Diverse new scale insects (Hemiptera: Coccoidea) in amber from the Cretaceous and Eocene with a phylogenetic framework for fossil Coccoidea

Coccoids are abundant and diverse in most amber deposits around the world, but largely as macropterous males. Based on a study of male coccoids in Lebanese amber (Early Cretaceous), Burmese amber (Albian-Cenomanian), Cambay amber from western India (Early Eocene), and Baltic amber (mid-Eocene), 16 new species, 11 new genera, and three new families are added to the coccoid fossil record: Apticoccidae, n. fam., based on Apticoccus Koteja and Azar, and including two new species A. fortis, n. sp., and A. longitenuis, n. sp.; the monotypic family Hodgsonicoccidae, n. fam., including Hodgsonicoccus patefactus, n. gen., n. sp.; Kozariidae, n. fam., including Kozarius achronus, n. gen., n. sp., and K. perpetuus, n. sp.; the first occurrence of a Coccidae in Burmese amber, Rosahendersonia prisca, n. gen., n. sp.; the first fossil record of a Margarodidae sensu stricto, Heteromargarodes hukamsinghi, n. sp.; a peculiar Diaspididae in Indian amber, Normarkicoccus cambayae, n. gen., n. sp.; a Pityococcidae from Baltic amber, Pityococcus moniliformalis, n. sp., two Pseudococcidae in Lebanese and Burmese ambers, Williamsicoccus megalops, n. gen., n. sp., and Gilderius eukrinops, n. gen., n. sp.; an Early Cretaceous Weitschatidae, Pseudoweitschatus audebertis, n. gen., n. sp.; four genera considered incertae sedis, Alacrena peculiaris, n. gen., n. sp., Magnilens glaesaria, n. gen., n. sp., and Pedicellicoccus marginatus, n. gen., n. sp., and Xiphos vani, n. gen., n. sp. Interpretation of fossil coccoids is supported by a parsimony phylogenetic analysis based on 174 morphological characters (both adult males and females) and 112 taxa (69 Recent and 43 extinct).