Thor anes AN ANALYSIS OF INTERSPECIFIC AND INTRASPECIFIC MORPHOLOGICAL DIFFERENCES IN TWO CLOSELY RELATED SPECIES OF AMBLYSEIUS BERLESE USING METHODS OF NUMERICAL TAXONOMY (ACARINA: PHYTOSEIDAE) D.A. CHANT, R.I.C. HANSELL, and H.J. RowELL Department of Zoology, University of Toronto, Toronto, Ontaric MSS TAI Abstract Can. Ent. 109: 1605-1618 (197) Morphological variation between two closely related species in the genus Amblyseius Berlese was examined by numerical taxonomic methods. Multivariate tests indicated that A. canadensis Chant and Hansell and A. novaescotiae (Chant) represent two separate and distinct morphological groups. This supports their taxonomic retention as valid. species. Intraspecific: morphological variation was also examined and found to be correlated with climatic and host plant variables. Introduction Phytoseiid mites of the genus Amblyseius Berlese are generalized, plant and litter inhabiting mesostigmatid mites which are largely predacious on other Acarina. The systematics of the family Phytoseiidae was reviewed by Chant (1965) and the biology and ecology of the family by Chant (1959) and by Huffaker et al. (1970). Phytoseiid mites have been used in the biological control of various pest acarines. The genus Amblyseius in Canada and Alaska was reviewed by Chant and Hansell (1971) and 48 species were described at that time. The genus contains a number of taxa which have been considered as separate species characterized by seemingly rather trivial differences in only a few morphological characters. Suspicions linger that at least some of these species are not valid and that they simply represent different populations of the same taxon, showing rather narrow variations in morphology. The true test of a species in the biological sense must include breeding experimentation. However, traditional as well as numerical taxonomic species are defined solely on phenotypic differentiation. Two ‘species’ over which we have harboured such suspicions are A. novaescotiae (Chant) and A. canadensis Chant and Hansell. The two species are separated by minor variations in the lengths of the dorsocentral and lateral setae on the dorsal shield, and in the shape of the spermatheca: A. canadensis (Figs. 1-4) has shorter dorsocentral setae, longer lateral setae, and a somewhat y-shaped spermatheca; A. novaescotiae (Figs. 5-8) has longer dorsocentral setae, shorter lateral setae, and a slightly bell-shaped spermatheca. There are also slight differences between the chelicerae of the two species (Figs. 3, 7). Both species are locally common: A. novaescotiae in Nova Scotia, northern Quebec, Manitoba, British Columbia, and the Northwest Territories; A. canadensis in northern Ontario, northern Quebec, Saskatchewan, the Yukon Territory, and Alaska. Collections from’a few locations contain specimens of both species (e.g. 10 miles west of Timmins, Ont., on willow) but more often do not. 4. novaescotiae is collected most frequently on alder, whereas A. canadensis is most common on willow. This suggests that they populate distinct geographical ranges and niches. The Canadian National Collection of Phytoseiidae contains about 200 specimens of the two species. It was impossible to determine using traditional methods of systematics whether or not these specimens represent two separate morphological species. This Paper reports an attempt to analyze the specimens using the techniques of numerical taxonomy for two purposes: to determine if A. novaescotiae and A. canadensis can be considered separate and distinct morphological groups; and to examine the extent of morphological variation between individual populations. An attempt was also made to determine if there are correlations between environmental (climatic and host plants) and morphological variables. 16051606 ‘THE CANADIAN ENTOMOLOGIST December 1977 Fios. 1-8. 1-4, A. canadensis, showing the chaetotactic nomenclature for the setae which were used in this study as morphological characters: (1) dorsal shield; (2) ventral surface; (3) chelicera; (4) spermatheca. 