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Figure 4 in Going underground: postcranial morphology of the early Miocene marsupial mole Naraboryctes philcreaseri and the evolution of fossoriality in notoryctemorphians

Figure 4. Comparison of ulnae of Naraboryctes philcreaseri and Notoryctes typhlops: a, right ulna of Naraboryctes philcreaseri (QM F57706) in cranial view; b, right ulna of Notoryctes typhlops (SAM M637) in cranial view; c, QM F57706 in medial view; d, SAM M637 in medial view; e, QMF 57706 in lateral view; f, SAM M637 in lateral view. Abbreviations: anp, anconeal process; cop, coronoid process; fls, flexor sulcus; ol, olecranon; rn, radial notch; sp, styloid process; trn, trochlear notch.

Figure 3 in Going underground: postcranial morphology of the early Miocene marsupial mole Naraboryctes philcreaseri and the evolution of fossoriality in notoryctemorphians

Figure 3. Comparison of radii of Naraboryctes philcreaseri and Notoryctes typhlops: a, left radius of Naraboryctes philcreaseri (QM F57679) in cranial view; b, right radius (reversed) of Notoryctes typhlops (SAM M637) in cranial view; c, QM F57679 in lateral view; d, SAM M637 (reversed) in lateral view. Abbreviations: gr, groove for interosseous membrane (tendinous sheet binding shafts of radius and ulna together); sp, styloid process.

Figure 7 in Going underground: postcranial morphology of the early Miocene marsupial mole Naraboryctes philcreaseri and the evolution of fossoriality in notoryctemorphians

Figure 7. Plot of first two canonical axes from quadratic discriminant analysis of degree of fossoriality (non-burrowing versus fossorial versus subterranean) in mammals based on an expanded version of the "limbs only" dataset of Hopkins and Davis (2009). Non-burrowing species are represented by green circles, fossorial species by pink squares and subterranean species by blue asterisks. Inner ellipses represent 95% confidence intervals for the means for each class, whilst the outer ellipses represent the 50% prediction intervals.Naraboryctes philcreaseri is represented by a black triangle and was treated as unknown, but falls among subterranean species and is predicted to be subterranean with very high probability (p = 1.0). Abbreviations: F, fossorial; N, non-burrowing; S, subterranean.

Figure 9 in Going underground: postcranial morphology of the early Miocene marsupial mole Naraboryctes philcreaseri and the evolution of fossoriality in notoryctemorphians

Figure 9. Part of the dated total evidence phylogeny shown in Figure 8, restricted to the clade Agreodontia (which includes Notoryctemorphia), with divergence dates compared to global temperatures and environmental change in Australia. The estimated time of inactivation of the RBP3 gene in the Notoryctes lineage is indicated: the black bar represents the point estimate (5.4 MYA), whilst the grey bars represent 95% HPDs (4.5-6.3 MYA). The global temperature curve is modified from Zachos et al. (2001). The date for the major increase in grass pollen is taken from Martin and McMinn (1994: fig. 2) whilst the date for the onset of major aridity in Australia (~1.4-1.5 MYA) is taken from McLaren and Wallace (2010).

Figure 2 in Going underground: postcranial morphology of the early Miocene marsupial mole Naraboryctes philcreaseri and the evolution of fossoriality in notoryctemorphians

Figure 2. Comparison of humeri of Naraboryctes philcreaseri and Notoryctes typhlops: a, left humerus of Naraboryctes philcreaseri (QM F57719) in cranial view; b, right humerus (reversed) of Notoryctes typhlops (SAM M637) in cranial view; c, QM F57719 in caudal view; d, SAM M637 (reversed) in caudal view. Abbreviations: bg, bicipital groove; cap, capitulum; dpc, deltopectoral crest; gtu, greater tuberosity; hh, humeral head; lsr, lateral supracondylar ridge; ltu, lesser tuberosity; mep, medial epicondyle; stf, supratrochlear foramen; tro, trochlea.

Figure 6 in Going underground: postcranial morphology of the early Miocene marsupial mole Naraboryctes philcreaseri and the evolution of fossoriality in notoryctemorphians

Figure 6. Comparison of tibiae of Naraboryctes philcreaseri and Notoryctes typhlops: a, left femur of Naraboryctes philcreaseri (QM F57686) in medial view; b, left femur of Notoryctes typhlops (SAM M637) in medial view; c, QM F57686 in lateral view; d, SAM M637 in lateral view. Abbreviations: mma, medial malleolus; pltp, posterolateral tibial process for articulation with lateral femoral condyle and fibula; sup, sulcus for patella; tc, tibial crest.

