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Parnaby, Harry E., Ingleby, Sandy, Divljan, Anja (2017): Type Specimens of Non-fossil Mammals in the Australian Museum, Sydney. Records of the Australian Museum 69 (5): 277-420, DOI: 10.3853/j.2201-4349.69.2017.1653, URL: http://dx.doi.org/10.3853/j.2201-4349.69.2017.1653

Zaglossus bartoni smeenki Flannery & Groves, 1998 Mammalia 62(3): 383, figs 6–7, appendix 1. (29 December 1998). Common name. Eastern Long-beaked Echidna. Current name. Zaglossus bartoni smeenki Flannery & Groves, 1998; following Groves (2005a), who notes that it might be a full species. Holotype. M.9682 by original designation. Male adult, study skin, skull, and skeleton, collected by Mr G. Leach, on 12 July 1972; registered September 1972. Condition. Cranium and both dentaries complete. Near complete skeleton, partly articulated. Study skin in good condition, with skin fractures on beak. Type locality. Collected from the confluence of Mau and Ugat Rivers [9°40'S 149°13'E], 12 miles SW of Marua Point, Collingwood Bay, Milne Bay Province, Papua New Guinea. Comments. Paratypes not designated. Registration number of holotype given as M. 96822 in the original description; this is a typographical error. Locality data including latitude and longitude taken from the M Register, type locality given as “near Marua Point, 12 miles SW of Collingwood Bay” by Flannery & Groves (1998)., Published as part of Parnaby, Harry E., Ingleby, Sandy & Divljan, Anja, 2017, Type Specimens of Non-fossil Mammals in the Australian Museum, Sydney, pp. 277-420 in Records of the Australian Museum 69 (5) on page 300, DOI: 10.3853/j.2201-4349.69.2017.1653, http://zenodo.org/record/5237800, {"references":["Groves, C. P. 2005 a. Order Monotremata. In Mammal Species of the World: A Taxonomic and Geographic Reference, ed. D. E. Wilson and D. M. Reeder, pp. 1 - 2. Baltimore, USA: The Johns Hopkins University Press."]}

The short-beaked echidna (Tachyglossus aculeatus) and the platypus (Ornithorhynchus anatinus) are iconic egg-laying monotremes (Mammalia: Monotremata) from Australasia. The aim of this study was to demonstrate the utility of diversity profiles in disease investigations of monotremes. Using small subunit (18S) rDNA amplicon deep-sequencing we demonstrated the presence of apicomplexan parasites and confirmed by direct and cloned amplicon gene sequencing Theileria ornithorhynchi, Theileria tachyglossi, Eimeria echidnae and Cryptosporidium fayeri. Using a combination of samples from healthy and diseased animals, we show a close evolutionary relationship between species of coccidia (Eimeria) and piroplasms (Theileria) from the echidna and platypus. The presence of E. echidnae was demonstrated in faeces and tissues affected by disseminated coccidiosis. Moreover, the presence of E. echidnae DNA in the blood of echidnas was associated with atoxoplasma-like stages in white blood cells, suggesting Hepatozoon tachyglossi blood stages are disseminated E. echidnae stages. These next-generation DNA sequencing technologies are suited to material and organisms that have not been previously characterised and for which the material is scarce. The deep sequencing approach supports traditional diagnostic methods, including microscopy, clinical pathology and histopathology, to better define the status quo. This approach is particularly suitable for wildlife disease investigation.

Recently, a novel species of the genus Borreliawas identified in Bothriocroton concolor and Ixodes holocyclus ticks from echidnas. Analyses of 16S rRNA and flaB genes identified three closely related genotypes of this bacterium (Borrelia sp. Aus A-C) that were unique and distinct from previously described borreliae. Phylogenetic analyses of flaB (763 bp), groEL (1537 bp), gyrB (1702 bp) and glpQ (874 bp) gene sequences and concatenated sequences (3585 bp) of three gene loci (16S rRNA, flaB and gyrB) were consistent with previous findings and confirm that this novel species of the genus Borrelia is more closely related to, yet distinct from, the Reptile-associated (REP) and Relapsing Fever (RF) groups. At the flaB locus, genotypes A, B and C shared the highest percentage sequence similarities (87.9, 88 and 87.9 %, respectively) with B.orrelia turcica (REP), whereas at the groEL and gyrB loci, these genotypes were most similar (88.2–89.4 %) to B.orrelia hermsii (RF). At the glpQ locus, genotypes A and B were most similar (85.7 and 85.4 % respectively) to Borrelia sp. Tortoise14H1 (REP). The presence of the glpQ gene, which is absent in the Lyme Borreliosis group spirochaetes, further emphasises that the novel species of the genus Borrelia characterized in the present study does not belong to this group. Phylogenetic analyses at multiple loci produced consistent topographies revealing the monophyletic grouping of this bacterium, therefore providing strong support for its species status. We propose the name ‘Candidatus Borrelia tachyglossi’, and hypothesize that this species of the genus Borrelia may be endemic to Australia. The pathogenic potential of this bacterium is not yet known.

We used magnetic resonance imaging to study the anatomy of cortical regions, nuclear groups, and major tracts in the brain of a monotreme, i.e., the short-beaked echidna (Tachyglossus aculeatus). Our specimens were from a collection held at the Australian Museum in Sydney and had been stored in formaldehyde solution for at least 70 years. Despite this, we were able to detect fine detail in the nuclear divisions of structures as well as in fiber tracts. In particular, we could detect the medial lemniscus as it approached the ventral posterior thalamic nucleus, subdivisions within the ventral posterior thalamic nucleus, lamination and subdivisions within the hippocampal formation, components of the olfactory pathways, and nuclei within the temporal amygdala. We were able to map the topography of subcortical white matter and relate it to cortical regions determined on the basis of physiology, as well as chemical and cytoarchitecture. As expected, dense aggregations of fibers were noted in association with the primary sensory areas of the isocortex (somatosensory, visual, and auditory) and connecting primary olfactory regions (intrabulbar anterior commissure and associated fibers). We found longitudinal fibers in the basal forebrain (medial forebrain bundle) and brainstem (corticopontine and corticospinal tracts), as well as a dense array of fibers associated with the vermal and paravermal zones of the anterior lobe of the cerebellum. We also observed previously unrecognized fiber systems, i.e., commissural connections between the paired frontal isocortical fields (dorsal Fr1), dense fibers to the retrosplenial association cortex, and prominent, paired longitudinal fiber bundles in the dorsal forebrain (longitudinal fasciculus) that intersected the dorsal anterior commissure. The connectome results are consistent with the known neuroanatomy of this monotreme and they extend our knowledge of the fiber topography within this unusual brain. Our findings demonstrate the feasibility of using this sort of imaging of archived brains to analyze the neuroanatomy of rare, endangered, and evolutionarily significant species.

