72-4557

MANISCHEWITZ, Jack Roger, 1942- - • A NUMERICAL PHENETIC STUDY OF THE SNAKE MITES OF THE FAMILY IXOD ORHYNCHIDAE (ACARI: MES0STI6MATA).

The Ohio State University, Ph.D., 1971 Entomology

| University Microfilms, A XEROX Company, Ann Arbor, Michigan

— a . •*- ' T

THTR T1TRRERTATION HAS BEEN MICROFILMED EXACTLY AS RECEIVED A NUMERICAL PHENETIC STUDY OF THE SNAKE MITES OF THE

FAMILY IXODORHYNCHIDAE (ACARI: MESOSTIGMATA)

DISSERTATION

Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University

By

Jack Roger Manischewitz, B.A., M.S

******

The Ohio State University 1971

Approved by

Adviser Department of Entomology PLEASE NOTE: Some Pages have Indistinct print. Filmed as received. UNIVERSITY MICROFILMS ACKNOWLEDGMENTS

i

The following individuals and institutions graciously provided

the specimens used in the present study: J. Camin (University of

Kansas), A. Fain (Institut de Medecine Tropicale, Antwerp, Belgium),

2. Feider (Universitatea "Al. I. Cuza," Jasi, Romania), W, Voss (Fort

Worth Children*s Museum), The Acarology Laboratory (Columbus, Ohio),

J. Cooreman (Institut Royal des Sciences Naturelles de Belgique,

Brussels), and M. Naudo (Museum National D'histoire Naturelie, Paris).

Drs. F. Rohlf and J. Kishpaugh (SUNY, Stony Brook) generously provided.the computer program NT-SYS. Thomas Kozlowski provided aid

in the use of computers. Free computer time was donated by the

Instructional and Research Computer Center of The Ohio State University.

I would like to acknowledge gratefully the important guidance and assistance provided by my adviser, Dr. Donald Johnston. This study was supported in part by a predoctoral traineeship from the National

Institutes of Health Training Grant 5 T01 A100216.

ii VITA i

March 31, 1942. . . . Born - Cincinnati, Ohio

1964...... B.A., Rutgers— The State University, New Brunswick, New Jersey

1964-1965 ...... Research Assistant, Department of Entomology and Economic Zoology, Rutgers— The State University, New Brunswick, New Jersey

1966...... M.S., Rutgers— The State University, New Brunswick, New Jersey

1966-1969 ...... Teaching Assistant, The Ohio State University, Columbus, Ohio

1969-1971 ...... Acarology Predoctoral Trainee, Acarology Laboratory, The Ohio State University, Columbus, Ohio

PUBLICATION

Manischewitz, Jack. 1966. Studies on Parasitic Mites of New Jersey. Jour. New York Ent. Soc. 74:189-97.

FIELDS OF STUDY

Major Fields: Zoology and Entomology

Studies in Acarology and Numerical Taxonomy. Professors D. E. Johnston and 6 . W. Wharton

iii TABLE OF CONTENTS

i

ACKNOWLEDGMENTS...... ii

VITA...... Ill

LIST OF TABLES...... v

LIST OF ILLUSTRATIONS...... Vi

INTRODUCTION...... 1

MATERIALS AND METHODS...... 4

RESULTS AND DISCUSSION...... 15

Predictiveness Determination and a Consideration of Weighting...... 15

The Ixodorhynchidae ...... 31

SUMMARY...... 53

APPENDIX

I. LIST OF O T U ' S ...... 55

II. CHARACTERS U S E D ...... 62

III. BASIC DATA MATRIX FOR 103 SPECIMENS...... 67

FIGURES...... 112

REFERENCES ...... 120

iv LIST OF TABLES

Table

1. List of Included Species and Their Code Names...... 5

2. Summarization of Characters...... 7

3. Stability Predictiveness and Character Predictiveness of Similarity Matrices...... 24

4. Stability Predictiveness and Character Predictiveness of Rienograms...... 25

v LIST OP ILLUSTRATIONS

Figure

1. Standardization M.C.D.. Phenogram...... 113

2. Graph Analysis of Ixodorhynchus. Ixobioides. and Ixodorhvnchoides, Based on the Standardization M.C.D. Similarity Matrix...... 115

3. Graph Analysis of Hemilaelaps, Scutanolaelaps. Strandtibbettsia . and Omentolaelaps. Based on the Standardization M.C.D. Similarity Matrix, ...... 117

4. Graph Analysis of Nearest Neighbors of Hemilaelaps triangulus, Based on the Standardization M.C.D. Similarity Matrix ...... 119

vi INTRODUCTION

The present study examines relationships within the family

Ixodorhynchidae Ewing, 1923, using the principles and techniques of numerical taxonomy. In addition, an attempt is made to provide infor­ mation relevant to numerical taxonomy in general.

A variety of transformations and similarity coefficients were used in producing a variety of similarity matrices and phenograms

(Sokal and Sneath, 1963). The techniques of principal components

(Harman, 1967) and graph analysis (Moss, 1967) were also used.

Prediction is often regarded as the major goal of classification

(Gilmour, 1940; Inglis, 1970; Sokal and Sneath; 1963). However, there has been little attempt to determine empirically the relative predictive success of similarity matrices or phenograms produced by alternative methods. Such an attempt is made in this study, primarily on the basis of randomly dividing characters into groups and finding the correlations between various matrices and phenograms.

As a result of a consideration of the meaning of character weighting, a new transformation, equalization, is proposed. It is designed to give characters equal weight.

In most numerical taxonomic studies, OTU's are species or other groupings of individuals. However, an alternative method, in which individual specimens are the OTU's, is also used (e.g., Moss, 1967, and

Herrin, 1969). The present study used both methods.

1 Ixodorhynchids are gamasine mites ectoparasitic on snakes..

Radovsky (1969) believes the Ixodorhynchidae arose from the Laelapinae,

and gave rise to the Omentolaelapidae and Eiitonyssidae.

Fain (1962) reviewed the Ixodorhynchidae, and readers interested

in the history of the family prior to 1962 should consult this work.

Fain's study was based on conventional taxonomic methods. Taxa were

determined on the basis of few characters, primarily those of chelicerae,

cornlculi, and coxal setae. He recognized the following genera:

Ixodorhynchus Ewing, 1923, Ixobioides Fonseca, 1934, Hemilaelaps Ewing

1933, Asiatolaelaps Fain 1961, and Strandtibbettsla Fain, 1961. He

further divided Hemilaelaps into 4 "groups" on the basis of coxal setae:

tri'angulus group, farrieri group, piger group, and ophidius group, the

. last corresponding to the genus Scutanolaelaps Lavoipierre, 1958, sunk

by Fain on the basis that one species, £». upembae Fain, 1961, was inter- l mediate between the two genera.

In 1962, • Johnston described Ixodorhynchus neodelphus. I, faini,

and the new genus and species Ixodorhynchoides truncatus. Voss and

Strandtmann (1962) described Ixodorhynchus uncatissimus, which,

according to Fain's criteria, would be placed in Ixobioides. Taufflieb

(1967), unaware of the Fain (1962) paper sinking Scutanolaelaps.

described Scutanolaelaps mehelyae. Voss (1967) described Hemilaelaps

phillippinensis. which because he believed it to be intermediate between Hemilaelaps and Asistolaelaps. caused him to synonymize

Asiatolaelaps with Hemilaelaps. H. lioheterodon was described by Fain

(1967). J. H. Gamin, University of Kansas, has discovered a new species, which, according to Fain's (1962) criteria, would belong in

Strandtibbettsia; however, Gamin has suggested that it represents a new genus.

The history of the family has been marked by the sole use of traditional taxonomic methods, including use of few characters in determining taxa. Due in part- to the emergence of papers during the same year (3 in 1962 and 3 in 1967), workers have sometimes been unaware of each other's work. Further, a preliminary examination of some characters not considered by Fain suggested that some generic changes would be appropriate, particularly with regard to Ixodorhynchus,

Ixobioides. Strandtibbettsia. and Asiatolaelaps. The above factors have contributed to the present state of generic uncertainty, which hopefully can be ameliorated by the present study. MATERIALS AND METHODS i

A total of 103 specimens, representing 33 taxonomic species were included in the study. All were adult females, as immature stages and males, were not generally available. Additional available specimens were omitted on the grounds that they had too much missing data, or that the species involved were already well-represented. Within each species an attempt was made to use specimens with differing host-locality data.

The number of specimens used per species varied from 22 to 1. Of the

32 currently recognized species of Ixodorhynchidae, all were available ' for inclusion except Hemilaelaps imphalensis (Radford, 1947). Also included were representatives of a new species found by Camin, and

Omentolaelaps mehelyae Fain, 1961, the single member of the family

Omentolaelapidae. Table 1 lists the species used and gives the code names used in the phenograms and graph analyses. Appendix I lists the specimens used, and gives host-locality data.

Specimens were observed by means of a phase contrast microscope

(American Optical). A micrometer disc in the eyepiece was used to obtain measurements.

A total of 190 characters were originally considered, and the states of these 190 characters in each of the 103 specimens were recorded. For continuous characters, the states correspond to the measurement in microns.

4 5

TABLE 1

List of Included Species and Their Code Names

Ixodorhynchus Ewing, 1923

£. liponyssoides Ewing, 1923, type species (LIP)

jl. neodelphus Johnston, 1962 (NEO)

I_. faini Johnston, 1962 (FAI)

I. leptodelrae Fain, 1962 (LEP)

1.. cubanensis Fain, 1962 (CUB)

I_. johnstoni Fain, 1961 (JOH)

Jt. uncatissimus Voss and Strandtmann, 1962 (UNC)

Ixobioides Fonseca, 1934

1., butantanensis Fonseca, 1934, type species (BUT)

1, fonsecae (Fain, 1961) (FON)

Ixodorhynchoides Johnston, 1962

£• truncatus Johnston, 1962, type species (TRU)

■ Hemilaelaps Ewing, 1933

H. lioheterodon Fain, 1967 (LIO)

H. philippinensis Voss, 1967 (PHI)

H. tanneri (Tibbetts, 1954) (TAN)

. H. evansi (Fain, 1961)(EVA)

triangulus group

H. trianguluB (Ewing, 1923), type species (TRI)

H. javanensis Fain, 1961 (JAV) TABLE 1 (continued)

farrlerl group r H. farrier! (Tibbetts, 1954) (FAR)

H. congolensis Fain, 1962 (CON)

H. causicola Fain, 1961 (CAU)

H. dipsadoboae Fain, 1962 (DIP)

H. radfordi (Feider and Solomon, 1959) (RAD)

H. feideri Fain, 1962 (FEI)

H. caheni Fain, 1961 (CAH)

P-iger group

H. piger (Berlese, 1918) (PIG)

H. novaeguineae Fain, 1961 (NOV)

ophidius group

QPhidius (Lavoipxerre, 1958) (OPH)

H. schoutedenj (Fain, 1961) (SCH)

H. upembae (Fain, 1961) (UPE)

H. mehelvae (Taufflieb, 1967) (MEH)

Strandtibbettsia Fain, 1961

S. gordoni (Tibbetts, 1957), type species (GOR)

£. brasiliensiB Fain, 1961 (BRA)

New species nearest to S.. brasiliensis (NSP)

Omentolaelaps Fain, 1961

0. mehelvae Fain, 1961 (OME) Fifty-eight characters were eliminated, due to correlations (see below).

The final 132 characters and their states are listed in Appendix II.

The Basic Data Matrix for the 103 specimens is Appendix III. An attempt was made to use as many different kinds of characters as possible, within the available realm of external morphology. Table 2 contains a

summarization of characters.

TABLE 2

Summarization of Characters

Area of Body Qualitative Meristic Continuous Totals

Gnathosoma 11 3 4 18 (14%)

Idiosomal Dorsum 14 6 4 24 (18%)

Idlosomal Venter 26 4 12 42 (32%)

Legs 28 18 2 48 (36%)

TOTALS 79 (60%) 31 (23%) 22 (17%) 132

The above table may be somewhat misleading. In practice, it

Is sometimes difficult to determine in which category a character

should be placed. And if some of the qualitative characters had been coded slightly differently, they would be considered quantitative.

Forty-four characters dealt with numbers of setae present

(including whether a particular seta was present or absent). All setal terminology used is that of Evans and Till (1965).

If a series of consecutive positive integers is used to code the states of a character, ordinarily it makes no difference what. integer is used as the starting point. However, when logarithms or the

Canberra metric (modified non-metric coefficient, Ducker, Williams, and

Lance, 1965)'is used, the lower the starting point, the greater will be the weight of that character (see Results and Discussion section for information on weighting). Therefore, for qualitative characters, consideration had to be given as to what series of integers (usually 2 ) was most appropriate. It was noted that most qualitative characters could be regarded as rough approximations of something quantitative.

Therefore, those consecutive integers which seemed most nearly to approximate the quantities involved were used. For example, if the character was a presence/absence character, the absence state could be regarded as a measurement of 0 microns. Therefore, presence/absence characters were given coded states of 1 and 0. However, if the charac­ ter dealt with whether or not an area was well-sclerotized, or whether or not a seta was spine-like, the states were regarded as representing different quantities of a property present in both states, and were therefore coded as 1 and 2. The decisions involved in the selection of consecutive integers were occasionally arbitrary and the seeming desirability of making them must be regarded as a disadvantage of methods calling for them.

A related problem caused 7 characters (15, 18, 19, 21, 25, 51,

52) to be basically unsuitable for logarithmic transformation or the

Canberra metric. If, for example, the states of a character are coded

1, 2, and 3, the difference between states 1 and 2 is equal to the difference between states 2 and 3 when standardization or condensation is used, but is greater than the difference between the original states 2 and 3 if logarithms or the Canberra metric is used. While this may be acceptable for most characters, it is unacceptable for those charac­ ters for which the original coding of states could have just as appro- i priately been reversed (e.g., 3, 2, 1, rather than 1, 2, 3). For such characters it is desirable to keep the differences between states coded t as consecutive integers equal. The easiest approach is to eliminate such characters, which is what was done during the predictiveness deter­ minations (see below). However, during the rest of the study these characters were included: when logarithms were used, the seven charac­ ters simply were not converted (the Canberra metric was only used during the predictiveness determinations).

Some characters which were found to be highly correlated

(r > .500) with other characters were not included in the final Basic

Data Matrix. Many measurement characters were highly correlated.

Those measurement characters that had a high correlation with the two characters regarded as the best indicators of overall size (23 and 38) were eliminated, so that overall size differences between OTU's would not have an inordinately large effect. Gnathosomal measurements were somewhat independent of others, and some were eliminated from the Basic

Data Matrix when it was found they were highly correlated with other gnathosomal measurements as a result of just one kind of local size difference.

If a number of leg setal characters were highly correlated as a result of serial homology, all but one were eliminated, following the advice of Sokal and Sneath (1963). Similarly, setal characters were eliminated if high correlations occurred involving the same row of 10

setae on different segments of the same leg, or if high correlations

occurred involving setae of the same segment of the same leg, the only

difference between the characters being that one involved an anterior t row and the other involved the equivalent posterior row. Determination

of character correlations, and the subsequent eliminations, was carried

out on the original basic data matrix with 103 OTU's, and the characters

involved remained eliminated during the rest of the study.

Ratios are often used in numerical taxonomy in order to reduce

the great effect of overall size. However, since those measurement

characters highly correlated with the indicators of size were elimi­

nated, use of ratios (dividing all measurements by some size indicator)

would be undesirable. Such conversion would most likely result in

characters becoming negatively correlated with size, which is no

improvement.

Some numerical taxonomists have deliberately eliminated highly

variable characters (^ss, 1967, Ehrlich, 1967). However, the judgment

of which characters are highly variable often presupposes knowledge of

the species or other taxa under study, which cannot be presupposed if a

purpose of the study is to determine such taxa. Even if such knowledge

is not presupposed, highly variable characters should not be eliminated,

according to Sokal and Sneath (1963). Characters for which most of the variation is known to be environmentally-induced would be legitimate exceptions. No attempt was made to eliminate highly variable charac­

ters in the present study.

To have excluded every specimen with missing data, as some have done (Herrin, 1969), would have resulted in excluding some species 11 entirely. Therefore, specimens with missing data were included, using the NC option (Sokal and Sneath, 1963). However, specimens with large numbers of NC's were excluded and no specimen was included that had more than 11 NC's. Sixty of the 103 specimens used had no NC's. The small number of NC's included is well within suggested limits (ibid.,

Crovello, 1968b, 1969).

In order to determine whether or not the current species are taxonomically valid, specimens were used as OTU's, using standardized data (Sokal and Sneath, 1963) with the correlation coefficient (ibid.), and with the average taxonomic distance coefficient (ibid.). Clustering was done using UPGMA (ibid.).

To provide insight into inter-specific relationships, further runs were made using species rather than specimens as OTU's. Currently recognized species, plus Camin's new species, were used. Data were obtained by taking averages of appropriate values.in the original basic data matrix based on 103 specimens. Three characters previously used

(75, 108, 117) became invariant, after averages were taken, and were excluded. Nine similarity matrices and phenograms were obtained, using all combinations of three transformations and three similarity coeffi­ cients: standardization, logarithms (to base 1 0 ) with subsequent standardization, condensation (Crovello, 1968a), correlation coefficient, average taxonomic distance coefficient, and M.C.D. (Cain and Harrison,

1958). As before, UFGMA was the clustering method used. Cophenetic correlation coefficients were obtained to determine how consistent the phenograms were with their corresponding similarity matrix (Sokal and

Rohlf, 1962). 12

A minor problem Involving use of logarithms is that the loga­

rithm of "0 " cannot be taken. Therefore, when logarithms were to be used, all "0 "s would first be placed by 0 .2 , an arbitrarily chosen i number which resulted in quantity differences between "0 " and other

states which seemed pleasing.

The exact form of the Canberra metric used was:

• X l k l s xl< + xlk jk n lxn ‘ xikl When both X.. and X.. were "0", the quantity - ■ J . 7 - was assigned ij ik the value "O'1.

Predictiveness determinations were performed on similarity matrices and phenograms produced by using the combinations of trans­ formations and similarity coefficients mentioned above, as well as other combinations including logarithms alone, no transformation, and the Canberra metric. This phase of the study was carried out to deter­ mine tentatively which methods are, in general, most predictive. The rationale behind the procedures involved is explained in the Results and Discussion section. The only basic data matrix used was that in which the OTU’s were species.

First, a table of random numbers (Rohlf and Sokal, 1969) was used to divide the characters randomly into two groups. To determine stability predictiveness, a particular combination of transformation and similarity coefficient was used independently on both subsets of 1 characters. Then the correlation (Pearson's product-moment correlation coefficient) was found between the resulting similarity matrices. 13

Sometimes, phenograms were obtained using UPGMA., and the correlation was then found between the two sets of cophenetic values taken from the two phenograms. The higher the correlation, the greater is the sta­ bility predictiveness of the combination of procedures involved. To determine character predictiveness, the procedures were identical except that one of the two similarity matrices was always produced by using standardized logarithms with the M.C.D. (see Results and Discus­ sion section).

Differences between the resulting correlations cannot be sub­ jected to conventional significance tests because such tests require an assumption of bivariate normal distributions which cannot be made in the present case. Therefore, the data set was divided into two subsets, as described above, a total of four times and most tests were done four times. This permits subjective judgments to be made con­ cerning significance of correlation differences (Rohlf, 1965),

It is possible to rank all the correlations which resulted from using different methods on the same pair of subsets, and to compare this ranking with a similar ranking based on another pair of subsets by i using Spearman's Rank Correlation Coefficient (Siegel, 1956). This coefficient was also used to determine the correlation between stability predictiveness and character predictiveness, for similarity matrices and for phenograms. Similarly, rank correlations were obtained which com­ pare predictiveness correlations of similarity matrices with those of phenograms.

Graph analysis (Moss, 1967) was employed on the similarity matrix resulting from standardization (no logarithms) and the M.C.D. 14

(OTU's equals species). Principal component procedure was used on the similarity matrix resulting from use of standardization and the corre­ lation coefficient (OTU's equals species). The first five factors were i extracted.

The computer program NT-SYS (prepared by F. Rohlf, J. Kishpaugh, and D. Kirk, all of SUNY, Stony Brook, N.Y.) was used in connection with standardization, correlation coefficients (other than Spearman's), average taxonomic distance coefficients, cophenetic correlation coeffi­ cients, UPGMA, and principal components. New programs were written for condensation, the M.C.D., and the Canberra metric. The IBM 360/75 was used. RESULTS AND DISCUSSION i

Predictiveness Determination and a Consideration of Weighting

A major goal of taxonomy is the production of the most predic­

tive matrices, phenograms, and taxa, possible. Numerical taxonomists

in particular have emphasized the importance of prediction. Therefore,

an important criterion (probably the most important) which should be

used in judging which alternative classificatory procedures are most

desirable, is predictiveness of results. Two basic approaches to deter­

mining relative predictiveness are possible: theoretical-mathematical

and empirical. Thus far, most justification for using particular methods has been theoretical (not counting simply preferring methods

because the results are subjectively most pleasing). Little attempt

has been made to determine objectively and empirically the most predic­

tive methods, or to devise means for doing so. That a variety of

transformations, similarity coefficients, and clustering techniques are

currently in use, each having their own proponents, attests to the

failure of theoretical-mathematical arguments to demonstrate convinc­

ingly which methods are most desirable.

A number of recent authors (Inglis, 1970, Mannetje, 1967, and

Lambert and Williams, 1966) have rightfully pointed out that empirically 16

examining the results from using alternative methods is the most appro­

priate approach to judging methods. A number of studies have made a I start in this direction (see below).

Two basic means of determining predictiveness are used in this

study. One, herein referred to as "stability predictiveness," attempts

to determine the likely relative stability of phenograms, etc., when new characters are added. Stability has been referred to by Williams and Dale (1965) as "the basic requirement" of taxonomy. While their

statement appears to be an exaggeration, stability is important from the point of view of information retrieval, and as an indicator of character predictiveness (see below). In determining stability predic­ tiveness, it would be possible to add new characters to the original

set, and thereby duplicate the conditions under which differing degrees of stability normally occur. However, the difference between the

stability ratings of alternative methods can be increased if two entire­

ly different sets of characters are used, as was done in this study.

The correlation between the two resulting matrices, or cophenetic values

of phenograms, then Indicates the relative stability predictiveness of matrices or phenograms resulting from particular alternative methods.

Rohlf (1965) appears to be the only previous worker who divided charac­ ters randomly into two groups and determined the correlation between the resulting similarity matrices (although he was not deliberately determining stability predictiveness).

Although in the present study characters were randomly divided into two'groups, there is no apparent reason why obtaining the two character subsets by dividing characters along morphological lines, 17

such as leg vs. non-leg, or adult vs. larval, is less valid. Six stud­

ies (that have come to my attention) have tested the Non-Specificity

Hypothesis (Sokal and Sneath, 1963) using both the distance coefficient and the correlation coefficient. The results can be used to show

stability predictiveness although this was usually not considered by

the author. However, Rohlf (1963), on finding a higher correlation between larval and adult characters when distance coefficients were used than when correlation coefficients were used, listed this finding as one reason why he preferred distance. In three of the six studies

(Rohlf, 1963, Ehrlich and Ehrlich, 1967, and Moss, 1968), the pair(s) of subsets on which taxonomic distance was used had a higher correla­ tion^) (stability predictiveness rating) than the pair(s) of subsets on which correlation was used. In the other three studies (Mlchener and Sokal, 1966, Hendrickson and Sokal, 1968, and Schnell, 1970) the opposite occurred. In the Schnell study, however, distance was at a disadvantage, due to a tremendous size factor (size affects distance more than correlation) which was usually not compensated for.

In some of the above studies more than one pair of stability I predictiveness values was obtained (e.g., when a non-specificity test was done using leg vs. non-leg characters, as well as dorsal vs. total characters). In such cases, the results were often consistent within a study. Therefore, in summing the results from different studies it is probably better to consider each study as a separate unit than to consider each individual test within a study as a separate unit.

As OTU's are basically treated in numerical taxonomy as con­ glomerates of characters, it would be expected that stability 18

predictiveness would also indicate relative character predictiveness.

However, to determine character predictiveness, it would be more

appropriate to use procedures which more closely approximate the way

in which character predictions are actually carried out.

At least four recent studies have considered character predic­

tiveness. Clifford, Williams, and Lance (1969), used the Canberra

metric and information statistic, and concluded, apparently subjec­

tively, that the phenogram resulting from the information statistic was more consistent with results of hybridization tests of the plant

taxa under study.

Mannetje (1967), studying legumes, used seven combinations of

\ correlation (with standardization), taxonomic distance (with standardi­

zation), non-metric coefficient (Canberra metric), probabilistic index

(of Goodall), nearest neighbor sorting, furthest neighbor sorting, centroid sorting, and flexible sorting. He determined how consistent

the resulting phenograms were with a previous division of the OTU's into two groups on the basis of successful symbiosis with various

Rhizobium. The phenograms were designated as satisfactory or unsatis­ factory. The satisfactory phenograms were those resulting from taxonomic distance with furthest neighbor sorting, taxonomic* distance with flexible sorting, and the non-metric coefficient with flexible sorting. The unsatisfactory phenograms were those of correlation with nearest neighbor sorting, distance with nearest neighbor sorting, correlation with furthest neighbor sorting, and the non-metric coeffi­ cient with centroid sorting. Eades (1970) used his Index of Matrix Reliability, which is based on determining how consistent a similarity matrix is with a presumably reliable division of the OTU's into groups, such as members of the same species or clone. He subtracted the mean of similarity coefficients of pairs of OTU's belonging to different groups (e.g., members of different species) from the mean of similarity coefficients of pairs of OTU's belonging to the same group, then dividing the difference by the overall standard deviation of similarity coefficients.

He used the correlation coefficient, taxonomic distance, and the M.C.D.

Taxonomic distance and the M.C.D. rated similarly, the results not consistently indicating which of the two was superior. However, both were superior to correlation.

The method of Eades appears acceptable, if the new character against which a matrix is being tested is either two-state, or quali­ tative multi-state, and if it is acceptable that the difference between all pairs of states is treated identically.

Rhodes et al. (1968) appear to be the only workers who used the correlation coefficient to evaluate how consistent a new character is with the previously determined similarity matrices and phenograms. The new character was cross-compatability (i.e., hybridization) ratings in the form of a matrix. The similarity coefficients used were the corre­ lation coefficient, taxonomic distance, and Clark's divergence coeffi­ cient (Sokal and Sneath, 1963). Clustering methods used were UPGMA,

WPGMA, UPGM, and WPGM. Similarity matrices resulting from use of

Clark's divergence coefficient, and taxonomic distance had correlations of 0.82 and 0.80 respectively, while that of the correlation matrix was 20 only 0.66. The phenograms showed the same trend. Evaluation of clus­ tering methods was more difficult, as their results were somewhat inconsistent. However, on the average, U P G M was superior to the other methods (my conclusion). Curiously, when the authors tried to subjec­ tively evaluate the phenograms on the basis of other previously unused

> considerations, such as geography and ecology, they regarded the corre­ lation phenograms as superior to those of distance and the divergence coefficient.

It may be noted that none of the above studies objectively con­ siders more than one new character at a time. However, no one character, including hybridization, dan be regarded as necessarily indicating which methods are most predictive, just as no one character is likely to yield as good a classification as one based on many characters. Therefore, it is better to use a method of determining character predictiveness which incorp9 rates many characters.