5-8, A. novaescotiae: (5) dorsal shield; (6) ventral surface; (7) chelicera; (8) spermatheca‘Volume 109 ‘THE CANADIAN ENTOMOLOGIST 1607 Methods Thirty-six female specimens from four populations of A. novaescotiae and 77 female specimens from six populations of A. canadensis were examined. The localities and population sizes are as follows. A. novaescotiae: Rupert River, Que. (n = 10); Timmins (7 = 14) and Sault Ste. Marie (x = 8), Ont.; and the four type specimens from Coldbrook, N.S. A. canadensis: North Bay (a = 12), Timmins (1 = 19) (including four type specimens), Hearst (1 = 15), and Ignace (n = 14), Ont.; Fairbanks (n = 12) and Circle (n = 5), Alaska. Figure 9 (map) shows the locations of the sites from which the specimens were collected. Twenty-five morphological characters were recorded for numerical analysis (Figs. 1, 2). These were: the distance from the posterior edge of the ventrianal shield to the para-anal setae, to the pore, to seta JV2, to seta ZV2, to seta JVI, and to the anterior edge of the shield; the maximum width of the ventrianal shield; the maximum and minimum widths of the genital shield; the lengths and widths of the sternal and dorsal shields; and the lengths of setae j3, j6, 22, 24, 84, J2, Z1, Z4, Z5, S2, S4, and JV4. All measurements were recorded in microns. Twenty-two climatic parameters were selected to examine climatic variability between the different collection sites and to determine if correlations between climatic and morphological variability were significant. The 13 climatic parameters relating to temperature were: the mean daily, mean maximum, and mean minimum temperatures @F) per annum and for the three summer months of June, July, and August; and the number of days with frost per annum. The nine climatic parameters relating to precipitation were: the mean rainfall (in in.) and the number of days with measurable fain per annum and for the three summer months of June, July, and August; and the number of days with measurable precipitation per annum. The climatic records were obtained from the Atmospheric Environment Services, Environment Canada. If a collection site did not have a weather recording station the A. canadensis OA. novaescot Fic, 9. Map of Canada and Alaska showing the collection sites for the specimens of 4, canadensis and A. novaescotiae used in this study.December 1977 ‘THE CANADIAN ENTOMOLOGIST 1608 se ‘wou uonemndod ysea punase s az aourapyoo 2566 wptm ‘suonendod z 2 “ ‘ jooszo%ou -y pur sisuapoUvD “y Jo 01d sts n suousods ad ano suontsod renparpur aps 19 jeo|uouea [2UOISUOUNP-OM, “OT “OI zu oo ' 4 , a OD8A cen ressaptey eis sug “#4 ww yb UO '@IDW'@4g 4[nOg Q woyesTeoney ie Vector2 (7 “yuo 'surwuyy s9dkyO ysD}y'9 124190 ie pysoly‘syueqsie40 suo'e>0u8) 4 WUO'ISIOOHY. ° ne yuo 'suww im “uo ‘4og iene Tswepoues vy se‘Volume 109 ‘THE CANADIAN ENTOMOLOGIST 1609 nearest station to that site was selected. These were: Kapuskasing for Hearst; Dawson City for Fairbanks; Port Radium for Circle; and Moosonee for Rupert River. The host plant from which each specimen was collected was recorded to examine the possibility of adaptation to host plants influencing morphological variability. ‘The taxonomic separability of the populations was studied in two ways: first, by canonical analysis (Cooley and Lohnes 1971) to discriminate betweeen populations, subsets of populations, and combined populations; second, by examining the factor pattern and structure revealed by Veldman’s principal axis factor analysis of the correlations between individual specimens and populations (Veldman 1967). The uniformity of the patterns resulting from these two techniques under the varying combinations of subsets of data is evidence of the robustness of the underlying taxonomic structure of the data. To examine the taxonomic significance of the population separations obtained by these techniques, the principal axis ordinations were compared with the minimum spanning tree based on single linkage nearest neighbour analysis using the same morphological data. The hierarchical structure of the data was examined by applying the unweighted pair group method analysis (UPGMA phenogram) applied to arithmetic averages of the initial correlations (Sneath and Sokal 1973). The relations between sets of environmental variables and sets of morphological variables were examined by canonical correlations (Cooley and Lohnes 1971) Each numerical technique is discussed in the following text where it is applied. The following programmes were run on the IBM 370-165 computer at the University of Toronto Computer Centre: Veldman’s principal axis factor analysis (1967) written D. Power and modified by R. Hansell; CANAN, canonical analysis after Cooley and Lohnes (1971) written by D. Power; MAGMDA, canonical analysis after Anderson (1958) and incorporating Box’s text of homoscedasticity (Cooley and Lohnes 1971) written by M.A. Gates; NTSYS, the numerical taxonomic package programme, written by J. Kishpaugh and F.J. Rohlf, State