The Tachyglossidae (long- and short-beaked echidnas) are a family of monotremes, confined to Australia and New Guinea, that exhibit striking trigeminal, olfactory and cortical specialisations. Several species of long-beaked echidna ( Zaglossus robusta , Zaglossus hacketti , Megalibgwilia ramsayi ) were part of the large-bodied (10 kg or more) fauna of Pleistocene Australasia, but only the diminutive (2–7 kg) Tachyglossus aculeatus is widespread today on the Australian mainland. We used high-resolution CT scanning and other osteological techniques to determine whether the remarkable neurological specialisations of modern echidnas were also present in Pleistocene forms or have undergone modification as the Australian climate changed in the transition from the Pleistocene to the Holocene. All the living and extinct echidnas studied have a similar pattern of cortical gyrification that suggests comparable functional topography to the modern short-beaked form. Osteological features related to olfactory, trigeminal, auditory and vestibular specialisation (e.g., foramina and cribriform plate area, osseous labyrinth topography) are also similar in living and extinct species. Our findings indicate that despite differences in diet, habitat and body size, the suite of neurological specialisations in the Tachyglossidae has been remarkably constant: encephalisation, sensory anatomy and specialisation (olfactory, trigeminal, auditory and vestibular), hypoglossal nerve size and cortical topography have all been stable neurological features of the group for at least 300,000 years.

Monotremes (platypus and echidna) are the descendants of the oldest ancestor of all extant mammals distinguished from other mammals by mode of reproduction. Monotremes lay eggs following a short gestation period and after an even briefer incubation period, altricial hatchlings are nourished over a long lactation period with milk secreted by nipple-less mammary patches located on the female’s abdomen. Milk is the sole source of nutrition and immune protection for the developing young until weaning. Using transcriptome and mass spectrometry analysis of milk cells and milk proteins, respectively, a novel Monotreme Lactation Protein (MLP) was identified as a major secreted protein in milk. We show that platypus and short-beaked echidna MLP genes show significant homology and are unique to monotremes. The MLP transcript was shown to be expressed in a variety of tissues; however, highest expression was observed in milk cells and was expressed constitutively from early to late lactation. Analysis of recombinant MLP showed that it is an N-linked glycosylated protein and biophysical studies predicted that MLP is an amphipathic, α-helical protein, a typical feature of antimicrobial proteins. Functional analysis revealed MLP antibacterial activity against both opportunistic pathogenic Staphylococcus aureus and commensal Enterococcus faecalis bacteria but showed no effect on Escherichia coli, Pseudomonas aeruginosa, Staphylococcus epidermidis, and Salmonella enterica. Our data suggest that MLP is an evolutionarily ancient component of milk-mediated innate immunity absent in other mammals. We propose that MLP evolved specifically in the monotreme lineage supporting the evolution of lactation in these species to provide bacterial protection, at a time when mammals lacked nipples.

The monotremes (platypuses and echidnas) represent one of only four extant venomous mammalian lineages. Until recently, monotreme venom was poorly understood. However, the availability of the platypus genome and increasingly sophisticated genomic tools has allowed us to characterize platypus toxins, and provides a means of reconstructing the evolutionary history of monotreme venom. Here we review the physiology of platypus and echidna crural (venom) systems as well as pharmacological and genomic studies of monotreme toxins. Further, we synthesize current ideas about the evolution of the venom system, which in the platypus is likely to have been retained from a venomous ancestor, whilst being lost in the echidnas. We also outline several research directions and outstanding questions that would be productive to address in future research. An improved characterization of mammalian venoms will not only yield new toxins with potential therapeutic uses, but will also aid in our understanding of the way that this unusual trait evolves.

This reflective review describes how a research model, which was originally established for the successful AI of the koala (Phascolarctos cinereus), is currently being developed and extended to the wombat (Lasiorhinus latifrons and Vombatus ursinus) and short-beaked echidna (Tachyglossus aculeatus). The research model centres around the establishment of an AI program and involves: (1) semen collection, evaluation and preservation, requiring an understanding of male reproductive physiology and gamete biology; (2) timing of insemination, based on a knowledge of oestrous cycle and gestation physiology and oestrous behaviour; and (3) the appropriate placement of semen, which relies on an accurate description of female reproductive anatomy. Published and unpublished studies of assisted breeding technology (ABT) development in all three species of Australian mammals reported in this review (koala, wombat and short-beaked echidna) clearly demonstrate the importance of recognising species-specific variation in reproductive biology. Even in closely related species, such as the koala and wombat, subtle differences in reproductive physiology can hinder the transfer of ABT across species. Significant progress in marsupial and monotreme ABT will also require adequate access to captive wildlife colonies (zoos, university collections and private partners) in sufficient numbers in order to conduct quality science.

