OF PSEUDOCOCCIDAE & ERIOCOCCID AE 75 



Pseudococcidae " e.g. the kinds of pores, the character of the anal ring ", and other 

 characteristics of Margarodidae " e.g. the number of antennal segments, the 

 absence of differentiated tarsal digitules ". At the same time it has certain characters 

 refuting any relationship with either of these families " e.g. absence of dorsal 

 ostioles disclaiming relationship with Pseudococcidae, and presence of anal ring 

 with setae and pores and absence of abdominal spiracles, disclaiming affinity with 

 Margarodidae". As already mentioned, Balachowsky included the Phenacole- 

 achiidae in the lecanoids (although admitting affinity to the margaroids mainly by 

 the possession of n-segmented antennae). Theron (1962), however, after detailed 

 study of the males concluded that Phenacoleachia zealandica " ostensibly belongs 

 to the margaroid group ", showing a particularly close relationship to the aberrant 

 margaroid Steingelia gorodetskia. He regarded them as annectant genera linking 

 the primitive Margarodidae with the more specialized lecanoids. Giliomee (1967), 

 on the basis of a simple numerical analysis of the characters available for comparison, 

 was able to demonstrate that Steingelia apparently represents a link between 

 Margarodidae and Coccidae, while Phenacoleachia a similar link between Margaro- 

 didae and Pseudococcidae. This suggestion was strongly supported by the recent 

 discovery and description by Beardsley (1964) of the new second Phenacoleachia 

 species, P. australis from Campbell Islands, which shows the presence of such 

 pseudococcid features as " ostioles and cellular anal ring ". These features tempted 

 Beardsley to suggest that the Phenacoleachiidae are definitely allied to the mealy- 

 bugs and probably represent a primitive subfamily of Pseudococcidae. The males 

 (apterous) also described by Beardsley, have two very distinctive pseudococcid 

 characters (as shown in the present work), i.e. the presence of disc pores and of 

 dorsal ostioles. 



The study of males has therefore revealed that there appear to be at least two 

 links between the primitive margaroids and the more specialized lecanoids. On the 

 other hand, a close relationship between Asterolecaniidae (lecanoid) and diaspidoids 

 was recently suggested on the basis of cytological studies and of some characters of 

 the female (Brown & McKenzie, 1962). Thus there may exist similar links between 

 lecanoids and diaspidoids. There is no detailed information about the males of 

 Asterolecaniidae,* but their study may perhaps also support this suggestion. 



Theron (1958), Ghauri (1962) and Giliomee (1967) each discussed the relationships 

 of various groups of Coccoidea based on their own observations and on the informa- 

 tion then available. Each contribution naturally allowed for more detailed and 

 comprehensive comparisons and conclusions. For the present author, Giliomee's 

 conclusions are of particular interest since he was dealing with the lecanoids, and 

 included in his discussion not only Coccidae (the main object of his studies) but also 

 males of 3 species of Pseudococcidae described in detail in his earlier paper (1961). 

 Incidentally, his data on these species have been included here in the tables, 

 calculations and discussion. Giliomee pointed out that the Coccidae are more 



* The males of an asterolecaniid (Cerococcus deklei Kosztarab and Vest) were later available to the 

 author. They completely agreed with the lecanoid type of males, as will be shown in a separate 

 publication. 



