674 0 1 9 9 1 1 9 11 1 1 1 11 11 5106002 0 2 061674 I 9 0 2 9 1 1 11 1 9 9 19 11 5106003 0 2 061674 1 9 1 1 9 1 O 11 1 9 9 11 2 2 5106004 0 2 061674 0 1 9 9 1 1 9 11 1 1 O 11 11 5108005 0 2 061674 1 9 1 2 9 1 i 0 0 1 9 9 11 2 2 5106006 0 2 061674 0 1 9 9 1 1 9 11 1 1 0 1 1 1 1 5108007 0 2 061674 0 1 9 9 1 1 9 91 1 1 9 11 01 5108008 0 2 061674 1 9 1 9 9 2 1 0 0 1 9 9 11 11 5106009 0 2 061674 1 9 1 1 9 1 O 0 0 1 9 9 11 22 5108010 0 2 052974 1 9 0 1 9 1 1 0 0 1 9 9 11 0 0 5108011 0 2 052974 0 1 9 9 1 1 9 1 1 1 1 1 11 11 5108012 0 2 05 2974 0 1 9 9 1 I 9 11 1 1 1 1 1 11 5108013 0 2 052974 I 9 1 1 9 1 1 0 0 1 9 9 11 2 2 5108014 0 2 052974 I 9 0 3 9 1 0 11 1 9 9 11 2 2 5108015 0 2 052974 I 9 1 2 9 1 1 0 0 1 9 9 11 22 5108016 0 2 052974 o 1 9 9 9 2 9 0 0 1 9 1 11 11 5108017 0 2 052974 1 9 1 i 9 i 1 0 0 1 9 9 11 2 2 5106018 0 2 052174 0 1 9 9 1 1 9 11 1 1 0 11 11 5108019 0 2 052174 0 1 9 9 9 3 9 11 1 1 1 11 11 5108020 0 2 052174 1 9 1 2 9 1 1 0 0 1 9 9 11 2 2 5106021 0 2 052174 1 9 1 1 9 1 O 11 1 9 9 11 22 5106022 0 2 052174 1 9 0 1 9 1 0 01 1 9 9 11 2 2 5108023 0 2 052174 1 9 0 X 9 1 1 11 1 9 9 11 2 2 5106024 0 2 052174 0 1 9 9 1 1 9 11 1 1 0 11 1 0 5108025 0 2 052174 1 9 0 1 9 1 0 11 1 9 9 11 22 5108026 0 2 052174 1 9 1 9 9 3 1 0 0 1 9 9 11 2 2 5106027 0 2 052174 1 9 1 1 9 1 0 o o 1 9 9 11 11 5109000 24 050974 0 1 9 9 1 1 9 11 1 O 0 11 91 O >"* O »■* fM N *■* »■* ***"< N •** •**1 C'i O O »"< •“< «-« •"•* fM■“*^ ** *■• *H «H fVJM * 4 H rM «H *H «“t h4 © 1^ Q >^{ \ f f \;r \j,^ r H M r H O f M '_f O ,_*<\(>--l<-«rHfHi-«r”tC\J--4-«r-tf-H("HrH0>J.'Hi--