Figure 5 in Going underground: postcranial morphology of the early Miocene marsupial mole Naraboryctes philcreaseri and the evolution of fossoriality in notoryctemorphians

Figure 5. Comparison of femora of Naraboryctes philcreaseri and Notoryctes typhlops: a, right femur of Naraboryctes philcreaseri (QM F57678) in cranial view; b, right femur of Notoryctes typhlops (SAM M637) in cranial view; c, QM F57678 in caudal view; d, SAM M637 in caudal view. Abbreviations: fh, femoral head; gtr, greater trochanter; icg, intercondylar groove; ltr, lesser trochanter; trf, trochanteric fossa; ttr, third trochanter.

Figure 8 in Going underground: postcranial morphology of the early Miocene marsupial mole Naraboryctes philcreaseri and the evolution of fossoriality in notoryctemorphians

Figure 8. Dated total evidence phylogeny based on 259 morphological characters and 9012 bp of sequence data from five nuclear genes (APOB, BRCA1, RBP3, RAG1, and VWF) analysed using MrBayes 3.2.2 assuming the Independent Gamma Rates (IGR) clock model, with topological and temporal constraints applied to selected internal nodes (see supplementary information). Notoryctes and Naraboryctes are indicated in bold. Blue bars represent 95% highest posterior density intervals (HPDs) on the divergence times. Nodes without Bayesian posterior probability (BPP) were constrained a priori.

Figure 1 in Going underground: postcranial morphology of the early Miocene marsupial mole Naraboryctes philcreaseri and the evolution of fossoriality in notoryctemorphians

Figure 1. Comparison of scapulae of Naraboryctes philcreaseri and Notoryctes typhlops in lateral view: a, left scapula of Naraboryctes philcreaseri (QM F57716); b, right scapula (reversed) of Notoryctes typhlops (SAM M637). Abbreviations: acr, acromion process; cau, "caudal" angle; cor, coracoid process; cra, "cranial" angle; inf, infraspinous fossa; psf, postscapular fossa; ssf, supraspinous fossa; ssp, scapular spine; sssp, secondary scapular spine.

Figure 3 in Systematics, evolution and phylogeny of Annelida - a morphological perspective

Figure 3. General organization of an annelid exemplified with Trypanosyllis coeliaca (Errantia, Syllidae). A. Entire animal. B. Enlargement of head region; arrowhead: pigmented eyes; arrow: pharynx tooth. C. Posterior end with growth zone (arrow). - ac = anal cirrus, dc = dorsal cirrus, dtc = dorsal tentacle cirrus, i = intestine, la = lateral antenna, ma = median antenna, pa = palp, pt = pharyngeal tube, pv = proventricle. Micrographs of living specimen.

Figure 2 in Systematics, evolution and phylogeny of Annelida - a morphological perspective

Figure 2. Phylogenetic hypotheses of annelid relationships. A. Cladistic analysis based on morphological data (modified from Rouse and Fauchald 1997). B. Phylogenetic tree based on phylogenomic data (modified after Struck et al. 2011; Weigert et al. 2014).

Figure 1 in Systematics, evolution and phylogeny of Annelida - a morphological perspective

Figure 1. Examples of annelid diversity. A-D. Members of the basal radiation; A. Owenia fusiformis, Oweniidae, length about 100 mm, Inset: part of the tube. B. Chaetopterus variopedatus, Chaetopteridae, length about 250 mm. C. Sipunculus nudus, Sipuncula, length about 350 mm. D. Eurythoe complanata, Amphinomidae, length about 140 mm. E-F. Former Archiannelida; E: Protodriloides chaetifer, Protodrilida, length about 13 mm; F. Diurodrilus subterraneus, length about 440 µm. G-H. Errantia; G. Platynereis dumerilii, Nereididae, length about 100 mm. H. Microphthalmus similis, incertae sedis, length about 18 mm. I-M. Sedentaria. I. Fabricia stellaris, Sabellidae, length about 4 mm. J. Pygospio elegans, Spionidae, length about 25 mm. K. Ophelia rathkei, Opheliidae, length about 8 mm. L. Lanice conchilega, Terebellidae, juvenile, length up to 300 mm. M. Enchytraeus sp. Clitellata, length about 15 mm. Originals B, C, D: W. Westheide, Osnabrück.

Figure 8 in Systematics, evolution and phylogeny of Annelida - a morphological perspective

Figure 8. Macrochaeta clavicornis (Acrocirridae). 2nd pair of pigmented eye, typical multicellular adult eye with converse oriented photoreceptive processes (arrows), lens absent. Pigment cup formed by a layer pigmented supportive cells (psc) penetrated by processes of rhabdomeric photoreceptor cells (prc), pupil formed by unpigmented supportive cells (usc). Cu = cuticle, ep = epidermis, prc = photoreceptor cell, psc = pigmented supportive cell, smv = sensory microvilli, usc = unpigmented supportive cells. Original: I. Dykstra, Osnabrück.

