Obligate parasitic plants in the Orobanchaceae germinate after sensing plant hormones, strigolactones, exuded from host roots. In Arabidopsis thaliana, the α/β-hydrolase D14 acts as a strigolactone receptor that controls shoot branching, whereas its ancestral paralog, KAI2, mediates karrikin-specific germination responses. We observed that KAI2, but not D14, is present at higher copy numbers in parasitic species than in nonparasitic relatives. KAI2 paralogs in parasites are distributed into three phylogenetic clades. The fastest-evolving clade, KAI2d, contains the majority of KAI2 paralogs. Homology models predict that the ligand-binding pockets of KAI2d resemble D14. KAI2d transgenes confer strigolactone-specific germination responses to Arabidopsis thaliana. Thus, the KAI2 paralogs D14 and KAI2d underwent convergent evolution of strigolactone recognition, respectively enabling developmental responses to strigolactones in angiosperms and host detection in parasites.

Highlights d The Striga genome reflects a three-phase model of parasitic plant genome evolution d A family of strigolactone receptors has undergone a striking expansion in Striga d Genes in lateral root development are coordinately induced in a parasitic organ d Host genes and retrotransposons are horizontally transferred into Striga

Neotropical skinks are unique among lizards and other vertebrates in their degree of convergence, in reproductive traits,with eutherian mammals. They have also been famously difficult to classify into species, largely because of a conservativebody plan and paucity of conventional diagnostic characters. Currently there are 26 recognized species, six of which occuronly on Caribbean islands. All are placed in a single genus, Mabuya. We conducted a systematic revision of Neotropicalskinks using both conventional and unconventional morphological characters, supplemented by DNA sequence analyses.We define 61 species grouped into 16 clades, recognized here as genera. They include three available generic names(Copeoglossum, Mabuya, and Spondylurus) and 13 new genera: Alinea gen. nov., Aspronema gen. nov., Brasiliscincusgen. nov., Capitellum gen. nov., Exila gen. nov., Manciola gen. nov., Maracaiba gen. nov., Marisora gen. nov., Noto-mabuya gen. nov., Orosaura gen. nov., Panopa gen. nov., Psychosaura gen. nov., and Varzea gen. nov. These 16 generaof skinks form a monophyletic group and are placed in the Subfamily Mabuyinae of the skink Family Mabuyidae. Sixother skink families are recognized: Acontidae, Egerniidae, Eugongylidae, Lygosomidae, Scincidae, and Sphenomorphi-dae. We describe three new subfamilies of Mabuyidae: Chioniniinae subfam. nov., Dasiinae subfam. nov., and Trachyl-epidinae subfam. nov. We describe 24 new species of mabuyines: Capitellum mariagalantae sp. nov., Capitellumparvicruzae sp. nov., Copeoglossum aurae sp. nov., Copeoglossum margaritae sp. nov., Copeoglossum redondae sp.nov., Mabuya cochonae sp. nov., Mabuya desiradae sp. nov., Mabuya grandisterrae sp. nov., Mabuya guadeloupae sp.nov., Mabuya hispaniolae sp. nov., Mabuya montserratae sp. nov., Marisora aurulae sp. nov., Marisora magnacornaesp. nov., Marisora roatanae sp. nov., Spondylurus anegadae sp. nov., Spondylurus culebrae sp. nov., Spondylurus caico-sae sp. nov., Spondylurus haitiae sp. nov., Spondylurus magnacruzae sp. nov., Spondylurus martinae sp. nov., Spondy-lurus monae sp. nov., Spondylurus monitae sp. nov., Spondylurus powelli sp. nov., and Spondylurus turksae sp. nov. Wealso resurrect 10 species from synonymies: Alinea lanceolata comb. nov., Alinea luciae comb. nov., Capitellum metalli-cum comb. nov., Mabuya dominicana, Marisora alliacea comb. nov., Marisora brachypoda comb. nov., Spondylurusfulgidus comb. nov., Spondylurus nitidus comb. nov., Spondylurus semitaeniatus comb. nov., and Spondylurus spilonotuscomb. nov. Of the 61 total species of mabuyine skinks, 39 occur on Caribbean islands, 38 are endemic to those islands,and 33 of those occur in the West Indies. Most species on Caribbean islands are allopatric, single-island endemics, al-though three species are known from Hispaniola, three from St. Thomas, and two from Culebra, St. Croix, Salt Island,Martinique, the southern Lesser Antilles, Trinidad, and Tobago. Co-occurring species typically differ in body size and be-long to different genera. Three ecomorphs are described to account for associations of ecology and morphology: terrestri-al, scansorial, and cryptozoic. Parturition occurs at the transition between the dry and wet seasons, and the number ofyoung (1–7) is correlated with body size and taxonomic group. Molecular phylogenies indicate the presence of many un-named species in Middle and South America. A molecular timetree shows that mabuyines dispersed from Africa to SouthAmerica 18 (25–9) million years ago, and that diversification occurred initially in South America but soon led to coloni-zation of Caribbean islands and Middle America. The six genera present on Caribbean islands each represent separate dis-persals, over water, from the mainland during the last 10 million years. Considerable dispersal and speciation alsooccurred on and among Caribbean islands, probably enhanced by Pleistocene glacial cycles and their concomitant sea lev-el changes. Based on IUCN Redlist criteria, all of the 38 endemic Caribbean island species are threatened with extinction.Twenty-seven species (71%) are Critically Endangered, six species (16%) are Endangered, and five species (13%) are Vul-nerable. Sixteen of the Critically Endangered species are extinct, or possibly extinct, because of human activities duringthe last two centuries. Several of the surviving species are near extinction and in need of immediate protection. Analysisof collection records indicates that the decline or loss of 14 skink species can be attributed to predation by the Small IndianMongoose. That invasive predator was introduced as a biological control of rats in sugar cane fields in the late nineteenthcentury (1872–1900), immediately resulting in a mass extinction of skinks and other reptiles. The ground-dwelling and diurnal habits of skinks have made them particularly susceptible to mongoose predation.