The importance of Glucagon like peptide 1 (GLP-1) for metabolic control and insulin release sparked the evolution of genes mimicking GLP-1 action in venomous species (e.g. Exendin-4 in Heloderma suspectum (gila monster)). We discovered that platypus and echidna express a single GLP-1 peptide in both intestine and venom. Specific changes in GLP-1 of monotreme mammals result in resistance to DPP-4 cleavage which is also observed in the GLP-1 like Exendin-4 expressed in Heloderma venom. Remarkably we discovered that monotremes evolved an alternative mechanism to degrade GLP-1. We also show that monotreme GLP-1 stimulates insulin release in cultured rodent islets, but surprisingly shows low receptor affinity and bias toward Erk signaling. We propose that these changes in monotreme GLP-1 are the result of conflicting function of this peptide in metabolic control and venom. This evolutionary path is fundamentally different from the generally accepted idea that conflicting functions in a single gene favour duplication and diversification, as is the case for Exendin-4 in gila monster. This provides novel insight into the remarkably different metabolic control mechanism and venom function in monotremes and an unique example of how different selective pressures act upon a single gene in the absence of gene duplication.

The short-beaked echidna (Tachyglossus aculeatus) is a monotreme and therefore provides a unique combination of phylogenetic history, morphological differentiation and ecological specialisation for a mammal. The echidna has a unique appendicular skeleton, a highly specialised myrmecophagous lifestyle and a mode of locomotion that is neither typically mammalian nor reptilian, but has aspects of both lineages. We therefore were interested in the interactions of locomotor biomechanics, ecology and movements for wild, free-living short-beaked echidnas. To assess locomotion in its complex natural environment, we attached both GPS and accelerometer loggers to the back of echidnas in both spring and summer. We found that the locomotor biomechanics of echidnas is unique, with lower stride length and stride frequency than reported for similar-sized mammals. Speed modulation is primarily accomplished through changes in stride frequency, with a mean of 1.39 Hz and a maximum of 2.31 Hz. Daily activity period was linked to ambient air temperature, which restricted daytime activity during the hotter summer months. Echidnas had longer activity periods and longer digging bouts in spring compared with summer. In summer, echidnas had higher walking speeds than in spring, perhaps because of the shorter time suitable for activity. Echidnas spent, on average, 12% of their time digging, which indicates their potential to excavate up to 204 m3 of soil a year. This information highlights the important contribution towards ecosystem health, via bioturbation, of this widespread Australian monotreme.

The early divergence of monotremes and therian mammals has resulted in considerable interest in the comparative physiology of the short-beaked echidna (Tachyglossus aculeatus), the most common and widespread living monotreme. However, there are many and varied interpretations of its physiology, reflecting the many and varied studies, limitations and uncertainties of aspects of some previous studies, and potential differences between the various subspecies. Consequently, we thoroughly examine here the standardized physiology of the most widely distributed subspecies of short-beaked echidna (T. aculeatus acanthion) over a wide range of ambient temperatures to definitively assess its physiology in a comparative context. We conclude that the low and variable body temperature of the short-beaked echidna is physiologically "primitive," but it also reflects adaptation to its myrmecophagous niche. Other aspects of its physiology are more typically mammalian. A low metabolic rate reflects its low body temperature, and ventilatory variables are matched to accommodate a modest gas exchange requirement. Thermal conductance is typical for a mammal of equivalent mass. In contrast to previous studies, we demonstrate that short-beaked echidnas can enhance evaporative water loss above thermoneutrality, like other mammals, with a similar capacity for evaporative heat loss. Cooling of their nasal blood sinus with nasal mucous may contribute to this enhanced evaporative cooling. Their capacity to evaporatively cool explains how their distribution can include habitats where ambient temperature, even in shelters, exceeds their supposed critical thermal limit.

Short-beaked echidnas (Tachyglossus aculeatus) are myrmecophages, or ant and termite insectivore specialists, and replicating their exact diet in captivity is problematic. Diets for captive animals often incorporate raw meat, eggs and cat food mixed together with water, and vitamin and mineral supplements. These diets have promoted a number of health problems in captive echidnas, such as gastritis, cystitis, gut impaction, obesity, and diarrhea. A manufactured diet was designed and three echidnas from two zoos were transitioned onto this diet to assess the acceptability and digestibility of this diet for echidnas. The new "test" diet was readily accepted by the echidnas with a 1 week transition period. Daily digestible energy intake was 280 kJ kg

Fires have occurred throughout history, including those associated with the meteoroid impact at the Cretaceous–Palaeogene (K–Pg) boundary that eliminated many vertebrate species. To evaluate the recent hypothesis that the survival of the K–Pg fires by ancestral mammals was dependent on their ability to use energy-conserving torpor, we studied body temperature fluctuations and activity of an egg-laying mammal, the echidna ( Tachyglossus aculeatus ), often considered to be a ‘living fossil’, before, during and after a prescribed burn. All but one study animal survived the fire in the prescribed burn area and echidnas remained inactive during the day(s) following the fire and substantially reduced body temperature during bouts of torpor. For weeks after the fire, all individuals remained in their original territories and compensated for changes in their habitat with a decrease in mean body temperature and activity. Our data suggest that heterothermy enables mammals to outlast the conditions during and after a fire by reducing energy expenditure, permitting periods of extended inactivity. Therefore, torpor facilitates survival in a fire-scorched landscape and consequently may have been of functional significance for mammalian survival at the K–Pg boundary.

Because monotremes are the earliest offshoot of the mammalian lineage, the platypus and short-beaked echidna were studied as model animals to assess the origin and biological significance of adaptations considered unique to therian mammals: epididymal sperm maturation and subsequent capacitation. We show that spermatozoa from both species assemble into bundles of approximately 100 cells during passage through the epididymis and that an epididymal protein—secreted protein, acidic, cysteine-rich (osteonectin; SPARC)—is involved in bundle formation. The bundles persisted during incubation in vitro for at least 1 h under conditions that capacitate therian spermatozoa, and then underwent a time-dependent dissociation to release spermatozoa capable of fertilization. Only after this dissociation could the spermatozoa bind to the perivitelline membrane of a hen's egg, display an altered form of motility reminiscent of hyperactivation, and be induced to undergo an acrosome reaction. It is concluded that the development of sperm bundles in the monotreme epididymis mandates that they require a time-dependent process to be capable of fertilizing an ovum. However, because this functional end point was achieved without overt changes in protein tyrosine phosphorylation (a hallmark of capacitation in therians), it is concluded that the process in monotremes is distinctly different from capacitation in therian mammals.