Figure 6. Head appendages and innervation. A in Systematics, evolution and phylogeny of Annelida - a morphological perspective

Figure 6. Head appendages and innervation. A. Syllis sp. (Syllidae). Anterior end with palps (pa), median (ma) and lateral antennae (la), nuchal organs (no), tentacular cirri on the right broken off. B. Parapionosyllis labronica (Syllidae). Dorsal view, nervous system labelled with antibody against acetylated α-tubulin, appendages supplied with prominent nerves, depth coding. C. Saccocirrus sp. (Saccocirridae). Ventral view, note ventral ciliated band (arrowheads), palps (pa) supplied with numerous ciliated sensory cells. D, E. Nereis sp. (Nereididae). Palp. D. Palp composed of palpophore (pph) and palpostyle (ps) the latter with numerous sensory cilia. E. Longitudinal section showing musculature and coelomic cavity inside palpophore (pph) and connection of palp nerve (pn) with the brain (b). - b = brain, dc = dorsal cirrus, dln = dorsolateral nerve, dn = dorsal nerve, ey = eye, la = lateral antenna, ma = median antenna, no = nuchal organ, pa = palp, pn = palp nerve, pph = palpophore, pr = prostomium, ps = palpostyle, rm = retractor muscle, vc = ventral cirrus. A, C, D: SEM micrographs, Originals S. Raabe & W. Mangerich, Osnabrück; B: cLSM micrograph, original M. Kuper, Osnabrück; E: Azan staining.

Figure 9. Nuchal organs. A in Systematics, evolution and phylogeny of Annelida - a morphological perspective

Figure 9. Nuchal organs. A. Schematic representation of nuchal organ in Nerillidium troglochaetoides (Nerillidae). After TEM observations, modified from Purschke (1997). B. Eusyllis (?) sp. (Syllidae). Nuchal organs (encircled) visible as ciliary patches in the posterior region of the prostomium, micrograph from living animal. C. Saccocirrus sp. (Saccocirridae). Nuchal organs form oval patches (encircled). D. Myrianida prolifera (Syllidae). Nuchal epaulettes form u-shaped ciliary band extending posteriorly on peristomium and 1st chaetiger. - ey = eye, la = lateral antenna, ma = median antenna, mc = motile cilium, mv = microvillus, oc = olfactory chamber, pa = palp, pr = prostomium, rm = retractor muscle, sd = sensory dendrite, so = soma of receptor cell, suc = supportive cell. C, D: SEM micrographs, W. Mangerich, S. Raabe, Osnabrück.

Figure 4 in Systematics, evolution and phylogeny of Annelida - a morphological perspective

Figure 4. Cuticle ultrastructure of annelids. A-B. Eurythoe complanata (Amphinomidae). A. Cross section of cuticle on the trunk. Cuticle made up of layers of parallel collagen fibres (cf) traversed by microvilli (mv), which branch apically above the epicuticle (ec, arrowhead), epicuticle (ec) with dense bodies (db). B. Tangential section showing arrangement of collagen fibres and microvilli. C. Polygordius appendiculatus (Polygordiidae). Microvilli extend far above epicuticle (ec). D. Sphaerodoropsis minuta (Sphaerodoridae). Cuticle with irregularly arranged hardly visible collagen fibres (cf), covered by dark disk-like structures (arrowhead); cuticle traversed by cilium (ci) of receptor cell. E. Ophiodromus pallidus (Hesionidae). Cuticle without collagen fibres, microvilli branch above cuticle (arrowhead). – bc = basal cuticle, cf = collagen fibre, ci = cilium, db = dense body, ec = epicuticle, mv = micovillus. TEM micrographs.

Figure 5. Parapodia and chaetae. A in Systematics, evolution and phylogeny of Annelida - a morphological perspective

Figure 5. Parapodia and chaetae. A. Eunice pennata (Eunicidae); parapodium comprised of dorsal cirrus (dc), neuropodium (nep), ventral cirrus (vc) and branchia (br); acicula invisible. SEM micrograph. B. Scoloplos armiger (Orbiniidae). Cross section showing parapodium with supportive chaetae and chaetal sac; arrowheads point to sectioned chaetae. TEM micrograph. C. Syllidia armata (Hesionidae); notopodium restricted to dorsal cirrus and acicula (ac). D. Streptosyllis websteri (Syllidae). Aciculae (ac) extending outside parapodial lobe (arraowheads). E. Sphaerodopsis minuta (Sphaerodoridae). Acicula with chaetoblast (chb); arrowheads point to junctional complexes. F. Fabricia stellaris (Sabellidae). Parapodium of thorax with capillaries and uncini. G. Lanice conchilega (Terebellidae). Uncini. - ac = acicula, br = branchia, ch = chaeta, cm = circular muscle, coe = coelom, dc = dorsal cirrus, dlm = dorsal longitudinal muscle, ep = epidermis, dbv = dorsal blood vessel, fc = follicle cell, i = intestine, mv = microvillus, nep = neuropodium, obm = oblique muscle, pm = protractor muscle, rm = retractror muscle, snv = subneural blood vessel, vbv = ventral blood vessel, vc = ventral cirrus, vlm = ventral longitudinal muscle, vnc = ventral nerve cord. C, D, F, G: micrographs from living specimens, slightly squeezed.