The α/β-hydrolases KAI2 and D14 are paralogous receptors for karrikins and strigolactones, two classes of plant growth regulators with butenolide moieties. KAI2 and D14 act in parallel signaling pathways that share a requirement for the F-box protein MAX2, but produce distinct growth responses by regulating different members of the SMAX1-LIKE/D53 family. kai2 and max2 mutants share seed germination, seedling growth, leaf shape, and petiole orientation phenotypes that are not found in d14 or SL-deficient mutants. This implies that KAI2 recognizes an unknown, endogenous signal, herein termed KAI2 ligand (KL). Recent studies of ligand-specificity among KAI2 paralogs in basal land plants and root parasitic plants suggest that karrikin and strigolactone perception may be evolutionary adaptations of KL receptors. Here we demonstrate that evolutionarily conserved KAI2c genes from two parasite species rescue multiple phenotypes of the Arabidopsis kai2 mutant, unlike karrikin-, and strigolactone-specific KAI2 paralogs. We hypothesize that KAI2c proteins recognize KL, which could be an undiscovered hormone.

A set of PpKAI2 - LIKE paralogs that may encode strigolactone receptors in Physcomitrella patens were identified through evolutionary, structural, and transcriptional analyses, suggesting that strigolactone perception may have evolved independently in basal land plants in a similar manner as spermatophytes. Carotenoid-derived compounds known as strigolactones are a new class of plant hormones that modulate development and interactions with parasitic plants and arbuscular mycorrhizal fungi. The strigolactone receptor protein DWARF14 (D14) belongs to the α/β hydrolase family. D14 is closely related to KARRIKIN INSENSITIVE2 (KAI2), a receptor of smoke-derived germination stimulants called karrikins. Strigolactone and karrikin structures share a butenolide ring that is necessary for bioactivity. Charophyte algae and basal land plants produce strigolactones that influence their development. However phylogenetic studies suggest that D14 is absent from algae, moss, and liverwort genomes, raising the question of how these basal plants perceive strigolactones. Strigolactone perception during seed germination putatively evolved in parasitic plants through gene duplication and neofunctionalization of KAI2 paralogs. The moss Physcomitrella patens shows an increase in KAI2 gene copy number, similar to parasitic plants. In this study we investigated whether P. patens KAI2-LIKE (PpKAI2L) genes may contribute to strigolactone perception. Based on phylogenetic analyses and homology modelling, we predict that a clade of PpKAI2L proteins have enlarged ligand-binding cavities, similar to D14. We observed that some PpKAI2L genes have transcriptional responses to the synthetic strigolactone GR24 racemate or its enantiomers. These responses were influenced by light and dark conditions. Moreover, (+)-GR24 seems to be the active enantiomer that induces the transcriptional responses of PpKAI2L genes. We hypothesize that members of specific PpKAI2L clades are candidate strigolactone receptors in moss.