The short-beaked echidna is believed to use olfactory cues from a cloacal scent gland to attract and locate mates during the breeding season. We investigated the chemical composition of echidna secretions, including cloacal swabs and solid, "waxy" exudates from the cloaca and spurs. Scent samples from 37 individuals were collected over a 1-year period and analyzed using a range of different analytical techniques. A total of 186 compounds were identified, including volatile carboxylic acids, aldehydes, ketones, fatty acids, methyl esters, ethyl esters, terpenes, nitrogen- and sulphur-containing compounds, alcohols, and aromatics. Long chain and very long chain monounsaturated fatty acids, sterols, and sterol esters were identified as the major constituents of solid exudates, some of which have not previously been described from any animal skin gland. There was a high degree of composition overlap between male and female cloaca swabs; however, there is significant variation, which could mediate echidna mating behavior. Many of the volatile and nonvolatile chemicals detected are used for communication in other species, suggesting that chemical signals have important and diverse functions in echidna social interactions.

Monotremes are the most basal egg-laying mammals comprised of two extant genera, which are largely nocturnal. Visual pigments, the first step in the sensory transduction cascade in photoreceptors of the eye, have been examined in a variety of vertebrates, but little work has been done to study the rhodopsin of monotremes. We isolated the rhodopsin gene of the nocturnal short-beaked echidna (Tachyglossus aculeatus) and expressed and functionally characterized the proteinin vitro. Three mutants were also expressed and characterized: N83D, an important site for spectral tuning and metarhodopsin kinetics, and two sites with amino acids unique to the echidna (T158A and F169A). Theλmaxof echidna rhodopsin (497.9 ± 1.1 nm) did not vary significantly in either T158A (498.0 ± 1.3 nm) or F169A (499.4 ± 0.1 nm) but was redshifted in N83D (503.8 ± 1.5 nm). Unlike other mammalian rhodopsins, echidna rhodopsin did react when exposed to hydroxylamine, although not as fast as cone opsins. The retinal release rate of light-activated echidna rhodopsin, as measured by fluorescence spectroscopy, had a half-life of 9.5 ± 2.6 min−1, which is significantly shorter than that of bovine rhodopsin. The half-life of the N83D mutant was 5.1 ± 0.1 min−1, even shorter than wild type. Our results show that with respect to hydroxylamine sensitivity and retinal release, the wild-type echidna rhodopsin displays major differences to all previously characterized mammalian rhodopsins and appears more similar to other nonmammalian vertebrate rhodopsins such as chicken and anole. However, our N83D mutagenesis results suggest that this site may mediate adaptation in the echidna to dim light environments, possiblyviaincreased stability of light-activated intermediates. This study is the first characterization of a rhodopsin from a most basal mammal and indicates that there might be more functional variation in mammalian rhodopsins than previously assumed.

Both lineages of the modern monotremes have distinctive features in the cerebral cortex, but the developmental mechanisms that produce such different adult cortical architecture remain unknown. Similarly, nothing is known about the differences and/or similarities between monotreme and therian cortical development. We have used material from the Hill embryological collection to try to answer key questions concerning cortical development in monotremes. Our findings indicate that gyrencephaly begins to emerge in the echidna brain shortly before birth (crown-rump length 12.5 mm), whereas the cortex of the platypus remains lissencephalic throughout development. The cortices of both monotremes are very immature at the time of hatching, much like that seen in marsupials, and both have a subventricular zone (SubV) within both the striatum and pallium during post-hatching development. It is particularly striking that in the platypus, this region has an extension from the palliostriatal angle beneath the developing trigeminoreceptive part of the somatosensory cortex of the lateral cortex. The putative SubV beneath the trigeminal part of S1 appears to accommodate at least two distinct types of cell and many mitotic figures and (particularly in the platypus) appears to be traversed by large numbers of thalamocortical axons as these grow in. The association with putative thalamocortical fibres suggests that this region may also serve functions similar to the subplate zone of Eutheria. These findings suggest that cortical development in each monotreme follows distinct paths from at least the time of birth, consistent with a long period of independent and divergent cortical evolution.

The living monotremes (platypus and echidnas) are distinguished from therians as well as each other in part by the unusual structure of the thalamus in each. In particular, the platypus has an enlarged ventral posterior (VP) nucleus reflecting the great behavioural importance of trigeminosensation and electroreception. The embryological collections of the Museum für Naturkunde in Berlin were used to analyse the development of the dorsal thalamus and ventral thalamus (prethalamus) in both species. Prosomeric organization of the forebrain emerged at 6 mm crown-rump length (CRL), but thalamic neurogenesis did not commence until about 8-9 mm CRL. Distinctive features of the dorsal thalamus in the two species began to emerge after hatching (about 14-15 mm CRL). During the first post-hatching week, dense clusters of granular cells aggregated to form the VP of the platypus, whereas the VP complex of the echidna remained smaller and divided into distinct medial and lateral divisions. At the end of the first post-hatching week, the thalamocortical tract was much larger in the platypus than the echidna. The dorsal thalamus of the platypus is essentially adult-like by the sixth week of post-hatching life. The similar appearance of the dorsal thalamus in the two species until the time of hatching, followed by the rapid expansion of the VP in the platypus, is most consistent with ancestral platypuses having undergone changes in the genetic control of thalamic neurogenesis to produce a large VP for trigeminal electroreception after the divergence of the two lineages of monotreme.

