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FIG. 2 in Craniodental Morphology And Phylogeny Of Marsupials

FIG. 2. Left lateral cranial and mandibular views of Marmosa murina showing principal osteological features mentioned in the text. Abbreviations: als, alisphenoid; ap, angular process; atp, alisphenoid tympanic process; conp, condylar process; corp, coronoid process; ect, ectotympanic; exo, exoccipital; fpj, frontal process of jugal; fr, foramen rotundum; fro, frontal; hpp, hamular process of pterygoid; iof, infraorbital foramen; ip, interparietal (coossified with supraoccipital); jug, jugal; lac, lacrimal; lc, lambdoid crest; lf, lacrimal foramina; maf, masseteric fossa; mas, mastoid exposure of pars canalicularis (of petrosal); max, maxillary; mef, mental foramina; nas, nasal; pal, palatine; par, parietal; pcf, paracanine fossa; pco, pars cochlearis (of petrosal); pop, postorbital process; pre, premaxillary; rmf, retromolar fossa; sq, squamosal; ssf, subsquamosal foramen; sup, supraoccipital (coossified with interparietal); zps, zygomatic process of squamosal.

FIG. 1 in Craniodental Morphology And Phylogeny Of Marsupials

FIG. 1. Dorsal and ventral cranial views of Marmosa murina showing principal osteological features mentioned in the text. Abbreviations: als, alisphenoid (coossified with basisphenoid); atp, alisphenoid tympanic process; bo, basioccipital (coossified with exoccipitals); bs, basisphenoid (coossified with alisphenoid); cc, carotid canal; ect, ectotympanic; fm, foramen magnum; fpj, frontal process of jugal; fro, frontal; gf, glenoid fossa; gpa, glenoid process of alisphenoid; ip, interparietal; jug, jugal; lac, lacrimal; max, maxillary; mp, mastoid process (of petrosal); nas, nasal; occ, occipital condyle (of exoccipital); of, orbital fossa; pal, palatine; par, parietal; pcf, paracanine fossa; pogp, postglenoid process (of squamosal); pop, postorbital process; pre(ap), premaxillary (alveolar process); pre(fp), premaxillary (facial process); prgp, preglenoid process (of jugal); pro, promontorium (of petrosal); ps, presphenoid; pt, pterygoid; rtp, rostral tympanic process (of petrosal); sq, squamosal; tcf, transverse canal foramen; tf, temporal fossa; za, zygomatic arch.

FIG. 18 in Craniodental Morphology And Phylogeny Of Marsupials

FIG. 18. Occlusal views of upper molars of Hypsiprymnodon moschatus (A, AMNH 160120 [right M1]), Caenolestes fuliginosus (B, BMNH 1954.295 [left M2, reversed]), and Acrobates pygmaeus (C, AMNH 37185 [right M2]). General features of upper molar crown morphology discussed in the text are illustrated, as are characters 132, 133, 136, 137, 140, 141, 143, 144, 145, and 146 (see main text for description of these characters and character states), as follows: Hypsiprymnodon moschatus 132(-), 133(0), 136(2), 137(1), 140(-), 141(0), 143(1), 144(1), 145(-), 146(-); Caenolestes 132(-), 133(1), 136(2), 137(-[but note that some Caenolestes specimens retain a paracone and can be scored as 1]), 140(-), 141(0), 143(1), 144(0), 145(-), 146(-); Acrobates 132(-), 133(1), 136(2), 137(-), 140(-), 141(0), 143(1), 144(0), 145(-), 146(-). Abbreviations: mec, metacone; nci, neomorphic cingulum; pac, paracone; stB, stylar cusp B; stD, stylar cusp D. Teeth are not shown to the same scale.