University of New York, Stoney Brook; canonical correlations, BMDX75, from the Biomedical package from the University of California; Univariate analysis; ZOOCAT written by M.A. Gates and implemented by J. Kornatowski, at the University of Toronto. Results and Discussion Morphological Variability Canonical analysis was applied first to determine the taxonomic separability of cach population. This technique finely discriminates between given groups of operational taxonomic units (OTU’s) on the basis of variability of the data. In this case, each OTU was defined as an individual specimen and each group was defined as a population of specimens at a given locality A clear separation of the populations identified as A. canadensis from those identified as A. novaescotiae is shown in the two-dimensional canonical analysis plot (Fig. 10). The type specimens appear with the others in their respective group. Interspecific separation is apparent along vector 1 which accounts for 85.4% of the trace. This indicates that most of the variability in the morphological data can be accounted for at this interspecific level. Vector 2 accounts for only 7.4% of the trace and separated the North Bay population from the other A. canadensis populations, and the Rupert River population from the other A. novaescotiae populations. The gaps separating the North Bay and Rupert River populations from their respective species clusters are much smaller than the gap separating the populations identified as separate species and are interpreted as intraspecific variations1610 ‘THE CANADIAN ENTOMOLOGIST December 197 The same clustering patterns, shown in Fig. 10, were observed in canonical analyses when the populations represented by five or fewer specimens were grouped with their closest neighbour (i.e. Circle with Fairbanks; the A. canadensis types from Timmins with the other A. canadensis specimens from Timmins; the A. novaescotiae types from Coldbrook with the 4. novaescotiae specimens from Timmins) and when the type specimens were removed. This indicates that the analyses were stable even though sample sizes were small in some cases. Principal axis factor analysis (PAFA) was used to determine if the clustering Pattern observed in the canonical analysis would remain stable when a different clustering technique was applied. PAFA was also used to determine which of the morphological characters were influential in obtaining the cluster pattern. This information was obtained from the factor score values of the PAFA; i.e. the characters which indicated either high positive or negative factor scores were influential in determining the ultimate outcome of the clustering pattern. ‘The three-dimensional principal axis plot (Fig. 11) using the mean morphological data for each population shows the same clear interspecific separation as was seen in Fig. 10. To examine the taxonomic significance of this clear separation the nearest neighbour linkage between the two species groups was calculated using the minimum spanning tree based on correlations. The nearest neighbour correlation between species was —0.262 which substantiates the morphological distinctness of the two species. Factor 1, which accounts for 66.6% of the trace, stresses this interspecific separation. Factor 2, which accounts for 13.7% of the trace, indicates some intraspecific separations. In the A. novaescotiae group, the Rupert River population is separated from the Sault~Timmins populations, and the type specimens appear medially between the Rupert River and the Sault-Timmins populations. In the A. canadensis group, the Circle and Fairbanks populations are separated from the others at a correlation of 0.597 (the correlation value is from the minimum spanning tree analysis). Factor 3, which accounts for 10.0% of the trace, indicates further intraspecific separations. In the A. novaescotiae group the type specimens are separated from the Sault-Timmins populations at a correlation of 0.630. Within the A. canadensis group the North Bay population is separated from the type specimens at a correlation of 0.569. ‘Table I. The morphological characters (in descending order of influence) for factor 1 (66.6% trace), showing interspecific separation. The indicated values are the mean ranges, measured in microns, for each morphological character A. canadensis A. novaescotiae Character All populations Type ‘All populations Type 6 length 12.2-14.3, 14.4 29.4-31.8 31.0 32 length, 17.6-19.6 21.0 32.8-34.5, 34.4 3V4 Iength 278-314 30.3, 33.1-37.1 