Diverse New Scale Insects (Hemiptera: Coccoidea) in Amber from the Cretaceous and Eocene with a Phylogenetic Framework for Fossil Coccoidea

Abstract

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).

Brood care in a 100-million-year-old scale insect

“… Hypothetical phylogeny based on Hodgson and Hardy (2013) and Vea and Grimaldi (2015) (extinct families omitted). Matsucoccidae, Ortheziidae, Margarodidae are commonly considered as the most primitive families ( Vea and Grimaldi, 2015 ), but their phylogenetic relationships are still unresolved (e.g., Gullan and Cook, 2007 ; Hodgson and Hardy, 2013 ). Thick lines indicate the known extent of the fossil record.…”

“…Furthermore, W. kotejai shares two potential synapomorphies with Recent and Tertiary, crown-group Ortheziidae: differentiated apical and subapical setae on the last antennal segment, and trochanter and femur fused ( Vea and Grimaldi, 2012 ). Two Cretaceous genera have been tentatively attributed to Ortheziidae: Burmorthezia Vea and Grimaldi in mid-Cretaceous Burmese amber is considered as an extinct sister group to the crown-group Ortheziidae ( Vea and Grimaldi, 2012 ), while Cretorthezia Koteja and Azar in Early Cretaceous Lebanese amber is probably a stem group of scale insects ( Koteja and Azar, 2008 ; Hodgson and Hardy, 2013 ) or an extinct group within Ortheziidae ( Vea and Grimaldi, 2015 ). Additionally, a putative female ( Cretorthezia sp.)…”

“…The other genus Cretorthezia Koteja and Azar was erected based on an adult male from Early Cretaceous Lebanese amber. It has been considered to be a stem group of scale insects ( Hodgson and Hardy, 2013 , Figure 3 ) or a group within Ortheziidae ( Vea and Grimaldi, 2015 , Figure 24). A putative female (‘ Cretorthezia sp .’, unnamed) was tentatively identified as an ortheziid by Koteja and Azar (2008) , and was further discussed by Vea and Grimaldi (2012) .…”

“…The reviewers also mentioned the genus Burmacoccus Koteja, which is based on an adult male from mid-Cretaceous Burmese amber. It belongs to a separate family Burmacoccidae (Koteja, 2004), and two recent cladistic analyses also show that it is not closely related to Ortheziidae ( Hodgson and Hardy, 2013 , Figure 3 ; Vea and Grimaldi, 2015 , Figure 24).…”

Scale insect (Hemiptera, Coccomorpha) survey of soil samples from southern Asia with description of two new species of Rhizoecidae

“…Rhizoecids are probably closely related to the pseudococcids and may have once fed aerially and moved to a hypogeal habit later. However, it is known that Pseudococcidae (a neococcoid family) have been found in Lower Cretaceous amber (Vea & Grimaldi 2015) and possibly the ancestors of rhizoecids as with present-day species have always lived underground or in leaf litter.…”