In developing a method for determining character predictiveness, it is appropriate to consider how prediction might actually take place.

If it is desired to predict the character state of an OTU from the known states of other OTU's, one approach would be to find the OTU (in which the state of that character is known) with which the original OTU has the greatest similarity, and predict that both OTU's have the same state. However, this is not the most desirable procedure. It should be apparent that if the original OTU has a similarity coefficient of

0.10000 with one OTU and 0.10001 with another, that both OTU's should make approximately equal contributions to determining what state should be predicted. The problem is rather similar to that of determining 21

mathematical expectation by multiplying a value for each possible

outcome by the probability.of its occurring, and summing. Thus in

I predicting a character state, each OTU for which the state is known

should make a contribution proportional to its similarity with the

original OTU. If a type of similarity coefficient is being used in

which low values indicate high similarity, the appropriate formula is:

Predicted Xt, - 3* “ ------’js J l . . ij n 1 S Sjx X*k JX

In practice, prediction is only likely to be used with regard

to character states that are difficult to determine. If the character

state is easy to determine, there is no need to predict— one can simply

observe the possessed state. Therefore, character prediction will occur

most often when the states of the character involved are known for only

a few, or a single, OTU in the study. As can be seen from the above

formula, if the character state is known for only one OTU, the predicted

state for the OTU whose character state is being predicted, is simply

the known character state value.

Ideally, when one is testing character predictiveness, tests would be conducted involving a varying number of known character states.

However, because there is no reason to expect that a different number of

known states would affect which methods produce roost predictive results,

and because it is simplest to conduct'such tests when only one character

state value is presumed known, only this case is considered further in

this study. 22

— — ^Prediction error" may be def ined-as .the absolute value of the

difference between the predicted value and the actual value. In a

character predictiveness test, the character state value for a particu­

lar OTU can be predicted anew using each other OTU individually; thus

a matrix of prediction errors for a character can be formed which has

the form of a similarity matrix.

If prediction is actually being carried out involving just one

character, and a value of 100 is predicted for a particular OTU, and

the actual value turns out to be 90, one would probably consider the

error to be.identical to the prediction error if 10 had been predicted

and the actual value were 9. Therefore, during character predictive­

ness studies it is appropriate that equal percent prediction errors

count equally. This may be accomplished by converting all basic data

to logarithms as was done in the present character prediction studies,

in producing prediction error matrices.

As was done by Rhodes et al. (1968), the correlation may be

found between the prediction error matrix (not regarded as that by

Rhodes) for a new character, and the original similarity matrix (which

did not take into account the new character). The same procedure could be repeated for many new characters, each new character resulting in a

new correlation. The results for all characters could then be combined by simply finding the mean correlation.

An alternative to the above is to reverse the last two proce­

dures, i.e., for each pair of OTU's add together the prediction error

for all new characters, obtain the mean prediction error for each OTU 23

pair, and then obtain the correlation between the mean prediction error matrix and the original similarity matrix. The latter approach involves

less effort, and as both seem equally valid, the latter was used.

However, if the prediction errors for different characters are

to be added together, a transformation is needed to give characters

equal weight. Therefore, the logarithms were standardized.

During a study of character predictiveness, "new" characters can be obtained by randomly dividing the character set into two subsets,

and using one subset to obtain the original similarity matrix, and the

other to be regarded as new characters to obtain the prediction error matrix. Table 3 shows the correlations indicating stability predictive­ ness and character predictiveness for similarity matrices.

To determine the character predictiveness of phenograms, the

same mean prediction error matrices were used as before, and the corre­

lation was then found between that matrix and the cophenetic values

from the appropriate phenograms. Table 4 shows stability predictiveness and character predictiveness of types of phenograms.

It may be noted that the mean prediction error matrix.is arrived at using the same procedures as are used to arrive at a similarity matrix using logarithms standardized, with the M.C.D. It is to be expected, therefore, when original similarity matrices are being tested,

that those matrices which were produced by methods including logarithms,

standardization, or the M.C.D. will have an advantage. However, this is certainly appropriate, if the rationale behind the procedures used

is valid. Corroborative evidence that it is valid comes from a compari­

son of rank order of results with those of the stability predictiveness TABLE 3

Stability Predictiveness and Character Predictiveness of Similarity Matrices

Correlation Distance M.C.D. Sta. Pr. Ch. Pr. Sta. Pr. Ch. Pr. Sta. Pr. Ch. Pr.

Raw Data .310 .434 .666 .787 .692 .808 .292 .379 .625 .566 .688 .623 Average .301 .407 .646 .677 .690 .716

Logarithms .541 .529 .615 .582 .757 .772 .496 ..709 .553 .729 .672 .708 Average .519 .619 .584 .656 .715 .740

S t andardiz at Ion .678 .436 .769 .759 .829 .823 .575 .293 .820 .828 .839 .831 .643 .432 .839 .825 .861 .860 .699 .432 .916 .873 .896 .887 Average .649 ■ .398 .836 .821 .856 .850

Logarithms with .690 .432 .753 .688 .832 .832 Standardization .589 .292 .826 .847 .839 .839 .653 .430 .847 .829 .861 .861 .709 .430 .902 .864 .879 .879 Average .660 .396 .832 .807 .853 .853

Condensation .767 .804 .780 .795 .808 .825 .639 .631 .699 .700 .747 .741 .801 .796 .801' .791 .816 .821 .801 .817 .802 .820 .823 .845 Average .752 . .762 .771 .777 .799 .808

Canberra Metric .771 .785 .678 .718 Average .725 .752 TABLE 4

Stability Predictiveness and Character Predictiveness of Phenograms

Correlation Distance M.C.D. Sta. Pr. Ch. Pr. Sta. Pr. Ch. Pr. Sta. Pr. Ch. Pi

Standardization .790 .461 .760 • .731 .855 .796 .662 .300 .812 .805 .851 .822 .695 .469 .836 .784 .893 .807 .755 .428 .903 .830 .938 .859 Average .726 .415 .828 .788 .884 .821

Logarithms with .736 .453 .745 .710 .872 .872 Standardization .622 .352 .825 .815 .848 .848 .680 .469 .868 .789 .888 .888 .756 .448 .908 .831 .915 .915 Average .699 .431 .837 .786 .881 .881

Condensation .844 .809 .864 .793 .897 .839 .671 .594 .724 .661 .757 .703 .843 .745 .864 .783 .886 .812 .842 .813 .854 .804 .876 .835 Average .800 .740 .827 .760 .854 .797 26

tests. The same two combinations, logarithms standardized with the

M.C.D., and standardization with the M.C.D. had the highest average

correlations in both studies, both for matrices and for phenograms.

Also, Spearman's Rank Correlation Coefficient comparing stability

predictiveness with character predictiveness (using the averages for

the nine combinations tested four times) was 0.95 for similarity matrices, and also 0.95 for phenograms.

However, because standardization does not give exactly equal weight to characters (see below), other transformations were possibly

at.a slight unfair disadvantage during the character predictiveness

tests. As the rank order of methods using standardization is as high

during the stability predictiveness tests as it was during the charac­ ter predictiveness tests, such a disadvantage presumably was small.

There is additional evidence from within the study that stan­ dardization is the most predictive transformation used (when distance or the M.C.D. is used). With regard to matrices, note that when

standardization was not used, the character predictiveness average values are always higher than the stability predictiveness values;

thus when neither subset of characters was treated with standardization, the correlation was less than if one subset was treated with standard­ ization and the other was not. Therefore, if during the character predictiveness tests, a transformation other than standardization had been used in deriving the prediction error matrix, standardization would still be demonstrated as more predictive than the transformation used in getting the mean prediction error matrix (with distance or the

M.C.D.). 27

Curiously, while standardization was shown to be superior when

distance or the M.C.D. was used, condensation was consistently shown to

be most predictive when correlation was used. In this connection, it

was noted that, subjectively, the condensatlon-correlation phenograms

appeared somewhat intermediate between the M.C.D. (and distance)

phenograms, and the standardization-correlation phenograms.

Comparing similarity coefficients, the M.C.D. consistently

proved more predictive than distance, which consistently proved more

predictive than correlation. Both of the above trends occurred both '

for matrices and phenograms, and for both kinds of predictiveness.

The Spearman Rank Order Correlation Coefficients comparing the

averages of the results from the first two random divisions of charac­

ters with those of the latter two, for similarity matrices and pheno­

grams (both stability and character predictiveness) were 0.90, 0.95,

0.87, 0.98.

Comparing the results for similarity matrices with those of

phenograms, for stability and character predictiveness, the Spearman's

Rank Order Correlation Coefficients were 0.91 and 0.97, respectively.

With regard to using standardization with logarithms or without

logarithms, when the M.C.D. is used, the results are not clear-cut.

With regard to matrices, there was virtually no difference in predic­

tiveness. With regard to phenograms, logarithms raised the character

predictiveness. However, this result is suspect, because ordinarily

one would expect the character predictiveness of a phenogram to be

somewhat lower than that of the matrix on which it is based, which was not the case here. The disadvantages of using logarithms (see Materials 28 and Methods) seem to indicate that unless use of logarithms can reli­ ably be demonstrated to raise predictiveness by a sizable amount, use of logarithms is less desirable than omitting them (except in character predictiveness studies in determining prediction error).

The M.C.D., which tentatively should be considered the most predictive similarity coefficient of those tested, unfortunately has not been used by the majority of numerical taxonomists. Among those who consider the M.C.D. to be inferior to distance are Sokal and Sneath

(1963). They claim the M.C.D. underestimates the true distance.

Johnson (1970), however, states that the M.C.D. is just as valid as a distance measurement as taxonomic distance.

Sokal and Sneath (1963) also note that the M.C.D. compares with taxonomic distance in largely the same way that average deviation com­ pares with standard deviation, and that taxonomic distance should be preferred for the same reasons that the standard deviation is preferred over average deviation. Eades (1970), however, explains that the preference for standard deviation is due to a number of conditions which ordinarily do not all occur in numerical taxonomy, and he believes the M.C.D. is more appropriate than distance (unless his kind of charac­ ter weighting is used).

Carmichael and Sneath (1969) prefer the M.C.D. to distance on the grounds that if the character state differences between two OTU's are "1" for one character and "3" for. another, that the similarity coefficient should be the same as for another pair of OTU's that have character state differences of "2" and "2” (the similarity coefficients • would be different with taxonomic distance). 29

Hall (1969) prefers the M.C.D. over taxonomic distance because taxonomic distance violates his rule that the contribution of one character should not be affected by the contribution of others.

Colless (1967) recently derived a coefficient which is virtually ‘ the same as the M.C.D. used with condensation.

Weighting of characters is generally regarded as undesirable by numerical taxonomists. However, it is impossible to determine if a transformation eliminates unintentional weighting, unless weighting is precisely defined, which, apparently, has not been done. Equal weight­ ing means equal effect in determining the similarity coefficients. If the M.C.D. is used, equal effect in determining the similarity coeffi­ cients occurs when each character makes an equal quantitative contribu­ tion to the totality of similarity coefficients. Therefore, for the

M.C.D., equal weight should be defined as the condition occurring when

2|x^ “ XjJ, over all OTU pairs is the same for all characters. For taxonomic distance, all - X ^ must first be squared. Accepting this definition, simple examples readily demonstrate that neither con­ densation nor standardization avoid weighting. For example, if two characters' original states are 0, 0, 1, 1, and 0, 1, 1, 2, and the

M.C.D. is to be used, and the characters are standardized, the first character will contribute a total of 6.9, while the second will con­ tribute 7.3. If the characters are condensed, the first character will contribute 4 and the second will contribute 3.

In order to achieve equal weighting while using the M.C.D., all that is necessary is to determine the relative weights in the original data, i.e., find S|x^ - X^j for each character and then divide 3°

all by 2|x^ - X.J. Then each character will give a total

contribution of 1. Because the division would result in extremely

small numbers, it may be appropriate to divide each - X^| by the

total number of OTU pairs, i.e. (n^-n)/2. Then the average Jx^ - X^jJ

for each character would be "1", as would be the average similarity

coefficient. Combining this transformation, for which an appropriate

name is "equalization," with the M.C.D. yields the formula:

K, ■ xikl /2lxi-XJ\ V- n)/2 m

Ideally,. equalization would have been among the transformations

used in this study. It would have been the most appropriate transfor­ mation for use in determining prediction error in the character predic­

tion tests. Unfortunately, it was arrived at too late to be used.

However, since it appears to be the only transformation which will

equally weight characters, it is hoped other workers will consider

using it. i . •

There are many questions remaining, at best, only partially

resolved in numerical taxonomy. Can a particular kind of deliberate weighting be demonstrated to be more predictive than equal weighting?

What kinds, if any, of empirically correlated characters should be

eliminated? Would it be better to somehow combine empirically corre­

lated characters (or give them less weight) than to eliminate them?

Are ratios the answer to the size problem? If a study involves a large

number of OTU's, should separate studies be done on subgroups within 31

the study? To what extent should NC's be permitted? What clustering

technique results in the most predictive phenogram? It is believed

that a proper approach to investigating all of the above questions

would include predictiveness determinations, such as those conducted

in the present study.

The Ixodorhynchidae

Eleven phenograms were produced to provide information pertain­

ing to relationships among the Ixodorhynchidae. Two phenograms were

produced using individuals as OTU's, one resulting from standardization

and correlation, the other resulting from standardization and taxonomic

distance. Nine other phenograms were produced resulting from combina­

tions of standardization, standardization of logarithms, condensation,

correlation, taxonomic distance, and the M.C.D., all involving use of

species as OTU's.

There is a strong similarity between the inter-specific rela­

tionships of the phenograms for which individuals are OTU's, and those

of the corresponding phenograms having species as OTU's. However, a

few notable differences occur.

Comparing the two phenograms based on standardization and correlation, it may be noted that the first division separates the

"Ixo group” (Ixodorhynchus. Ixobioides. and Ixodorhynchoides) from all else, in both phenograms. In the phenogram in which individuals are

OTU's, the first division of the non-Ixo group results in three groups, one being Omentolaelaps mehelyae (OME), another containing Strandtlbbettsla gordoni (GQR), Hemtlaelaps radfordi (RAD), and

H. feideri (FEI), and the third containing the remainder. In the corresponding phenogram in which species are OTU's, the first division of the non-Ixo group results in one group containing 0. mehelyae (OME),

Hemilaelaps ophidlus (OPH), H. schoutedeni (SCH), and H. mehelyae (MEH), and a second group containing the remainder. In the former phenogram,

H. evansi (EVA) joins H. ophidlus (OPH), H. schoutedeni (SCH), and

H, mehelyae (MEH), whereas in the latter phenogram H. evansi joins

H. tanner! (TAN) and H. philippinensis (PHI),

The- two phenograms resulting from standardization and taxonomic distance have fewer major differences. However, Ixodorhynchus cubanensis (CUB) is separated from other members of Ixodorhynchus in the phenogram using species as OTU's, whereas in the other phenogram

I, cubanensis (CUB) joins with other members of the genus Ixodorhynchus

(FAI, LIP, NEO, LEP).

It was found that, with regard to delimiting species, the stan­ dardization correlation phenogram gave results more consistent with past studies, and contained fewer borderline cases than the distance phenogram. This finding is somewhat surprisihg, in view of the earlier finding that distance is more predictive than correlation (when stan­ dardization is used). Perhaps the explanation is that correlation is more predictive at the tips of phenograms, where species delimitations are made, while distance is more predictive closer to the base. At the tips, qualitative characters are most likely to have the same state, thus increasing the importance of measurement characters, which are often a reflection of size (the effect of size was largely, but not entirely, eliminated earlier in the present study, as explained in

Materials and Methods). Thus in a separate study using only the 45

OTU'b belonging to I_. liponyssoides (LIP), I.* neodelphus (NEO), and

I. faini (FAI), 43% of the varying characters were measurements, while

only 17% were measurements when all 103 OTU's were used. As size

differences affect placement of OTU's in distance phenograms more than

in correlation phenograms (Sokal and Sneath, 1963, Moss, 1968), pheno­ gram differences between OTU's that are quite similar might largely reflect size differences when distance is used, but not when correla­

tion is used. The possibility that different kinds of similarity coefficients might be most predictive in different parts of their pheno­ grams appears to warrant further investigation.

Because of the above-mentioned finding, only the correlation phenogram was used in attempting to resolve the few problems that arose concerning the clustering of Individuals into species.

Hemilaelaps schoutedeni (SCH) and H. mehelyae (MEH) appear to be one species, both in terms of the phenogram and subjective judgment. In the phenogram, the holotype of H. mehelyae (MEH) joins the one specimen of H. schoutedeni (SCH) used, and then the paratype of H. mehelyae (MEH) joins the other two, at the high correlation level of 0.9250. This is a higher correlation than usually occurs between member of the same species in the phenogram.

Taufflieb, when he described H. mehelyae (MEH), was clearly unaware of H. schoutedeni (SCH), as indicated by his statement that

H. ophidius was the only prior species in the genus Scutanolaelaps 34

(Scutanolaelaps = ophidlus group), and by his comparing H. mehelyae

(MEH) with H. ophldius (OPH) rather than with its closer neighbor,

H . schoutedeni (SCH). “ j Comparing the descriptions of H. mehelyae and H. schoutedeni five differences are apparent. Taufflieb does not show the bilobed membranous pouch in front of the sternal shield, or the pointed projection extending anteriorly from coxa II, both of which are included in the H. schoutedeni description. Both structures are present in both species; they are somewhat difficult to observe in

H. mehelyae. apparently because the specimens are over-cleared.

H. schoutedeni is shown with a slightly longer peritreme, whereas, actually, the length of the peritreme of the paratype of

H, schoutedeni. and that of the holotype of H. mehelyae. are the same length (both absolutely and relative, to coxae), while those of the paratype of H. mehelyae are slightly longer.

The anal plate of H. mehelyae is shown with two slight lateral concavities, anteriorly, and with the anterior margin well-sclerotized both laterally and medially, whereas in the description of

H. schoutedeni no such lateral concavities are shown, and the medial portion of the anterior margin is not well-sclerotized. These differ­ ences do occur in the three specimens available to me. However, in other groups of OTU's, which are regarded as species, these characters vary. It is concluded that these characters often show intra-specific variation, and should not be regarded as indicating two species are involved. 35

H. schoutedeni has been collected from two different host genera, and H. mehelyae was collected from a third host genus. The areas the species were collected from are about 1000 miles apart. In i view of this wide separation, and host difference, it is probably best

to retain H. mehelyae until more information is available.

Fain synonymized Hemilaelaps (=Ophidilaelaps)capensis Till,

1957, from Africa, with H. farrieri (FAR) from Korea. The present

study tends to confirm the synonomy, as one specimen from each locality was used, and they emerged as nearest neighbors, highly correlated

(0.7750). •

Five specimens of H. philippinensis (PHI), four from Cyclcorus lineatus, and a fifth from an unidentified host, are all joined at a correlation of 0.9000. A sixth H. philippinensis. the only one used

from Oligodon modestus. joins the others at a correlation of 0.5250, a * 4 low correlation for members of the same species. However, examining another specimen (not originally used, as a result of some missing data) from the same Oligodon. it was found that of the six characters

for which the states were considerably different for the original two groups, only for two of the characters were the states for this speci­ men the same as they were for the other specimen from Oligodon. the four other characters being as they were in the five specimens not from

Oligodon. It is therefore tentatively concluded that intra-specific variation is involved, and that only one species should be recognized.

There is some indication that Hemilaelaps radfordi (RAD) and

H. feideri (FEI) are the same species. Two H. radfordi join the one

specimen used of H. feideri at a high correlation, and are then joined 36

by the third H. radfordi at a level still consistent with conspecificity

(0.5750). The third H. radfordi is unlike the other H. radfordi and the

H* feideri in that the former is missing ad2 and pd2 of genu 4, both of which the latter two possess.

There are several obvious ways in which H. feideri and H.

radfordi are consistently different with regard to the four available

specimens. Concerning dorsal setae, H. radfordi has but one pair of

(presumed) j6 setae, while H. feideri has two (character 92). H.

feideri has 8 podonotal setae not on the podonotal shield, while all

three H. radfordi have 14. As pointed out by Fain (1962), many measure­ ments of H. feideri are somewhat shorter than those of H. radfordi. and

the peritreme of H. radfordi extends to coxa 1 while that of H. feideri

extends' only to coxa 11. Fain (1962) claims that most of the dorsal

setae of H. radfordi are slightly expanded near the tip, while those of

E* feideri are not. I believe these setae are slightly expanded in both cases, although less so in H. feideri. Fain's drawing of H. feideri

shows all coxal spurs with rounded tips, while in the specimen I have,

they are bifid (as they are in H. radfordi). Ordinarily, this character does not vary intra-specifically.

In view of the different hosts involved, the observed differ­ ences, the small number of specimens used, and the lack of opportunity to examine the holotypes, the species are not synonymized here.

Three Ixodorhynchus neodelphus (NEO) were isolated from the other 1. neodelphus and from each other. The correlations from the

similarity matrix confirm the isolation. All three, NE0017, NE0018, and NE0019, are from the same Kansas Storeria dekayi. Another ..I, neodelphus, from another Kansas Storeria dekavl. clustered with the

majority of neodelphus (a mixed group, but primarily from Ohio

Thamnophls sirtalis). To further investigate the situation, another

standardization correlation phenogram was produced, using only the 45

specimens of 31. liponyssoides (LIP), _I. neodelphus (NEO), and faini

(FAI), as OTU's. Also, a principal components analysis was done using

the same 45 OTU's. The results of these two analyses should be more

reliable than those of the original phenogram for the following reason:

originally, the characters were standardized for all 103 OTU’s; there­

fore, with regard to the three species being considered, some characters

probably received considerably more weight than others. Standardizing

the characters for just these three species, the characters should come

close to having equal weight for these three species.

In the new phenogram, no 3^. neodelphus were isolated, i.e., all

X* neodelphus were joined together before 3^. neodelphus joined 1^. faini.

All three species emerged in accordance with present concepts (a new

distance phenogram, using the same 45 OTU's, was almost as inconsistent with current concepts as the original distance phenogram based on 103

OTU's).

A graph of the first two factors of the principal components

analysis gave a clear-cut separation into the present three species,

although the more western I. neodelphus (Kansas and Texas) were slightly

closer to Jt. faini than the more eastern 1, neodelphus (Ohio). It is

concluded that there is insufficient evidence in the present study for

dividing I. neodelphus. 38

In standardizing the original data, the number of OTU's of one

species, relative to the numbers of OTU's of other species, will have an effect. If one is interested in inter-specific relationships, it

seems undesirable that the number of specimens used per species should vary, particularly if the number of available specimens of a species

largely reflects the location of the study, , and if the percent of speci­ mens used of specimens possessed of a species varies greatly (both of the latter two conditions occurred in the present study). Therefore, all phenograms mentioned in the remainder of the study, in which inter­ specific relationships are considered, are those in which OTU's are species.

The order of phenograms with regard to the amount of chaining, from the most to the least, determined by inspection, was standardization

(with or without logarithms) with taxonomic distance, standardization

(with or without logarithms) with the M.C.D., all three condensation phenograms, and standardization (with or without logarithms) with correlation.

Considering first the genera Ixodorhynchus. Ixobioides. and

Ixodorhynchoides. it was noted that all 3 emerge as a single group (here termed Ixo group) in the standardization correlation phenogram. However, in the standardization M.C.D. phenogram. Ixodorhynchus lohnstoni (JOH) and Ixobioides fonsecae (FON) are isolated from the rest of the Ixo group, as well as from each other, and all else. The standardization taxonomic distance phenogram not only isolates 1^. lohnstoni (JOH) and

I. fonsecae (FON) from each other and all else, but similarly isolates

Ixodorhynchus cubanensis (CUB) and Ixobioides butantanensis (BUT). 39

The standardization correlation phenogram divides the Ixo group

into two subgroups, the first consisting of Ixobioides (FON, BUT),

Ixodorhynchus uncatissimus (UNO), Ixodorhynchoides truncatus (TRU), and

Ixodorhynchus lohnstoni (JOH), and the second consisting of the remainder of Ixodorhynchus (LIP, FAI, NEO, LEP, CUB). Within the first

subgroup, I,, lohnstoni (JOH) and X» fonsecae (FON) are added last, individually, after the others have all joined together.

The standardization M.C.D, phenogram, after isolating 1,

lohnstoni (JOH) and X» fonsecae (FON) (as indicated above), divides the

Ixo group first by separating I . butantanensis (BUT) from the remainder.

The remainder is then divided into 2 groups, one consisting of _I. uncat issimus (UNC) and X» truncatus (TRU), and the other consisting of the remainder of Ixodorhynchus (LIP, FAI, NEO, LEP, CUB).

The standardization distance phenogram, after isolating the 4 species indicated above, divides the Ixo group first by separating

X. leptodeirae (LEP) from the remainder. The remainder is then divided into two groups, the first consisting of I. uncatissimus (UNC) and

I_. truncatus (TRU), and the second consisting of 1. liponvssoides (LIP),

X. neodelphus (NEO), and X* faini (FAI).

Further information concerning relationships within the Ixo group came from principal component analysis, based on correlation coefficients. When the first two factors were graphed, the Ixo group emerged together, with X* lohnstoni (JOH) and 1, fonsecae (FON) closest to the origin (all species quite isolated from other species tended to be placed near the origin). The second factor divided (although the division was not great) the Ixo group into the same two basic divisions 4°

seen in the standardization correlation phenogram. This same division was quite pronounced in the graph of the third and fourth factors, in i which most of Hemilaelaps was in between the two main groups of the Ixo group.

The graph analysis shown in Figure 2 (based on standardization and the M.C.D.) indicates that the first and second nearest neighbors of every member of the Ixo group are other members of the Ixo groups.

Also, it is apparent that, with regard to nearest neighbors, the Ixo group does not divide* well into subgroups, with the exception of the substantial isolation of I. lohnstoni (JOH) and I. fonsecae (FON).

The former genus Scutanolaelaps (OFH, SCH, MEH) Lavolpierre,

1958 (exclusive of S>. upembae, which is always well-separated from the other Scutanolaelaps) emerges consistently as a compact group isolated from all else. In the standardization M.C.D. phenogram the first division (not considering Omentolaelaps) separates Scutanolaelaps from all else.

> The graph analysis of Figure 3 shows that the closest relation­ ship of Scutanolaelaps (exclusive of £. upembae) with other ixodor- hynchids is that of £5. mehelyae (MEH) with H. triangulus (TRI) (Ewing,

1923). The nearest neighbor of £>. upembae (UPE) is also H. triangulus

(TRI). The standardization M.C.D. phenogram places S^. upembae in a large mixed group of Hemilaelaps. Strandtibbettsia brasiliensis. and the former genus Asiatolaelaps (TAN, EVA). The standardization dis­ tance phenogram places S. upembae with Gamin*s new species (NSP). The standardization correlation phenogram shows £>. upembae first joining

H. triangulus (TRI). 4 1

The principal components analysis (using the first two factors)

shows Scutanolaelaps quite isolated from all else, with £[. upembae t (UPE) among a mixed group. The nearest ixodorhynchid neighbor of the i isolated Scutanolaelaps is S. upembae.

Asiatolaelaps Fain, 1961 (including TAN, EVA, and synonymized with Hemilaelaps by Voss, 1967) and its presumed relative H.

philippinensis (PHI) only rarely emerged intact. In the standardi­

zation correlation phenogram, the group emerged together. In the other phenograms, all three species are separated from each other. However,

the graph analysis (Figure 3) indicates the nearest neighbor of H.

philippinensis is A. tanneri. No other first or second nearest neighbor relationships connect the three species.