Cornu Saigae Tataricae (antelope horn), Manis Squama (pangolin scale), Cornu Cervi Pantotrichum (velvet antler) and Cornu Bovis grunniens (yak horn) are valuable medicinal animal horns and shells (MAHS). As the major source of biological agents and ethnodrugs, MAHS show pretty good bioactivities. However, with the increased demand for MAHS, some of the medicinal resources are endangered, and there has been a concomitant increase in the prevalence of adulterated or impostor MAHS. It is of great significance to exploit the substitutes for endangered medicinal animal resources. This study is going to provide a new mode for the exploitation of the substitutes of MAHS.Plasma recalcification time, thrombin time and thrombin consumption were recorded to evaluate the anticoagulation effect of MAHS. Dissolution rate of thrombus in vitro and whole blood-gore were observed to appraise the antithrombotic effect of MAHS.All the MAHS involved in this study except Cornu Procaprae Gutturosae (argali horn), Cornu Saigae Tataricae and Cornu Bovis (cattle horn) could not only prolong recalcification time (P0.01) and thrombin time (P0.01), but also dissolve the thrombus in vitro (P0.01) and whole blood-gore (P0.01). The bioactivities among MAHS differed greatly from each other.The results indicate that Cornu Caprae Hircus (goat horn), Cornu Bubali (water buffalo horn) and Trionycis Carapax (turtle shell) are rational to be explored as the substitutes of Cornu Saigae Tataricae, Cornu Bovis grunniens and Manis Squama, respectively. On the contrary, velvet antler is not suitable to be substituted by Cornu Cervi (deerhorn). We presume that the bioactive evaluation methods are effective means of seeking substitutive resources of endangered medicinal animals with the advantages of close correlation to drug action, low dosage, and high sensitiveness.

To trace the emergence of modern IgA isotypes during vertebrate evolution we have studied the immunoglobulin repertoire of a model monotreme, the platypus. Two highly divergent IgA-like isotypes (IgA1 and IgA2) were identified and their primary structures were determined from full-length cDNAs. A comparative analysis of the amino acid sequences for IgA from various animal species showed that the two platypus IgA isotypes form a branch clearly separated from their eutherian (placental) counterparts. However, they still conform to the general structure of eutherian IgA, with a hinge region and three constant domains. This indicates that the deletion of the second domain and the formation of a hinge region in IgA did occur very early during mammalian evolution, more than 166 million years ago. The two IgA isotypes in platypus differ in primary structure and appear to have arisen from a very early gene duplication, possibly preceding the metatherian eutherian split. Interestingly, one of these isotypes, IgA1, appears to be expressed in only the platypus, but is present in the echidna based on Southern blot analysis. The platypus may require a more effective mucosal immunity, with two highly divergent IgA forms, than the terrestrial echidna, due to its lifestyle, where it is exposed to pathogens both on land and in the water.

The portion of echnida sublingual glycoproteins which could be solubilized was fractionated by electrophoresis in polyacrylamide gel and by filtration on a column of Sephadex G-100. Analyses for carbohydrate and amino acid are given and are compared with those of the submaxillary mucins of cattle, dog, pig and sheep. The molar ratio of galactosamine to sialic acid was roughly 1:1 in the echidna fractions, the sialic acid was N-acetyl-neuraminic acid, and there was very little neutral sugar. Proline was relatively high in amount, and isoleucine, half-cystine, methionine, leucine, tyrosine, and phenylalanine were absent or present to a very small extent. The material was weakly antigenic in the rabbit.

SUMMARY The N-terminal region of human HOXA13 has seven discrete polyalanine tracts. Our previous analysis of these tracts in multiple major vertebrate clades suggested that three are mammal-specific. We now report the N-terminal HOXA13 repetitive tract structures in the monotreme Tachyglossus aculeatus (echidna). Contrary to our expectations, echidna HOXA13 possesses a unique set of polyalanine tracts and an unprecedented polyglycine tract. The data support the conclusion that the emergence of expanded polyalanine tracts in proteins occurred very early in the stem lineage that gave rise to mammals, between 162 and 315Ma.

Specimens of Amblyomma oudemansi (Neumann, 1910) were collected in Papua New Guinea from an endangered monotreme, Zaglossus bruijni (Peters & Doria), the western long-beaked echidna. These ticks were compared morphologically and molecularly with species formerly assigned to Aponomma Neumann, 1899 (now included in Bothriocroton Keirans, King, & Sharrad, 1994 or Amblyomma Koch, 1844), and a phylogeny was generated. Based on our results, we reassign this tick to Bothriocroton, as B. oudemansi (Neumann, 1910) n. comb. Original descriptions are provided for the female and the nymph of this species and the male is redescribed. A revised list of all Bothriocroton records and holdings in the US National Tick Collection is also provided.

The genomes of the egg-laying platypus and echidna are of particular interest because monotremes are the most basal mammal group. The chromosomal distribution of an ancient family of short interspersed repeats (SINEs), the core-SINEs, was investigated to better understand monotreme genome organization and evolution. Previous studies have identified the core-SINE as the predominant SINE in the platypus genome, and in this study we quantified, characterized and localized subfamilies. Dot blot analysis suggested that a very large fraction (32% of the platypus and 16% of the echidna genome) is composed of Mon core-SINEs. Core-SINE-specific primers were used to amplify PCR products from platypus and echidna genomic DNA. Sequence analysis suggests a common consensus sequence Mon 1-B, shared by platypus and echidna, as well as platypus-specific Mon 1-C and echidna specific Mon 1-D consensus sequences. FISH mapping of the Mon core-SINE products to platypus metaphase spreads demonstrates that the Mon-1C subfamily is responsible for the striking Mon core-SINE accumulation in the distal regions of the six large autosomal pairs and the largest X chromosome. This unusual distribution highlights the dichotomy between the seven large chromosome pairs and the 19 smaller pairs in the monotreme karyotype, which has some similarity to the macro- and micro-chromosomes of birds and reptiles, and suggests that accumulation of repetitive sequences may have enlarged small chromosomes in an ancestral vertebrate. In the forthcoming sequence of the platypus genome there are still large gaps, and the extensive Mon core-SINE accumulation on the distal regions of the six large autosomal pairs may provide one explanation for this missing sequence.

