From roots to myriad leaves The legacy of Norman Platnick in spider systematics Martín Ramírez. Museo Argentino de Ciencias Naturales. AAS 2020 Virtual Summer Symposium See a very nice biography prepared by Lorenzo Prendini (AMNH) A personal perspective Of his achievements and influence in spider systematics How it developed in the work of others and in my work Izquierdo Matías Photo Photo 1. The root of true spiders Finding the roots Early ‘70s – hot time for phylogenetic systematics Platnick mastered Hennig’s tricks Chunk of spider taxa Photo Gerd Hennig Photo Willi Hennig. 1966 English edition Finding the roots Ample knowledge of • Setting relevant outgroups other spiders Finding the roots • Setting relevant outgroups • Finding synapomorphies / monophyly Finding the roots • Setting relevant outgroups • Finding synapomorphies / monophyly • Detecting the groups with primitive characters. Usually small clades of restricted distribution. • Inferring the early splits Finding the roots • Setting relevant outgroups • Finding synapomorphies / monophyly • Detecting the groups with primitive characters. Usually small clades of restricted distribution. • Inferring the early splits • Inferring main clades Very quick and accurate approach To figure out the big picture Norman’s candy Small groups with primitive characters and restricted distribution From Chile or New Zealand? Much better! Focus on the roots hypochilids 1977. Hypochilids, atypids. Spiders with primitive characters (of Mygalomorphae and Araneomorphae) Hypochilus A rare cladogram with spiders + outgroups. Charles Griswold Warren Savary Evert Schlinger Norman Platnick (25 years) Lenny Vincent 1976. American Arachnological Society Western meeting (photo Charles Griswod) Around the root of Araneomorphae Many primitive characters Hypochilidae. USA, China Austrochilidae, Gradungulidae. Australia, Ray Forster New Zealand, Chile Amazing fauna of superb phylogenetic and biogeographic importance Mike Gray Opened the field to others Spelungula – Nelson cave spider Hickmania – Tasmanian cave spider Primitive characters for interesting biological transitions Sexual selection Muscle-operated male genitalia (instead of hydraulic) Silk and webs Many cribellate (vs. ecribellate) Ecophysiology 4 book lungs (instead of 2 + tracheae) Michalik, Piacentini, Lipke, Ramírez 2015 Book lungs → tracheae Complex story – according to genomic data Bond et al Fernández et al Garrison et al 2014 2014 2016 (Wheeler et al Fernández et al Shao and Li 2017) 2018 2018 Kallal et al., see Monday Book lungs → tracheae Araneomorphae Uloborus (Uloboridae) from Schmitz 2016 Tracheae. Different conformation and ontogeny, Book lungs similar end-product Complex tracheae passing to prosoma and legs What is the function of tracheae? Book lungs in opisthosoma Complex tracheae passing to prosoma Locomotion • Interference between hydraulic and legs pump and hemolymph • Contribute to aerobic metabolism circulation during periods of high activity (Anke Schmitz 2005). Salticidae Pedicel • Low efficiency for locomotion • Help muscular action to monitor the web (Brent Opell 1987). Uloboridae O2 intake CO2 release O2 supply for muscles What is the function of tracheae? Simple tracheae limited to opisthosoma Not known function Locomotion Blocking tracheae has no impact in • locomotory metabolism • CO2 release Pedicel (Schmitz & Perry 2002; Schmitz 2005) • Most common tracheal system O intake 2 • Extremely conserved CO release 2 • Broad range of body sizes and ecologies (Ctenidae, Theridiidae, Araneidae, Gnaphosidae, Lycosidae…) Unknown O2 supply for muscles UCE phylogenomic dataset Denser sample of araneomorphs with 4 book lungs + early splits + varied tracheal systems Hypochilidae Filistatidae Synspermiata Austrochiloidea Leptonetidae Congruent with the recent Palpimanoidea phylogenomic analyses 534 loci, ∼120.000 bp 6 independent origins of tracheae 1 2 3 4 5 6 4 book lungs 6 independent origins of tracheae 1 Convergences reflected in different ontogeny and details 2 3 4 5 6 4 book lungs 4 independent losses of posterior respiratory system 1 All from simple tracheal systems limited to the opisthosoma 2 3 4 book lungs 4 5 independent origins of extensive tracheal systems passing to prosoma (many more in entelegynes) All originated from simple tracheal systems limited to the opisthosoma Tracheae did not originate for high locomotory metabolism Extensive tracheae are a subsequent elaboration Image C. Griswold What could be the function of simple tracheae? muscles median tracheae ampullate glands Parawixia - Araneidae What could be the function of simple tracheae? ampullate glands tracheae Althepus - Psilodercidae Hypothesis Prediction O2 supply for spinning organs Blocking tracheae detrimental to (muscles and / or