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^ OOOOOOOOCOOOHHHHHHOOOOOOOOOO^H.^iHHHtHHrHHNNOKMNNNMNNnOOOOOOOOOOOHHHHHH S © ooooooooooooooooooocoooooocooooooooooocooooeeooooocooooocoooooooo g o © O O O O O O O O O O O O O OOrMrM«M»MrMrM#MrMrMfHrMf«fHfMtMrM»M.MrMrMfM.MrMrMfM*MrMrHrMiMfMCMtA«nkniAiniA(A>n«n(A>niniAtAiAtAiAtrktAtAtAiAiA«AtAtn«n«n«niXMniAiAUNkAtAU\ 94 mite g date trait 5112016 05 061674 0 1 9 9 1 9 11 1 1 0 11 11 5112017 05 061674 1 9 1 1 9 1 0 0 1 9 9 11 2 2 5112018 05 061674 O 1 9 9 1 9 11 1 1 0 11 11 5112019 05 061674 0 1 9 9 1 9 11 1 1 1 11 11 5112020 05 061674 1 9 1 1 9 1 0 1 1 9 9 11 11 5112021 05 061674 1 9 X* 1 9 1 0 0 1 9 9 0 0 1 1 5112022 05 05 2274 0 1 9 9 1 9 11 1 1 0 11 11 5112023 05 052274 1 9 0 1 9 1 0 0 1 9 9 0 1 11 5112024 05 052274 1 9 0 1 9 1 0 0 1 9 9 11 11 5112025 05 052274 1 9 0 1 9 1 0 0 1 9 9 11 lO 5112026 05 052274 1 9 0 2 9 0 0 0 1 9 9 11 21 5112027 05 05 2274 o 1 9 9 1 9 11 1 0 0 1 1 11 5112028 05 052274 1 9 1 1 9 1 11 1 9 9 11 0 0 5112029 05 052274 1 9 1 1 9 0 0 0 1 9 9 11 11 5112030 05 052274 0 1 9 9 1 9 11 1 0 1 11 11 5112031 05 052274 1 9 1 1 9 1 0 0 1 9 9 1 1 11 5112032 05 050974 1 9 1 1 9 1 0 0 1 9 9 11 0 0 5113000 0 1 050874 0 1 9 9 1 9 11 1 1 0 1 1 0 0 5113000 0 1 050874 1 V 0 1 9 0 0 0 1 9 9 11 0 0 5114000 0 1 050874 o I 9 9 1 9 11 1 1 0 11 0 0 5114000 0 1 0508 74 1 9 1 3 9 O 0 0 1 9 9 11 0 0 5114001 0 2 061674 0 1 9 9 1 9 11 1 1 0 11 0 0 5115000 14 051774 0 1 9 9 1 9 11 1 1 0 1 1 11 5115000 24 051774 1 9 1 1 9 1 11 1 9 9 11 0 0 5115001 05 061674 1 9 1 3 9 1 11 1 9 9 11 2 2 5115002 05 061674 o 1 9 9 1 9 11 1 9 1 11 11 5115003 05 061674 1 9 1 1 9 1 11 1 9 9 11 2 1 5116000 14 051774 0 1 9 9 1 9 11 I 1 1 11 0 0 5116000 24 051774 1 9 1 3 9 1 11 1 9 9 11 2 2 5116001 05 061674 0 1 9 9 1 9 1 1 1 1 0 11 11 5116002 05 061674 1 9 1 1 9 0 0 0 1 9 9 GO 2 2 5116003 05 061674 o I 9 9 1 9 11 1 1 O 11 91 5116004 05 061674 1 9 1 1 9 0 0 0 1 9 9 11 2 2 5117000 24 051774 1 9 1 1 9 1 0 0 1 9 9 11 2 2 5117001 05 051774 0 0 9 9 9 9 99 9 9 9 99 99 5200000 0 0 022774 0 2 9 9 1 9 11 1 1 0 91 90 5200000 0 0 022774 1 9 1 2 9 0 11 1 9 9 11 99 5201000 14 042474 o 0 9 9 1 9 11 1 1 0 11 11 5201000 24 042474 1 9 1 3 9 1 11 1 9 9 11 2 0 5202000 14 050974 0 0 9 9 1 9 0 0 1 9 1 11 0 1 5202000 24 050974 1 9 1 0 9 0 11 1 9 9 1 1 0 1 5203000 0 1 050874 0 1 9 9 9 9 11 1 1 0 11 11 5203000 01 0508 74 1 9 1 1 9 0 0 0 1 V 9 11 2 2 5204000 14 051774 0 1 9 9 1 9 11 1 1 1 11 11 5204000 24 051774 1 9 1 1 9 1 11 1 9 9 11 2 2 5205000 14 051774 0 1 9 9 9 9 11 1 0 1 11 09 5205000 24 051774 1 9 1 1 9 1 11 1 9 9 11 11 5501000 14 042474 0 1 9 9 1 9 11 1 1 0 11 11 5501000 24 042474 1 9 1 3 9 0 0 0 1 9 9 11 1 0 5501001 05 050774 1 9 1 1 9 0 11 1 9 9 11 90 5501002 05 050774 1 9 1 9 9 1 11 1 9 9 11 Ol 5502000 14 042474 0 1 9 9 1 9 11 1 1 0 99 0 0 5502000 24 042474 1 9 1 1 9 0 11 1 9 9 99 99 5502001 05 061474 1 9 1 3 9 1 11 1 9 9 11 2 2 5502002 05 061474 1 9 1 1 9 0 11 1 9 9 11 2 2 5502003 05 061474 1 9 1 1 9 O 11 1 9 9 11 2 2 5502004 05 061474 1 9 1 1 9 1 11 1 9 9 11 2 2 5502005 05 061474 1 9 1 1 9 1 11 1 9 9 0 1 2 2 5502006 05 052374 1 9 1 1 9 0 11 1 9 9 0 0 2 2 5502007 05 052374 o 1 9 9 1 9 11 1 1 0 1 1 11 5502008 05 052374 1 9 1 1 9 1 11 1 9 9 11 2 2 5502009 05 052374 1 9 1 1 9 0 Ol 1 9 9 11 2 0 5502010 05 052374 1 9 1 1 9 0 11 1 9 9 11 2 2 5502011 05 050774 1 9 1 2 9 0 11 1 9 9 11 92 5502012 05 050774 0 1 9 9 1 9 11 1 1 0 11 09 5502013 05 050774 1 9 1 1 9 0 11 1 9 9 11 2 2 95 mite g date trait 550201* 05 05077* 0 1 9 9 1 1 9 11 1 1 0 11 0 0 5502015 05 05077* 0 1 9 9 1 1 9 11 1 1 0 11 11 5502016 05 C5077* 1 9 0 1 9 1 1 11 1 9 9 11 2 2 5502017 05 05077* 1 9 1 9 9 2 1 11 1 9 9 11 11 5502018 