FIG. 17 in Craniodental Morphology And Phylogeny Of Marsupials

FIG. 17. Occlusal views of left upper molars of Distoechurus pennatus (A, AMNH 104058 [M1]), Hypsiprymnodon moschatus (B, AMNH 160120 [M2]), Trichosurus vulpecula (C, AMNH 65547 [M2]), and Lagostrophus fasciatus (D, AMNH 155105 [M2]). General features of upper molar crown morphology discussed in the text are illustrated, as are characters 132, 133, 136, 137, 140, 141, 143, 144, 145, and 146 (see main text for description of these characters and character states), as follows: Distoechurus 132(-), 133(1), 136(2), 137(-), 140(-), 141(0), 143(1), 144(1), 145(0), 146(0); Hypsiprymnodon moschatus 132(-), 133(0), 136(2), 137(-[but note that this can be scored in lightly worn specimens of H. moschatus; see fig. 18A]), (140(-), 141(0), 143(1), 144(1), 145(-), 146(-); Trichosurus vulpecula 132(-), 133(0), 136(2), 137(-), 140(-), 141(0), 143(1), 144(1), 145(0), 146(0); Lagostrophus 132(-), 133(0), 136(2), 137(-), 140(-), 141(0), 143(1), 144(2), 145(1), 146(1). Note that in the illustrated specimen of D. pennatus the neomorphic labial cingulum is restricted to the region between the major labial cusps, but in other specimens it extends along the entire labial margin of the upper molars. Abbreviations: mec, metacone; mecl, metaconule; mel, metaloph; prc, protocone; prl, protoloph; stB, stylar cusp B; stD, stylar cusp D; urcr, urocrista. Teeth are not drawn to the same scale.

FIG. 6 in Craniodental Morphology And Phylogeny Of Marsupials

FIG. 6. Posterior braincase of Dromiciops gliroides (A, UWBM 78641), Marmosa murina (B, AMNH 266418), and Echymipera kalubu (C, AMNH 190977). Alternative states for characters 30 and 31 (see main text for descriptions of these characters and character states) are illustrated as follows: Dromiciops 30(1), 31(0); Marmosa 30(1), 31(1); Echymipera 30(0), 31(-). Note that the incompletely mineralized lambdoid sesamoids (see Character 89) have fallen away in this juvenile specimen of Echymipera (the mature morphology is illustrated in fig. 11), fully exposing the parietal-supraoccipital suture. Abbreviations: exo, exoccipital; ip, interparietal; mas, mastoid; mf, mastoid fenestra; par, parietal; so, supraocciptal; sq, squamosal; ssf, subsquamosal foramen. Specimens are not drawn to the same scale.

FIG. 5 in Craniodental Morphology And Phylogeny Of Marsupials

FIG. 5. Posterior orbital region of Potorous tridactylus (A, AMNH 65293) and Perameles gunnii (B, MVZ 127070). Alternative states for characters 15–17 (see main text for descriptions of these characters and character states) are illustrated as follows: Potorous 15(0), 16(1), 17(1); Perameles 15(1), 16(0), 17(0). Abbreviations: als, alisphenoid; atp, alisphenoid tympanic process; for, foramen rotundum; fro, frontal; jug, jugal; lac, lacrimal; max, maxillary; os, orbitosphenoid; pal, palatine; plpf, posterolateral palatal foramen; pt, pterygoid; sf, sphenorbital fissure; sq, squamosal. Specimens are not drawn to the same scale.

FIG. 7 in Craniodental Morphology And Phylogeny Of Marsupials

FIG. 7. Palatal morphology of Thylamys venustus (AMNH 261254) illustrating nomenclature for fenestrae, foramina, and other features described in the text (reproduced from Voss and Jansa, 2003: fig. 5; 2009: fig. 14). Dental loci (I1–M4) provide convenient landmarks for defining the size and position of palatal structures. Abbreviations: if, incisive foramen; m, maxillary fenestra; mp, maxillopalatine fenestra; p, palatine fenestra; plpf, posterolateral palatal foramen.