The two species of Strandtibbettsia (GOR, BRA) and CaminTs new species (NSP) were all separated from each other in all 3 standardiza­ tion phenograms, although graph analysis (Figure 3) reveals that the nearest neighbor of NSP is S. brasiliensis (BRA). No other first or second nearest neighbor relationships connect the three species.

Interestingly, the standardization correlation phenogram joins S.. gordoni (GOR) to H. radfordi (RAD) and H, feideri (FEI) although the standardization distance phenogram and the standardization M.C.D. phenograms do not. Graph analysis (Figure 3) indicates that the nearest neighbors of gordoni are H. radfordi and H, feideri. although the nearest neighbors of the latter two species are among > the Hemilaelaps. 42

H. lioheterodon (LIO) is indicated in all phenograms as being rather isolated. Graph analysis (Figure 4) indicates its nearest neighbor is H. triangulus (TRI).

Fain (1962) divided Hemilaelaps into 4 "groups" (the ophidius group, equivalent to Scutanolaelaps is discussed above). The piger group, triangulus group and farrieri group generally do not hold together, although the piger group (PIG, NOV) remains together in the standardization distance phenogram. It is concluded that these groups do not provide a desirable division of Hemilaelaps. .

Principal component analysis does not seem suitable for deter­ mining genera within the Ixodorhynchidae. There is a strong tendency for individually isolated species to appear together near the origin, and much of the variation within the Ixodorhynchidae is of this nature.

The approach used in determining genera and subgenera was to draw phenon lines (Sokal and Sneath, 1963) across the most desirable kind of phenogram used (see above), the standardization M.C.D. pheno­ gram (Figure 1).

The determination of where the phenon lines were to be placed was arbitrary. Among the criteria taken into consideration were:

1. How many taxon changes would occur if the lines were moved slightly.

2. To what extent do the taxa correspond to past taxa. 3. How con­ venient is the number of resulting taxa.

The generic phenon line was placed at 0.6500. Six genera result, 3 of which are monotypic. The genera, listed in decreasing order of the cophenetic values at which they join another taxon, are:

1. Scutanolaelaps (OPH, SCH, MEH), 2. X-us (JOH), 3. Y-us (FON), 43

4. Strandtibbettsia (GOR), 5. Hemilaelaps (BRA, NSP, and all current

Hemilaelaps. see below), 6. Ixodorhynchus (LIP, NEO, FAI, LEP, CUB,

BUT, TRU, UNC).

The subgeneric phenon line was placed at 0.5000. The genus

Ixodorhynchus was thereby divided into 3 subgenera: 1., Ixodorhynchus

(LIP, NEO, FAI, LEP, CUB), 2. Ixobioides (BUT), 3. Ixodorhvnchoides

(TRU, UNC).

The genus Hemilaelaps is divided into 7 groups which warrant the status of subgenus; however, as 5 of the groups are monotypic, and a sixth possibly will become monotypic, it was decided best not to name the groups. The groups are 1. LIO, 2. NSP, 3. EVA, 4. UPE, 5. PHI,

6. RAD, and FEI, 7. CAH, CON, CAU, NOV, TAN, PIG, FAR, DIP, JAV, TRI,

BRA, and possibly H. imphalensis (not included in present study, but based on Fain's description is close to PIG).

Omentolaelaps (OME) is usually separated from all ixodorhynchids by the first division in phenograms. However, in the standardization correlation phenogram, Omentolaelaps mehelyae (OME) is placed with

Scutanolaelaps. 0. mehelyae has no close neighbors among the ixodorhyn­ chids.. It is concluded that the present status of Omentolaelapidae

Fain, 1961 as a separate family is justified.

Ixodorhynchus Ewing, 1923

Chelicerae completely lacking fixed digit; movable digit with large recurved teeth; corniculi with 1 or 2 barbs; 5-12 deutosternal teeth. Dorsum with j6 longer than jl, and j5 longer than 25; podonotal shield separated or not separated from oplsthonotal shield; opisthonotal 44

shield without lateral incisions in posterior part (exception: I,.

butantanensis). Venter with Zvl setae; endopodal sclerite between

coxa I and II attached to sternal shield; 3 pairs of sternal setae on

sternal shield; peritremal shield not extending posteriorly as far as

anal shield; venter of opisthogaster (not considering genito-ventral

and anal shield) without reticulate sclerotization; genital setae on

genito-ventral shield; sternal shield without anterior medial pointed

projection; bilobed pouch in front of sternal shield absent; peritreme

extending anteriorly as far as coxa I or II. Basal seta of coxa I

spur-like with rounded tip (exception: I. butantanensia. in which it

is setiform); coxa II without anterior pointed projection; posterior

seta of coxa II spur-like, with rounded tip; coxa III without scale­

like apophysis. Trochanter I without pd seta; trochanter II with 5

setae; genu I with 2 ventral setae; genu II lacking ad3; genu IV lack­

ing pl2; tibia I with 3 ventral setae; tibia II lacking pl2; tibia IV

lacking pd3; following setae not spine-like: pl3 of tarsus II, all and" al2 of tarsus IV, and pll of genu III.

Ixodorhynchus sensu strictu

Corniculi with 1 barb. Podonotal hexagon enclosing at least

2 setae; podonotal shield not separate from opisthonotal shield; opisthonotal shield without lateral incisions in posterior part; opisthonotal shield with at least 45 setae. Metasternal setae present.

Basal seta of coxa I spur-like with rounded tip. Genu I lacking ad3 and pd3; genu I with pl2; genu III lacking pd2; tibia I lacking pd3;

tarsus IV lacking pd3. £. liponvssoides Ewing, 1923

£• neodelphus Johnston, 1962 t faini Johnston, 1962

Jt. leptodeirae Fain, 1962

cubanensis Fain, 1962.

Subg, Ixobioides Fonseca, 1934

Corniculi.with 2 barbs. Podonotal hexagon enclosing no setae; podonotal shield separate from opisthonotal shield; opisthonotal shield with lateral incisions in posterior part; opisthonotal shield with 20 setae. Metasternal setae absent. Basal seta of coxa I setiform.

Genu I lacking ad3 and pd3; genu 1 lacking pl2; genu XIX with pd2; tibia I lacking pd3; tarsus IV with pd3.

butantanensis (Fonseca, 1934)

•Subg. Ixodorhynchoides Johnston. 1962

; Corniculi with 2 barbs. Podonotal hexagon enclosing no setae; podonotal shield separate from or attached to opisthonotal shield; opisthonotal shield without lateral Incisions in posterior part; opisthonotal shield with 22-33 setae. Metasternal setae present.

Basal seta of coxa I spur-like with rounded tip. Genu X with ad3 and pd3; genu I with pl2; genu III with pd2; tibia X with pd3; tarsus XV lacking pd3.

£• truncatus (Johnston, 1962)

I,, uncatissimus Voss and Strandtmann, 1962 0 4 6

X-us. new genus

Chelicerae completely lacking fixed digit; movable digit with large recurved teeth; corniculi with 1 barb; about 30 deutosternal teeth. Dorsum with j6 longer than jl, and j5 longer than Z5; podonotal shield separated from opisthonotal shield; opisthonotal shield with lateral incisions in posterior part. Venter without Zvl setae; endopodal sclerite between coxa I and II attached to sternal shield;

3 pairs of sternal setae on sternal shield; peritremal shield not extending posteriorly as far as anal shield; venter of opisthogaster

(not considering genito-ventral and anal shield) without reticulate sclerotization; genital setae on genito-ventral shield; sternal shield without anterior medial pointed projection; no bilobed pouch in front of sternal shield; peritreme extending anteriorly as far as coxa I,

Coxa II without anterior pointed projection; coxa III without scale-like apophysis. Trochanter I without pd seta; trochanter II with 6 setae; genu I with 2 ventral setae; genu II with ad3; genu IV lacking pl2; tibia I with 2 ventral setae; tibia II lacking pl2; tibia IV lacking pd3; pl3 of tarsus II and pll of genu III spine-like; all and al2 of tarsus IV not spine-like.

Type species: Ixodorhynchus johnstoni Fain. 1961 Y-us, new genus

Chelicerae completely lacking fixed digit; movable digit with

large recurved teeth; corniculi with 2 barb^; fewer than 10 deuto-

sternal teeth. Dorsum with j6 longer than jl and j5 longer than Z5;

podonotal shield attached to opisthonotal shield; opisthonotal shield without lateral incisions in posterior part. Venter with Zvl setae;

endopodal sclerite between coxa I and II attached to sternal shield;

2 pairs of sternal setae on sternal shield; peritremal shield not

extending posteriorly as far as anal shield; venter of opisthogaster

(not considering genito-ventral and anal shields) without reticulate

sclerotization; genital setae on genito-ventral shield; sternal shield without anterior medial pointed projection; no bilobed pouch in front of sternal shield; peritreme extending anteriorly as far as coxa I.

Coxa II without anterior pointed projection; coxa III without scale­ like apophysis; basal seta of coxa I spur-like with rounded tip.

Trochanter I without pd seta; trochanter II with 5 setae;, genu I with

2 ventral setae; genu II lacking ad3; genu IV lacking pl2; tibia I with 2 ventral setae; tibia II lacking pl2; tibia IV lacking pd3; pl3 of tarsus II setiform; spine-like setae: pll of genu III, all and al2 of tarsus IV.

Type species: Ixodorhynchus fonsecae Fain, 1961 Strandtlbbettsia Fain, 1961

Chelicerae with vestigial fixed digit; movable digit without large recurved teeth; corniculi without barb's; about 12 deutosternal teeth. Dorsum with j6 longer than jl and j5 longer than 25; podonotal shield attached to opisthonotal shield; opisthonotal shield with con­ cavities in posterior part. Venter with Zvl setae; endopodal sclerite between coxa X and XI not attached to sternal shield; 1 pair of sternal setae on sternal shield; peritremal shield extending posteriorly as far as anal shield; venter of opisthogaster (not considering genito-ventral and anal shields) with reticulate sclerotization; genital setae off genito-ventral shield; sternal shield without anterior medial pointed projection; no bilobed pouch I n front of sternal shield; peritreme extending anteriorly as far as coxa I or II. Basal seta of coxa I spur-like with rounded tip; c o x a II without anterior pointed projec­ tion; coxa III without scale-like spophysis. Trochanter I without pd seta; trochanter II with 5 setae; genu I with 2 ventral setae; genu II with ad3; genu IV lacking pl2; tibia I with 2 ventral setae; tibia II with pl2; tibia IV lacking pd3; following setae not spine-like: pl3 of tarsus II, all and al2 of tarsus IV, and pll of genu III.

£!. gordoni (Tibbetts, 1957)

Scutanolaelaps Lavolplerre, 1958

Chelicerae with fixed digit subequal in length to movable digit; movable digit without large recurved teeth; corniculi without barbs;

36-66 deutosternal teeth. Dorsum with jl longer than j6, and 25 longer 49

than j5; podonotal shield attached to opisthonotal shield; opisthonotal

shield without incisions in posterior part. Venter with Zvl setae;

endopodal sclerite between coxa I and II attached to sternal shield; i sternal shield with 3 pairs of sternal setae; peritremal shield not

extending posteriorly as far as anal shield; venter of opisthogaster

(not considering genito-ventral and anal shields) without reticulate

sclerotization; genital setae on genito-ventral shield; sternal shield

with anterior inedial pointed projection; bilobed pouch immediately

anterior to sternal shield; peritreme extending anteriorly only as far

as coxa III, Basal seta of coxa I setiform; coxa II with anterior

pointed projection; coxa III with scale-like apophysis. Trochanter I. with pd seta; trochanter II with 5 setae; genu I with 3 ventral setae;

genu II with ad3; genu IV with pl2; tibia I with 3 ventral setae;

tibia II with pl2; tibia IV with pd3; following setae not spine-like:

pl3 of tarsus II, all and al2 of tarsus IV, and pll of genu III.

£5. ophidius Lavoipierre, 1958

£>• schoutedeni Fain, 1961

Hemilaelaps Ewing, 1933

Chelicerae with fixed digit, rarely vestigial; movable digit without large recurved teeth; corniculi without barbs; €-60 deuto­ sternal teeth. Dorsum with jl sometimes longer than j6, sometimes j6 longer than jl; Z5 sometimes longer than j5, sometimes j5 longer than

Z5; podonotal shield attached or not attached to opisthonotal shield; opisthonotal shield without incisions in posterior part. Venter with Zvl setae; endopodal sclerite between coxa I and II attached to sternal

shield; sternal shield with 2 or 3 .pairs of sternal setae; peritremal

shield not extending posteriorly as far as anal shield; venter of

opisthogaster (not considering genito-ventral and anal shields) without

reticulate sclerotization; genital setae on genito-ventral shield;

sternal shield without anterior medial pointed projection; no bilobed

pouch anterior to sternal shield; peritreme extending anteriorly to

coxa I, II, or III. Basal seta of coxa I setiform, or spur-like with

rounded or bifid tip; coxa II without anterior pointed projection;

coxa III without scale-like apophysis. Trochanter I with or without

pd seta; trochanter II with 5 setae; genu I with 2 ventral setae

(exception: H, evansi with 3 ventral setae); genu II with ad3 (excep­

tions: H. philippinensis. H. congolensis and Camin's new species);

genu IV lacking pl2; tibia I with 2 ventral setae; tibia II with pl2

present or absent; tibia IV lacking pd3; following setae not spine-like

pl3 of tarsus II, all and al2 of tarsus IV, and pll of genu III.

Group 1

Chelicerae with fixed digit distally flagelliform; palpal

trochanter lacking V2. Dorsal shield with "cribrum" (microtrichae) at

posterior end. Sternal shield with 2 pairs of sternal setae. Coxa I with only setiform setae. Genu I lacking pd3 and al2; tibia I lacking

pd3.

H. lioheterodon Fain, 1967 Group 2

Chelicerae with vestigial fixed digit; palpal trochanter

lacking V2; palpal genu lacking d2 and al. Genu I lacking ad3 and al2;

genu II lacking ad3; tarsus IV lacking pd3.

New species of Catnin

. Group 3

About 60 deutosternal teeth. Dorsal setae Z5 at least twice as

long as j5. Laterally well-sclerotized parts of sternal shield not

connected medially; peritreme not extending anteriorly beyond coxa III.

Basal seta of coxa I triangular, barbed, and fused with coxa. Genu I

with av2.

H. evansi (Fain, 1961)

Group 4

Dorsal jl setae greater than twice the length of j6; dorsal

setae j3 and j4 absent. Basal seta of coxa I basally' spur-like,

distally flagelliform; coxa II with scale-like apophysis. Tibia IV

with pd3.

H. upembae (Fain, 1961)

-■■■■ Group 5

Palpal trochanter lacking V2, Podonotum lacking jl; podonotum with j4 and j5 broad basally, suddenly narrowing. Lateral well-

sclerotized parts of sternal shield not connected medially. Basal seta

of coxa I triangular, but with lateral lobe. Genu X lacking ad3 and

pd3; genu II lacking ad3; tibia I lacking pd3; tibia IV with al2.

H. philippinensis Voss, 1967 * Group 6

Dorsum with many setae slightly expanded near tip; opisthonotum with more than 5 pair of setae in J column; podonotum with over 5 setae not on shield (adding setae of both sides together). Sternal shield with 2 pairs of setae.

H. radfordi (Feider and Solomon, 1959)

H. feideri Fain, 1962

Group 7

Palpal genu with d2 and al. Podonotum with fewer than 5 setae

(adding both sides together) off shield; dorsal shield lacking a

’’cribrum" at posterior end. If sternal shield has well-sclerotized areas laterally (not considering endopodal sclerites), they are con­ nected medially by well-sclerotized area. If basal seta of coxa I is basally spur-like, it is not distally flagelliform.

H, tanneri (Tibbetts, 1954)

H. triangulus (Ewing, 1923)

S* iavanensis Fain, 1961

H. farrieri (Tibbetts, 1954)

H. dinsadoboae Fain, 1962

H. cahent Fain, 1961

B* Piget (Berlese, 1918)

H. novaeguineae Fain, 1961

H. congolensis Fain, 1962

B* causicola Fain, 1962

H. brasiliensis (Fain, 1961)

H. imphalensis (Radford, 1947) SUMMARY

A numerical taxonomic study was carried out using 103 speci­ mens belonging to 31 described, and one undescribed, species of

Ixodorhynchidae and one described species of Omentolaelapidae. One- hundred and thirty-two characters were used, following elimination of many of the highly correlated characters (r > .5) from the original

190 characters recorded. Transformations used were logarithms, logarithms standardized, standardization, and condensation. Similarity coefficients used were the correlation coefficient, taxonomic distance, the M.C.D, and the Canberra metric. UPGMA was the clustering method used. Principal component analysis and graph analysis were also con­ sidered in the interpretation of similarity matrices.

■ Predictiveness determinations were carried out on the similarity matrices and phenograms to determine what combination of methods would provide the most predictive (and therefore, most desirable) results.

Stability predictiveness and character predictiveness were determined using procedures which included randomly dividing the set of characters into subsets, producing similarity matrices and phenograms, and finding correlations between similarity matrices (and phenograms) based on different character subsets. The most predictive combination used was found to be standardization (with or without logarithms) with the

M.C.D.

53 Character weighting is defined, and a new transformation,

"equalization" is suggested. It is regarded as the only transformation which consistently weights characters equally.

Taxa were determined by applying phenon lines to the standardi­ zation M.C.D. phenogram. Two new monotypic genera, X-us (based on

Ixodorhynchus johnstoni Fain, 1961), and Y-us (based on Ixobioides fonsecae Fain, 1961) are proposed, Asiatolaelaps Fain, 1961 is con­ firmed as a synonym of Hemilaelaps Ewing, 1933. Strandtlbbettsia brasiliensis Fain, 1961 is transferred to Hemilaelaps. Hemilaelaps is regarded as the proper genus for a new species from Diadophis punctatus in Kansas, found to be most similar to £. brasiliensis. Scutanolaelaps

Lavoipierre, 1958 is revived, although H. upembae (Fain, 1961) remains in Hemilaelaps. Ixobioides Fonseca, 1934, and Ixodorhynchoides

Johnston, 1962, are regarded as subgenera of Ixodorhynchus Ewing, 1923.

Ixodorhynchus uncatissintus Voss and Strandtmann, 1962 is placed in the subgenus Ixodorhynchoides. The genus Hemilaelaps Ewing, 1933 is divided into 7 groups. Omentolaelapidae Fain, 1961, with the single species

Omentolaelaps mehelyae Fain, 1961, is regarded as a valid family.

Diagnoses of the genera, subgenera, and groups are given. APPENDIX I

LIST .OF O T U ' S

OTU HOST O C A L I T Y COLLECTOR MUSEUM NUMBER LIPOOl THAMNOPHIS SIRTALIS WAYNE CO OHIO D E JOHNSTON OSU IA 137 9 L3P002 THAMNOPHIS SIRTALIS WAYNE CO OHIO D E JOHNSTON OSU IA 137 13 LIP003 MATRIX SIPEDON EMMET CO MICHIGAN W J W R E N N OSU 65 0 7 3 0 7 3 L3P004 THAMNOPHIS SIRTALIS CHEBC YGAN CO MICHIGAN W J WRENN OSU 6 5 0 7 1 3 2 4 LI POOS MATRIX SIPEDON EMMET CO F ICHIGAN W J WRENN OSU 6 5 0 7 3 0 7 5 L3P006 THAMNOPHIS SIRTALIS OAKLAND CO MICHIGAN D E JOHNSTON OSU 64(43 R) 20 LIP007 THAMNOPHIS SIRTALIS PT PELEE NAT PK ONTARIO D E JOHNSTON OSU 46(25 R) 40 L1P008 THAMNOPHIS SIRTALIS PT PELEE NAT PK ONTARIO D E JOHNSTON OSU 33(14 R) 45 LIP009 THAMNOPHIS SIRTALIS PT PELEE NAT PK ONTARIO D E JOHNSTON OSU 39(18 R) 60 L3P030 THAMNOPHIS SIRTALIS PT PELEE NAT PK ONTARIO D E JOHNSTON OSU 18(2 R) 69 2 LIP011 THAMNOPHIS SIRTALIS ARKONA ONTARIO D E JOHNSTON OSU 67(46 R) 93 L3P032 THAMNOPHIS SIRTALIS MOBILE CO ALABAMA D E JOHNSTON OSU USNM 55803 5 55 56

APPENDIX I - CONTINUED

OTU HOST LOCALITY COLLECTOR MUSEUM * NUMBER LIP033 THAMNOPHIS SIRTALIS MOBILE CO ALABAMA D E JOHNSTON OSU USNM 55803 6 LIPO 14 THAMNOPHIS SIRTALIS MOBILE CO ALABAMA D E JOHNSTON OSU USNM 55803 7 LIP015 THAMNOPHIS SIRTALIS WAYNE CO OHIO D E JOHNSTON OSU IA 137 11 LIP016 THAMNOPHIS SIRTALIS WAYNE CO OHIO D E JOHNSTON OSU IA 137 14 L3POT4 THAMNOPHIS SIRTALIS OAKLAND CO MICHIGAN D E JOHNSTON OSU 65{44-R) 73 LIP075 THAMNOPHIS SIRTALIS OAKLAND CO MICHIGAN D E JOHNSTON OSU 53(32~R) 43 L3 POT6 THAMNOPHIS RADIX BROOKINGS SOUTH DAKOTA J CAMIN OSU 61-2 L1P077 MATRIX SIPEDON PT PELEE N PK ONTARIO CANADA D E JOHNSTON OSU 4l(20-R) 50 LJP078 THAMNOPHIS SIRTALIS PT PELEE N PK ONTARIO CANADA D E JOHNSTON OSU 42I21-R) 66-3. LI POT 9 NATRIX SIPEDON INSULARUM LAKE ERIE ISLANDS OHIO J CAMIN OSU 62-4 1 NE0017 STORERIA DEKAYI FRANKLIN CO KANSAS H K GLOYD OSU KU 55179 1 NE0018 STORERIA DEKAY I FRANKLIN CO KANSAS H K GLOYD OSU KU 55179 2 ME003 9 STORERIA DEKAYI FRANKLIN CO KANSAS H K GLOYD OSU KU 55179 3 NE0020 THAMNOPHIS SIRTALIS WAYNE CO OHIO D E JOHNSTON HOLOTYPE USNM 2813 NE0021 THAMNOPHIS SIRTALIS WAYNE CO OHIO D E JOHNSTON OSU IA 137 8 57

APPENDIX I - CONTINUED

OTU HOST LOCALITY COLLECTOR MUSEUM NUMBER NE0022 HALDEA STRIATULA TRAVIS CO TEXAS FIELDS COLL OSU RWS I 48 1 NE0085 STORERIA DEKAYI .DOUGLAS CO KANSAS C D BUNKER OSU KU 2345 NE0086 TROPJDOCLONION LINEATUM DICKINSON CO KANSAS D H TAYLOR OSU KU16363 NE0087 THAMNOPHIS SIRTALIS NEAR WOOSTER OHIO D E JOHNSTON ' OSU .VENTRAL NEOOF8 THAMNOPHIS SIRTALIS NEAR WOOSTER OHIO D E JOHNSTON OSU DORSAL NE0089 THAMNOPHIS SIRTALIS WASHTENAW CO MICHIGAN D E JOHNSTON OSU 29(11-R) 63 FA 3023 THAMNOPHIS SIRTALIS PT PELEE NAT PK ONTARIO D E JOHNSTON OSU 42(21 Z> 66 2 FA 1024 THAMNOPHIS SIRTALIS OAKLAND CO MICHIGAN D E JOHNSTON OSU 53(32 R) 4 F A 3 0 2 5 NATRIX SIPEDON MARION CO FLORIDA D E JOHNSTON OSU 74(52 R) 58 FA 10 26 MATRIX SIPEDON MARION CO FLORIDA D E JOHNSTON OSU 68(47 RJ 3 FA ]027 THAMNOPHIS SIRTALIS PT PELEE NAT PK ONTARIO D E JOHNSTONHOLOTYPE USNM . 2812 FA 1028 THAMNOPHIS SIRTALIS OAKLAND CO MICHIGAN D E JOHNSTON OSU 64(43 R) 8 FA 3029 THAMNOPHIS SIRTALIS OAKLAND CO MICHIGAN D E JOHNSTON OSU 64(43 R) 13 FA 1030 THAMNOPHIS SIRTALIS OAKLAND CO MICHIGAN D E JOHNSTON OSU 64(43 RJ 16 FA 3021 THAMNOPHIS SIRTALIS PT PELEE NAT PK ONTARIO D E JOHNSTON . . OSU 42(21 R) 67 2 58

APPENDIX I - CONTINUED

OTU HOST LOCALITY COLLECTOR MUSEUM NUMBER FAI09O THAMNOPHIS SIRTALIS PT PELEE N PK ONTARIO CANADA D E JOHNSTON OSU 56(35-R) 6 FAlOcl T H A M N O P H I S SIRTALIS PT PELEE NAT PK ONTARIO CANADA D E JOHNSTON OSU 20(4~R) 4 FA3092 THAMNOPHIS SIRTALIS OAKLAND CO MICHIGAN D E JOHNSTON OSU 77(56-R) PH1067 HOST UNIDENTIFIED TANJAY NEGROS IS. PHILIPPINES VOSS PARATYPE BIS. MUS.? 5122 1 PH3068 OL3GODON MODESTUS NEGROS IS. PHILIPPINES VOSS PARATYPE BIS. MUS.? 1664 1 PHI069 CYCLOCORUS LlNEATUS NEGROS IS. PHILIPPINES VOSS PARATYPE BIS. MUS.? 18758. PHI070 CYCLOCORUS LlNEATUS NEGROS IS. PHILIPPINES VOSS PARATYPE BIS. MUS.? 18758.. PH1071 CYCLOCORUS LlNEATUS NEGROS IS. PHILIPPINES VOSS PARATYPE BIS. MUS.? 18758... PH 1072 CYCLOCORUS LlNEATUS NEGROS IS. PHILIPPINES VOSS HOLOTYPE BIS. MUS. 18758 G0R035 NATRIX TIGRINA LATERALIS SEOUL KOREA TIBBETTS PARATYPE OSU . G0R036 NATRIX SUBMINI AT A SUBM INIATA MANO SOMAR INDONESIA A FAIN OSU ON LEFT G0R037 NATRIX SUBMINIATA SUBMINIATA MANO SOMAR INDONESIA A FA JN OSU ON RIGHT GORO? 8 N A T R I X T IGRINA LATERALIS SEOUL KOREA H S AH OSU SQUARE COVERSLP G0R039 NATRIX T IGRINA LATERALIS SEOUL KOREA H S AH OSU ROUND COVERSLIP TPIO^O COLUBER CONSTRICTOR COLUMBIA MISSOURI P J SPANGLER OSU FROM 43 a p p e n d i x i - c o n t i n u e d

o t u h o s t LOCALITY COLLECTOR MUSEUM NUMBER TR3051 COLUBER CONSTRICTOR FDXI COLUMBIA MISSOURI N E ESKEW OSU SQ 5166-1 TR1052 COLUBER CONSTRICTOR WASHTENAW CO MICHIGAN D E JOHNSTON . OSU . 24<8-R> TP3053 THAMNOPHIS SIRTALIS WASHTENAW CO MICHIGAN D E JOHNSTON OSU 3 K 1 3 - R ) 77 RAD0P2 HOST-LOCALITY DATA MISSING INFORMATION MISSING RAD0F3 HOST-LOCALITY DATA MISSING INFORMATION MISSING RAD084 HOST-LOCALITY DATA MISSING INFORMATION MISSING LEP032 LEPTODEIRA MACULATA MEXICO A FAIN PARATYPE OSU IRSNB LEP033 LEPTODEIRA MACULATA MEXICO A FAIN PARATYPE OSU 1RSNB JOH034 HETERODON PLATYRHINOS MARION CO FLORIDA A FAIN PARATYPE BRA040 SYPHLOPHIS PULCHER JUQUIA BRAZIL A FAIN PARATYPE OSU BPA097 SYPHLOPHIS PULCHER JUQUIA BRAZIL A FAIN HOLOTYPE IRSNB M904 TRU041 ELAPHE VULPINA PT PELEE NAT PK ONTARIO D E JOHNSTON HOLOTYPE USNM 2811 TFU045 ELAPHE VULPINA BARABOO WISCONSIN D E JOHNSTON OSU ON RIGHT BUT046 TOMODON DORSATUS HAUTE MAR INGA BRAZIL FAIN OSU 0PH047 CAUSUSLICHTENSTEINII BRITISH CAMEROONS A FAIN PARATYPE OSU APPENDIX I - CONTINUED