05 05077* 0 1 9 9 9 2 9 11 1 1 1 11 11 5502019 05 05077* 1 V 1 1 9 1 1 0 0 1 9 9 11 22 5503000 2 * 05097* O 1 9 9 1 1 9 99 1 9 9 11 10 5503000 1 * 05097* 1 9 1 1 9 1 0 11 1 9 9 11 11 5503001 05 C61*7* 0 1 9 9 1 1 9 11 1 1 0 11 11 5503002 05 061*7* 1 9 1 1 9 1 0 11 1 9 9 11 2 2 5503003 05 061*7* 0 1 9 9 1 1 9 11 I 1 0 1 1 11 550300* 05 061*7* 1 9 1 1 9 1 O 11 1 9 9 11 2 2 5503005 05 052*7* 1 9 1 1 9 1 1 11 1 9 9 11 11 5503006 05 052*7* 0 1 9 9 1 1 9 11 1 0 1 11 10 5503007 05 052*7* 1 9 1 2 9 1 1 11 1 9 9 11 11 5503008 05 052*7* 0 1 9 9 1 1 9 11 1 0 1 11 11 5503009 05 052*7* 1 9 0 3 9 1 0 11 1 9 9 11 1 2 5503010 05 052*7* 0 1 9 9 1 1 9 11 1 1 0 11 11 5503011 05 052*7* 1 9 1 3 9 1 O 11 1 9 9 11 22 6000000 0 0 02277* 0 1 9 9 1 1 9 11 1 1 0 19 91 6 0 0 0 0 0 0 0 0 02277* 1 9 1 1 9 1 1 11 1 9 9 99 0 0 6001000 1 * 0*2*7* 0 1 9 9 1 1 9 21 1 1 0 11 0 0 6001000 2 * 0*2*7* 1 9 1 0 9 1 0 11 2 9 9 11 22 6001001 05 061*7* 1 9 1 1 9 1 0 11 1 9 9 11 22 6001002 05 061*7* 1 9 1 1 9 1 0 11 1 9 9 11 2 2 6001003 05 061*7* 0 1 9 9 1 1 9 11 1 1 0 11 0 0 600100* 05 061*7* 1 9 1 3 9 1 1 11 1 9 9 11 2 2 6001005 05 061*7* 0 1 9 9 1 1 9 11 2 1 1 11 0 1 6001006 05 061*7* 1 9 1 O 9 1 I 11 1 9 9 2 2 2 2 6001007 05 061*7* 1 9 1 1 9 1 0 11 I 9 9 11 22 6001008 05 05237* 1 9 1 O 9 1 1 11 1 9 9 12 22 6001009 05 05237* 1 9 1 2 9 1 0 11 1 9 9 11 2 2 6001010 05 05237* 1 9 1 1 9 1 1 11 1 9 9 11 2 2 6002011 05 05237* 0 1 9 9 1 1 9 21 1 1 0 11 11 6 0 0 1 0 1 2 05 05237* 1 9 1 3 9 1 9 11 1 9 9 21 2 2 6001013 05 05237* 0 1 9 9 1 1 9 11 1 1 0 11 11 600101* 05 05 237* 1 9 O 1 9 1 1 OO 1 9 9 IO 2 2 6001015 05 05237* 1 9 1 3 9 1 1 11 1 9 9 11 2 2 6001016 05 05077* 0 1 9 9 1 1 9 11 1 1 1 11 0 0 6002027 05 05077* 1 9 1 9 9 3 0 XI 1 9 9 2 0 22 6001018 05 05077* 1 9 1 1 9 1 1 11 1 9 9 11 2 2 6001019 05 05077* 0 1 9 9 1 1 9 11 1 1 0 11 91 6002000 2 * 05097* 0 1 9 9 9 9 9 11 1 O 0 11 11 6002000 1* 05097* 1 9 1 3 9 1 0 11 1 9 9 11 0 0 6003000 2 * 05177* 0 1 9 9 1 1 9 11 1 0 0 11 11 6003000 1 * 05177* 1 9 1 2 9 1 1 11 2 9 9 11 29 6100000 0 0 02277* 0 1 9 9 1 1 9 11 1 1 1 11 09 6100000 0 0 02 277* 1 9 1 O 9 1 1 11 1 9 9 99 09 6101000 Ol 05087* 0 1 9 9 9 0 9 11 1 0 0 11 0 0 6101000 0 1 05087* 1 9 1 3 9 1 0 11 1 9 9 11 10 6101001 0 2 06167* 0 1 9 9 1 1 9 11 1 1 0 11 11 6201002 0 2 06167* 1 9 1 1 9 1 1 11 1 9 9 1 1 2 2 6101003 0 2 06167* 1 9 1 1 9 1 O 0 0 1 9 9 11 11 610100* 0 2 06167* 1 9 1 1 9 1 1 0 0 1 9 9 11 01 6101005 0 2 06167* 1 9 1 9 9 3 O 11 1 9 9 11 0 0 6101006 0 2 06167* 1 9 1 1 9 1 0 11 1 9 9 11 0 1 6101007 0 2 06167* 0 1 9 9 1 1 9 11 1 1 0 11 0 0 6102000 0 1 05067* I 9 1 1 9 i 1 11 1 9 9 11 0 0 6200000 0 0 02277* 0 1 9 9 1 1 9 11 1 1 0 11 1 1 6200000 OO 02277* 1 9 1 3 9 1 1 11 1 9 9 11 19 6201000 0 1 05087* 0 O 9 9 9 3 9 11 1 O 1 11 0 0 6201000 0 1 05067* 1 9 1 9 9 2 1 11 1 9 9 11 11 6201001 0 2 05297* 0 1 9 9 1 1 9 11 1 1 1 11 0 0 6201002 0 2 05217* 0 1 9 9 9 9 9 99 1 9 9 21 90 6201003 0 2 05217* 1 9 1 9 9 3 1 11 2 9 9 21 90 6202000 0 1 05087* 0 1 9 9 9 0 9 11 1 1 0 11 11 96 m ite g d ate t r a i t 6202000 01 050874 1 9 1 9 9 0 0 11 11 9 9 11 22 6202001 02 061674 0 1 9 9 1 1 9 11 11 1 0 11 11 6202002 02 061674 1 9 1 9 9 2 0 11 11 9 9 11 10 6202003 02 061674 0 1 9 9 1 1 9 11 11 1 O 11 19 6600000 00 022774 0 1 9 9 1 1 9 11 11 1 O 99 00 6600000 00 022774 1 V 1 2 9 1 1 11 11 9 9 11 22 6601000 24 050974 1 9 1 2 9 1 1 11 11 9 9 11 22 660x001 05 052474 1 9 1 9 9 3 0 11 11 9 9 11 11 6601002 05 061474 1 9 1 2 9 1 I 11 11 9 9 11 22 6601003 05 052474 1 9 1 1 9 1 0 11 11 9 9 11 22 6601004 05 052474 1 9 1 1 9 1 1 11 11 9 9 11 22 6601005 05 052474 0 1 9 9 9 O 9 11 11 1 1 11 11 6601006 05 052474 1 9 1 1 9 1 1 01 11 9 9 11 22 LIST OF REFERENCES