317 Genital shield ‘width (min.) 50,5-58.3 54.7 64.3-75.6 65.0 ZAlength 67.9-75.5 75.0 49.4-52.2 51.9 ZS length 53.0-58.0 573 42.0-43.0 46.5 s4 length 53.0-63.9 63.9 42.8-45.6 44.5 Genital shield ‘width (max.) 85.1-94.1 88.5 96.6-109.3 94.7 Zi length 45.6-48.6 50.7 39.1-40.3 4nl S2 length 47.7-54.6 53.3 36.2-41.1 38.9 ‘Sternal shield length 58.8-62.9 58.5 52.4-54.3 34.5Volume 109 ‘THP CANADIAN ENTOMOLOGIST 1611 Table Hl. The morphological characters (in descending order of influence) for factor 2 (13.7% trace), showing intraspecific separations. The indicated values are the mean ranges, measured in microns, for each morphological character. V/A is the ventrianal shield A. canadensis A. novaescotiae Circle & Sault & Characters Fairbanks Others Type Rupert. Timmins ‘Type VIA posterior toJVI 137.2-139.0 122.5-133.0 127.3 138.9 125.9-126.5 131.3 VIA posterior to pore 83.8- 86.5 76.9- 80.0 79.2 86.5 781-785 B16 V/A length 1392-1401 125.0-134.9 128.7 142.0 128.9-129:1 134.6 VIA posterior to para seta 44.35 47.1 39.8 43.3 41.0 45.0 39.3-39.5 40.0 VIA posterior toIV2 94.1- 98.4 85.8- 91.7 89.0 101.7 92.0- 92.1 96.3 Tables I-III list the morphological characters which were influential in obtaining the above clustering pattern for factors 1-3 respectively. There is no overlap at the species level in the 11 characters in Table I, which indicates interspecific separation. These characters are related to genital and sternal shield measurements, and to the lengths of certain setae, dorsocentrally, dorsolaterally, and ventrally. There is no overlap in the five characters in Table II or the three characters in Table III, which also indicates intraspecific separation. All of the characters in Table II are related to ventrianal shield measurements, whereas the characters in Table III are related to dorsal shield measurements and to the lengths of the anterior lateral setae. ‘The gaps separating populations or groups of populations, and their magnitudes, indicated by the above computer analyses, are supported by observing the influential character values as well as the number of influential characters (Tables I-IID. As the size of the trace removed by each factor decreases, the number of influential characters and the significance of the separations also decrease. To examine the hierarchical structure of the data the unweighted pair group method analysis was applied. The UPGMA phenogram (Fig. 12) using the same data as above corresponds to the clustering of the populations found in the three-dimensional principal axis analysis (Fig. 11). The interspecific separation is again obvious, as are the subgroups occurring within each species. Within A. canadensis the Circle and Fairbanks populations are clustered, those from Timmins, Hearst, Ignace and the type specimens are clustered, and the North Bay population remains separate from the latter. Within A. novaescotiae the Sault and Timmins populations are clustered, and the Rupert River and the type specimens are both separated from this cluster and from each other. ‘Table II. The morphological characters (in descending order of influence) for factor 3 (10% trace), showing, further intraspecific separations. The indicated values are the mean ranges, measured in microns, for each morphological character ‘A. canaden: A. novaescotiae Characters North Bay Others Type Type Others iB length 24.5 418 44.3 43.7 26.7 33.3- 33.8 Z2 length 13.5 21.7- 23.3 20.6 19.4 22.2- 23.6 Dorsal shield width 260.8 297.7-315.9 266.6 261.0 306,2-354.81612, THE CANADIAN ENTOMOLOGIST December 197 ‘When the type specimens were omitted from the analysis the clustering patterns observed in both the principal axis analysis and the phenogram remained stable. With all of the three numerical techniques described above the interspecific separation between specimens identified as A. novaescotiae and A. canadensis is ‘obvious. This clearly indicates that they are distinct and separate morphological groups, and as such are considered valid taxonomic species. In the above analyses the interspecific separation accounted for most of the variability in the morphological data. This may have influenced the intraspecific separations. To examine intraspecific variations more closely each species was examined separately. The canonical analyses of the A. canadensis group alone and of the A. novaescotiae group alone did not differ substantially from the intraspecific clustering found when the combined groups were analyzed using the same technique (Fig. 10). The two-dimensional