FIG. 8 in Craniodental Morphology And Phylogeny Of Marsupials

FIG. 8. Postpalatal region of Caluromys philander (A, AMNH 267002), Dromiciops gliroides (B, UWBM 78641), Dactylonax palpator (C, AMNH 191042), and Potorous tridactylus (D, AMNH 66168). Alternative states for characters 43 and 45–50 (see main text for descriptions of these characters and character states) are illustrated as follows: Caluromys 43(0), 45(0), 46(0), 47(1; but note that C. philander is polymorphic for this character), 48(0), 49(0), 50(0); Dromiciops 43(1), 45(1), 46(3), 47(0), 48(1), 49(0), 50(0); Dactylonax 43(1), 45(1), 46(2), 47(1), 48(0), 49(0), 50(1); Potorous 43(-), 45(2), 46(1; but note that the modal condition for P. tridactylus is 0), 47(1), 48(0), 49(1), 50(0). Abbreviations: als, alisphenoid (purple, coossified with basisphenoid); apf, anterior pterygoid foramen; atp, alisphenoid tympanic process; bk, basisphenoid keel; bs, basisphenoid (purple, coossified with alisphenoid); cc, carotid canals; ecr, ectopterygoid crest; fo, foramen ovale; pal, palatine (green); plfp, posterolateral palatal foramen; ps, presphenoid (blue); pt, pterygoid (yellow); ptf, pterygoid fossa; tcf, transverse canal foramen; v, vomer (orange). Specimens are not drawn to the same scale.

FRONTISPIECE. The honey opossum (Tarsipes rostratus, illustrated by Gould, 1863) has aptly been described as "a paragon of autapomorphic specialization" (Aplin and Archer, 1987). in Craniodental Morphology And Phylogeny Of Marsupials

FRONTISPIECE. The honey opossum (Tarsipes rostratus, illustrated by Gould, 1863) has aptly been described as "a paragon of autapomorphic specialization" (Aplin and Archer, 1987).

FIG. 15 in Craniodental Morphology And Phylogeny Of Marsupials

FIG. 15. Occlusal views of left upper second molars (M2) of Metachirus nudicaudatus (A, AMNH 266453), Caenolestes fuliginosus (B, UMMZ 155575), Echimipera kalubu (C, AMNH 221654), and Notoryctes typhlops (D, AMNH 198651). General features of upper molar crown morphology discussed in the text are illustrated, as are characters 132, 133, 136, 137, 140, 141, 143, and 144 (see main text for description of these characters and character states), as follows: Metachirus 132(1), 133(0), 136(0), 137(1), 140(3), 141(0), 143(0), 144(0); Caenolestes 132(-), 133(1), 136(2), 137(-), 140(-), 141(0), 143(1), 144(0); Echimipera 132(1), 133(0), 136(0), 137(1), 140(4), 141(0), 143(1), 144(0); Notoryctes 132(1), 133(0), 136(0), 137(2), 140(-), 141(0), 143(0), 144(0). Note, however, that some C. fuliginosus specimens retain a paracone and can be scored as 137(1). Abbreviations: alci, anterolabial cingulum; ccr, centrocrista; mec, metacone; mecl, metaconule; nci, neomorphic cingulum; pac, paracone; prc, protocone; stB, stylar cusp B; stD, stylar cusp D. Teeth are not drawn to the same scale.