Figure 11. Nervous system and brain. A in Systematics, evolution and phylogeny of Annelida - a morphological perspective

Figure 11. Nervous system and brain. A. Nervous system of the trunk with longitudinal and segmental circular nerves exemplified by Parapodrilus psammophilus (Dorvilleidae). Ventral cord consists of unpaired median (mn) and main paired nerves (mvn). B-D. Anti α-tubulin immunoreactivity; dotted lines indicate segment borders. B. Polygordius appendiculatus (Polygordiidae), ventral nerve cord (green) comprising three closely apposed neurite bundles, serotonergic perikarya (red) in a repetitive pattern although distinct ganglia are absent (medullary cord). Note high number of segmental nerves. C-D. Brania clavata (Syllidae); depth coding images. C. Brain (b) and ventral nerve cord in ventral view, ventral cord consists of several closely apposed nerves forming 3 bundles behind 1st ganglion (g1), 4 segmental nerves (arrowheads, ppn) in each segment; brain gives rise to several stomatogastric nerves (sn). D. Ventral cord in the trunk region. F. General diagram of the cephalic nervous system in polychaetes, numerals refer to palp nerve roots, somata stippled. E-H. Nereis sp. (Nereididae). E Ventral nerve cord in basiepithelial position (arrowheads refer to epidermal extracellular matrix). F. Parasagittal section with mushroom bodies (mb), note subepithelial position of brain; arrowheads point to cerebral ganglia. H Enlargement of anterior part of mushroom body with stalks of globuli cells (gc). – br = brain, cc = circumoesophageal connective, dcdr = dorsal commissure of drcc, dcvr = dorsal commissure of vrcc, dlln = dorsolateral longitudinal nerve, drcc = dorsal root of cc, ecm = extracellular matrix, ep = epidermis, g1 = 1st ganglion, gc = globuli cell, in = intestine, lln = lateral longitudinal nerve, mb = mushroom body, mn = median nerve of ventral cord, mvn = main nerve of ventral cord, nla = nerve of lateral antenna, nma = nerve of median antenna, no = nuchal organ, np = neuropil, obm = oblique muscle, pn = palp nerve, ppn = parapodial nerve, sn = stomatogastric nerve, so = somata of neurites, sog = suboesophageal ganglion, vbv = ventral blood vessel, vcdr = ventral commissure of drcc, vcvr = ventral commissure of vrcc, vlm = ventral longitudinal muscle, vrcc = ventral root of cc. A, F: modified from Müller and Orrhage (2005). Micrographs; B C: Lehmacher, C, D: M. Kuper, Osnabrück.

Figure 7. Pigmented eyes. A in Systematics, evolution and phylogeny of Annelida - a morphological perspective

Figure 7. Pigmented eyes. A. Platynereis dumerilii (Nereididae). Two pairs of adult eyes (ey) situated on the prostomium. B. Microphthalmus similis (Errantia, incertae sedis). Arrowheads point to small prostomial eyes. C. Nicolea zostericola (Terebellidae). Numerous small pigmented eyes below tentacular crown (arrowheads). D. Nereis sp. (Nereididae). Section showing pigmented eye with lens (le); arrowhead points to zone with rhabdomeres, arrow: marks layer of cell bodies of photoreceptor cells below pigment cell layer (psc). E. Piscicola geometra (Clitellata). Pigmented eye with phaosomous photoreceptor cells (prc), arrowhead points to phaosomes. F. Saccocirrus papillocercus (Saccocirridae). Small pigmented eye, structurally indistinguishable from larval eye; arrow indicates inverse orientation of photoreceptive structures, eye cup communicates with exterior via small pore (arrowhead). G. Gyptis propinqua (Hesionidae). Multicellular-pigmented eye with lens, arrows indicate converse orientation of photoreceptive processes. - br = branchia, cu = cuticle, ep = epidermis, ey = eye, la = lateral antenna, le = lens, pa = palp, prc = photoreceptor cell, psc = pigmented supportive cell, smv = sensory microvilli, tc = tentacular cirri, te = tentacle. A-C: micrographs from living animals; D, E: histological sections, Azan staining; F, G: TEM micrographs.