canonical analysis plot of the A. canadensis populations alone stresses the separation of the North Bay population from all other populations along vector 1, which accounts for 58.1% of the trace. The other populations, including the type specimens, were clustered together and it was difficult to distinguish clinal differences between these populations. Table IV lists the characters which were influential in obtaining the vector 1 separation, in descending order of influence, as well as their mean values for the indicated populations. These chasacters are related to the length of seta j3, as well as to some genital and ventrianal shicld measurements, and clearly show the morphological differences between the North Bay population and the 28 as A-canadensis Dsoousremorie.ont. Qnorth say,Onr Bram non Qin non topesn Rows Que, @uewrsiont ryee.cetdbreck.n.s, Qtanees.ont ee tS Orarvvantuataske OeirciaAtene @ryoe.timmi ne-Ons Yennye oe Fig. 11, Three-dimensional principal axis plot of all populations, including the type specimens, using the mean morphological data for each character and each population. Nearest neighbour correlations are shown linking the most similar population means.Volume 109 ‘THE CANADIAN ENTOMOLOGIST. 1613, other A. canadensis populations. The two-dimensional canonical analysis of the A. novaescotiae populations alone stresses the separation of the Rupert River population and the Sault-Timmins populations in one direction and of the type specimens from the Sault—Timmins populations in the opposite direction, along vector 1, which accounts for 87.9% of the trace. Table V lists the characters which were influential in obtaining the vector 1 separation, in descending order of influence, as well as the mean values for the indicated populations. These characters are related to the ventrianal and sternal shield measurements and clearly show the differences between the populations. The above analyses indicate that geographical subgroups do occur within each species. The North Bay population is shown to be a subgroup distinct morphologically from the other A. canadensis populations. The Rupert River population, which located in northern Quebec, and the type specimens from Nova Scotia are shown to be morphologically distinct from the Ontario populations of A. novaescotiae. In our mn, however, the taxonomic distances between these subgroups within the species do not warrant subspecific designation (see Fig. 12). z 2 oe 6 6 ¢ ci FA 7 223.9%? 2622233222 ifsc beigie $f 5 22 32 255 EF wf ees ge 2 3 Fic. 12. Phenogram of all populations, including the type specimens, using mean morphological data based ‘on correlations. The co-phenetic correlation is 0.978.1614 ‘THE CANADIAN ENTOMOLOGIST December 1977 Table IV. The discriminating morphological characters (in descending order of influence) used to obtain vector 1 Separations within A. canadensis. The indicated values are the mean ranges, measured in microns, for each morphological character. V/A is ventrianal shield Characters North Bay Others Type j3 length 24.5) 41.8-44.3 37 Genital shield ‘width (min.) 50.5 55.0-58.3 54.7 VIA posterior to pore 76.9 79.9-86.5 79.3 VIA posterior to V2 85.8 88.0-98.4 89.0 Relation between Morphological and Climatic Variations To examine the relationship between intraspecific morphological variability found in both A. canadensis and A. novaescotiae and specific environmental variables, canonical correlation analyses were applied. The morphological characters were selected from the scaled vectors of the canonical analysis which indicated that these characters were influential in discriminating between the populations. The selected characters are shown in Tables IV and V, for A. canadensis and A. novaescotiae respectively. ‘The correlation plots showing the relationship between the discriminating morphological characters of A. canadensis and 13 temperature variables and nine precipitation variables are both very similar, with a correlation of 0.925 in both cases. Because identical patterns were observed in both plots, only the plot showing the correlation between morphological and temperature variables is shown (Fig. 13). The plot suggests that the wide separation of the North Bay population from the other populations on morphological bases may be related to the differences between the climatic conditions found at North Bay and at the other sites. Table VI shows the climatic parameters which were influential in obtaining the above correlations, in descending order of influence, for