Brody, A. R., ed. 1971. Colloquiums Entomology of house dust allergy. Proc. North Central Branch - E.S.A. 26:57-71.

Fain, A. 1971. Deux nouvelles especes de Dermatophagoidinae de la region de Kivu (Republique Democratique du Congo) (Acarinas Sarcoptiformes). Bull. Inst. Roy. Sci. Nat. Belg. 47:1-5.

Fain, A., and J. W. J. Lowry. 1974. A new pyroglyphid mite from Australia (Acarina: Sarcoptiformes, Pyroglyphidae). Acarologia 16:331-339.

Fain, A., and J. E. M. H. van Bronswijk. 1973. On a new species of Dermatophagoides (j>. n?ftte*igalia) from house dust, producing both normal and heteromorphic males (Sarcoptiformes: Pyroglyphidae). Acarologia 15s181-187.

Filipponi, A. 1964. Experimental taxonomy applied to the Macrochelidae (Acari: Mesostigmata). Pages 92-100 ia G. 0. Evans, ed. Ftoc. 1st Int. Congr. Acarology. Akad. Kiado, Budapest.

Furumizo, R. T. 1973. The biology and ecology of the house-dust mite Dermatophagoides farinae Hughes, 1961 (Acarina: Pyroglyphidae).Ph.D. Thesis. University of California, Riverside, California. 142 pp.

Gridelet, D., and P. LeBrun. 1973. Contribution a 1*etude ecologique des acariens des poussieres de maisons. Acarologia 15:461-476.

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