FIG. 9 in Craniodental Morphology And Phylogeny Of Marsupials

FIG. 9. Ventral view of left ear region of Marmosa murina (A, AMNH 267368), Phascogale tapoatafa (B, AMNH 160267), and Petaurus breviceps (C, AMNH 154468; note that the pterygoid bone, which when intact extends posteriorly as far as the carotid canal in P. breviceps [char. 47], is missing in this specimen). Alternative states for characters 23, 55, 60, 68, 79, 84, and 87 (see main text for descriptions of these characters and character states) are illustrated as follows: Marmosa 23(1), 55(1), 60(0), 68(0), 79(0), 84(0), 87(0); Phascogale 23(1), 55(2), 60(0), 68(2), 79(3), 84(1), 87(0); Petaurus 23(0), 55(3), 60(2), 68(1[not visible on intact skulls]), 79(4), 84(1[not visible on intact skulls]), 87(1; not visible on intact skulls). Abbreviations: als, alisphenoid; atp, alisphenoid tympanic process; bo, basioccipital; bs, basisphenoid; cc, carotid canal; cf, condyloid foramen; ctp, caudal tympanic process (of pars canalicularis of petrosal); ect, ectotympanic; fgpn, foramen for greater petrosal nerve; fips, foramen for inferior petrosal sinus; gf, glenoid fossa; gpa, glenoid process of alisphenoid; hf, hypoglossal foramen; jf, jugular foramen; jug, jugal; mas, mastoid exposure of pars canalicularis of petrosal); mf, mastoid fenestra; oc, occipital condyle; pes, posterior epitympanic sinus (of squamosal); pfo, primary foramen ovale; pgf, postglenoid foramen; pogp, postglenoid process (of squamosal); pp, paroccipital process (of exoccipital); pro, promontorium (of pars cochlearis of petrosal); pt, pterygoid; ptps, postympanic process (of squamosal); rtp, rostral tympanic process (of pars cochlearis of petrosal); smf, stylomastoid foramen; smn, stylomastoid notch; sq, squamosal; ssf, subsquamosal foramen; tcf, transverse canal foramen. Specimens are not drawn to the same scale.

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The evolution of marsupial social organization

<p>It is generally believed that marsupials are more primitive mammals than placentals and mainly solitary living, representing the ancestral form of social organization of all mammals. However, field studies have observed pair- and group-living in marsupial species, but no comparative study about their social evolution was ever done. Here we describe the results of primary literature research on marsupial social organization which indicate that most species can live in pairs or groups and many show intra-specific variation in social organization. Using Bayesian phylogenetic mixed-effects models with a moderate phylogenetic signal of 0.18 we found that solitary living is the most likely ancestral form (35% posterior probability), but has high uncertainty, and the combined probability of partly sociable marsupial ancestors (65%) should not be overlooked. For Australian marsupials, group-living species were less likely to be found in climates representing tropical rainforest, and species with a variable social organization were associated with low and unpredictable precipitation representing deserts. Our results suggest that modern marsupials are more sociable than previously believed and that there is no strong support that their ancestral state was strictly solitary living, such that the assumption of a solitary ancestral state of all mammals may also need reconsideration.</p>

Fig. 55 in Revision of MaCroponema Mawson, 1978 (Nematoda: Strongylida) from macropodid marsupials with the description of two new species

Fig. 55 Distribution of Macroponema obendorfi n. sp. in Osphranter robustus and O. antilopinus in northern Australia. Closed circles represent localities at which the parasite has been collected from O. robustus; open circles indicate locations at which common wallaroos have been examined but the nematode has not been found; closed squares represent localities at which the parasite has been collected from O. antilopinus; open squares indicate locations at which antilopine wallaroos have been examined but the nematode has not been found

Figs. 38–44 Macroponema comani Mawson, 1978 from Macropus giganteus. 38 in Revision of MaCroponema Mawson, 1978 (Nematoda: Strongylida) from macropodid marsupials with the description of two new species

Figs. 38–44 Macroponema comani Mawson, 1978 from Macropus giganteus. 38 Posterior end of male, ventral view, showing lateral (L) and central cordate (C) thickenings of spicule sheaths. 39 Posterior end of male, left lateral view, showing thickenings of spicule sheaths. 40 Bursa, apical view. 41 Genital cone, dorsal view. 42 Female tail, right lateral view. 43 Vagina and ovejector, left, lateral view. 44 Female genital system, showing uteri and seminal receptacles. Scale-bars: 0.1 mm