both temperature and precipitation. The table indicates that North Bay represents the warmest and wettest collection site. The correlation plots showing the relationship between the discriminating morphological characters of A. novaescotiae and 13 temperature and nine precipitation variables are very similar, with a correlation of 0.924 in both cases. Again, only the plot showing the correlation between morphological and temperature variables is shown ig. 14). The pattern suggested by the correlation plot, with the Rupert River population separated by a large gap in one direction from the Sault-Timmins population, and the type specimens separated by a smaller gap from Sault-Timmins Table V. The discriminating morphological characters (in descending order of influence) used to obtain vector I separations within A. novaescoriae. The indicated values are the mean ranges, measured in microns, for each morphological character. V/A is ventrianal shield Sault & Characters Type Timmins Rupert ‘VIA posterior toJV2 96.3 92.0-92.1 101.7 ‘Sternal shield ‘width 65.4 69.3-70.3 714 V/A posterior to pore 31.6 78.1-78.5 86.5Volume 109 ‘THE CANADIAN ENTOMOLOGIST 1615 Table VI. Climatic parameters (in descending order of influence) used to obtain a correlation between ‘morphological and temperature and precipitation variables, for A. canadensis, Characters North Bay Others (Characters NorthBay Others No. of days with, ‘Mean daily temp. frost per annum 174 208-244 CF) in June oe 48-72 Mean daily max ‘Mean daily max. temp. CF) per temp. CF) in annum 50 26- 46 June 2B 56-72 ‘Mean daily min. ‘Mean rain (in.) temp. (°F) in June st 40- 48 per annum 27 5-22 populations in an opposite direction, agrees quite well with the plots based on morphological characters alone. Table VII shows the climatic parameters which were influential in obtaining the above correlations, in descending order of influence, for both temperature and precipitation. The table suggests that the climatic parameters may influence intraspecific morphological variability in A. novaescotiae as well. Relation between Morphological and Host Plant Variations An attempt was made to correlate the host plants from which the mites were collected with their morphological variations. The minimum spanning tree for the 113 specimens showed interspecific separation. Within the A. canadensis populations, morphological variation had only a tenuous geographical basis: variation within most collection sites, in most cases, was as great as variation between sites. The North Bay population, which has been shown to be slightly different morphologicatly in the above analyses, is again the exception. Eleven specimens collected on willow clustered together while the twelfth specimen collected on poplar was linked to the type specimen. In general, specimens from a given site tended mostly to fall in the same general broad area of the minimum spanning tree. The mites from common host plants showed some clustering effects but no simple correlations were apparent. The populations of A. novaescotiae (Fig. 15) showed a clustering of the Rupert River specimens, found mostly on Prunus sp., but also on mountain ash and Labrador tea, and a clustering of the other populations, including the type specimens, found mainly on alder, but also on willow and Rubus sp. The canonical correlation analysis between the discriminating morphological characters of A. novaescotiae and the host plants produced a correlation of 0.936 (Fig. 16). The Rupert River population was again separated from the other populations by a large gap. This suggests that host plant adaptation may also play a role in accounting for intraspecific variability in this inseance. ‘The canonical correlation analysis between the discriminating morphological characters of A. canadensis and its host plants produced a correlation of only 0.416. ‘Table VII. Climatic parameters (in descending order of influence) used to obtain a correlation between ‘morphological and temperature and precipitation variables, for A. novaescotiae Sault & Sault & Characters Rupert Timmins Type | Characters Rupert Timmins Type Mean daily max. No. of days with temp. (F)forJune 65 70-7271 frost per annum = 221, 211-178 157 ‘Mean daily max No. of days with temp. CF) for precip. per August 6 73-7276 annum 182 274-163 158ture Vor‘ 1 ompe 1616 THE CANADIAN ENTOMOLOGIST December 1977 This was expected because 68 of the 77 specimens of this species were found on willow. Little intraspecific morphological variation can be