OTU ' HOST LOCALITY COLLECTOR MUSEUM NUMBER DPH048 CAUSUS LICHTENSTEIN11 CONGO (KINSHASA) A FA)N OSU R G 703 SCH049 BOAEDON FULIGINOSUS KlVU CONGO HIERNAUX PARATYPE OSU FAR054 BOTHROPHTHALMUS LlNEATUS LlNEATUS KIVU CONGO A FAIN OSU FARO55 DINODON RUFOZONATUM SEOUL KOREA H S AH OSU CAU056 CAUSUS RHOMBEATUS MAYUMBE CONGO A FAIN PARATYPE OSU ON LEFT CAU057 CAUSUS RHOMBEATUS MAYUMBE CONGO A FAIN PARATYPE OSU ON RIGHT DJP058 D1PSAD0B0A UNICOLOR LlSALA CONGO A FAIN PARATYPE OSU FE1059 NATRIX NATRIX HELVETICA NAPLES ITALY A FAIN PARATYPE OSU 3221 PIGO60 COLUBER GEMONENSlS DALMATIA YUGOSLAVIA A FAIN' OSU 545 1 PIG061 COLUBER GEMONENSlS DALMATIA YUGOSLAVIA A FAIN OSU 545 2 NCV062 DENDROPHIS CALLIGASTER BOUGAINVILLE SOLOMON A FAIN PARATYPE OSU 2614 TAN063 NATRIX TIGRINA LATERALIS SEOUL KOREA TIBBETTS PARATYPE OSU 1 TAN064 NATRIX TIGRINA LATERALIS SEOUL KOREA TIBBETTS PARATYPE OSU 2 E.VA065 ELAPHE FLAVOLINEATA INDIA A FAIN PARATYPE OSU 336 1 i ‘EVA066 ELAPHE FLAVOLINEATA INDIA A FAIN PARATYPE OSU 336 2 APPENDIX I - CONTINUED

OTU - HOST LOCALITY COLLECTOR MUSEUM NUMBER OME073 MEHELYA POENSlS KlVU CONGO A f a i n PARATYPE OSU

NSP080 DIADOPHIS PUNCTa TUS DOUGLAS CO KANSAS J CAMIN UNIV KANSAS 4A-67 NSPOfl DIADOPHIS PUNCTATUS DOUGLAS CO KANSAS J CAMIN UNIV KANSAS 4J-67 TRI0O3 HOST NOT. KNOWN LOCALE NOT KNOWN D E JOHNSTON OSU ‘ 29(11-R) CUB0S4 LIOPHIS ANDREAE CUBA A FAIN HOLOTYPE IRSNB M909 FONOS5 XENEDON GUENTHERI MATTO GROSSO BRAZIL A FAIN HOLOTYPE IRSNB M892 FCMOS6 XEKEDON GUENTHERI MATTO GROSSO BRAZIL A FAIN PARATYPE IRSNB JAVOS8 LYCDDON SUBClNCTUS JAVA INDONESIA A FAIN HOLOTYPE IRSNB M889 CCNOS9 CAUSUS RHOMBEATUS KATANGA CONGO A FAIN ' PARATYPE FAIN COLL 9654 CAH100 NAJA MELANOLEUCA LUTUNGURU CONGO A FAIN PARATYPE FAIN COLL L 1GI01 LIOHETERODON MODESTUS AMPIJOROA MADAGASCAR A FAIN PARATYPE FAIN COLL RML45847 DORSAL L‘10102 LIOHETERODON MODESTUS AMPIJOROA MADAGASCAR A FAIN PARATYPE FAIN COLL RML45847 VENTRL UFE103 BOAEDON FULIG1N0SUS ABERCORN N RHODESIA A FAIN PARATYPE FAIN COLL 8901 MEH104 MEHELYA POENSlS BRAZZAVILLE CONGO TAUFFL1EB HOLOTYPE PARIS MUS 2139 MEH105 MEHELYA POENSlS BRAZZAVILLE CONGO TAUFFLIEB PARATYPE PARIS MUS 2139 UNC106 PSEUSTES POEClLONOTUS CANAL ZONE VOSS PARATYPE OSU 1 APPENDIX II

CHARACTERS USED

1. FIXED DIGIT ABSENT (0), VESTIGIAL (1), DISTALLY FLAGELLIFORM (2), OR ROBUST (3). 2. TEETH ON MOVABLE DIGIT RECURVED AND LARGE (1), OR NEITHER (0). 3. LENGTH OF MOVABLE DIGIT. 4. DORSAL LENGTH OF SECOND CHELICERAL SEGMENT (NOT INCLUDING PORTION WHICH IS ANTERIOR TO ANY PART OF MOVABLE DIGIT). 5. V2 OF PALPAL TROCHANTER PRESENT (1) OR ABSENT (0). • 6. BOTH Dl AND D3 PRESENT ON PALPAL FEMUR (2) OR NOT (I). 7. D3 OF PALPAL GENU PRESENT (X) OR ABSENT (0). 8. TWO OR MORE TINES ON PALPAL CLAW PRESENT (2) OR ABSENT (1). 9. BARB ON CORNICULUS WHICH IS OVER 25 MICRONS FROM TIP OF CORNICULUS. PRESENT (1) OR ABSENT (0). 10. DISTANCE BETWEEN GS1 AND GS3. 11. DISTANCE BETWEEN GS4 AND POSTERIOR MARGIN OF GNATHOSOMA. 12. NUMBER OF DEUTOSTERNAL TEETH. 13. NUMBER OF ROWS OF DEUTOSTERNAL TEETH. 14. NUMBER OF SETAE ON PODOSOMAL PART OF DORSAL SHIELD (NOT INCLUDING SETAE OF CHARACTERS 15, 92, 93). 15. jl SETAE MISSING (0), LESS THAN ONE-HALF THE LENGTH AND OF j6 SETAE (1), BETWEEN ONE-HALF THE LENGTH AND EQUAL IN LENGTH TO j6 SETAE (2), GREATER IN LENGTH THAN j6 SETAE, BUT LESS THAN TWICE THEIR LENGTH (3), MORE THAN TWICE THE LENGTH OF j6 SETAE (4). 16. LENGTH OF z5 SETA. 17. PODOSOMAL HEXAGON RECOGNIZABLE (AT LEAST j5 AND z5) (1) OR NOT (0). 18. EXTENT OF DIVISION BETWEEN PODONOTAL AND OPISTHONOTAL SHIELDS: NO DIVISION (0), CONCAVITY (1), "PINCHED-OFF" INCISION (2), INCISION (3) , COMPLETE SEPARATION (4). 19. BOTH LATERAL EDGES OF POSTERIOR HALF OF OPISTHONOTAL SHIELD CONTAINING AN INCISION (2), A CONCAVITY (1), OR NEITHER (0). 20. NO DORSAL SHIELD SETA (NOT COUNTING Z5) IS POSTERIOR TO J5 (1) OR NOT (0). 21. Z5 SETAE LESS THAN ONE-HALF THE LENGTH OF j5 (0), BETWEEN ONE-HALF THE LENGTH AND EQUAL IN LENGTH TO j5 (1), LONGER THAN j5 BUT NOT TWICE AS LONG (2), OR MORE THAN TWICE AS LONG (3). 22. AT LEAST SOME DORSAL SETAE EXPANDED NEAR TIP (1) OR NOT (0). 23. LENGTH OF OPISTHONOTAL SHIELD. 24. DISTANCE BETWEEN Z5 SETAE. 25. OVER ONE-HALF THE SETAE OF OPISTHONOTAL SHIELD LESS THAN ONE-HALF THE LENGTH OF z5 SETAE (NOT CONSIDERING J5 AND Z5 SETAE) (0) OR NOT (1). 62 63

APPENDIX II - CONTINUED

26. BILOBED POUCH IN FRONT OF STERNAL SHIELD PRESENT (1) OR NOT (0). 27. ENDOPODAL SCLERITE BETWEEN COXAE I AND II IS CONTINUOUS WITH STERNAL SHIELD (1) OR NOT (0). , 28. STATE OF HEAVY LATERAL SCLEROTIZATION CONTAINING STERNAL SETAE AND PORES (ON AT LEAST ONE SIDE): NO SUCH SCLEROTIZATION (1), CONTAINING ONLY STERNAL SETAE I (2), CONTAINING STERNAL SETAE I AND FIRST PAIR OF STERNAL PORES (3), CONTAINING STERNAL SETAE I AND II (4), EXTENDING FROM STERNAL SETAE I TO SECOND PAIR OR PORES (5), OR EXTENDING FROM STERNAL SETAE I TO STERNAL SETAE III (6). 29. NUMBER OF PAIRS OF SETAE ON STERNAL SHIELD. 30. METASTERNAL SETAE (AT LEAST ONE) PRESENT (1) OR ABSENT (0). 31. DISTANCE BETWEEN STERNAL SETAE I. 32. DISTANCE BETWEEN STERNAL SETA 2 AND STERNAL SETA 3. 33. DISTANCE BETWEEN STERNAL SETA 3 AND STERNAL SETA 4. 34. DISTANCE BETWEEN GENITAL SETAE. 35. GENITO-VENTRAL SHIELD SCALY (1) OR NOT (0). 36. POSTERIOR MARGIN OF SCLEROTIZED PORTION OF GENITO-VENTRAL PLATE IS POSTERIOR TO POSTERIOR MARGIN OF BOTH Jvl SETAE (1) OR NOT (0). 37. LENGTH OF STERNAL SETA I. 38. DISTANCE BETWEEN COXAE III. 39. PERITREME EXTENDING (ON AT LEAST ONE SIDE) TO COXA I (3), TO COXA II (2), OR ONLY AS FAR AS COXA III (1). ) 40. STIGMA ANTERIOR TO MIDDLE OF COXA IV (ON AT LEAST ONE SIDE) (1), OR POSTERIOR (ON BOTH SIDES) (0). 41. EXTENSION OF PERITREMAL PLATE POSTERIOR TO STIGMA: NO SIGNIFICANT EXTENSION (0), MODERATE EXTENSION (POSTERIOR EDGE OF PLATE IS CLOSER TO TRANSVERSE LINE EXTENDING FROM POSTERIOR EDGE OF COXA IV THAN TO STIGMA) (1), LENGTHY EXTENSION (EXTENDING PAST ANTERIOR EDGE OF ANAL PLATE ON AT LEAST ONE SIDE) (2). 42. Jv3 AND Zv2 RECOGNIZABLE ON AT LEAST ONE SIDE (Jv3 MUST BE BOTH POSTERIOR AND MESAD TO Zv2 AND ANTERIOR TO MIDDLE THIRD OF WELL- SCLEROTIZED PART OF ANAL SHIELD) (2), OR NOT, DUE TO HYPOTRICHY (1), OR NOT, DUE TO HYPERTRICHY (3). . 43. VENTER OF OPISTHOGASTER (NOT CONSIDERING GENITO-VENTRAL AND ANAL SHIELDS) WITH RETICULATE SCLEROTIZATION (1), OR NOT (0). 44. NUMBER OF SETAE IN Zvl POSITION ON BOTH SIDES. 45. DISTANCE BETWEEN Jvl SETAE. 46. DISTANCE BETWEEN Jv2 SETAE. 47. WIDEST WIDTH OF WELL-SCLEROTIZED PART OF ANAL PLATE. 48. DISTANCE BETWEEN ADANAL SETA AND POSTANAL SETA. 49. DISTANCE BETWEEN ADANAL SETA AND ANTERIOR MARGIN OF WELL- SCLEROTIZED PART OF ANAL SHIELD. 50. ANTERIOR MARGIN OF CRIBRUM IS POSTERIOR TO ANTERIOR MARGIN OF POSTANAL SETA (1), OR ANTERIOR TO ANTERIOR MARGIN OF POSTANAL SETA, AND STATES "3" AND "4" DO NOT APPLY (2), OR ANTERIOR TO POSTERIOR MARGIN OF ANAL VALVES, AND STATE "4" DOES NOT APPLY (3), OR ANTERIOR TO ANTERIOR MARGIN OF ANAL VALVES (4). 64 APPENDIX II - CONTINUED

51. POSITION OF ADANAL SETAE RELATIVE TO ANAL VALVES: ANTERIOR EDGE OF AT LEAST ONE ADANAL SETA IS ANTERIOR TO ANAL VALVES (1), CHARACTER STATES "1" AND "3" DO NOT APPLY (2), OR (POSTERIOR EDGE OF AT IEAST ONE ADANAL SETA IS POSTERIOR TO ANAL VALVES (3). 52. MEDIAL ONE-THIRD OF ANTERIOR MARGIN OF SCLEROTIZED PART OF ANAL SHIELD HAS CENTRAL CONCAVITY (1), IS STRAIGHT (2), OR IS CONVEX (3). 53. CONCAVITY (NOT ASSOCIATED WITH CRIBRUM) IN BOTH LATERAL MARGINS OF ANAL SHIELD (1) OR NOT (0). 54. MEDIAL PORTION OF ANTERIOR MARGIN OF WELL-SCLEROTIZED PART OF ANAL SHIELD AS WELL-SCLEROTIZED AS LATERAL PORTION (2), OR LESS WELL- SCLEROTIZED (1). 55. TWO LATERAL CONCAVITIES IN ANTERIOR MARGIN OF WELL-SCLEROTIZED PART OF ANAL SHIELD (1) OR NOT (0). 56. POSTERIOR MARGIN OF BODY WITH MANY SPATULATE SETAE (1) OR NOT (0). 57. BASAL SETA OF COXA I SETIFORM (1), BASALLY SPUR-LIKE, BUT DISTALLY SETIFORM (2), SPUR-LIKE WITH ROUNDED TIP (3) , TIP WITH TWO SUBEQUAL LOBES SIDE BY SIDE IN DORSAL VIEW (4), INTERMEDIATE BETWEEN STATES "4" AND ”611 (5), TRIANGULAR, WITH SINGLE TIP, BACKWARDS-POINTING BARBS, AND NOT FUSED W I T H COXA (6), OR AS STATE "6", BUT FUSED WITH COXA, WITH NO APPARENT LINE OF DEMARCATION (7). 58. POSTERIOR SETA OF COXA II CLOSER TO BASAL (1) OR DISTAL (0) END OF COXA. 59. POSTERIOR SETA OF COXA II POINTING FORWARD (0) OR BACKWARD (1). 60. POINTED PROJECTION FROM ANTERIOR EDGE OF COXA II PRESENT (1) OR ABSENT (0). 61. COXA III WITH SCALE-LIKE APOPHYSIS (1) OR NOT (0). 62. TROCHANTER I WITH PD PRESENT (1) OR ABSENT (0). 63. TROCHANTER II WITH ALL AND AV SPINE-LIKE (2) OR NOT (1). ' 64. AL1 OF TROCHANTER IV WITH THIN POINTED LATERAL PROJECTION (1) OR NOT (0). 65. NUMBER OF AL SETAE ON FEMUR II. 66. MD OF FEMUR III LONGER (2) OR SHORTER . (1) THAN HALF THE LENGTH OF GENU III. 67. MD OF FEMUR IV LONGER (2) OR SHORTER (1) T H A N HALF THE LENGTH OF GENU IV. 68. NUMBER OF AD SETAE ON GENU I. 69. NUMBER OF PL SETAE ON GENU I. 70. GENU II WITH AV PRESENT (1) OR ABSENT (0) . 71. NUMBER OF AL SETAE ON GENU III. 72. GENU III WITH AV PRESENT (1) OR ABSENT (0). 73. GENU III WITH PV PRESENT (1) OR ABSENT (0) . . 74. DISTANCE BETWEEN ADI AND AD2 OR GENU III. 75. NUMBER OF AD SETAE ON GENU IV. 76. NUMBER OF PD SETAE ON GENU IV. 77. NUMBER OF PD SETAE ON TIBIA I. 78. NUMBER OF PL SETAE ON TIBIA I. 65

APPENDIX II - CONTINUED

79. NUMBER OF VENTRAL SETAE ON TIBIA I. 80. NUMBER OF AD SETAE ON TIBIA II. 81. NUMBER OF PL SETAE ON TIBIA II. , 82. DISTANCE BETWEEN PDl AND PD2 OF TIBIA II. 83. NUMBER OF PL SETAE ON TIBIA III. 84. NUMBER OF AL SETAE ON TIBIA IV. 85. SECOND-MOST BASAL PV SETA OF TARSUS I IS DISTAL (1) OR BASAL (0) TO THE LENGTHY PL PRIOR TO TARSUS TIP (THE LENGTHY PL IS ALMOST ALWAYS THE THIRD-MOST BASAL PL, RARELY THE SECOND-MOST BASAL). 86. NUMBER OF AD AND AL SETAE ON TARSUS I WHICH ARE BASAL TO THE AL SETA WHICH IS CLOSEST TO THE SECOND-MOST BASAL PL SETA (IN THOSE RARE CASES FOR WHICH STATE "1" OCCURS FOR CHARACTER 125, USE THE MOST BASAL PL SETA RATHER THAN THE SECOND-MOST BASAL). 87. AV3 (= MV) OF TARSUS II PRESENT (1) OR ABSENT (0). 88. PLl OF TARSUS II SPINE-LIKE (2) OR NOT (1). 89. TARSUS IV WITH PD3 PRESENT (1) OR ABSENT (0). 90. TARSUS IV WITH AD2 PRESENT (1) OR ABSENT (0). 91. PERITREME EXTENDING POSTERIOR TO STIGMA (1) OR NOT (0). 92. NUMBER OF PAIRS OF POSSIBLE j6 SETAE (DISTINGUISH BETWEEN j6 AND z6 SETAE ON BASIS OF WHETHER OR NOT DISTANCE BETWEEN MEMBERS OF PAIR IS GREATER OR LESS THAN DISTANCE BETWEEN z5 SETAE). 93. NUMBER OF SETAE ENCLOSED BY PODONOTAL HEXAGON (NOT INCLUDING j5, z5, OR POSSIBLE j6 SETAE). 94. TROCHANTER II WITH AL2 OR AD (1) OR NOT (0). 95. NUMBER OF PODONOTAL SETAE NOT ON PODONOTAL SHIELD. 96. STERNAL SHIELD HAVING MEDIAL POINT AT ANTERIOR END (1) OR NOT (0). 97. DISTANCE BETWEEN SECOND-MOST POSTERIOR PAIR OF SETAE ON OPISTHONOTAL SHIELD LESS THAN DISTANCE BETWEEN MOST POSTERIOR PAIR (2), OR NOT (0), OR SIDES DIFFERING (1). 98. MOST VENTRAL SETAE WHICH ARE NOT ON PLATES ARE SUDDENLY NARROWING (2) OR NOT (1). 99. LENGTH OF GENITO-VENTRAL SHIELD. 100. MEMBRANOUS POUCH COVERING MOST OF VENTRAL OPISTHOSOMAL SURFACE (1) OR POUCH ABSENT (0). 101. CRIBRUM INCLUDING WELL-SCLEROTIZED BAND (2) OR NOT (1). 102. NUMBER OF AV SETAE ON FEMUR I. 103. GENU IV WITH AV PRESENT (1) OR ABSENT (0). 104. GENU IV WITH PV PRESENT (1) OR ABSENT (0). 105. NUMBER OF PL SETAE ON GENU IV. 106. GS3 AT LEAST 3 TIMES LONGER THAN GS2 (2) OR NOT (1). 107. AT LEAST SOME DORSAL j SETAE RELATIVELY BROAD BASALLY, SUDDENLY NARROWING (2) OR NOT (1). 108. NUMBER OF AL SETAE ON FEMUR I. 109. ANTERIOR EDGE OF PODONOTUM ANTERIOR TO BOTH GS1 SETAE (1) OR NOT (0). 110. CLAWS NORMAL (1), OR NOT CLAW-LIKE (0). 111. WELL-SCLEROTIZED VENTRAL SHIELD COVERING MUCH OF VENTER OF OPISTHOGASTER (NOT CONSIDERING GENITO-VENTRAL PLATE AND ANAL PLATE) (1), OR NO SUCH VENTRAL SHIELD (0). 66 APPENDIX II - CONTINUED

112. PALPAL GENU WITH Dl PRESENT (1) OR ABSENT (0). 113. PALPAL GENU WITH AL PRESENT (1) OR ABSENT (0). 114. J1-J4 CLEARLY RECOGNIZABLE ON BOTH SIDES (2), OR NOT, DUE TO HYPOTRICHY (1), OR NOT, DUE TO HYPERTRICHY (3). 115. BOTH j3 AND j4 SETAE RECOGNIZABLE (ON AT LEAST ONE SIDE) (2), OR NOT, DUE TO HYPOTRICHY (1), OR NOT, DUE TO HYPERTRICHY (3). 116. HEAVY LATERAL SCLEROTIZATION OF STERNAL SHIELD WHICH IS SEPARATE FROM, AND POSTERIOR TO, SCLEROTIZATION OF CHARACTER 28 PRESENT (2) OR ABSENT (1). 117. BARB WITHIN 25 MICRONS OF CORNICULUS TIP PRESENT (1) OR ABSENT (0). 118. NUMBER OF SETAE ON OPISTHONOTAL SHIELD. 119. PERITREMAL PLATE FUSED WITH DORSAL SHIELD (1) OR NOT (0). 120. Z5 RECOGNIZABLE ON AT LEAST ONE SIDE (1) OR OBSCURED BY HYPERTRICHY (2). 121. PALPAL FEMUR WITH AL PRESENT (1) OR ABSENT (0). 122. LATERAL SIDES OF ANAL SHIELD NEARLY PARALLEL (1) OR NOT (0). 123. GENITAL SETAE ON GENITO-VENTRAL SHIELD (1) OR OFF SHIELD (0). 124. ON TARSUS I, THE LENGTHY PL PRIOR TO TARSUS TIP (INCLUDING AREA OF DENSE SETATION) IS THE THIRD-MOST BASAL PL SETA (2) OR THE SECOND-MOST BASAL PL SETA (1). 125. SETA OF COXA IV SPINE-LIKE (2) OR NOT (1). 126. PL3 OF TARSUS II SPINE-LIKE (2) OR NOT (1). 127. ALl OF TARSUS IV SPINE-LIKE (2) OR NOT (1). 128. ' PL OF GENU III SPINE-LIKE (2) OR NOT (1). 129. AREA(S) OF HEAVY SCLEROTIZATION ON STERNAL SHIELD, WHICH IS ENTIRELY ANTERIOR TO STERNAL SETAE I PRESENT (2) OR ABSENT (1). 130. "CRIBRUM" (MECROTRICHAE) AT POSTERIOR END OF OPISTHONOTAL SHIELD (1) OR NO CRIBRUM ON OPISTHONOTAL SHIELD (0). 131. AV SURFACE OF TROCHANTER IV SCALE-LIKE (1) OR NOT (0). 132. COXA II WITH PROMINENT SCALE-LIKE APOPHYSIS PRESENT (1) OR ABSENT (0). APPENDIX III 1

basic data matrix for103 spec-imens NC = 99.

otu character 1 2 3 4 5 6 7 8 9 L3POOX 0. 1. 75. 47. 1. 1. 1. 1. 1. LIP002 0. 1. 79. 49. 1. 1. X. 2. 1. L 3PC03 0. 1. 80. 46. 1. 1. 1. 1. 1. LI POO4 0. 82. 50. 1. 1. 1. 1. 1. L J POOS 99. 99. 99. 99. 1. 1. 1. 1. LIP006 0. 1. 80. 50. 1. r. 1. 2. 1. L 3PC07 0. 1. 77. 48. 1. i. 1. 2. LIP008 0. 1. 76. 50. 1. i. 1. 2. i ! L3P009 0. 1. 74. 53. 1. i. 1. 1. LIPOIO 0. 1. 81. 49. 1. i. X. 1. i. L3P031 0. 1. 77. 47. 1. X. 1. 1. i. LIPO 12 0. 1. 77. 49. 1. 1. 1. L • i. L3P033 0. 1. 74. 50. 1. 1. X. 1. i. LIP0X4 0. 1. 74.: 49. 1. 1. 1. 1. i. L3P035 0. 76. 47. 1. 1. 1. i. LIP016 0. l\ 81. 58. 1. 1. 1. 1. i. 1 3 P074 0. 1. 81. 50. 1. 1. 1. 1. i. LIPO75 0. 1. 85. 34. 1. 1. 1. 1. i. LIP076 0. 1. 76. 46. 1. £ • 1. 1. x. LIP077 99. 99. 99. 99. 1. 1. 1. 1. X. L3P078 0. 1. 80. 46. 1. 1. 1. £ • 1. LIP079 o . 1. 81. 45. 1. 1. 1. X. 1. NEC'OI 7 0. 1 . 68. 44. 1 . 1 . C. I 1 .

NE0018 0 . 1 . 69. 41. 1 . i ! X. 1 . 1 . NE0019 0 . 1 . 66. 39. 1 . 1 . 1 . C 1 1 .

NE0020 0 . 1 . 66. 43. 1 . 1 . 1 . 1 . 1 .

NE0021 0 . 1. 66 . 44. 1 . 1 . 1 . 1 . 1 .

NE0022 0 . 1 . 63. 31. 1 . 1 . 1 . 1 . 99. N E O O f 5 0 . 1. 73. 44. 1 . 1. 1 . 1 . 1 . NE0086 0 . 1 . 72. 40. 1 . 1. 1 . 1 . 1 . NEOOf7 0 . 1 . 71. 45. 1 . 1 . 1 . C | 1 .

NE OOf-8 0 . 1 . 69. 45. 1 . 1. 1 . 1. 1 . NE0089 0 . 1. 64. 42. 1 . 1. 1 . 1. 1 .

FA 1023 o . 1 . 77. 42. 1 . 1. X. 1 . 1 . FA 3 024 0 . 1'. 78. 38. 1 . 1 . 1 . 1 .