We have examined the organization of the pretectal area in two monotremes (the short beaked echidna-Tachyglossus aculeatus, and the platypus-Ornithorhynchus anatinus) and compared it to that in the Wistar strain rat, using Nissl staining in conjunction with enzyme histochemistry (acetylcholinesterase and NADPH diaphorase) and immunohistochemistry for parvalbumin, calbindin, calretinin and non-phosphorylated neurofilament protein (SMI-32 antibody). We were able to identify distinct anterior, medial, posterior (now called tectal gray) and olivary pretectal nuclei as well as a nucleus of the optic tract, all with largely similar topographical and chemoarchitectonic features to the homologous regions in therian mammals. The positions of these pretectal nuclei correspond to the distributions of retinofugal terminals identified by other authors. The overall size of the pretectum in both monotremes was found to be at least comparable in size, if not larger than, the pretectum of representative therian mammals of similar brain and body size. Our findings suggest that the pretectum of these two monotreme species is comparable in both size and organization to that of eutherian mammals, and is more than just an undifferentiated area pretectalis. The presence of a differentiated pretectum with similar chemoarchitecture to therians in both living monotremes lends support to the idea that the stem mammal for both prototherian and therian lineages also had a differentiated pretectum. This in turn indicates that a differentiated pretectum appeared at least 125 million years ago in the mammalian lineage and that the stem mammal for proto- and eutherian lineages probably had similar pretectal nuclei to those identified in its descendants.

Although it is generally understood that the testes recruited kidney ducts for reproductive function during the evolution of vertebrates, little is understood of the biological significance of the adaptation. In the context of the evolution of the mammalian epididymis, this report provides evidence that a major role of the epididymis is to enhance a male's chance of achieving paternity in a competitive mating system. A unique example of sperm cooperation in monotremes is used as evidence that the epididymis produces sperm competition proteins to form groups of 100 sperm into bundles that have a forward motility nearly thrice that of individual spermatozoa. As it required 3-h incubation in vitro under capacitation conditions to release motile sperm from the bundles, it is suggested that the monotremes provide an example of capacitation that is quite different from capacitation in higher mammals. It is suggested that variation between species in the intensity of sperm competition could explain the variation that occurs between species in the amount of post-testicular sperm maturation and storage in the epididymis, an explanation of why the human epididymis does not play as important a role in reproduction as the epididymis of most mammals.

Background: In the few marsupial species studied to date that possess a retinal vasculature, the arterial and venous segments, down to the smallest calibre capillaries, have been shown to occur in pairs. This unique pattern of retinal vasculature has been described in only a few marsupial species. It is a pattern seen in the marsupial central nervous system (CNS) but not in other tissues in this group or in any tissues in eutherian mammals. The aim of the present study was to investigate the presence of retinal vessels in a range of non-eutherian mammalian species (marsupials and monotremes) and to determine if the pattern of paired vessels was a widespread phenomenon within this animal group. Methods: Species studied included a monotreme, the short-billed echidna (Tachyglossus aculeatus) and a range of Australian marsupials, the honey possum (Tarispedidae rostratus), fat-tailed dunnart (Sminthopsis crassicaudata), grey-bellied dunnart (Sminthopsis griseoventer), numbat (Myrmecobius fasciatus), broad-footed marsupial mouse (Antechinus godmani) and the North American opossum (Didelphis virginiana). Eyes were fixed in glutaraldehyde or paraformaldehyde and retinae were embedded in resin for light and electron microscopic analysis. Results: Examination revealed that in those species with retinal vessels (fat-tailed dunnart, grey-bellied dunnart, numbat, marsupial mouse, North American opossum) the pattern of vessels differs from the conventional plexus-like arrangement of mammalian retinal vasculature (i.e. anastomotic networks of capillaries between arterioles and venules). By contrast, in marsupials retinal vessels always occur in closely related pairs with the arteriolar limb usually situated on the vitread aspect. Vessels penetrate the retina and branch to form layers of paired capillaries as far as the outer nuclear layer in some species, which is considerably deeper than in normal mammalian holangiotic retinae. The capillaries form blind-ended hairpin loops and display classical morphological features of CNS capillaries. The vascular endothelium is supported by a distinct basal lamina that splits to envelop pericytes. The latter, although abundant, are invariably interposed between the two vessels that form each vascular unit. Conclusion: The phylogenetic relations of this vascular pattern in the marsupial CNS and retina, and in the CNS of a few other classes of non-mammalian vertebrates, suggest that retinal vascularisation may have evolved independently in marsupial and eutherian mammals and that the former may have evolved from a common primitive mammal-like reptilian ancestor which possessed paired vasculature in the CNS. Eutherian mammals may have evolved from an ancestor with anastomotic networks in the CNS or this pattern may have evolved later in eutherian mammal evolutionary radiation. The possible functional and physiological significance of the paired vessels is discussed.