accounted for by host plant adaptation in this case. - wou | a Ot ypetmales, Oot rele ee oe cf co 13 ae ~ yet ee te oe *” : FE Rupert River. Que. Senin Ste. mortes Ont eet Timmins-Ont BE Type coldbrock ts nate Sf A SER Re veal ke te Be 14 Fics. 13-14. Canonical correlation plot of (13) the A. canadensis and (14) the A. novaescotiae populations, including the type specimens, using the discrimi jing morphological characters against temperature variables. The correlation is 0.925 and 0.924 respectively.Host Plants ‘Volume 109 ‘THE CANADIAN ENTOMOLOGIST 1617 pa THA sa mip (2A 1 (21414)? (1M Tpe(aya RIL TQ) Ris) t OAs (2A— Tw c TIS) i sna sie THA Fig. 15. Minimum spanning tree of 4. novaescoriae specimens showing nearest neighbour linkages between of individuals which differ in both location and host plant. In the format X (Y) Z: the X represents the ion from which the specimens were found, i.e. R—Rupert River, Que.; S—Sault Ste. Marie, Ont.; ‘T—Timmins, Cat.; Type—type specimens: (Y) is the number of specimens which were linked together from the same location and host plant: Z is the host plant, i.e. W—willow; A—alder; MA—mountain ash; LT—Labrador tea; P—Prunus sp. R—Rubus sp. 7 & OW tote * x FE Rupert River. Que, ue a Ste. Marie, Ont Joe dept ono Fe tyes, col duroet,N.s Morphological Characters Fic. 16, Canonical correlation plot of A. novaescotiae populations, including the type specimens, using the discriminating morphological characters against host plants. The correlation is 0.936.1618 THE CANADIAN ENTOMOLOGIST December 197 The above canonical correlation analyses suggest that the interspecific morpholog- ical variability may be related to climatic parameters and in the case of A. novaescotiae, to host plant adaptation. These correlations, however, should be treated with caution until experimental evidence can confirm these results General Summary The interspecific taxonomic structure of our data is verified because all three numerical taxonomic methods, canonical analysis, principal axis factor analysis, and the phenogram analysis, stressed the separability of those specimens identified as A. canadensis from those identified as A. novaescotiae. Geographical subgroups are shown to occur within each species. The North Bay. population, which represents the warmest and wettest collection site, is shown to be distinct morphologically from the other A. canadensis populations. The Rupert River population, of northern Quebec, representing the coldest collection site, and the type specimens from Nova Scotia, representing the warmest site, are shown to be morphologically distinct from the Ontario populations of A. novaescotiae. This intraspecific morphological variability was shown to have a high correlation with climatic parameters when the canonical correlations were analyzed, and in the case of A. novaescotiae a high correlation between morphological variability and host plant was observed. Acknowledgments We are grateful to the following staff and former staff for their assistance with this paper: Mrs. Eiko Yoshida Shaul and Ms. Judith Currelly, for preparation of the illustrations; Mr. Michael Gates, for providing computer assistance. Dr. G.F, Bennett, Department of Biology, Memorial University, Newfoundland, provided the mic- rovideomat microscope for the initial measurements used in this study. ‘The research reported in this paper was supported by grants from the National Research Council of Canada. References Anderson, R.W. 1958, An introduction to multivariate statistical analysis. Wiley, New York. 365 pp. Chant, D.A. 1959. Phytoseiid mites (Acarina: Phytoseiidae). Part I. Bionomics of seven species in southern England. Part II. A taxonomic review of the family Phytoseiidae, with descriptions of 38 new species. Can. Ent. Suppl. 12. pp. 41-166. 1965. Generic concepts in the family Phytoselidae (Acarina: Mesostigmata). Can. Ent. 97: 351-374, Chant, D.A. and RIC. Hansell. 1971. The genus Amblyseius (Acarina: Phytoseiidae) Alaska. Can. J. Zool. 49(5): 703-758. Cooley, W.W. and P.R. Lohnes. 1971. Multivariate data analysis. Wiley, New York. 365 pp. Huffaker, C.B., M. van de Vrie, and J.A. McMurtry. 1970. Ecology of the tetranychid mites and their possible control by predators: an evaluation. Hilgardia 40(11): 391-458 Sneath, P.H.A. and R.R, Sokal. 1973. Numerical taxonomy. Freeman, San Francisco. $73 pp. Veldman, D.J. 1967. Fortran programming for the behavioral sciences. Holt, Rinehart and Winston, New York. pp. 190-245. Canada and (Received 29 June 1976) a