FA 10 2 5 0 . 1 . 80. 45. 1 . 1 . X. 1 . 1 . 67 68

a p p e n d i x i i i - c o n t i n u e d

o t u c h a r a c t e r 3 2 3 4 5 6 7 8 9 • 1 F A 3026 0. 1. 80. 43. 1. 1. £ • 1. £ # FA 10 2 7 0. 1. 81. 47. 1. 1. 1 • 1. £ • FA 3 028 0. 1. 77. 43. 1. 1. £ • 1. 1 • FA 1029 0. 1. 77. 42. 1. £ * 1. £ • FA 3 020 0. 1. 76. 42. 1. 1. £ • 1. £ • FA 10 31 0. 1. 75. 46. 1. 1. £ • ' 2. £ • FA 3 090 0. 1. 81. 42. 1. 1. £ » 99. £ • FA 10 91 99. 99. 99. 99. 1. 1. £ • 1. £ • FA 3 092 0. 1. 80. 40. 1. 1. £ • 1. £ * PH 1067 3. 0. 35. 46. 0. 1. 0. 2. 0. PHI 068 2. 0. 35. 48. 0. 1. 0. 2. 0. PH 1069 3. 0. 34. 48. 0. 1. 0. 2. 0 . PH3070 3. 0. 35. 48. 0. 1. 0. 2 . 0. PH 1071 3. 0. 34. 46. 0. 1. 0. 2. 0 . PH 1072 3. 0. 36. 49. 0. 1. 0. 2. 0. G0R035 1. 0. 43 . 49. 0. 1. 0. 2. 0. G0R026 1. 0. 45. 41. 0. 1. 1. 2. 0. G O R O B 7 1. o. 41. 40. 0. 1. 0. 2. 0. G0R028 3. 0. 38. 53. 0. 1. 1. 2. 0. G0R039 1. o. 41. 47. 0. 1. 0. 2. 0. TP 305 0 3. 0. 38. 67. 1. 1. 1. 2. 0. TRI051 3. o. 35. 67. 1. 1. 1. 2. 0. TP3052 3. 0. 35. 56. 1. 1. 1. 2. 0. T R 10 33 3. o. 35. 64. 1. 1. 1. 2. 0. TR 3 093 3. 0. 37. 62. 1. 1. 1. 2. 0. RAD0 82 3. o. 38. 88. 1. 1. 0. 1. 0. R A D O f 3 • 0. 40. 85. 1. 1. 0. 1. 0. RAD0E4 3. o. 39. 88. 1. 1. 0. 1. 0. LEP022 0 . 1. 84. 44. 1. 1. 0. 1. 1. LEP033 0. 1. 85. 45. 1. 1. 1. 2. 1. J0H034 0. 99. 99. 51. 1. 1. 1. 1. BRA040 1. 0. 33. 50. 0. 1. 0. 2. 0. BRA097 1. 0. 31. 48. 0. 1. 0. 2. 0. TRU041 0. 1. 123. 45. 1. 1. 1. 1. 1. TRU045 0. 1. 115. 47. 1. 1. 1. 1. 1. BUT046 0. 1. 105. 49. 1. 1. 1. 1. 1. 0PH047 3. 0. 38. 69. 0. 1. 1. 2. 0. 0PH048 3. 0. 36. 62. 0. 1. 1. 2. 0. SCH049 3. 0. 44. 66. 0. c, • 1. 2. 0. FARO 54 3. o. 38. 78. 1. 1. 1. 2. 0. FAR055 0. 37. 74. 1. 1* 1. 2. 0. CAU056 99. 99. 99. 99. 1. 1. 1. 1. 0. CAU057 3. 0. 59. 111. 1. 1. 1. 1. 0. DIP058 3. 0. 33. 56. 1. 1. 1. 1. 0. 69

APPENDIX III - CONTINUED

OTU CHARACTER 1 2 3 4 5 6 7 8 9 FE1059 3. 0. 39, 80. 1. 2. 0. 1. 0. PIG060 3. 0. 37. 75. 1. 1. 0. 1. 0. PIG061 3. 0. 38. 71. 1. 1. 0. 1. 0. N0V062 3. 0* 35. .66. 0. 1. 0. 2. 0. TAN063 3, 0. 32. 70. 1. 1. 0. 2. 0. TAN064 3. 0. 32. 71. 1. 1. 1. 99. 0. EVA065 3, 0. 36. 44. 1. 1. 1. 2. 0. EVA0 66 3. 0. 38. 46. 1. 1. 1. 2. 0. 0KE073 3. 0. 47. 67. 0. 1. 0. 1. 0. NSP0 80 1. 0. 32. 40. 0. 1. 0. 2. 0. NSPOfl 1. 0. 32. 40. 0. 1. 0. 2. 0. CUB094 0. 1. 100. 48. 1. 1. 1. 1. 1. FCN095 0. 1. 136. 62. 1. 1. 0. 1. 1. F0N0 96 o. 1. 135. 43. 1. 1. 0. 1. 1. JAV0*8 3 « 0. 32. 52. 0. 1. 1. 2. 0. C0N099 3. 0. . 45. 106. 1. 1. 1. 2. 0. CAH300 3* 0. 49. 88. 1. 1. 1. 99. 0. LI 0101 2. 0 . 60. 60. 0 . 1. 1. 2. 0. L10]02 2. 0 . 55. 68. 0. 1. 1. 1. 0 . UPE103 3. 0 . 42. 56. 0. 1. 99. 99. 0. MEH104 3. 0. 41. 66. 0 . 2. 1. 2. 0. MEH105 3. 0 . 43. 67. 0. 2. 1. 2. 0. UN C 106 0. 1. 128. 53. 1. 1. 1. 99. 1. 70

a p p e n d i x i i i - c o n t i n u e d

OTU c h a r a c t e r 10 11 12 13 14 15 16 17 18 L 3 P001 11. 44. 11. 6. 32. 1. 68 LIP002 12. 48. 99. 99. 32. 1. 57 L ]P003 13. 51. 10. 6 . 33. 1. 68 LIP004 11. 52. 11. . 6. 32. 1. 66 L)POOS 12. 48 . 8. 6. 32. 1. 66 LIP006 10. 44 . 7. 6. 33. 1. 60 L3P007 12. 48. 8.- 6. 34. 1. 59 LIP008 14. 54. 9. 6. 32. 1. 55 L3P009 13. 48. 10. 6. 32. 1. 51 LIP010 15. 46. 8. 6. 32. 1. 65 L3P031 13. 46. 99. 99. 32. 1. 59 LIP012 14. 44 . 11. 6. 32. 1. 62 L3P033 16. 47. 10. 6. 32. 1. 67 LIPO 14 14. 47. 9. 6. 33. 1. 65 L3P035 13. 43. 12. 6. 33. 1. 66 LIPO 36 13. 48. 9. 6. 33. 1. 64 L 3 POT 4 13. 46. 7. 6. 33. 1. 57 U P 0 7 5 13. 52. 9. 6. 32.- 1. 59 L3P076 12. 44. 7. 6. 32. 1. 66 LIP077 15. 53. 8. 6. 32. 1. 62 L3P078 13. 50. 7. 6. 32. 1. 59 LIP079 15. 52. 8. 6. 32. 1. 67 NEG017 11. 39. 8. 6. 34. 1. 45 NEO018 12. 40 . 8. 6. 34. 1. 45 NE0019 31. 41. 7. 6. 32. 1. 43 NE0020 9. 33. 99. 99. 33. 1. 59 N EDO 11 33. 40. 8. 6 . 32. 1. 54 NE0022 11. 41. 8. 6. 30. 1. 40 NE0085 12. 41. 10. 6. 32. 1. 49 NE00 86 9. 35. 10. 6. 32. 1. 56 NEQ087 8. 37. 9. 6. 30. 1. 64 NE0088 13. 36. 6. 5. 32. 1. 63 N EOOf9 31. 36. 9. 6. 31. 1. 43 FA 1023 9. 44. 8. 6. 29. 1. ’38 FA 3 024 9. 50. 10. 6. 27. 1. 44 FA’102 5 9. 52. 8. 5. 26. 1. 39 FA 3026 33. 4 7 ‘. 5. 5. 24. 1. 44 FA 1027 12. 39. 9. 6. 28. 1. 51 FA 3 028 12. 41. 9. 6. 30. 1. 45 FA ]029 10. 43. 5. 5. 29. 1. 40 FA10?0 10. 38. 7. 6. 29. 1. 47 FA 1031 12. 50. 7. 6. 30. 1. 41 FA 3090 12. 49. 8. 7. 31. 1. 45 FA 1091 13. 46. 6. 6. 28. 1. 38 71

APPENDIX III - CONTINUED

OTU CHARACTER *• 10 11 12 13 14 15 16 17 18

FA 3 042 12. 43. 6 • 6 . 29. 1 . 45. 1. W PH 1067 13. 35. 1 2 . 6 . 40. 0 . 13. 1.

PH3068 13. 30. 15. 6. 36. 0 . 1 6 . 1. h h PHI069 13. 36 . 1 8 . 7. 37. 0 . 15. 1. PH3070 14. 36. 19. 6 . 39. 0 . 14. 1. PH1071 1 2 . 33. 14. 6 . 40. 0 . 15. 1. PHI 072 15. 36. 18, 7. 40. 0 . 14. 1. G0R035 1 0 . 40. 1 2 . 10. 26. 1 . 43. 1. G0R036 12. 38. 1 1 . 1 0 . 24. i; 37. 1. G0R037 1 1 . 39. 99. 99. 2 1 . l . 37. 1. GOP038 12. 39. 1 1 . 11. 24. l . 44. 1.

G0R039 1 1 . 38. 13. 13. 26. l . 39. 1. WWWWWOOOO TP3050 19. 24. 1 2 . 6 . 30. 3. 1 1 . 1.

TRIO 51 2 1 . 41. 1 0 . 6 . 33. 3. 1 0 . 1. h h

TR1052 19. 36. 8 . 6 . 32. 3. 8 . 1. O TR1053 1 8 . 41. 8 . .6. 31. 3. 1 1 . 1. TR]043 23. 40. 8 . 6 . 33. 3. 1 1 . 1.

RADO 82 16 , 34. 14. 6 . ‘29. 1 . 50. 1. h h h RADOf3 . 15. 39. 1 2 . 6 . 30. 1. 65. 1. RAD084 16. 34. 16. 6 . 29. 1 . 66 . 1. LEP022 15. 37. 1 1 . 6 . 75. 1. 44. 0. LEP033 1 2 . 41. 7. 6 . 71. 1 . 44. 0. J0H034 17. 53. 32. 15. 29. 1 . 60. 1. BRA040 1 2 . 34. 8 . 6 . 33. 3. 8 . 1. BRAO97 13. 32. 7. 5. 31. 2 . 8 . 1. TRU041 16. 60. 8 . 6 . 35. 1 . 37. 1. TPU045 16. 59. 8 . 7. 36.. 1 . 37. 1. BUT046 99. 99, 99. 99. 2 2 . 1 . 1 0 . 1. 0PHO47 25. 37. 48. 6 . 35. 3. 1 6 . 1. 0PH048 24. 35. 66 . 6 . 37. 3. 16. 1. SCH049 33.. 52. 53. 7. 34. 4. 6 . 1. FAR0E4 1 8 . 37. 17. 6 . 35. 1 . 25. 1. FAROE5 2 0 . 37. 9. 6 . 35. 1. 36. 0. CAU056 2 1 . 31. 50. 7. 35. 1 . 73. 1. CAU0E7 2 2 . 34. 54. 6 . 30. 1 . 83. 1.

DIP058 17. 34. 19. 7. 56. 1 . 38. 0. OWWWWOOO^'WWOO'P'WWOO

FE 3 059 16. 36. 8 . 6 . 34. 1. ■ 64. 1. h PIG060 2 0 . 38. 1 6 . 6 . 41. 1 . 35. 1. P3G061 20. 39. 15. 6 . 42. 1 . 36. 1. NOVO62 2 0 . 39. 1 0 . 8 . 34. 1 . 25. 1. T A NO 6 3 16. 32. 16. 7. 37. 2 . 1 2 . 1.

TAN0 64 15. 31. 1 2 . 6 . 34. 2 . 13. 1. WWWWW EVA065 15. 28. 58. 7. 32. 1 . 63. 1.

EVA066 17. 27. 56. 7. 32. 1 . 6 8 . 1. h h 72

appendix i i i - continued

o t u CHARACTER i’l 1 0 11 1 2 13 14 15 1 6 18

ONE073 24. 35. 62. 8 . 89. 99. 2 NSP0 80 1 1 . 29. 9. 8 . 50. 1 . 1 1 NSPOF1 33. 29. 7. 6 . 47. 1 . 14 CUB0 94 14. 52. 1 0 . 6 . 40. 1 . 53 FCNOc. 5 19. 64. 8 . 6 . 36. 1 . 15 F0N0 96 17. 36. 99. 99. 33. 1 . 16 JAV098 17. 37. 30.- 7. 42. 1 . 38 CONO9 9 23. 43. 8 . 6 . 28. 1 . 65 CAHIOO 2 1 . 49. 24. 6 . 33. 1 . 38 LI DIO 1 14. 40. 1 0 . 6 . 33. 1 . 13 L ]03 02 14. 43. 6 . 6 . 32. 1 . 1 2 UPE103 2 0 . 34. 32. 8 . 33. 4. 6 MEH104 38. 58. 37. 6 . 32. 4. 6 MEH10 5 39. 52. 36. 6 . 34. 4. 5 UNC 3 06 17. 69. 99. 99. 31. 1 . 67 APPEND IX III - CONTINUED

* * OTU t character 19 20 21 22 23 24 25 26

L1P001 0 . 1. 0 . o. 307. 118. 0. LIP002 0 . 1. 0 . 0 . 317. 118. 0. L3P003 0 . 1. 0 . 0 . 332. 146. 0. LIP004 0 . 1. 0 . 0 . 346. 141. 0. L) POOS 0 . 1. 0 . 0 . 348. 147. 0. LIP006 0 . 1. 0 . 0 . 333. 138. 0. L] P007 0 . 1. 0 . 0 . 336. 138. 0. LIP008 o. 1. 0 . 0 . 328. 132. 0. LJP009 0 . 1. 0 . 0 . 316. 131. 0. LIP010 0 . 1. 0 . 0 . 332, 148. 0. L3P031 0 . 1. 0 . 0 . 328. 143. 0* L IPO 12 0 . 1. 0 * 0 . 332. 130. 0. L3P013 0 . 1. 0 . 0 . 354. 140. 0. L IPO 14 o. 1. 0 . 0 . 350. 140. 0. L3P035 0 . 1. 0 . 0 . 319. 114. 0. L IPO 16 0 . 1. 0 . 0 . 335. 132. 0. L 1 P0*V4 0 . 1. 0 . 0 . 319. 127. 0. LIP075 o. 1. 0 . 0 . 349. 159. 0. L3P076 0 . 1. 0 . 0 . 304. 136. 0. LIP077 0 . 1. 0 . 0 . 336. 156. 0. L 3P078 o. 1. o # 0 . 350. 172. 0. LIP079 0 . 1. 0 . 0 . 329. 137. 0. NE0017 0 . 1. 0 . 0 . 282. 1 2 0 . 0. •NE0018 0 . 1. 0 . 0 . 281. 1 1 2 . 0. NED03 9 o. 1. 0 , 0 . 266. 108. 0. NE00 20 0 . 1. 0 . 0 . 274. 113. 0. NEGO21 0 . 1. 0 . 0 . 276, 118. 0. NE0022 0 . 1. 0 . 0 . 249. 104. 0. NEDOf5 0 . 1. 0 . 0 . 313. 123. 0. NE0086 0 . 1. 0 . 0 . 228. 1 0 0 . 0. N E 00 {■ 7 0 . 1. 0 . 0 . 299. 1 2 0 . 0. NEOOf 8 o. 1. 0 . 0 . 305. 115‘. 0. NE0089 0 . 1. 0 , 0 . 288. 138. 0, FA 102 3 o. 1. 0 . 0 . 310. 1 0 0 . 0, FA 3 024 0 . 1. 0 . 0 . 307. 108. 0. FA 102 5 0 . 1. 0 . 0 . 340. 138. 0. FA J026 0 . 1. 0 . 0 . 343. 127. 0. FA 1027 0 . 1. 0 . 0 . 328. 119. 0. FA 3 028 0. 1. 0 . 0 . 302. 109. 0. FA 102 9 0 . 1. 0 . 0 . 302. 1 0 2 . 0. FA 3 020 0 . 1. 0 . 0 . 312. 98. 0. FA 10 31 0 . 1. 0 . 0 . 294. 97. 0. FA ]090 0 . 1. 0 . 0 . 314. 105. 0. FA 1091. 0 . 1. 0 . 0 . 324. 94. 0. APPENDIX III - CONTINUED

,sO T U ' c h a r a c t e r . " 19 20 21 22 23 24 25 26 27

FAI092 0 . 0 . 0 316. 112. 1 . 0 . 99 PH 1067 0 . • 2 . 0 203. 71. 1 • 0 . 1 PHI 068 0 . 1 . 0 204. 58. 1 . 0 . 1 PH 10 6 9 0 . 1. 0 213. 60. 1 . 0 . 1 PHI070 0 . • 99, 0 207. 66. 1 . 0 . 1 PH1071 0 . 1. 0 197. 60. 1 . 0 . 1 PH)07 2 0 . 2 * 0 203. 72. 1 . 0 . 1 G0R03 5 X • 1. 0 369. 121. 1 . 0 . 0 G0R026 1 9 0 . 0 326. 131. 1 . 0 . 0 G0R037 X » 0 . 0 309. 136. 1 . 0 . 0 GCR038 1 * 0 . 0 344. 152. 1 . 0 . 0 G0R039 X 9 0 . O 349. 128. 1 . 0 . 0 TRIOEO 0 , 2 . 0 269. 74. 1 . 0 . TRI051 0 . . 2 . 0 245. 64. 1 . 0 . TR1052 0 . 2 . O 271. 68. 1 . 0 . TR10 53 0 . . 2 . 0 238. 74. 1 . 0 . TRI093 0 , 2 . 0 254. 72. 1 . 0 . RAD0P2 0 . 0 . 1 319. 112. 1 . 0 . RADOf-3 0 . 0 . 1 311. 94. 1 . 0 . RAD0f4 0 . 0 . 1 299. 115. 1 . 0 . LEPO-2 0 . 0 . 0 288. 135. 1 . 0 . LEP023 0 . 0 . 0 310. 144. 1 . 0 . JOHO34 2 . 0 . 1 330. 132. 1 . 0 . BRA040 0 . 2 . 0 281. 103. 1 . 0 . BRA097 0 . 2 . 0 265. 121. 1 . 0 . TRU041 1 . 0 . 0 308. 98. 0 . TRU045 1. 0 . o 329. 95. 1 . 0 . BUT046 2 . 1. 0 356. 99i 1 . 0 . 0PH047 0 . • 2 • 0 460. 72. 1 . 1. 0PH048 0 . . 99. o 462. 67. 1 . 1. SCH049 0 . 2 . 0 410. 33. 1 . 1. FARO 54 0 . 1. 0 238. 69. u • 0 . FAR055 0 . 1 . 0 253. 81. u . 0 . CAU056 1 . 0 . 0 299. 102. 1 • 0 . CAU057 0 . 0 . 0 283. 107. 1 . 0 . DIP058 0 . 0 . o 278. 65. 1 • 0 . FE1059 0 . 0 . 1 344. 118. 1 . 0 . PI GO60 0 . 0 . 0 243. 77. 0 . 0 . PIG061 0 . 0 . 0 259. 72. 0 . 0 . NDV062 0 . 1. 0 310. 128. 0 . 0 . T A NO 6 3 0 . 1 . 0 330. 67. 1 . 0 . TAN064 0 . 1 . 0 334. 60. 1 . 0 . EVA065 0 . 3. o 319. 79. 1 . 0 . EVA066 0 . . 3. 0 299. 84. 0 . 0 . APPENDIX III - CONTINUED

OTU character Vt - ^ . 19 20 21 22 23 24f 25 26 27 CWE073 0. 99. 99. 99 « 380. 99. 1. 0 . 1 NSPOfcO 1. ■ 1.. 2. 0 • 172. 67. 1. 0 . 1 NSPOfl 1. 1. 1. 0 • 189. 76. 1. 0 . 1 CUBO«T4 0 . 1. 0. 0 • 319. 140. 1 . 0 . 1 Ft NO?5 0. 0. 2. 0 • 385. 128. 1. 0 . 1 FDN0 96 0 . 0. 2. 0 • 420. 160. 1 . 0 . 1 * J A V 0 * 8 0. 1. 0. 0 • 288. 67. 1. 0 . CONOC9 0 . 1. 0. 0 » 324. 81. 1 . 0 . 1 CAHJOO 0. 1. 1. 0 • 395. 96. 1. 0 . 1 LI0101 0 . 1. 2. 0 • 287. 46. 1. 0 . 1 LJ0102 0. 1. 2 . 0 • 268. 45. 1. 0 . 1 UPE103 0 . 1. 2 . 0 • 248. 48. 1 . 0 . 99 MEH104 0 . 1. 2. 0 • 364. 40. 1. 1. 99 MEH105 0 . 1. 2. 0 • 369. 46. 1. 1. 1 UNC3 06 0. 1. 0 . 0 • 415. 91. 1. 0 . 99 76

APPENDIX I I I - CONTINUED

OTU character i*j 28 29 30 31 32 33 34 3 5 36 i LIPOOl 1 . 3. 1 6 6 , 33. 35. 90. 1. 0 . L3P002 1. 3, 1 72. 36. 35. 97. 1. 0 . LIP003 • 1 . 3. 1 74. 44. 37. 105. 1. 0 . L3P004 1. 3. 1 74. 41. 45. 114. 1. 0 . LI POO5 1 . 3. 1 77. 40. 37. 100 . 1. 0 . LIP006 1. 3. 1 69. 37. 40. 96 . 1 . 0 . LIP007 1 . 3. 1 ' 85. 40. 43. 95. 1. 0 . L3P008 1. 3. 1 6 6 . 40. 37. 101. 1. 0 . LIP009 1 . 3. 1 74. 40. 37. 101. 1. 0 . L3P030 1. 3. 1 74, 41. 35. 112. 1. 0 . LIPOll 1 . 3. 1 69. 42. 34. 100. 1. 0 . L3P012 1. 3. 1 73. 42. 42. 100. 1. 0 . LIPO 13 1 . 3. 1 77. 44. 35. 105. 1. 0 . L3P014 1. 3, 1 71. 44. 32. 103. 1. 0 . LIPO 15 1 . 3. 1 71. .46. 36. 105. 1. 0 . L3P016 1. 3. 1 78. 42. 28. 112.. 1. 0 . LIP074 .1. 3. 1 69. 44. 40. 101. 1. 0 . L3P075 1. 3. 1 72. . 45. 37. 113. 1. 0 . L1P076 1 . 3 • 1 6 6 . 41. 45. 96. 1. 0 . LDP077 1. 3. 1 76. 41. 36. 103. 1. 0 . LIP078 1 . 3. 1 80. 43. 36. 114. 1. 0 . LI POT9 1. 3, 1 80. 39. 37. 98. 1. 0 . NE0017 1 . 3. 1 59. 35. 26. 98. 1. 0 . NE00 3 8 1. 3. 1 54. 31. 34. 90. 1. 0 . NEOO19 1 . 3. 1 54. 35. 31. 95. 1. 0 . NE0020 1. 3. 1 55. 24. 35. 111. 1. 0 . NE0021 1 . 3. 1 52. 26. "36. 98. 1. 99. NE0022 1. 3. 1 59. 28. 28. 90. 1. 0 . NE0085 1 . 3. 1 60. 34. 32. 104. 1. 1 . NEOO£6 1. 3. 1 59. 2 1 . 26. 83. 1. 0 . NEOO67 1 . 3, 1 49. 25. 38. 97. 1. 1. NE0O68 1. 3. 1 52. 29. 37. 104. 1. 1. NEOO89 1 . 3. 1 64. 29. 39. 110. 1. 0 . FA 3 023 1. 3. 1 45, 44. 36. 87. 1. 0 . FA 1024 1 . 3. 1 46. 44. 39. 76. 1. 0 . FA 3 025 1, 3. 1 6 6 . 39. 43. 93. 1. 0 . FA 1026 1 . 3. 1 55. 45. 43. 80. 1. 0 . FA 3 027 1. 3. 1 52. 42. 44. 84. 1. 0 . FA 10 2 8 1 . 3. 1 44. 46. 41. 83. 1. 0 . FA 1029 1. 3. 1 40. 42. 27. 69. 1. 0 . FA 1030 1 . 3. 1 43. 47. 23* 85 . 1 • 0 . FA] Or 1 1. 3. 1 51. 35. 38. 85. 1. 0 . FA 1090 1 . 3. 1 49. 44. 40. 72. 1. 0 . FA 1091 1. 3. 1 49. 49. 31. 67. 1. 0 . 77

APPENDIX III - CONTINUED

. ’ I OTU CHARACTER 28 29 30 31 32 33 34 35 36

FA 1092 1. 3. 1. 50. 47. 24. 73. 1. 0 . PH3067 6 » 3, 1. 30. 31. 2 1 . 62. 1 . O'. PH1068 2 . 3. 1. 29. 29. 21. 54. 1 . 1. PH3069 6 . 3. 1. .31. 31, 22. 59. 1 . 1. PH 1070 6 • 3m 1. 30. 32. 23. 57. 1. 0 . PHJ071 6 . 3m 1. 30. 33. 18. 59. 1 . 0 . PHI072 6 . 3m 1. 29. 30. 23. 60. 1 . 0 . GPR035 ^ • 1 • 1. 64. 56. 34. 76. 0 . 0 . G0R036 3. X • 1. 71. 54. 26. 80. 1 . 0 . G0R037 w • X • 1. 70. 47. 42. 76. 0 . 0 . G0R038 3. x • 1. 74. 50. 45. 108. 0 . 0 . G0R039 3. X i 1. 63. 53. 32.. 95. 0 . 0 . TRIO 50 X • 3. 1. 54. 40. 39. 73. 99. 99. TR]051 'X • 3. 1. 55. 39. 41. 69. 1 . 99. TR10 52 x • 3. 1. 58. 34. 38. 80. 1 . 0 . T P1053 i t 3. 1. 58. 34. 43. 77. 1 . 0 . TRI093 * • 3. 1. 62. 35. 43. 64. 1 . 0 . RAD0F2 4* 2 . 1. 54. 43. 40. 100. 0 . 0 . RAD0R3 3. 2 . 1. 54. 48. 25. 95. 0 . 0 . RADOf4 3. 2 . 1. 49. 31. 23. 100 . 0 . 0 . LEP032 1 . 3. 1. 72. 28. 35. 1 1 1 . 1 . 1 . LEPO-3 3. 3. 1. 70. 30. 30. 109. 1. 1 . JOH034 1 . 3. 1. 7 4 . 69. 48. 72. u • 0 . BRA040 3. 3m 1. 59. 27. 48. 91. 1. 0 . BRAO97 3. 3rn 1. 72. 30. 34. 77. 1 . 0 . TRU0A1 ]. 3rn 1. 67. 31. 36. 76. 1 . 0 . TRU045 1 . 2 . 1. 63. 39. 21 . 63. 1. 0 . BUT 046 1. 3rn 0. 64. 35. 99. 82. 1. 0 . 0PH047 1 . 3. 1. 3 5 . 30. 50. 8 8 . 1 . 1 • 0PH048 1. 3m 1. 37. 34. 52. 93. 1. 1 • SCH0A9 1 . 3m 1. 32. 38. 29. 87. 1. 0 * FAR054 6 . 3. 1. 54. 26. 29. 67. 1 . X • FARO55 4. 3. 1. 54. 36. 39. 76. 1. X • CAU056 6 . 3rn 1. 72. 35. 26. 96. 1 . X • CAU057 5. 3m 1. 7 7 . 39. 19. 91. 1. X • DIP058 2 . 3m 1. 55. 32. 25. 55. 1 . X • FE 1059 3. 2 . 1. 42 . 26. 38. 85. 0 » 0 • P3G060 3. 3. 1. 43. 34. 29. 72. 1. X • PI GO61 5. 3. 1. 4 8 . 31. 37. 71. 1. X • NCV062 6 . 3. 1. 59. 32. 39. 82. 1. 0 . TAN063 6 . 3. 1. 4 0 . 48. 26. 53. 1 . 0 . TAN064 6 . 3. 1. 40. 50. 32. 49. 1 . 0 . EVA065 5. 3. 1. 4 7 . 35. 26. 1 0 2 . 1. 0 . ElAO66 6 . 3. 1. 52. 37. 26. 101 . 1. 0 . 78