This study reports the development and application of techniques to assess the reproductive status of male echidnas. The pattern of testosterone secretion over a 24-h period in five echidnas was documented. Testosterone secretion after injection i.m. of either 1000 IU hCG (n = 4) or 4 mu g GnRH agonist (n = 6) was determined to establish whether this could be used as a practical index of the prevailing steroidogenic capacity of the testes. hCG (1000IU) was also used to assess seasonal changes in testosterone secretion in six echidnas over a 13-month period. Seasonal changes in testicular Volume were examined by transabdominal ultrasonography. Electroejaculation was attempted to monitor seasonal changes in sperm production, which was also determined by spermatorrhea. There was no apparent diurnal pattern of testosterone secretion in echidnas and circulating concentrations of testosterone remained relatively low (maximum 1.2 ng/mL) and stable over 24 h. Injection of hCG resulted in an increase (P < 0.01; n = 4) in testosterone concentration with a peak (2.9 +/- 0.3 ng/mL) approximately 4 h after injection. GnRH also induced an increase (P < 0.01; n = 6) in circulating testosterone that was apparent after 1 h (2.6 +/- 0.3 ng/mL) and concentrations remained elevated (3.4 +/- 0.3 ng/mL) for up to 8 h after injection. Seasonal changes in testosterone secretion determined after injection of hCG, increased (P = 0.03; n = 6) from late-autumn, peaked in late-winter, and decreased by early-spring. Testicular volume followed a similar seasonal pattern (P < 0.01; n = 6) with an increase from late-autumn, peak in winter and a decline in mid-spring. There was no seasonal change in live weight. Electroejaculation was attempted throughout two breeding seasons but no semen was obtained. Spermatorrhoea in the echidna was described for the first time and was subsequently used to assess seasonal sperm production. Spermatozoa were found in the urine from June to September. This study has demonstrated that exogenous hormones can be used to obtain an index of the prevailing steroidogenic capacity of the testes in echidnas, which is not apparent with repetitive non-stimulated samples over 24 h. The assessment of testosterone secretion after injection of trophic hormones provides a valuable and practical procedure for the assessment of reproductive status. Testicular ultrasonography and spermatorrhea are useful in assessing reproductive status and in this study were successfully used to determine seasonal reproduction in captive echidnas. (c) 2006 Elsevier B.V. All rights reserved.

Echidnas have evolved separately from other mammalian groups for around 200 million years and incorporate a mixture of reptilian and mammalian features. Because of these attributes, they have historically been considered "primitive" animals. However, they have successfully adapted to a wide variety of ecological niches and their neurophysiology demonstrates a number of unusual and apparently sophisticated characteristics, including a relatively large brain and cerebral cortex and a comparatively massive frontal cortex. Studies of learning in the echidna have thus far been limited to only a handful of experiments which demonstrated relatively basic abilities such as forming a position habit in a T-maze, successive habit-reversal learning, and simple visual and instrumental discrimination. This study aimed to expand on these results and test the "primitive" echidna on what are generally considered more advanced cognitive tasks-same/different and conditional same/different concept learning. The results demonstrated that echidnas are able to discriminate on the basis of a relational same/different concept, using simultaneously presented multi-element stimuli, and transfer that discrimination to novel stimuli. After further training, they were then able to repeat the performance when the correct choice was conditional on the background color of the stimulus panels.

Russell A. Mittermeier, Don E. Wilson (2015): Tachyglossidae. In: Handbook of the Mammals of the World – Volume 5 Monotremes and Marsupials. Barcelona: Lynx Edicions: 34-56, ISBN: 978-84-96553-99-6, DOI: http://doi.org/10.5281/zenodo.6670846

4. Short-beaked Echidna Tachyglossus aculeatus French: Echidné a nez court / German: Ameisenigel / Spanish: Equidna de hocico corto Other common names: Australian Echidna, Short-nosed Echidna, Spiny Anteater; Bristly Echidna (setosus), Kangaroo Island Echidna (multiaculeatus) Taxonomy. Myrmecophaga aculeata Shaw, 1792, Sydney region, Australia. Although Shaw’s original description simply states “it is a native of New Holland,” the accompanying illustration is a copy ofa drawing now in the Thomas Watling Drawings Collection at the Natural History Museum in London, along with the text “This animal was found on a large red ant-hill; it seems to live on them, therefore we call it the ant-eating porcupine of N.S. Wales.” Five subspecies recognized. Subspecies and Distribution. T. a. aculeatus Shaw, 1792 — E Australia (E New South Wales, Victoria, and S Queensland). T. a. acanthion Collett, 1884 — arid regions of most of Australia (Northern Territory, Queensland, Western Australia, and South Australia). T. a. lawesii Ramsay, 1877 — S & SE New Guinea and N Australia (NE Queensland). T. a. multiaculeatus Rothschild, 1905 — Kangaroo I (South Australia). T. a. setosus E. Geoffroy Saint-Hilaire, 1803 — Tasmania and Bass Strait Is. Descriptive notes. Head-body 30-45 cm; weight 2-6 kg. Beak of the Short-beaked Echidna is c.6 cm. The five subspecies of the Short-beaked Echidna are distinguishable by degree of hairiness of pelage and by length of grooming claws on second and third digits of hindleg. There are differences in metabolic rate and thermoregulation associated with differences in pelage, but there are also significant differences in maternal care and length oflactation (e.g. 147 days in the “Tasmanian Echidna,” 1. a. setosus; 210 days in the “Kangaroo Island Echidna,” 1. a. multiaculeatus, and in the subspecies acanthion). There are no published studies on genetic differences between the populations. The Kangaroo Island Echidna is the most distinctive of the mainland subspecies; it has very long fine pelage obscured by a large number of long, thin spines and its third claw is short. The Tasmanian Echidna has soft thick fur that may completely hide spines;it is often light brown, and a population on Flinders Island is completely blond. Blond Echidnas are also seen on Kangaroo Island. Claws on second and third toes of the Tasmanian Echidna are long and nearly equal in size, and the hindfoot resembles a coarse comb. The subspecies lawesii has long stout spines and thick fur. The form acanthion has sparse coarse fur and long spines. In some echidnas from northern New South Wales (probably acanthion), the second claw is nearly three times the length of the third claw. Habitat. All Australian terrestrial habitats and a variety of habitats in New Guinea, including open woodlands and forests. Food and Feeding. Dietary data from across Australia suggests that in areas where sufficient termites are available they form the majority of the diet of the Short-beaked Echidna, with ants making up a smaller proportion. Where termites are absent or in low availability, ants are the most common dietary item, but these are supplemented with pasture grubs. Breeding. Short-beaked Echidnas are seasonal breeders, and breeding times vary with latitude: breeding begins later in more northerly populations. Mating occurs between early June and September in Tasmania, between early June and early August on Kangaroo Island, between late June and early August at Mount Kosciuszko, and between mid-July and early September in south-eastern Queensland. Activity patterns. Short-beaked Echidnas are commonly seen in daytime, particularly in colder areas, but the majority oftheir activity is nocturnal. Movements, Home range and Social organization. Outside of the mating period, male and female Short-beaked Echidnas are solitary, and males do not assist with rearing young. During the mating season, roving males range widely in search of females. Home ranges of females are 0-15 km® to 0-7 km? depending on habitat quality. Home ranges of males are on average twice as large (0-3-1-5 km?®), with size dependent on availability of females. Status and Conservation. Classified as Least Concern on The IUCN Red List. Shortbeaked Echidnas are protected throughout Australia. They are threatened by overhunting in New Guinea, and the Kangaroo Island Echidna, although still common and widespread on the island, is showing some decline in numbers. Bibliography. Augee et al. (2006), Griffiths (1978, 1989), Morrow & Nicol (2009), Morrow et al. (2009), Nicol (2013), Nicol & Morrow (2012), Nicol et al. (2011), Sprent & Nicol (2012)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2015, Tachyglossidae, pp. 34-56 in Handbook of the Mammals of the World – Volume 5 Monotremes and Marsupials, Barcelona :Lynx Edicions on page 56, DOI: 10.5281/zenodo.6670846