silk glands) spinning performance Aerial webs Spider hangs from web, holding individual threads Requires thin, strong threads and firm anchorages “Junction” webs (Blackledge et al. 2009), special feet Ampullate fibers Piriform glue Substrate webs Walking on top “Brushed” webs (Blackledge et al. 2009) General feet Hexura – Antrodiaetidae Photo Marshal Hedin Aglaoctenus – Lycosidae Photo Gonzalo Useta Araneomorphae: origin of aerial webs, Amp + Pi spigots aerial webs Ampullate + piriform gland system substrate webs webless Correlation with tracheae Robust to alternative codings p = 0.05 – 0.08 No correlation with tracheae Discovering amazing fauna Outgroups to highly diverse clades Key for important morphological / behavioral transitions 1983. Photo Charles Griswold 2. Spigots, silk, phylogeny Photo Rob Raven Pioneering work Jacqueline Kovoor – histology of silk glands Hans Peters – SEM, origin of threads Kovoor & Peters 1988 Spigots in systematics Jonathan Coddington 1986 orb web in Shear’s book 1989 spigots araneoids Interpreting and linking previous work systematics + behavior + diversity + cladistics Identification criteria. Glands – spigots in SEM – ontogeny Norman picking up Coddington 1989 Platnick 1990 1990. Gnaphosoid spigots: bizarre and diverse What these possibly do?? Gnaphosidae Molycriinae Sticky tape to entangle spiders!! Jonas Wolff et al. 2017 The rest – still a mystery A dream team Jonathan Coddington, Charles Griswold and Norman Platnick Hennig Meeting 1989 – Cornell Norman brought his binders of Polaroids… Polaroids by Darrell Ubick “One of the most exciting and productive times in my life” (Charles Griswold) 1991. Haplogyne (Synspermiata) spigots Mailing floppy disks with text and matrices Adding taxa, refining characters Now with genomic data, the tree is different… SPR Considerably: few groups intact Here and there: only 10 SPR moves The characters are still excellent Loxosceles ribbon silk Pholcidae viscid silk Coddington et al. 2002 Japyassu &Macagnan 2004 Spigots and silk Most variable and biologically meaningful character system in spiders Dragline Attachment disks Viscid silk Viscid tape Dry sticky tape Pheromone bolas Cribellate nanofibrils Reserve warp … Photo Peter Michalik Spigots are now standard data in spider systematics Griswold et al. 2005 Wood 2008 Agnarsson 2004 Alvarez & Hormiga 2011 Magalhaes et al 2017 Miller et al. 2012 Kuntner 2006 The good characters My posdoc at AMNH 2000-2002 5000 SEM images The good characters Me: “Hi Norm, I analyzed my matrix and here is the tree” (three pages) Norman: “No way. You are using too many characters that are full of homoplasy” (Next day) Me: “Hi again, this is the tree when I inactivate all my new characters” Norm: “Um… good luck with that” See on Monday! Azevedo et al. Platnick 2002 DNA UCE+Sanger Morphology Ramírez 2014 Morphology Morphology Wheeler et al. 2017 DNA Sanger See on Monday! Azevedo et al. Platnick 2002 Same story again DNA UCE+Sanger Morphology Ramírez 2014 Trees changes, more or less Morphology Morphology The morphology tells biologically fascinatingWheeler et storiesal. 2017 DNA Sanger New life for morphology: fossils and time calibrations Giribet 2015 Wood et al. 2012, 2013 Magalhaes et al. 2020 Azevedo et al. submitted Learning modesty from morphology hard bones Examples. Hard/impossible to do phylogeny with morphology only Norman thought: Ammoxenidae I thougth: Prodidomidae Also looks like: Zodariidae DNA: Deep RTA lineage Myrmecicultoridae, new family. Paula Cushing, Norman Horner et al., excavating ant nests Learning modesty from morphology hard bones Examples. Hard/impossible to do phylogeny with morphology only Malenella Sparassidae Looks like Anyphaenidae Looks like Dionycha Macrobuninae (a marronoid) Deep RTA lineage Fossil calibrations – better be cautious Imagine that these were fossils… A legacy Amazing spider diversity Beautiful morphology In a phylogenetic context Many evolutionary stories to be told 3. The myriad leaves Photo Matías Izquierdo Norman Platnick – Describing species 158 new genera and 2023 new species of spiders Broad knowledge In about 50 of the 120 families currently recognized Mostly single or first author. Spider catalogs Three books (1989, 1993, 1998) World Spider Catalog, online from 2000 to 2014. Ghost spiders of USA and Canada PhD thesis 1973, published 1974 Assistant Curator in AMNH in 1973 (22 years old) Directed by Herbert Levi Museum of Comparative Zoology, Harvard University His own drawings A small effective team Mohammad Shadab Louis Sorkin Norman Platnick Illustrations Curation, loans Coauthor of 36 papers since Since 1978 1974, species delimitation Shadab illustrator Ocular grid – No camera lucida Some context 1988. Platnick, N. I., and M. U. Shadab. A revision of the American