APPENDIX III - CONTINUED

• OTU CHARACTER 28 29 30 31 32 33 34 35 36

0ME073 6 « 3. 1. 104. 49. 48. 6 0 . 0 . 0 . NSPOfO 3. 3. 1 . 22. 29. 20. 6 8 . 1 . 1. NSP081 1. 3. 1. 25. 32. 2 2 . 79. 1 . 1. CUB094 1. 2 . 1. -77. 43. 17. 102,. 1 . 0 . F0N095 1 . 2 . 1. 93. 56. 20 . 67. 1 . 0 . FTNOf 6 3. 2 . .1. 95. 48. 27. 69. 1 . 0 . JAV098 1. 3. 1. 45. 23. 22. 67. 1 . 0 . CCN0C9 3. 3. 1. 50. 43. 31. 83. 1 . 0 . CAHIOO 4 • 3. 1. 79. 49 . 44. 73. 1 . 0 . L J0301 5, 2 . 1. 67. 56. 35. 73. 1 . 0 . LI0102 5. 2 . 1. 64. 52. 31. 77. 1 . 0 . UFE303 3. 3. 1. 49. 40. 30. 81. 1 . 0 . MEH104 1 . 3. 1. 31. 46 . 29. 109. 1 . 0 . MEH105 3. 3. 1. 29. 43. 29. 112. 1 . 99. UNC106 1 . 3, 1. 79. 53. 17. 69. 1 . 0 . 79

APPENDIX III - CONTINUED

OTU ' CHARACTER 37 38 39 40 41 42 43 44 45

LIP001 35. 165. 3. 1. 1 . 2 . 0 . 2 . 91. LJP002 30. 197. 3. 1. 1 . 2 . 0 . 2 . 89. LIP003 35. 209. 3. 1. 1 . 2 . 0 . 2 . 88. L3P004 37. 2 0 0 . 3. 1. 1 . 2 . 0 . 2 . 107. LIP005 36. 219. 3. 1. 1 . 2 . 0 . 2 . 79. L1P006 31. 195. -3. 1. 1 . 2 . 0 . 2 . 90. LIP007 34. 214. 3. 1. 1 . 2 . 0 . 2 . 92. L] P008 31. 179. 3. 1. 1. 2 . 0 . 2 . 85. LIP009 2 8 . 184. 3. ,1. 1 . 2 . 0 . 2 . 94. LJP030 35. 186. 3. 1. 1. 2 . 0 . 2 . 96. LIPOll 29. 2 0 2 . 3. 1. 1 . 2 . 0 . 2 . 92. L3P032 34. 204. 3. 1. 1 . 2 . 0 . 1 . 107. LIP033 35. 192 . 3. 1. 1 . 2 . 0 . 2 . 102. L3P014 34. 195. 3. 1. 1 . 2 . 0 . 2 . 101. LIP015 32. 173. 3. 1. 1 . 2 . 0 . 2 . 96. L3P036 34. 197. 3. 1. 1 . 2 . 0 . 2 . 115. LI POT4 38. 205. 3. 1. 1 . 2 . 0 . 2 . 95. L 3 POTS 37. 187. 3. 1. 1 . 1. 0 . 2 . 118. LIP076 34. 205. 3. 1. 1 . 2 . 0 . 2 . 85. L 3P077 33. 178. 3.. 1. 1 . 2 . 0 . 0 . 101. LIP078 31. 2 1 0 . 3. 1. 1 . 2 . 0 . 2 . 101. L3P079 36. 184. 3. 1. 1 . 2 . 0 . 2 . 91. NED017 33. 138. 3. 1. 1 . 1. 0 . 3. 53. NE0018 37. 171. ' 3. 1. 1 . 1. 0 . 3. 81 . NEOO19 36. 149. 3. 1. 1 . 1. 0 . 4. 43. NE0020 41. 125. 3. 1. 1 . 1. 0 . 2 . 106. NEOO21 43. 137. 3. 1. 1.. 1. 0 . 2 . 84. NE0022 41. 1 6 1 . 3. 1. 1 . 1. 0 . 2 . 76. NEOO85 38. 169. 3. 1. 1 . 1 . 0 . 2 . 111. NEOOf6 46. 146. 3. 1. 1 . 1. 0 . 2 . 65. NEOO87 50. 155. 3. 1. 1 . 1. 0 . 2 . 111. NEOOf8 41. 148. 3. 1. 1 . 1. . 0 . 2 . 113. NE0089 34. 148. 3. .1. 1 . 1 . 0 . 2 . 89. FA 3023 145. 3. 1. 1 . 1. 0 . 2 . 47. FA 1024 29. 131. 3. 1. 1 . 1 . 0 . 2 . 35. FA 3 025 29. 2 0 0 . 3. 1. 1 . 1. 0 . 3. 41. FA 1026 31. 173. 3. 1. 1 . 1. 0 . 2 > 36. FA 3 027 — • 148. 3. 1. 1. 1. 0 . 3. 36. FA 1028 33. 130. 3. 1. 1 . 0 . 3. 29. FA 3 029 31. 120 . 3. 1. i! 0 . 0 . 47. FA 1030 35. 134. 3. 1. i . 0 . 2 . 37. FA 3031 32. 142. 3. 1. i. ll 0 . 2 . 39. FA 1090 35. 127. 3. 1. i . 1. 0 . 2 . 34. FA3091 36. 145. 3. 1. i . 1. 0 . 2 . 47. APPENDIX III - CONTINUED

OTU CHARACTER 37 38 39 40 41 42i 43 44 4 5 FA 10 92 36. 126. 3. 1. 1 . 1 . 0 . 2 . 3 3 . PHI067 34. 145. 3. 1. 0. 2 . 0 . 2 . 4 7 . PH 1068 27. 138. 3. 1. 0. 2 . 0 . 2 . 52 . PHI069 30. 138. 3. 1. 0. 2 . 0 . 2 . 4 5 . PH 1070 28. 145. 3. 1. 0. 2 . 0 . 2 . 4 3 . PH1071 28. 152. 3. 1. 0. 2 . 0 . 2 . 4 4 . PH 1072 32. 143. 3. 1. 1 . 2 . 0 . 2. 4 4 • G0R025 34. 305. 2 . 1. 2 . 2 . 1 • 2 . 8 2 . G0R036 40. 271 . 3. 1. 2 . 2 . X i 1. 9 2 . G0R037 25. 250. 2 . 1. 2 . 2 . X • 2 . 90 • G0R038 41. 2 6 0 . 2 . 1. 2 . 2 . X • 2. 8 4 . G0R039 40. 283. 2 . 0. 2 . 2 . X • 2. 8 9 . TR1050 29. 164. 3. 1. 1 . 2 . 0 . 2. 83 . T R J 0 51 28. 138. 3. 1. 1 . 2 . 0 . 2 . 91 . TR1052 25. 135. 3. 1. 1 . 2 . 0 . 2 . 7 6 . TPI0E3 24. 146. 3. 1. 1 . 2 . 0 . 2 . 7 1 . TR1093 27. 165. 3. 1. 1 . 2 . 0 . 2. 6 8 . RAD082 47. 243. 3. 1. 1 . 2 . 0 . 2 . 1 0 6 . RAD0 83 56. 229. 3. 1. 1 . 2 . 0 . 2 . 1 1 5 . RAD084 60. 215. 3. 1. 1 . 2 . 0 . 2 . 9 7 . LEP032 58. 172. 2 . 1. 1 . 1 . 0 . 5. 1 0 9 . LEP033 58. 174. 2 . 1. 1 . 2 . 0 . 6 . 91 . J0H034 34. 240. 3. 1. 0. 3. 0 . 0 . 81 . BPA040 23. 183. 2 . 1. 0 . 2 . 0 . 2 . 5 9 . BRA097 32. 197. 2 . 1. 0 . 2 . 0 . 2. 5 9 . TRUO^l 38. 2 0 0 . 2 . 1. 0 . 3. 0 . 2. 7 7 . TRU045 40. 172. 3. 1. 0. 2 . 0 . 2. 7 1 . BUT 04 6 22. 183. 3. 1. 1 . 1 . 0 . 1. 8 0 . OPH047 19. 246. 1. 1. 0. 3. 0 . 2 . 1 0 2 . GPH048 24. 242. 1. 1. 0. 3. 0 . 2 . 1 1 6 . SCH049 27. 246. 1. 1. 0. 2 . 0 . 2. 7 8 . FAR054 28, 151. 2 . 1. 1 . 2 . 0 . 2 . 7 7 . FARO 55 43. 1 8 2 . 3. 1. 1 . 2 . 0 . 2. 1 0 3 • CAU056 45. 179. 3. 1. 1 . 2 . 0 . 2 . 8 0 • CAU0 57 43. 156. 3. 1. 1 . 2 . 0 . 2. lOO . D] P058 37. 172. 3. 1. 1 . 1. 0 . 2 . 103 . FE1059 56. 2 1 8 . 2 . 1. 1 . 2 . 0 . 2. 1 1 2 . PI GO60 45. 2 1 0 . 3. 1. 1 . 2 . 0 . 2 . 131 . PI GO61 50. 197. 3. 1. 1 . 2 . 0 . 2 . 1 2 2 . NCV062 29. 2 0 0 . 3. 1. 1 . £ « 0 , 1. 9 2 . TA NO 63 25. 149. 3. 1. 1 . 1 t 0 . 2 . 4 3 . TAK064 31. 131. 3. 1. 1 . x • 0 . 2 . 3 3 . EVA065 99, 205. 1. 1. 1. 1 • 0 . 2 . 7 4 . EVA066 28. 183. 1. 1. 1 . x • 0 . 2 . 5 8 . 8 L

APPENDIX III - CONTINUED

OTU CHARACTER 37 38 39 40 41 V 43 44 45

CMEOT3 36 • 455. 2 . 0 . 0 . 2 . 0 . 0 . 109. NSPOfO 32, 137. 3. 1. 1. 2 . 0 . 2 . 94. NSPO&l 36 • 1-38. 3. 1. 1. 2 . 0 . 2 . 114. CUB094 45. 197. 2 . 1. 1. ■ 1. 0 . 2 . 92. F0N095 40. 251. 3. 0 . 0 . 1. 0 . 3. 32. F0N096 39. 283. .3. 99. 0 . 1. 0 . 2 . 50. JAVOS8 48. 192 . 3. 1. 0 . 1 . 0 . 3. 56. CCMOS9 61. 2 0 0 . 3. 1. 1. 1 . 0 . 2 . 92. cahioo 38. 2 1 0 . 3. 1. 1 . 2 . 0 . 2 . 69. L10101 39. 2 0 0 . 3. 1. 1. 2 . 0 . 2 . 57. LI0102 38. 2 0 0 . 3. 1. 1. 2 . 0 . 2 . 63. UPE103 21. 163. 3. 1. 1. 2 . 0 . 1 . 56. MEH104 28. 248. 1. 1. 0 . 2 . 0 . 2 . 73. MEH105 25. 218. 1. 1. 0 . 2 . 0 . . 2 . 75. UNC106 24. 2 2 8 . 2 . 1. 0 . 2 . 0 . 1 . 80. 82

APPENDIX III - CONTINUED

OTU CHARACTER r> 46 47 48 49 50 51 52 53 54 i LIPCOl 65. 182. 52. 64. 2 . 2 . 1. 1 . 1 LIP002 58. 187. 46. 74. 2 . 2 . 1. 1 . 1 L3P003 57. 202. 56. 61. 3. 2 . 1. 1 . 1 LIP004 61. 210 . 43. 80. 2 . 2 . 1. 1 . 1 L)POOS 130. 207. 52. 72. 2 . '2. 1. 1 . 1 LI POO6 51. 189. 44. 71. 2 . 2 . 1. 0 . 2 L1P007 50. 179. 48. 76. 3. 2 . 1. 1 . 2 LIP008 64. 208. 56. 75. 3. 2 . 1. 1 . 2 L]P009 62. 169. 46. 73. 3. 2 . 1. 0 . 2 LIPOIO 8 1 . 212. 46. 72. 2 . 2 . 1. 1 . 1 L3P031 72. 193. 53. 69. 3. 2 . 1. 1 . 1 LIPO12 8 1 . 234. 49. 72. 2 . 2 . 1. 1 . 1 L3P013 1 0 2 . 241. 58. 78. 2 . 2 . 1. 1 . 1 LIPO 34 97. ,215. 50. 75. 2 . 2 . 1. 1 . 1 ' L3P015 65. 189. 44. 72. 3. 2 . 1. 0 . 1 LIPO16 80. 208. 57. 61. 3. 2 . 1. 1 . 1 L3P074 55. 192. 39. 73. 2 . 2 . 1. 0 . 2 LIPO75 78. 212. 52. 81. 3. 2 . 1. 0 . 1 L3P076 50. 168. 44. 73. 3. 2 . 1. 1 . 2 LIP077 67. 197. 47. 83. 2 . 2 . 3. 0 . 99 L 3 POT 8 50. 223, 43. 74. 3. . 2. 1. 1 . 2 LIP079 77. 195. 48. 80. 2 . 2 . 1. 0 . 2 NEC03 7 50. 146. 39. 90. 3. 2 . 2. 0 . 2 NEOO18 59. 150. 37. 75. 2 . 2 . 2. 0 . 2 NEOO]9 45. 137. 31. 91. 3. 3. 1. 1 . 1 NEOO20 58. 143, 26. 81. 2 . 3. 1. 1 . 2 NE0021 57. 139. 32. 77. 2 . 2 . 1. 0 . 1 NEOO 2 2 57. 152. 40. 63. 3. 1 . 1. 0 . 2 NEOOf5 55. 151. 36. 83. 3. 2 . 1. 0 . 2 NE0086 33. 105. 31. 52. 2 . 2 . 1. 1 . 2 NEOOf7 56. 160. 34. 90. 2 . 2 . 1. 0 . 2 NEOOf8 66 • 157. 29. 82. 2 . 3. 1. 1 . 2 NE0089 76. 147. 32. 75. 3. 2 . 1. -1. 2 FA 1023 33. 109. 37. 75. 2 . 2 . 2. 0 . 2 FA]024 52. 109. 40. 71. 2 . 2 . 1. 0 . 2 FA 102 5 42. 146. 50. 70. 2 . 2 . 1. 0 . 2 FA3026 32. 128. 43. 73. 2. 2 . 1. 0 . 2 FA 10 2 7 25. 114. 44. 75. 2 . 2 . 1. 0 . 1 FA 3028 31. 1.10. 41. 83. 2 . 2 . 1. 0 . 1 FA 1029 23. 110. 37. 67. 2 . 2 . 2. 0 . 2 FA ]0-0 26. 116. 47. 69. 2. 2 .. 1. 0 . 2 FA )031 31. 106. 41. 6 8 . 2 . 2 . 1. 0 . 1 FA 3 090 32. 113. 56. 63. 2 . 2 . 1. 0 . 1 FA 10 91 90. 120. 43. 71. 2 . 2 . 1. 0 . 1 APPENDIX III - CONTINUED

■a OTU CHARACTER 46 47 48 49 50 51 52 53 54

FA 3 092 33. 108. 37. 76. 3. 2. 1. 0. 2. PHI067 75. 148. 40. 51. 2. 3. 2. 1. 2. PH] 06*8 67. 133. 50. 39. 2. 3. 1. 0. 1. PHI069 73. 156. 39. .57. 2. 3. 2. 1. 2. PH]070 77. 156. 45. 54. 2. 3. 3. 0. 2. PH107X 75. 146. 42. 54. 2. 3. 2. 1. 2 • PH]072 80. 152. 38. 55. 2. 3. 3. 1. 2. GOR03 5 50. 125. 54. 72. 4. 2. 3.. 0. X • GOR036 55. 128. 67. 65. 3. 1. 3. 0. 1 • GOR037 63. 122. 62. 74. 3. 2. 3. 0. X • G0R028 53 • 119. 64. 84. 3. 1. 3. 0. X • G0R039 48. 110. 65. 71. 3. 1. 3. 0. X • TPIOtO 8 6 . 114. 44. 64. 2. 3. 2. 0. 2. TR1051 1 0 0 .' 109. 38. 60. 2. 3. 3. 0. 2. TR1052 1 0 ]. 128. 48. 57. 2. 3. 3. 0. 2. TRI053 93. 121. 49. 52. 2. 3. 3. 0. 2. TP 1093 80. 120. 46. 58. 2. 3. 3. 1. 2. RAD0P2 57. 128. 58. 30. 2. 1. 2. 1. RAD0F3 59. 133. 6 6 . 2 8 . 2. 1. 2. 1. 1. RAD0 84 84. 133. 56. 30. 2. 1. 2. 1. 1. LEP032 81. 159. 47. 63. 2. 2. 2. 1. 2. LEP033 90. 166. 47. 6 6 . 3. 2. 2. 1. 2 • J0H034 132. 150. 61. 45. 4. 1. 3. 1. i. BRA040 8 6 . 121. 74. 49. 2. 3. 3. 1. 2. BRA097 8 6 . 115. 8 0 . 51. 2. 3. 3. 1. 2. TRU041 94. 120. 46. 81. 3. 2. 1. 0. 1. TRU045 72. 136. 38. 72. 3. 2. 2. 0. 1. BUT046 86 . 175. 33. 67. 3. 99. 99. 0. 1. OPH047 82. 179. 69. 78. 2. 2. 2. 1. 1. 0PH048 1 0 8 . 203. 65. 76. 1. 2. 1. 1. 1 . SCH049 161. 112. 6 8 . 59. 2. 2. 1. u • 1. FARO 54 88 . 128. 40. 45. 2. 2. 1. 1. 2. FAR0£5 95. 148. 99. 55. 99. 3. 2. 1. 2. CAU056 53. 174. 46. 87. 2. 3. 1. 1. 2. CAU057 54. 159. 40. 90. 2. 3. 1. 1. 1. DIP058 90. 119. 49. 31. 1. 2. 3. 1. 1. FEJ059 83. 133. 51. 36 . 2. 1. 3. 1. 1. PIGO60 53. 136. 46. 48. 2. 3. 3. 1. 2. P ]GOf 1 67. 138. 53. 51. 2. 3. 3. 1. 2. NOVO 62 94. 169. 6 6 . 48. 2. 2. 2. 0 • 1. TANO63 63. 118. 56. 90. 2. 3. 2. 1. 2. TAN064 46. 108. 6 6 . 81. 2. 2. 2. 1. 2. EVA065 47 • 116. 53. 26. 1. 2. 3. 0. 2. EVA066 46. 119. 46. 26. 1. 2. 3. 0. 2. appendix i i i - continued r> ' ■ . ■ 1 OTU CHARACTER 46 47 48 49 50 51 52 53 54

OMEOT3 244. 241. 44. 101 • 99. 3. 3. 0 . i . NSPOfO 69. 101. 41. 22 • 2 . 2 . 3. 0 . 1 . NSPOfil 90. 107. 40. 25 m 2 . 2 . 3. 0 . 1 . CUB0^4 46. 138. 40. 63• 2 . 2 . 3. 0 . 2 . FONOcs 6 8 . 174. 6 6 . 29 • 3. 1 . 1 . 0 . 1 . FCN0S6 93 • 182. 59. 35 • 3. 1. 2 . 0 . 1 . JAV098 78. 112. 53. 44 • 2 . 2 . 3. 1 . 2 . CC>N0<9 106. 168. 44. 94• 2 . 2 . 1 . 1 . 1 . CAH100 63. 133. 54. 50 • 2 . 2 . 3. 1 . 2 . L 30101 48. 91. 32. 43 • 2 . 2 . 3. 1 . 2 . L10102 56. 93. 30. 41 * 2 . 3. 3. 0 . 2 . UFE103 53. 106. 36. 55 • 2 . 3. 3. 1 . 2 . MEH104 143. 125. 56. 52 • 2 . 2 . 1 . 1 • 2 . MEH105 143. 128. 69. 53 • 2 . 2 . 1 . 1 . 2 . UNC106 59. 140. 73. 85 • 2 . 2 . 3. 0 . 1 . m 71 71 71 71 71 ■n *n 71 71 71 2 2 2 2 2 2 2 2 2 2 Z r - r* r - r~ r~ r~ r* r- r~ f—r - 1“ 1— i—r~ r~ r - I— i“ r—r~ ^ t> >> J> t> x> rn m rn m m m m m m m m *_ * M UJ i—» *—r U4 «_l M u_l *-4 ■_ i t—4 UJUl UJM UMM w Uf w »—»w w LJa c? O n o o o n o © o .-o *o "d "o "o "a 73 73 73 73 73 73 73 TJ■o -o -o "O TJ "O TJTJ o O o o O o o o o o o © o © o O o © o © o © o o o o o o o o o © O O o © o n o o o o o o o -t *f» *0 111 INJro T" M M ro *0 7) 7* rc>7ft no M M UJ>—• -J UJU-* UJMM H UJ a o o o O o o o o c O h b lO 00 -J O' Ul ^ WiO CO -0 O' Ul l\) J-* O vOCD -u!o 00 -4 O' Ul -t' O' Ul -P* LOfVJ!-» © vO 00 -J O' Ul -t' W M H 0

0 0 Ul 0 0 0 0 0 0 0 0 0 0 o o o o o o o o o © o o o OOOOOOOOO 0 0 o c o o o o o ui n • • ft • • • * • • ft • • ft ft ft ft • • • ft ft • • • ft ft ft ft ft ft • • ft ft • ft • • • ft ft * • ■ T > 73 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 Ul J> 0 0 0 0 0 o o o o o o o o o o 0 o o 0 o o O' n ft • • • • • • • ft • • ft ft • ft • • • « *ft • • • ft ft ft ft ft ft « • ft ft ft • ft • • • ft ft ft ft • -i m 73

3 3 3 Ul 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 UJ uj UJ UJUJ u> (jJ 3 3 UJUJ UJw -j • • • ft * * • • • ft ft • ft ft ft ft ft • ft • ft ••• • ft ft ft ft ft ft ft ft • ft • • • • • 9 ft ft ft • > * -o Ul -U 00 m z o t—1 Ul X vO M . M 1-4 0 0

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0 m 0 0 o o o o o o o ooO©o©ooo©o o o o © o © 0 0 o O O o o o o o o o o o © 0 o ro o • ••• • • • ft ft ft ft • ft ft ft ft ft ft • ft m • • • ft ft ft ft ft • • . ft ft ft • • • • • ft ft • ■ •

O' UJ

00 Ul APPENDIX III - CONTINUED

OTU CHARACTER 55 56 57 58 59 60 61 62

FA 1092 0 . 0 . 3. 1. 0 . ‘ 0 . 0 PHI067 0 . 0 . 5.. 1 . l l 0 . 0 . 0 PH 1068 0 . 0 . 5. 1. I. 0 . 0 . 0 PH]069 0 . 0 . 5. 1. . 1 . 0 . 1. 0 PH1070 0 . 0 . 5. 1. 0 . 0 . 0 PH]071 0. 0 . 5. 1. l l 0 . 1 . 0 PH 1072 0 . 0 . 5. 1 . 1 . 0 . 1 . 0 G0R035 0 . 0 . 3. I. 1 . 0 . 0 . 0 G0R036 0 . 0 . 3. 1. 1 . 0 . 0 . 0 G0R037 0 . 0 . 3. 1 . 1 . 0 . 0 . 0 G0R038 0 . 0 . 3. 1 . 1 . 0 . 0 . 0 G0R039 0 . 0 . 3. 1. 1 . 0 . 0 . 0 TRI050 0 . ' 0. 3. 1. 1 . 0 . 0 . 1 TRJ051 0. 0 . 3. 1 . 1 , 0 . 0 . 1 TR10 52 0 . 0 . 3. I. 1 . 0 . 0 . 1 TP 1053 0 . 0. 3. 1 . 1 . 0 . 0 . 1 TR1093 0 . 0 . 3. 1 . I. 0 . 0 . 1 RADOf2 0 . 0 . 4. 1 . 1 . 0 . 0 . 0 RAD083 0 . 0 . 4. 1. 1 . 0 . 0 . 0 RAD054 0 . 0 . 4. 1. 1 . 0 . 0 . 0 LEP032 0 . 0 . 3. 1. 1 . 0 . 0 . 0 LEP0r3 0 . 0 . 3. 1 . 1 . 0 . 0 . 0 JOH034 0 . 0 . 3. u • 0 * 0. 0. 0 BRAO^O 0 . 0 . 3. 1. 1 . 0. 0. I BRA057 0. 0. 3. 1. 1. 0. 0. 1 TRU.041 0, 0. 3. 1. 1.. 0. 0. 0 TRU045 0. 0. 3. 1. 1. 0. 0. 0 BUT.04 6 0. 0. I. 1. 0. 0. 0 0PH047 0. 0. 1. 0. 99. 1. 1. 1 0PH048 1. 0. I. 0. 99. 1. 1. 1 SCH049 1. 0.- 1. 0. 99. 1. 1. 1 FAR054 1. 0. 4. 1. 1. 0. 0. 0 FARO 5 5 1. 0. 4. I. 1. 0. 0. 0 CAU056 0. 0. 4. 1. 1. 0. 0. 0 CAU057 0. 0. 4. 1. 1. 0. 0. 0 D1P058 ]. 0. 4. 1. 1. 0. 0. 0 FE 1059 1. 0. 4. 1. 1. 0. 0. 0 PJGOfO 0. 0. 4. 1. 0. 0. 0 PIG061 0 . 0. 4. 1. l ! 0. 0. 0 Nrvot2 0. ■0. 4. 1. I. 0. 0. 0 TAN063 0. 0. 6 . 1. , 0. 0. 0 TAN064 0. 0. 6. i! 1 . 0. 0. 0 EVA065 0. 0. 7. 1. 1. 1. 0. 0 EYA066 0. 0. 7. 1. 1. 1. 0. 0 87

APPENDIX III - CONTINUED

OTU CHARACTER 55 56 57 58 59 60 61 62

OME073 0. 1. 3 . 0. 1. 1. 0. 0. NSPOf0 1. 0. 4 . 1. 1. 0 . 0. 0. NSP081 0. 0. 4 . 1. 1. 0 . 0. 0. CUB094 0. 0, 3 . 1. 1. 0 . 0. 0. FONO95 0. 0. 3 . 1. 1. 0 . 0. 0. FCM096 0. 0. 3. 1. X. 0 . 0. 0. JAVO 9 8 0. 0. 3 . 1. 1. 0 . 0. 0. CCN099 0. 0. 4 . 1. 1. 0 . 0. 0. cahioo 0. 0. 4 . 1. 1. 0 . 0. 0. L ]0101 0. 0. 1. 1. 99. 0 . 0. 0. L10102 0. 0. 1. 1. 99. 0 . 0. 0. AJPE103 0. 0 . 2. 1. 1. 0 . 0. 0. MEH104 1. 0. 1. 0. 99. 1. 1. 1. MEH105 0. 0. 1. 0. 99. 1. 1. 1.