Figs. 29–37 Macroponema comani Mawson, 1978 from Macropus giganteus. 29 Anterior region, left lateral view. 30 Buccal capsule, lateral view. 31 Buccal capsule, ventral view. 32 Anterior extremity, apical view. 33 in Revision of MaCroponema Mawson, 1978 (Nematoda: Strongylida) from macropodid marsupials with the description of two new species

Figs. 29–37 Macroponema comani Mawson, 1978 from Macropus giganteus. 29 Anterior region, left lateral view. 30 Buccal capsule, lateral view. 31 Buccal capsule, ventral view. 32 Anterior extremity, apical view. 33 Mouth opening, apical view, showing detail of cephalic papillae and amphids. 34 Transverse optical section through oesophagus at level of oesophageal bulb, showing anterior extension of intestine. 35 Oesophageal corpus, showing bead-like sclerotisations of lining. 36 Transverse section of oesophageal corpus, showing bead-like sclerotisations of lining. 37 Spicule tip, left lateral view. Scale-bars: 29–32, 34–37, 0.1 mm; 33, 0.01 mm

Fig. 45 in Revision of MaCroponema Mawson, 1978 (Nematoda: Strongylida) from macropodid marsupials with the description of two new species

Fig. 45 Distribution of Macroponema comani Mawson, 1978 from Macropus giganteus in eastern Australia. Closed circles represent localities at which the parasite has been collected; open circles indicate locations at which kangaroos have been examined but the nematode has not been found. The stippled area indicates the known geographical range of M. giganteus

Fig. 28 in Revision of MaCroponema Mawson, 1978 (Nematoda: Strongylida) from macropodid marsupials with the description of two new species

Fig. 28 Distribution of Macroponema beveridgei Mawson, 1978 in Osphranter robustus and O. antilopinus in northern and eastern Australia. Closed circles represent localities at which the parasite has been collected from O. robustus; open circles indicate locations at which common wallaroos have been examined but the nematode has not been found; closed squares represent localities at which the parasite has been collected from O. antilopinus; open squares indicate locations at which antilopine wallaroos have been examined but the nematode has not been found

Figs. 46–54 Macroponema obendorfi n in Revision of MaCroponema Mawson, 1978 (Nematoda: Strongylida) from macropodid marsupials with the description of two new species

Figs. 46–54 Macroponema obendorfi n. sp. from Osphranter robustus and O. antilopinus. 46 Anterior region, right lateral view. 47 Buccal capsule, lateral view. 48 Buccal capsule, ventral view. 49 Anterior extremity, apical view. 50 Oesophageal corpus, showing bead-like sclerotisations of lining. 51 Bursa, apical view. 52 Spicule tip, left lateral view. 53, Female tail, left lateral view. 54 Vagina and ovejector, left lateral view. Scale-bars: 46–48, 50, 51, 53–54, 0.1 mm; 49, 52, 0.01 mm

Figs. 23–27 Macroponema beveridgei Mawson, 1978 from Osphranter robustus and O. antilopinus. 23 Bursa, apical view. 24 Spicule tip, left lateral view. 25 Genital cone, apical view. 26 Female tail, right lateral view. 27 in Revision of MaCroponema Mawson, 1978 (Nematoda: Strongylida) from macropodid marsupials with the description of two new species

Figs. 23–27 Macroponema beveridgei Mawson, 1978 from Osphranter robustus and O. antilopinus. 23 Bursa, apical view. 24 Spicule tip, left lateral view. 25 Genital cone, apical view. 26 Female tail, right lateral view. 27 Vagina and ovejector, right lateral view. Scale-bars: 23, 26, 27, 0.1 mm; 24, 25, 0.01 mm

Fig. 14 in Revision of MaCroponema Mawson, 1978 (Nematoda: Strongylida) from macropodid marsupials with the description of two new species

Fig. 14 Distribution of Macroponema arundeli n. sp. from Macropus giganteus in eastern Australia. Closed circles represent localities at which the parasite has been collected; open circles indicate locations at which kangaroos have been examined but the nematode has not been found. The stippled area indicates the known geographical range of M. giganteus