UNCI06 0. ' 0. 3 . I. 1. 0 . 0. 0. 00 oo

APPENDIX III - CONTINUED ■ Otn < ' — o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o i— < Q.Q.0_G-0_Q-Q.CLQ_O.G.Q.0Q '■ C c tu C£ O o r- ' o o >0 O C\ I C \ JO < \ ] P O C V I t \ J < \ ) r M r \ i r \ l C V | C \ l < \ | £ \ J C \ l r \ ] C \ j r \ i r M C \ J ( \ J C \ l ( \ l C \ ] ( \ l { \ I C \ J C \ J < \ l f \ | { \ | ( \ J C \ l t \ J C \ J C \ J C \ J ( \ I C \ J C M C \ j r \ l ( \ J C \ J OvO r- r- cm oooooooooooooooooo'oooooooooooooooooooooooooo o o o o o o o o o o o o o o o o o o o o o o o o o o ' o o o o o o o o o o o o o o o o o o o

• t t tt HHHHHHHHHHHrHHHHHHHHrHH rtH H H H H H H H rtH H H H H H H H H H H lH rtrtF H rtH H H H H H H H H rtH H H H H H OO oo o o o o o o o o o o o o o o o o o o o o o o o o o O o o O O G O O G O O O O O O O O O O fSf\jmsi‘iA l . C L 0 - Q - Q . O - 0 - Q - C 1 - C 1 - C JJJJJZZZZZZZZZZZIi.lLU.U.IiU.iLU.U.U.11 1 . U . U . U L i . U i I . U . U L l . i I Z Z Z Z Z Z Z Z Z Z Z J J J J J j OO QpOOCi - ' i-,n n w n H " ''H m '-> O Q iQ C O O p Q m m - 1 - 1 2 * c ■0 X 000*mit/>00 CP H H 03 03 t- r 1- 7 3 7 3 7 3 - 1 - 1 —1-1 HOOOOO'OTJ'O'D'D'DTl < 1> > O M I—I m wl>t»l>t>0x'0 X 73 30 7373 O m mT>>l>7!xi3373 73oaoooxxxxxx> ■*> 3> X x < r O 0 1—4 ■oCC^>S»XxX -» cl cr J> ■*> x 73■ O O O O * - 1*—''— >—< 73 73 7J73 73*-<'—••—*■—••—•■—>•—1 o 0 O O 0 0 O 0 O OOOOOOOO O o o oo o o O O O O O O O O O o O O O O O O O 0 O 0 O —1 O'O' O' O ' O ' O' O ' U l ui ui in i,n in -P* -fr- .p--IN -P“ .IS >0 > w *v1 iiimis xi vO ' n ui 1 n Ul W W U W (>l -J ->i ^ O' o vfl c O ' U l -P* UJ l\J 0 00 -J O' Ul -S' '0 CD -J O' Ul -J O -(> w Ov005-JO'tnMt-*OvOOO-JfO

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—J

00 10 APPENDIX III - CONTINUED

OTU ' CHARACTER 64 65 66 67 68 69 70 71

OME073 1. 1. 2. 2. 3. X. 0. 2 NSPOf 0 0. 1. 2. 2. 2. 2. 0. 1 Nspoei 0. 1. 2. 2. 2. 2. 0- 1

CUB0S4 ■ 0. 1. 1, 1. 2. 2. 0. 1 F0N095 0. 1. 1. 1. 3. 2. 0. 1 FGN096 0. I. 1. 1. 3. 2. 0. 1 JAV098 0. 1. 2. 1. 3. 2. 0 •' 1 C CN099 0. 1. 2. 2. 3. 2. 0, 2 CAHlOO 0. 1. 1. 1. 3. 2, 0. 2 LIOIOI 0. 1. 1. 1. 3 • 1. 0. 1 LI 0102 0. 1. 1. 1. 3. 1. 0. 1 UFE103 0. 1. 2. 2. 3. 2. 0. 2 MEH104 0. 2. 2. 2. 3. 2. 1. 2 MEH105 0. 2. 2. 2. 3. 2. 1. 2 UNC106 0. 1. 1. 1. 3. 2. 0. 1 APPENDIX III - CONTINUED

ClTU ■ ■ CHARACTER 73 74 75 76 77 78

LI POO 1_ 0. 14. 2. ‘ 1. 2. 2. L ]P002 0. 13. 2. 1. 2. 2. LIP003 0. 14. 2. 1. 2. 2. LIP004 0. 15. 2. 1. 2. 2. LI PO O 5 0. 13. 2. 1. 2. 2. LJPC06 0. 13. 2. 1. 2. 2. LI POO7 0. 16. 2. 1. 2. 2. L )P008 0. 12. 2. 1. 2. 2. LIP009 0. 13. 2. 1. 2. 2. LIPOIO 1. 10. 2. 1. 2. 2. LIPOll 0. 11. 2. 1. 2. 2. LIP012 0. 12. 2. 1. 2. 2. L I P O 13 0, 15. 2. 1. 2. 2 • L3P014 0. 12. 2. 1. 2. 2. LIP015 0. .12. 2. 1. 2. 2. L3P036 0. 11. 2. 1. 2. 2. L1P074 0. 12. 2. 1. 2. 2. L 3 POT5 0. 13. 2. 1. 2. 2. LIP076 0, 8. 2. 1. 2. 2. L 3 POT7 0. 13. 2. 1. 2. 2. LIP078 0. 14. 2. 1. 2. 2. L 3 POT9 0. 12. 2. 1. 2. 2. N E O O 17 0. U. 2. 1. 2. 2. N EOO]8 0. 11. 2. 1. 2. 2. N E O O 19 0. 11. 2. 1. 2. 2. N E O O 20 0. 12. 2. 1. 2. 2. NE0.021 0. 11. 2. 1. 2.. 2. N E O O 22 0. 9. 2. 0. 2. 2. NEOOfi5 0. 14. 2. 1. 2. 2. NEOOf6 0. 10. 2. 1. 2. 2. N EOOf7 0. 10. 2. 1. 2. 2. NEOOfS 1. 10. 2. 1. 2. 2. N E O O 89 o. 10. 2. 1. 2. 2. FA 1023 0, 13. 2. 1. 2. 2. FA 1024 0. 10. 2. 1. 2. 2. FA 1025 0. 13. 2. 1. 2. 2. FA 1026 0. 14. 2. 1. 2. 2. FA 3027 0. 10. 2. 1. 2. 2. FA 10 2 8 0. 12. 2. 1. 2. 2. FA 1029 0. 13. 2. 1. 2. 2. FA 1030 0. 11. 2. 1. 2. 2. FA 1031 0. 13. 2. 1. 2. 2. FA 1090 0. 11. 2. 1. 2. 2. FA 3 091 0. 12. 2. 1. 2. 2. 92

APPENDIX III - CONTINUED

OTU ' CHARACTER 73 74 75 76 77 78 79 80

FA 1092 0 . 11. 2 . 1. 2 . . 2. 2 . 1 PH3067 0 . 12. 2 . 2 . 2 . 2. 2. 1 PH 1068 0 . 12. 2 . 2. 2 . 2. 2 . 1 PHI069 0 . 16. 2 . 2 . 2. 2. 2. 1 PH 1070 0 . 15. 2 . 2. 2 . 2. 2 . 1 PH1071 0 . 13. 2 . 2. 2. 2. 2 . 1 PH 1072 0 . 16. 2 . 2. 2 . 2. 2 . 1 G0R025 0. 16. 2 . 2 . 3. 2. 2. 1 G0R036 1 . 18. 2 . 2. 3. 2. 2 . 1 G0R037 ]. 17. 2 . 2. 3. 2. 2. 1 G0R038 0 . 18. 2 . 2. 3. 2. 2 . 2 GGR039 0 . 19. 2 . . 2 . 3. 2. 2 . 2 TRI050 0 . 18. 2 . 2. 3. 2. 2 . 2 TR1CJ1 0 . 18. 2 . 2 . 3. 2. 2. 2 T R 10 52 0 . -17. 2 . 2. 3. 2. 2 . 2 TRI0£3 ' 0 . 16. 2 . 2 . 3. 2. 2. 2 TRI093 o. 18. 2 . 2. 3. 2. 2 . 2 RAD0F2 0 . 16. 1. 1. 3. 2. 2. 2 RAD083 0 . 14. 2 . 2. 3. 2. 2 . 2 RADOf4 0 . 15. 2 . 2 . 3. 2. 2 . 2 LEP032 0 . 17. 1. 1. 2 . 2. 2 . 1 LEP033 0 . 20. 2 . 1. 2. 2. 2. J0H034 0 . 12. 2 . 1. 3. 2. 2 . 1 BRA040 0. 20. 2 . 2 . 3. 2. 2. 1 BRA097 0 . 22. 2 . 2. 3. 2. 2 . 1 ■TRU041 0 , 16. 2 . 1. 3. 2. 2 . 1 TRU045 0 . 15. 2 . 1. 3. 2. 2 . 1 BUI 04 6 0 . 18. 2 . 2. 2. 1. 2 . 1 OPH047 1 . 35. 2 . 3. 3. 2. 3. 2 0PH048 1. 36. 2 . 3. 3. 2. 3. 2 SCH049 1 . 31. 2 . 3. 3. • 2. 3. 2 FAR054 0 . 14. 2 . 2. 3. 2. 2 . 2 FARO 55 0 . 15. 2 . 2. 3. 2. 2 . 2 CAU056 0 . 21. 2 . 2. 3. 2. 2. 2 CAU057 0 . 18. 2 . 2. 3. 2. 2 . 2 D3P058 0 . 22.. 2 . 2. 3. 2. 2. 1 FE 1059 0 , 16. 2 . 2 . 3. 2. 2 . 2 P3G06O 0 . 19. 2 . 2 . 3. 2. 2. 2 PIGO61 0 . 17. 2 . 2. 3. 2. 2 . 2 N CV 0 6 2 0 . 24. 2 . 2 . 3. 2. 2. 2 TAN063 0 . 15. 2 . 2. 3. 2. 2 . 2 T A NO 64 0 . 15. 2 . 2 . 3. 2. 2 . 2 EVA065 1 . 26. 2 . 2. 3. 2, 2 . 1 EVA066 1. 28. 2 . 2 . 3. 2. 2. 1 APPENDIX III - CONTINUED

OTU 1 CHARACTER 73 74 75 7 6 77 78 7 9 80 81 OME073 1 . 68. 2 . 2 . 3. 2. 2 . 2. 1. NSPOfO 0 . 12. 2 . 2 . 3. 2. 2 . 1. 1. NSP0 81 0 . 12. 2 . 2 . 3. 2. 2 . 1. 1. CUB094 0 . 12. 2 . 1 . 2. 1. 2 . 1'. 1. FONO 95 1 . 12. 2 . 2 . 3. 2. 2 . 1. 1. FCN096 1. 16. 2 . 2 . 3. 2. 2 . 1. 1. JAV098 0 . 17. 2 . 2 . 3. 2. 2 • 2. 1. C O M O 99 0 . 28. 2. 2 . 3. 2. 2 . 2. 2. CAHIOO 0 . 20. 2 . 2 . 3. 2. 2 . 2. 2. L JOIOI 0. 16. 2. 2 . 2. 2. 2 . 1. 1. LI0102 0 . 17. 2 . 2 . 2. 2. 2 . 1. 1. UPE303 0 . 19. 2 . 3. . 3. 2. 2 . 2, 1. MEH10 4 1 . 31. 2 . 2 . 3. 2. 3 . 2. 2. MEH305 1. 35. 2 . 3. 3. 2. 3 . 2. 2. UNC106 0 . ,21. 2 . 1 . 3. 2. 2 . 1. 1. APPENDIX III - CONTINUED

OTU CHARACTER 82 83 84 85 86 871 88 89 90 LIPOOl 19. 1. 0 . 2 . 0 . LIP002 IT. 1. l! 0 . 2 . i! u 0 . LIP003 19. 1. 1. 0 . 2 . i . i . 0 . L3P004 20. 1. 1. 1. 2 . 0 . LIP005 17. 1. 1. 0 . 2 . i. i. 0 . L1P006 15. 1. 1. 0 . 2. i. i. 0 . LIP007 18. 1. 1. 0 . 2 . i. i . 0 . LIP008 18. 1. 0 . 2. i. i. 0 . LIP009 18. 1. i! ■0 . 2 . i. i. 0 . L3P030 18. 1. i. 0 . 2 . i. i. 0 . LIP011 17. 1. i. 0 . 2 . X 0 j X • 0 . L3P032 20. 1. i. 0 . 2. 1. 1. 0 . L I P O 13 21 . 1. i. 0 . 2 . 0 . L3P014 20. 1, 0 . 2 . i ! i. 0 . LIPO 15 17. . 1 . i! 0 . 2 . i. i. 0 . L3P016 20. 1. i. 0 . 2. i. i. 0 . LIP074 17. 1. i. 0 . 2 . 0 . L 3 PO T 5 20. 1. i. 0 . 2. i. i. 0 . LIP076 17. 1. i. 0 . 2 . i. i. 0 . L3P0T7 18. 1. i. 0 . 2 . i. i. 0 . LIP078 18. 1. i. 0 . 2 . 0 . L3 POT9 16. 1. i. 0 . 2. U 11 0 . NE0017 15. 1. i. 0 . 2 . 0. 1 . 0 . NE0038 14. 1. i. 0 . 2 . 0. 1. 0 . NEOO 3 9 13. 1. i. 0 . 2 . 0. 1 . 0 . N E C O 20 16. 1. i. 0 . 2 . 0, 1. 0 . N E O O 21 15. 1. 0 . 2 . 0. 1 . 0 . ME0022 13. 1. il 0 . 2. 0. 1. 0 . N E O O 85 16. 1. 0 . 2 . 0. 1 . ' 0 . NEOOF6 12. 1. i! 0 . 2. 0. 1. 0 . 0 N E O O 87 15. 1. i. 0 . 2. 0. 1 . 0 . 0 NEOO £8 14. 1. i. 0 . 2. 0. 1. 0 . 0 N E O O 89 14. 0. i. 0 . 2. 0. 1 . 0 . 0 F A 3 023 15. 1. i. 0 . 2. 0. 1. 0 . 1 FA 1024 14. 1. i. 0 . 2. 0. 1 • 0 . 1 FA 3 025 IP. 1. i. 0 . 2. 0. 1. ■ 0 . 1 FA 1026 18. 1. i. 0 . 2. 0. 1 • 0 . 1 PA 3027 16. 1. i. 0 . 2. 0. 1. 0 . FA 10 2 8 16. 1. i. 0 . 2. 0. 1 . 0 . 1 FA 3029 15. 1. 0 . 2. 0. 1. 0 . 1 FA 1030 15. 1. l\ 0 . 2. 0^ 1 . 0 . 1 FA 3031 16. 1. i. 0 . 2. 0. 1. 0 . 1 FA 1090 16. 1. i. 0 . 2. 0. 1 . 0 . 1 FAJ091 16. 1. i. 0 . 2. 0. 1. 0 . 1 95

APPENDIX III - CONTINUED

OTU character 82 83 84 85 86 88 89 90

FA 1092 16. 1. 1. 0 . 2 . 0 PHI 067 17* 1. 2. 0 . 2. PH 106 8 15. 1. 2. 0 . 2 . PH 1069 17. 1. 2. 0 . 2. PH 1070 18. 1 . 2. 0 . 2 . PH3 071 16. 1. 2. 0 . 2.

PH 1072 18. 1 . 2. . 0 . 2 . G 0 R 0 3 5 18. 1. 2. 0 . 2. G0R036 19. 1. 2. 0 . 2 . G 0 R 0 3 7 17. 1. 2. 0 . 2. G 0 R 0 3 8 20. 1 . 2. o.. 2 . G 0 R 0 3 9 16. 1. 2. 0 . 2. TR 10 50 21. 1. 2 . 0 . 3. T R 3 051 20. 1. 2. 0 . 3. TR 10 52 19. 2. 2. 0 . 3. T R1 0 5 3 19. 1. 2, 0 . 2. T R I 0 9 B 20 . 1. 2. 0 . 3. RADOf-2 14. 1. 1. 0 . 2. RA DO E 3 13. 2. 1. 0 . 2 . R A D 0 8 4 14. 1. 1. 0 . 2. LEP 0 3 2 18. 1. 1. 0 . 1. LEP033 19. 1. 0 . 2. JOH034 15. 1. i! 0 . 99. B R A 0 4 0 20. 1. i. 0 . 2. B R A 0 9 7 20 . 1. i. 0 . 2 . TRU041 17. 1. 0 . 2. o o o TRU045 17. 1. i! 0 . 2 . BUI 046 22. 1. i. 0 . 2. O P H 0 4 7 29. 1. 2. 0 . 3. 0PH048 27. 1. 2. 0 . 3. SC HO 4 9 32. 1. 2. 0 . 3. F A R O 54 15. 1. 2. 0 . 2. F A R 0 S 5 19. 1. 2. 0 . 2 . C A U 0 5 6 16. 1. 1. 0 . 2. C A U 0 5 7 16. 1. 1. 0 . 2 . D3 P 0 5 8 18. 1. 2. 0 . 2.

F E 1059 11. 1. 1. . o. 2 . P 3 G 0 6 0 15. 1. 1. 0 . 2. PI GO 6 1 13. 1. 1. 0 . 2 . N O V O 62 18. 1. 1. 0 . 2. T A N 0 6 3 19. 1. 2, 0 . 2 . T A N O 64 16. 1. 2, 0 . 2. E V A 0 6 5 22 . 1 . 1, 0 . 3. EYA066 22. 1. 1. 0 . 3. 96

APPENDIX III - CONTINUED

OTU CHARACTER 82 83 84 85 86 87 88 89 90

0ME073 40. 0. 2. 1. 2. 1. 1. 1 NSPOfO 12. 1. 1. 0. 2. 1. 1. 0 NSPOfU 13. 1. 1. 0. 2. *1. 1. 0 CUB 09 4 16. 1. 1. ' o. 1. 0. 1. 0 FON095 19. 1. 1. 1. 2. 1. 2. 1 F0N096 17. 1. 1. 0. 2. 1. 2. 1 JAV0S8 18. 1. 1. 0. 3. 1. 1. 1 CCNO99 22. 1. 1. 0. 2. 1. 1. 1 CAH100 18. ^ • 0. 2. 1. 1. 1 L 30101 23. 1. 0. 2. 1. 1. 1 L 10102 23. 1*. 1. 0. 2. 1." 1. 1 UPE103 19. 1. 99. 99. 1. 1. 1 MEH104 27. ll 2. 0. 3. 1. 1. 1 MEH105 30. 1. 2. 0. 3. 1. 1. 1 UNCI06 18. 1. 1. 0. 2. 1. 1. 0 97

APPENDIX III - CONTINUED

OTU ' CHARACTER 91 ?2 93 94 95 96 97 98 99 L 1 POO 1 0. 2. 5. 0. 0. 0. 1. 1 • 2 1 9 U P 0 0 2 0. 2. 5, 0. 0. 0. 2. X ft 2 2 5 LIP003 0. 2. 6. 0. 0. 0. 2. X ft 2 3 6 L1P004 0, 2. 5. 0. 0. 0, 2. 1 ft 2 4 1 LIP005 0. 2. 5. 0. 0. 0. 0. 1 ft 2 3 3 U P 0 0 6 0. 2. 5. 0. 0. 0. 2. 1 * 2 1 8 LIP007 0. 2. 5. 0. 0. 0, 2. 1 • 2 3 0 L ] P008 0. 2. 4. 0. 0. 0. 2, 1 * 2 2 3 LIP009 0. 2. 4. 0. 0. 0, 2. 1 ft 2 1 7 LJPOJO 0. 2. 5; 0. 0, 0. 1. 1 ft 2 3 1 LI POll 0. 2. 6. 0. 0. 0. 2 * 1 ft 2 3 7 L 1 PO] 2 0. 2. 5. 0. 0. 0. 2. 1 ft 2 2 8 LIPO 13 0. 2. 4. 0. 0. 0. 2. 1 ft 2 3 3 L1P014 0. 1. 4. 0. 0. 0. 1. 1 ft 2 2 3 LI PO 15 0. 2. 5. 0. 0. 0. 2. 1 ft 2 2 3 L1P016 0. 2. 5. 0, 0. 0, 2. J. ft 2 3 8 LI POT 4 0. 2. 5. 0. 0. 0. • 2. 1 ft 2 1 5 L J POT 5 0. 2. 3. 0. 0. 0. 1. 1 ft 2 3 0 LIPO76 0. 1. 6. 0. 0. 0. 0. X • 2 2 4 LJP077 0. 2. 5. 0. 0. 0. 2. X • 2 1 9 LIP078 0. 2. 4, 0. 0. 0. 0. X • 2 4 7 L )P07 9 0. 2. 5. 0. 0. 0, 2. X • 2 3 8 N E O O 17 0. 1. 2. 0. 0. 0. 2. X • 2 0 0 N E O O 18 0. 1. 4. 0. 0. 0, 2. X • 2 1 5 N E O O 19 0. 1. 2. 0. 0. 0. 2 • X • 1 8 9 NE0020 0. 1. 3. 0. 0. 0. 2. X ft 2 3 0 N E O O 21 0. 1. 3. 0. o.. 0. 2. X • 2 3 2 N E O O 22 0. 1. 3. 0, 0. 0. 2. X • 191 NE00B5 0. 1. 4. 0. 0. 0. 1. X • 2 1 8 NE00E6 0. 1. 3. 0. 0. 0. 2. X ft 1 8 4 NE00F7 0. 1. •4. 0. 0. 0. 2. X • 2 4 6 NEO O f 8 0. 1. 4. 0. 0. 0. 2. X • 2 4 5 N E O O 89 0. 1. 2. 0. 0. 0. 1. X • 2 2 5 F A ]023 0. 1. 3. 0. 0. 0. 2. X ft 195 FA 1024 0. 1. 2. 0. 0. 0. 2. X • 1 9 7 FA J 025 0. 1. 2. 0. 0. 0. 1. X • 2 1 3 FA 1026 0. 4. 0. 0. 0. 1. X • 2 2 3 FAJ027 0. 1.* 4. 0. 0. 0. 1. X • 2 0 2 FA 1028 0. 1. 3. 0. 0, 0. 99. X • 1 7 9 FA J 029 0. 1. 3. 0. 0, 0. 1. X • 192 FA 1030 0. 1. 3. 0. 0. 0. 2. X • 1 8 5 FA JO-1 0. 1. 2. 0. 0; 0. 2. X ft 1 9 7 FA 1090 0. 1. 2. 0. 0. 0. 1. X ft 2 0 5 FA 1091 0. 1, 3. 0. 0. 0. 1. 1 ft 1 9 4 98

APPENDIX III - CONTINUED

OTU CHARACTER 91 92 93 94 95 96 97 98 99 1 FAI092 0 . * ft ‘ 2. 0. 0. 0. X. X. X 89. PHI 067 0, A • 0. 0 . 0 . 0. 2. 2. 159. PH 1068 0 . x • 0. 0. 0. 0. 2. 2. 164* PH 1069 0. x • 0. 0 . 0. 0. 2. 2. 160. PHI070 0. x « 0. 0. 0. 0. 2. 2. 159. PH 1071 0 . X • 0. 0. 0 . 0 . 2. 2. 156. PHI072 0. x • 0 . 0. 0. 0. 2. 2. 164. G 0 R 0 3 5 0. 2. 0. 0 . 4. 0 . 2. 1 m 200. G O R 0 3 6 0. 3. 0. 0 . 4 . 0. 2. 1 • 2 0 5 . GGR0 3 7 0. 3. 0. 0 . 4. 0 . 2. 1 « 201. G O R 0 3 8 0. 3. 0 . 0. 4. 0*, 2. 1 • 180 . G 0 P 0 3 9 0 . 2. 0 . 0. 4. 0 . 2. 1 • 184. TRI0 30 0. 1. 0. 0. 0. 0. 2. 1 «230. T R 1 0 M 0. X. 0. 0 . X. 0. 2. 1 • 2 3 6 . TRI052 0. 1. 0 . 0. 0. 0. 2. 1 t 2 1 0 . T P ]053 0. 1. 0. 0. 2. 0 . 2. 1 • 216. T R I 0 9 3 0. 1. 0 . 0 . X. 0. 2. 1 • 2 2 5 . R A D O f 2 0. 1. 0. 0 . X4. 0 . 2. 1 • 3 0 9 . RA DO 83 0 . 1. 0 . 0. X4. 0. 2. JL • 3 0 9 . R A D O f 4 0, X. 0. 0 . X4. 0 . 2. 1 * 288. LEP032 0. 99. 5. 0. 0. 0. 0. X •222. LEP0'3 0. 99. 4. 0 . 0. 0 . 0. 1 * 222. J O H 0 3 4 0. 2. 0. X. 5. 0. 2. 1 • 2 0 7 . BPA040 0 . X. 0. 0. 0. 0 . 2. X • 2 1 0 . BRA097 0. X. 0 . 0. 0. 0. 2. X • 2 1 3 . TRU041 0. 2. 0. 0, 5. 0 . 2. 1 • 2 1 2 . T R U 0 4 5 0. 2. 0. 0 . 3. 0. 2. X • 2 1 5 . BUT 046 0 . X. 0 . 0. 9. 0 . 2. X # 220. OPH047 0. X. 0 . 0. 0. I. 0. d • 2 6 7 . OPH048 0 . 0. 0. 0 . 0. I. 0. c • 279 . S C H 0 4 9 0. X. 0. 0. 0. X. 0. X • 2 4 6 . FAP0£4 0 . 2. 0. 0. 2. 0. 2. X ft 202 . FAR055 0. 2. 99. 0. 2. 0. 2. X • 2 2 2 . CAU0f>6 0 . X. 0. 0 . 0. 0. 2. X • 221. CAU057 0. X. X. 0. 0. 0. I. X ft 2 1 2 . DJP 0 5 8 0 . 99. 99. 0. 0. 0. 2. X * 195. FEI059 0 . 2. 0. 0. 8. 0. 2. X • 3 2 0 . P3G060 0. X. 0. 0 . 0. 0. 2. X • 2 5 6 . PIG061 0. X. 0. 0. 0. 0. 2. X • 2 6 2 . .NCV062 0. X. 0 . 0. 0. 0. 2. X • 253. TAN063 0. X. 0 . 0. 0. 0. 2. X • 2 0 5 . TAN064 0. X. 0 . 0 . 0. 0. 2. X • 213. EVA065 0. X. 0. 0. 0. 0. 2. X • 2 0 5 . E V A 066 0 . X. 0 . 0. 0. 0. 2. 2. 2 0 3 . / 99

APPENDIX III - CONTINUED

.. OTU CHARACTER 91 92 93 94 95 9jb 97 98 99 0 M E 0 7 3 1. 99. 99. 0. 0. 0. 99. 1. 148.' N S P O f O 0. 1. 0. • o. 2. 0. 2. 1. 182. ns po e 1 0. 1. 0. 0. 2. 0. 2. 1. 178. C U B 0 S 4 0. 1. 4. 0. 0. 0. 2. 1. 233. F 0 N 0 9 5 0. 1. 0. 0. 0. 0. 2. . 1. 213. F O N O C 6 0. 1. 0. . 0. 1. 0. 2. 1. 207. J A V 0 9 8 0. 2. 4. -.o,. 0. 0. 2. 1. 202. C 0 N 0 9 9 0. 1. 0. 0. 0. 0. 2. - 1. 252. c a h i o o 0. 2. 0. 0. 3. 0. 2. 1. 260. L3CU 0 1 0. 1. 0. 0. 1. 0. 2. 1. 252. LIOX02 0. 0. 0. 0. 2. 0. 0. 1. 238. U P E 3 0 3 0. 1. 0. 0. 0. 0. 2. 1. 177. M E H 1 0 4 0. 1. 0. 0. 0. 1. 0. 1. 221. M E H 1 0 5 0. 1. 0. 0. 0. 1. 2. 1. 218. U M C 1 0 6 0. 2. 0. 0. 0. 0. 0. 1. 223. APPENDIX III - CONTINUED

OTU c h a r a c t e r 100 101 102 103 1 0 4 108 L3P001 0. 2. 1. 1. o. LIP002 0. 2. 1. 1. 1. L ]P003 0. 2. 1, 1. 1. LIP004 0. 2. 1. . 1. 0. L ]POOS 0. 2. 1. 1. o. L1P006 0. 2. 1. 1. 1. U P 0 0 7 0. 2. 1. 1. 1. LIP008 0. 2. 1. 1. 0. L3P009 0. 2. 1. -1. 1. LIPOIO 0. 2 . 1. 1. 0. LIPO 31 0. 2. 1. 1. 1. LIP012 0. 2. 1. 1. 0. L3P013 0. 2 • 1. 1. 1. L3P014 0. 2. 1. 1. 0. L3P035 0. 2. 1. 1. 0. LIPO 3 6 0. 2. 1. 1. 0. L 3 POT4 0. 2. 1. 1. 1. LIP075 0. 2. 1. 1. 0. L3P076 0. 2. 1. 1. 0. LIP077 0. 2. 1. 1. 0. L3P078 0. 2. 1. 1. 1. LIP079 0. 2. 1. 1. 1. N E O O 37 0. 99. 1. 1. 0. N E O O 18 0. 2. 2. 1. 0. N E O O 19 0. 99. 1. 1. 0. NEOO20 0. 2. 1. 1. 0. N E O O 21 0. 2. 1. 1. 0. NE0022 0. 2. 1. 1. 0. N E O O 85 0. 2. 1. 1. 0. NEOOf6 0. 2. 1. 1. 0. NEOOf7 0. 2. 1. 1. 0. N E O O 88 0. 2. 1. 1. 0. NE0089 0. 2. 1. 1. 0. FA 10 2 3 0. 2. 1. 1. 0. 99 FA 3024 0. 2. 1. 1. 0. FA'102 5 0. 1. 1. 1. 0. F A ]026 0. 2. 1. 1. 0. FA 1027 0. 2. 1. 1. 0. FA 3 028 0. 2. 1. 1. 0. FA 1029 0. 2. 1. 1. 0. FA 3 020 0. 2. 1. 1. 0. FA 10 31 0. 2. 1. 1. 0. FA 3 090 0. 2. 1. 1. 0.

FA 10 91 0. 2. 1. 1. 0. (\jroro(vrvjr\)r\>r\>!\j(\»i\)ivjf\jiV3ror\j(\>i\3Wi\)rv)fv»roPOMi'jr\jrv>rv)rN»rororv)i\)i\>fotv)(\)iv)ivi\>rNjr\>(\» APPENDIX III - CONTINUED

OTU CHARACTER 100 101 102 103 104 106 107 108

FA 1092 0. 99. 1. 1. ' o. 1 PH 1067 0 . 2 . 1. 1. 0 . 2 PHI 068 . o. 2 . 1. 1. 0 . 2 PHI069 0 . 2 . 1. . 1. 0 . 2 PHI070 0. 1. 1. 1. 0 . 2 PH 1071 o. 2 . 1. 1. 0 . 2 PH1072 0 . 2 . 1. 1. 0 . 2 G0R03 5 0 . 2 . 1. 1. 1 • G0R036 0 . 2 . 1. - 1 . A « GDR037 0 . 2 . 1. 1. A • G0R038 0 . 2 . 1. 1. A • G0R039 0 . 2 . 1. 1. A • T R ] 0 5 0 0 . 1 • 1 * i ♦ o. TRI051 0 . A • 1 • !• 0. TR 10-52 0. A • 1 « A • 0. TRI053 0. A * 1 « A * 0. TPI093 0. A • A • 1 « 0. RAD082 0. 2 . 1. 1. 1. RADOf3 0. 2. 1. 1. 1. RAD0 84 0. 2 . 1 . 1 . 1. LEP022 0. 2. 1. 0. 0. LEP033 0. 2. 1. 1. 0. J0H034 0. 1. 1. 0. 0. BRAOAO 0. 2. 1. 1. 0. BPA097 0. 2. 1. 1. 0. TRU041 0. 2. 1. 1. 0. TRU045 0. 2. 1. 1. 0. BUT046 0. 2. 1. 1. 0. 99 0PH047 0. 2. 1. 1. 0. 2 0PH048 0. 2. 1. 1. 0. 2 SCH049 0. 2. 1. 1. 0. 2 FARO 34 0. 1. 1. 1. 1. FAR055 0. 1. 1. 1. 0. . CAU056 0. 2. 1. 1. 0. CAU057 0. 2. 1. 1. 0. DIPO 58 0. 1. 1. 1. 1. FE]059 0. 2. 1. 1. 1. PIG060 0. 1. 1. 1. 0. PJGOf1 0. 1. 1. 1. 0. NOVO62 0. 2. 1. 1. 0. TAN063 0. 2. 1. 1. 0. TAN064 0. 2. 1. 1. 0. EVA065 0. 1. 1. 1. 0. 2 EVA066 0. 1. 1. 1. 0. 2 APPENDIX III - CONTINUED

OTU ' character 1 0 0 .1 0 1 1 0 2 103 104 105

0ME073 3. 99. 2. 1 . 1 . 1 NS PO 80 0 . 2 . 1 . 1 . 1 . 1 NSP0F1 0 . 2 . 1 . 1 . 1 . 1 CUB094 0 . 2 . 1 . 1 . 0 . 1 FDW0«5 0 . 1 . 1 . 1 . 1 . 1 F0N096 0 . 1 . 1 . 1 . 1 . 1 JAV058 0 . 1 . 1 . 1 . 0 . 1 C0N099 0 . 2 . 1 . 1 . 0 . 1 CAH300 0 . 1 . 1 . - 1 . 0 . 1 LI0101 0 . 1 . 1 . 1 . 0 . 1 L 3 0102 0 . 1 . 1 . 1 . 0 . 1 UPE103 0 . 1 . 1 . 1 . 0 . 1 MEH304 0 . 2 . 1 . 1 . 0 . 2 MEH105 0 . 99. 1. 1 . 0 . 2 UNC3 06 0 . 99. 1. 1 . 0 . 1 APPENDIX III - CONTINUED

OTU ' c h a r a c t e r 109 110 111 112 113 114 115 116 117

L 3P001 0. 1 0* 3. 2. 0 . ‘ LI POO2 0. 1 1 If ® " 3. 2 • 0. L3P003 0. 1 0. 3. 2. 0. LIP004 0. 1 0. 3. 2. 0. L ]P005 0. 1 0. 3. 2. 0. LIP006 o. 1 0. 3. 2. . ’ 0. L3P007 0. 1 0. 3. 2. 0. LIP008 o. 1 0. 3. 2. 0. LIP009 0. 1 0, 3* 2. 0. LIPOIO o. 1 0. 3. 2. 0. LIPOll 0. 1 0. 3. 2. 0. LIPO 12 0. 1 0. 3. 2. 0. LJPOI3 0. 1 0. 3. 2. 0. LIP014 0. 1 0. 3. 2. 0. LIPO15 . o. 1 0. 3. 2. 0. LIPO16 0. 1 0. 3. 2, 0. L3P074 0, 1 0. 3. 2. 0. LIP075 0. 1 0. 3. 2. 0. L3P076 0. 1 0. 3. 2. 0. LIP077 0. 1 0. 3. 2. * 0 • LIP078 0. 1 0. 3. 2. 0. LIP079 0. 1 0. 3. 2. 0. NE0017 0. 1 0. 3. 2. 0. NEDOI8 o. 1 0. 3. 2, 0. NEOO] 9 0. 1 0. 3. 2. 0. NEOO20 o. 1 0. 3. 2 • 0. NEOOH 0. 1 0. 3. 2. 0. NEOO 22 0. 1 0. 3. 2. . 99. NEOO I-5 0, 1 0. 3. 2. 0. NEOO86 0. 1 0. 3. 2. 0. NEOOf7 0. 1 0. 3* 2. 0. NEOO 88 0. 1 0. 3. 2. 0. NE0089 0. 1 0. 3. 2. 0. FA 1023 0. 1 0. 3. 2. 0. FA 1024 0. 1 •* 0. 3. 2. 0. FA 1025 0. 1 0. 3. 2. 0. FA 3026 0. 1 0. 3. 2* • 0 • FA 1027 0. 1 0. 3. 2. 0. FA]028 0. 1 0. 3. 2. 0. FA 10 2 9 0. 1 0. 3. 2. 0. FA30-0 0. 1 0. 3. 2, . 0. FA 1031 0. 1 0. 3. 2. 0. FA 1090 0. 1 0. 3. 2. 0. FA 10 91 o. 1 0. 3. 2. • 0 * 104

app endi x h i - continued

o t u - c h a r a c t e r 109 110 111 112 113 114 115 116 117

FA 3 092 0. 1. 0. 1. 1. 3. 2. 1. O O O O O O O O O O O O O O O O O C O O O O O O O i - t ' H ^ H O O O O O O O C O O O O O O O O PH 1067 0. 1. 0. 1. 1. 2. 2. 1. P H 3 068 . 0. 1. 0. 1. 1. 2. 2. 1. PH 1069 0. 1. 0. 1. 1. 2. 2. 1. P H 3 0 7 0 0. . 1. 0. 1. 1. 2. 2. 1. PH 1071 0. 1. 0. . 1. 1. 2. 2. 1. P H 3 072 0. 1. 0. ' 1. 1. 2. 2. 1. G O R 0 3 5 0. 1. 0. 1. 1. 3. 2. 1. G O R 0 3 6 0. 1. o. 1. 1. *3. 2. 1. G 0 R 0 3 7 0. 1. 0. 1. 1. 3. 2. 1. G 0 R 0 3 8 0. 0. 1. 1. 3. 2. 1. G 0 R 0 3 9 0. 1* 0. 1, ■1. 3. 2. 1. T R 3 0 5 0 0. 1. 0. 1. 1. 2. 2. 1. T R 10 51 0. 1. 0. 1. 1. 2. 2. 1. T R 1052 0. 1. 0. 1. 1. 2* 2. 1. T R 1053 0. 1. 0. 1. 1. 2. 2. 1. TP 1093 0. 1. 0. 1. 1. 2. 2. 1. RA DO f 2 o. 1. 0. 1. 1. 3. 2 » 2 • R A D 0 6 3 0. • 1. 0. 1. 1. 3. 2. 1. R AD O 84 0. 1. 0. 1. 1. 3. 2. 1, LEP0'2 0. 1 • 0. 1. 1. 3. 3 • 1, LEP 0 3 3 0, 1. 0. 1. 1. 3. 3 , 1. J O H 0 2 4 0. u • 0. 1. 1. 3. 2. 1. B R A 0 4 0 0. 1. 0. 1. 1. 2. 2. 1. BRA0 9 7 0. 1. 0. 1. 1. 3. 2. 1. T R U 0 4 1 0. o. 0. 1. 1. 2. 2. 1. TRU0 4 5 0. o. 0. 1. 1. 2. 2. 1. B U T 0 4 6 0. 0. 0. 1. 1. 2. 2. 1. □ P H 0 4 7 0. 0. 0. 1. 1. 2. 2. 1. 0 P H 0 4 8 1. 0. 0. 1. 1. 2. 2. 1. S C H 0 4 9 0. 0. 0. 1. 1. 2. 2. 1. FARO 54 0. 1. 0. 1. 1. 2. 2. 1. FAR055 0. 1. 0. 1. 1. 2. 2. 1. CAU0 56 0. 1. 0. 1. 1. 2. 2. 1. CAU057 0. 1, 0. 1. 1. 2. 2. 1. DIP058 0. 1. 0. 1. 1. 3. 2. 1. FE 3 059 0. 1. 0. 1. 1. 3. 2. 1. PIG060 0. 0. 1. 1. 2. 2. 1. P 3G061 0. 1 1 0. 1. 1. 2. 2. 1. N0V062 0. 1. 0. 1. 1. 2. 2. 1. T A N O 63 1. 1. 0. 1. 1. 2. 2. 1. T A M O 64 1. 1. 0. 1. 1. 2. 2. 1. EVA065 0. 0. 0. 1. 1. 2. 2. 1. EVA066 0. o. 0. 1. 1. 2, 2. 1. APPENDIX III - CONTINUED

OTU ' CHARACTER 109 110 Ill 112 113 114 115 116 117

ONE013 1. 0, 1. 1. 1. 3. 3. 1. 0. NSPORO 0. 1. 0. 0. 0. 2. 2. 1. 0. NSPOf1 0. 1. 0. 1. 0. 2. 2. 1. 0. CUB094 0. 1. 0. 1. 1. 3. 2. 1. 0. FCN0<5 0. 0. 0. 1. 1. 3. 2. 1. 1. FOM096 0. 0. 0. 1. 1. 3. 2. 1. 1.

JA VO98 ■ 0. 1. 0. ~ 1 • 1. 2. 2. 1. 0. CONO 99 0. 1. 0. 1. 1. • 2. 2. 1. 0. CAH300 0. 0. 0, 1. 1. 2. 2. 1. 0. LI0101 0. 1. 0. 1. 1. 2. 2. 1. 0. L 30102 0. 1. 0. 1. 1. 1. 2. 1. 0. UPEI03 0. 0. 0. 99. 99. 2. 1. 1. 0. M E M 04 0. 0. 0. 1. 1. 2. 2. 1. 0. MEH105 0. 0. 0. 1. 1. 2. 2. 1. 0. UAC106 0. 0. 0, 1. 3. • 2. 1. 1. APPENDIX III - CONTINUED

OTU ' CHARACTER 118 119 120 121 122 123 124 125 L I P 0 0 1 55. 1. . ‘ 1 0. 1. L J P 0 0 2 50. 1. 1 0. 1. L I P 0 0 3 63 . 1. 1 0. 1. L J P 0 0 4 50. 1. 1 0. 1. L I P 0 0 5 57. 1. 1 0. 1. L 1 P 0 0 6 58. 0. 1. L I P 0 0 7 55. l! 1 0. 1. U P 0 0 8 57. 1. 0. • 1. L 1 P 0 0 9 52. 1. 1 0. 1. L 1 P 0 3 0 53. 1. 1 0. 1. L I P 0 1 1 55. 1. 1 0. 1. L 1P O ] 2 60. 1. 1 0. 1. LI P O 3 3 58. 1. 1 0. 1. L I P 0 3 4 58. 1. 1 0. 1. LIPO 3 5 52. 1. 1 0. 1. L 3 P03 6 54. 1. 1 0. 1. L I P O 74 54. 1. 1 0. 1. L 3 P 0 7 5 53. 1. 1 0. 1. L I P 0 7 6 56. 1. 1 0. 1. L 3 P 0 7 7 54. 1. 1 0. 1* L I P 0 7 8 54. 1. 1 0 . 1 . L 3 P 0 7 9 55. 1. 1 0. 1. N E 0 0 T 7 52. 1. 1 0. 1. N E 0 0 1 8 55. 1. 1 0. 1. N E O O 3 9 51. 1. 0. 1. N E 0 0 2 0 46. 1. ! i 0. 1. N E O O 21 49. 1. i 0.. 1. N E O O 22 45. 1. i 0. 1. N E O O 85 61. 1. i 0. 1. N E O O 86 45. 1. i 0. 1. N E O O 87 47. 1. i 0. 1 • N E O O 8 8 45. 1. i 0. 1. NE O O 89 47. 1. i 0. 1. FA 3 023 52. 1. i 0. 1. FA 1024 51. 1. i 0. 1. FA 3025 45. 1. i 0. 1. FA 1026 54. 1. i 0. 1. FA 3027 50. 1. i 0. 1. FA 1028 52. 1. i 0. 1. FA 3 029 45. 1. i 0. 1. FA 10 3 0 54. 1. i 0. 1. FA 30-1 55. 1. i 0. 1. FA 10 9 0 48. 1. i 0. 1. FA 3 091 50. 1. i 0. 1. 107

APPENDIX III - CONTINUED

OTU CHARACTER 118 119 120 121 122 123 124 125 126 FA 1092 47. 1. 1 0. 1. 2. P H I 067 23. 0. 1 0. 1. 2. PHI068 26. o. 1 0. 1. 2. PHJ 069 23. 0. 1 0. 1. 2. PH 1070 24. 0. 1 0. 1. 2. PH 1071 24. 0. 0. 1. 2. PHI072 24. 0. 1 0. 1. 2. G0R035 39. 0. 1 O'. 0. 2. G0R0 36 38. 0. 1 0. 0. 2. G0R037 38. 0. 1 0. 0. 2. GOR038 41. 0. 1 0. 0. 2. G 0R 0 3 9 39. 0.' 1 0. 0. 2. T R 10 50 31. 1. 1 0. 1. 2. TR1051 24. 1. 1 0. 1. 2. T R 1052 31. 1. 1 o. ■ 1. 2. TFI053 30. 1. 1 0. 1. 2. TRI093 33. 1. ■ 1 0. 1. 2. RAD082 29. 0. 1 0. 1. 2. R A D O f 3 29. 0. 1 0. 2. RA DOE4 30. 0. 0. ll 2. LEP032 79. 1. I 1 0. 1. 2. LEP033 81. 1. 1 0. 1. 2. J 0 H 0 3 4 28. 1. 1 0. 1. 2. BRA040 23. 1. 1 0. 1. 2. BRA097 22. 1. 1 0. 1. 2. TRU041 23. 1. 0. 1. 2. TRU045 24. 1. o.. 1. 2. BUI 046 20. 1. ; i 0. 1. 2. OP HO 4 7 39. 1. 1 0. 1. 2. 0PH048 39. 1. 1 0. 1. 2. SCH049 23. 0. 1 0. 1. 2. FAR054 26. 0. 1 0. 1. 2. FARO 55 26. 0. 1 0. 1. 2. CAU056 20. 1. 1 0. 1. 2. CAU057 18. 1. 1 0. 2. DJP058 37. 0. 1 0. ll 2. F E 1059 34. 0. 1 0. 1. 2. P1G060 26. 0. 1 0. 1. 2. PIG061 27. 0. 1 0. 1. 2. NCV062 21. 1. 1 0. 2. TAN063 23. 1. 1 0. ll 2. T A N 064 23. 1. 1 0. 2. EVA065 28. 0. 1 0. ll 2. EVA066 28. 0. 1 0. 1. 2. 108

APPENDIX III - CONTINUED

OTU 1 CHARACTER 118 119 120 121 122 123 124 125 126 OH EOT 3 81. 0. 2. 1. 0. 0. 2. 1. 1 N S P O F O 33. 1. 1. 1. 0. 1. 2. 1. 99 NS PO 81 34. 1. 1. 1. 0. 1. 2. 1. 99 CUB094- 62. 1. 1. o. 1. 1. 1. r. 1 FONO 95 39. 0. 1. 1. 0. 1. 2. 2. 1 F O N G 96 41. 0 . 1 . . . 1. 0. 1. 2. 2. 1 JA VO 9 8 27. 0. 1. 1. 0. 1. 2. 1. 1 C r H 0 9 9 20. 1. 1. 1. 0* 1. 2. 1. 1 c a h i o o 27. 0. 1. 1. 0. 1. 2. 1. 1 L10 1 0 1 23. 1. 1. 1. 0. 1. 2. 1. 1 L I 0 1 0 2 22, 1. 1. 1. 0. 1. 2. 1. 1 U F E 1 0 3 30. 0. 1. 1. 0. 1. 99. 1. 1 MEH104- 23. 0. 1. 1. 0. U « 2. 1. 1 M E H 1 0 5 24. 0. 1. 1. 0. l. 2. 1. 1 U N C 1 0 6 33. 0. 1. 1. 0. l. 2. 1. 1 APPENDIX III - CONTINUED

OTU - c h a r a c t e r 127 128 129 130 131 132

L ]POOl 3. 1. 1. 0 . 0 . 0 . LIP002 1. 1. 1. 0 . 0 . 0 . L3P003 0 . 0 . 0 . LIP 0 0 4 1* 1*. l! 0 . 0 . 0 . L ]P005 1. 1. 1. 0.. 0 . 0 . LIP006 1. 1. 1. - o. 0 . 0 . L3P007 1. 1. 1, 0 . 0 . 0 . LIP008 0 . 0; 0 . L ]P009 ].* i! il 0 . 0 . 0 . LIP010 1 * . 1 • 1 • 0 . 0 . 0 . L ]P O ] 1 i. i. i. 0 . 0 . 0 . L I P O 12 i. i. i. 0 . 0 . 0 . L1P013 0 . 0 . 0 . L I P O 14 r. ii i! 0 . 0 . 0 . L 1PO ] 5 0 . 0 . 0 . LIP016 i* i! i! 0 . 0 . 0 . LJP074 i. i. i. 0 . 0 . 0 , LIP075 i. i. i. 0 . 0 . 0 . L ]POT6 0 . 0 . 0 , LIP077 r. i! ii 0 . 0 . 0 . LJP078 1. 1. 1. 0 . 0 . 0 . LIP079 I. 1. 1, 0 . 0 . 0 . NECi0].7 1. 1. 1. 0 . 0 . 0 . N E O O 18 1. 1. 1. 0 . 0 , 0 . N E O O ] 9 1. 1. 1. 0 . 0 . 0 . N E O O 20 1. 1. 1. 0 . 0. 0 . NE0021 3. 1. 1, 0 . 0.. 0 . N E O O 22 1. 1. 1. 0 . 0 . 0 . NE0065 0 . 0 , 0 , N E O O 86 x* i! il 0 , 0 , 0 . NE00E7 i. i. i. 0 . 0 . 0 . NE0088 i. i. i. 0 . 0 , 0 , NE0069 i. i, i. 0 . 0 . 0 . FA 1023 i. i. i. 0 * 0 . 0 . F A ] 024 i. i. i. 0 . 0 . 0 . FA 102 5 i. i. i. 0 . 0 . 0 . FA] 026 3. 1. 1. 0 . 0 . 0 . FA 1027 1 . 1. 1. 0 . 0 . 0 . F A ] 0 2 8 1. 1. 1. 0 . 0 . 0 . FA 1029 1 . 1. 1. 0 . 0. 0 . FAJC20 3, 1. 1. 0 . 0 . 0 . FA 1031 1 . 1. 1. 0 . 0 . 0 . F A ]09 0 3• 1. 1. 0 . 0 . 0 . FA 1091 1 . 1. 1. 0 . 0 . 0 . 110

APPENDIX III - CONTINUED

o t u ' c h a r a c t e r 127 128 129 130 131 132

FA J092 1. 1. 1. o. o. 0. PH 1067 1, 1. 1. 0. 0. 0. PH1068 1. 1. 1. o. i . 0. PH 1069 1. 1. 1. 0. 0. 0. PH3070 1. a• i . o. o. 0. PH 1071 1. a. l. o. 0. .0* PHI072 1. 1. i. ~ o. ‘ o. 0. G0R035 1. 1. 1. 0. 0. 0, G0R036 ’ 1. 1. i. o. o. 0. G0R037 1. l. a. o. o. 0. G0R038 1. a. l. o. o. 0. G0R039 1. a. l. o. 0. 0. TP]Of0 1. i . a. o. o. 0. TR10 51 1. a. l. o. 0. 0. TRI0f2 1. ’ a. i. o. o. 0. TR10 53 1. a. i . o. 0. 0. TRI053 a. l. o. o. 0. RAD082 1. a. l. o. 0. 0. RAD0f3 3. a. l. o. 0. 0. RAD084 1. a. a. o. 0. 0. LEP032 a. 1. 1. 0. 0. 0. LEP033 1. a. l. o. o. 0. J0H034 a. 2. 1. 0. 0, 0. BRA040 1. a. l. o. 0. 0. BRA 097 1. a. i . o. 0. 0. TRU041 1. a. a. o. 0. 0. TPU045 1. a. a. o. 0. 0. BUT046 1. a. l. o. 0. 0. 0PH047 a. a. l. o. 0. 0. 0PH048 1. a. l. o. 0. 0, SCH049 a. i. a. o. 0. 0. FARO54 1. l. a. o. 0. 0. FAROf5 1. a, a. o. 0. 0. CAU056 1. a. l, o. 0. 0. CAU057 1. a. l. o. 0. 0. DIP058 X. a. a. o. 0. 0. fe a 0£9 a. 1. a, 0. 0. 0. PIG060 1. i. a. o. 0. 0. P ]G06X a. l. a. o. 0. 0. NOVO62 1. i . a. o. 0. 0. T A. NO 63 1. a. l. o. 0. 0. TAN064 1. a. a. o. 0. 0. EVA065 X. 1. 1. o. 0. 0. EVA066 1. i . a. o. 1. 0. APPENDIX III - CONTINUED

OTU ’ CHARACTER 127 1 2 8 129 130 131 132 ON EO"/3 1. 1. 1 0. 0. 0 NSP080 1. 99. 1 0. 0. 0 NSPOfl 1. 99. 1 0. 0. 0 C U B 0 9 4 1. 1. 1 0. 0. 0 F t N O 95 2. 2. 1 0. 0. 0

FON096 2. 2. 1 _ o. 0. 0 JAV098 1. 1. 1 0. 0. 0 C 0N0 9 9 1. 1. 1 0. 0. 0 CAHIOO 1. 1. 2 0. 0. 0 LIOlOl 1. 1. 1 1. 1. 0 L 3 01 C'2 1. - 1. 1 1. 1. 0 UPE1 0 3 1. 1. 1 0. 1. 1 M E H 1 0 4 1. 1. 1 0. 0. 0 MEH10 5 1. 1. 1 0. 0. 0 UNC106 1. 1. 1 0. 0. 0 Figure 1. Standardization M.C.D. Phenogram. Cophenetic Correlation Coefficient =0,93. y—FAI M- neo I LIP BUT UPE CAH CON -CAU NOV TAN PIG FAR DIP JAV TRI BRA PHI | RAD I fei EVA NSP LIO GOR FON JOH MEH □ SCH OPH OME 114

Figure 2 Graph Analysis of Ixodorhynchus, Ixobioides. and Ixodorhynchoides. Based on the Standardization M.C.D. Similarity Matrix. 115

FON

BUT

TRU

UNC NEO- CUB LIP FAI

JOH

2 116

Figure 3. Graph Analysis, of Hemilaelaps, Scutanolaelaps., Strandtibbettsia, and Omentolaelaps, Based on thp Standardization M.C.D. Similarity Matrix. OPH

// SCH //

MEH 118

Figure 4 Graph Analysis of Nearest Neighbors of Hemilaelaps triangulus. Based on the Standardization M.C.D. Similarity Matrix. 119

UPE

TRI

BRA .5n

JAV

NSP REFERENCES CITED

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