Russell A. Mittermeier, Don E. Wilson (2015): Tachyglossidae. In: Handbook of the Mammals of the World – Volume 5 Monotremes and Marsupials. Barcelona: Lynx Edicions: 34-56, ISBN: 978-84-96553-99-6, DOI: http://doi.org/10.5281/zenodo.6670846

Family TACHYGLOSSIDAE (ECHIDNAS) * Medium-sized, egg-laying mammals, completely lacking teeth, adapted for insectivorous or myrmecophagous diet, with robust, dome-shaped, spine-covered bodies and slender beaks. e 30-65 cm. * Australasian Region. * Mainly montane forests and alpine grasslands in New Guinea, but almost all terrestrial habitats in Australia. s 2 genera, 4 species, 11 taxa. * 3 species Critically Endangered; none Extinct since 1600., Published as part of Russell A. Mittermeier & Don E. Wilson, 2015, Tachyglossidae, pp. 34-56 in Handbook of the Mammals of the World – Volume 5 Monotremes and Marsupials, Barcelona :Lynx Edicions on page 34, DOI: 10.5281/zenodo.6670846

We studied the interaction between torpor and reproduction in free-ranging female Tasmanian echidnas using a combination of techniques including urogenital smears, hormone analysis, ultrasonography, external temperature loggers and camera traps. Male echidnas initiated mating activity by locating hibernating females. All females that mated or were disturbed by males prior to July 27 re-entered hibernation, including many that were pregnant. Pregnant females only entered hibernation in early pregnancy when plasma progesterone concentrations were about twice basal and progesterone then remained constant during torpor. By re-entering hibernation pregnant females extended their gestation period and delayed egg-laying. Progesterone peaked 4–6 days before egg-laying, then dropped rapidly.

In this study, temperature-related structural changes were investigated in human, duck-billed platypus (Ornithorhynchus anatinus, body temperature T(b) = 31-33 degrees C), and echidna (Tachyglossus aculeatus, body temperature T(b) = 32-33 degrees C) hemoglobin using circular dichroism spectroscopy and dynamic light scattering. The average hydrodynamic radius (R(h)) and fractional (normalized) change in the ellipticity (F(obs)) at 222 +/- 2 nm of hemoglobin were measured. The temperature was varied stepwise from 25 degrees C to 45 degrees C. The existence of a structural transition of human hemoglobin at the critical temperature T(c) between 36-37 degrees C was previously shown by micropipette aspiration experiments, viscosimetry, and circular dichroism spectroscopy. Based on light-scattering measurements, this study proves the onset of molecular aggregation at T(c). In two different monotremal hemoglobins (echidna and platypus), the critical transition temperatures were found between 32-33 degrees C, which are close to the species' body temperature T(b). The data suggest that the correlation of the structural transition's critical temperature T(c) and the species' body temperature T(b) is not mere coincidence but, instead, is a more widespread structural phenomenon possibly including many other proteins.

This study was undertaken to determine whether the olfactory tubercles of two monotremes (platypus and echidna) showed cyto- or chemoarchitectural differences from the tubercles of therian mammals. Nissl staining was applied in conjunction with enzyme reactivity for NADPH diaphorase and acetylcholinesterase, and immunoreactivity for calcium binding proteins (parvalbumin, calbindin and calretinin) and tyrosine hydroxylase (echidna only). Golgi impregnations of the tubercle were also available for the echidna. The olfactory tubercle is a poorly laminated structure in the echidna, despite the pronounced development of other components of the echidna olfactory system, and the dense cell layer of the olfactory tubercle was found to be discontinuous and irregular. Granule cell clusters (islands of Calleja) were present, but were small, poorly defined and did not show the intense NADPH diaphorase activity seen in marsupial and placental mammals. A putative small island of Calleja magna was seen in only one echidna out of four. In Golgi impregnations of the echidna olfactory tubercle, the most abundant neuron type was a medium-sized densely spined neuron similar to that seen in the olfactory tubercle of some therians. Large spine-poor neurons were also seen in the polymorphic layer. In the platypus, the olfactory tubercle was very small but showed more pronounced lamination than the echidna, although no granule cell clusters were seen. In both monotremes, the development of the olfactory tubercle was poor relative to other components of the olfactory system (bulb and piriform cortex). The small olfactory tubercle region in the platypus is consistent with poor olfaction in that aquatic mammal, but the tubercle in the echidna is more like that of a microsmatic mammal than other placentals occupying a similar niche (e.g., insectivores).

The monotremes are an intriguing group of mammals that have major differences in their reproductive physiology and lactation from therian mammals. Monotreme young hatch from leathery skinned eggs and are nourished by milk secreted onto areolae rather than through nipples. Parturition and lactation are in part controlled through the paraventricular and supraoptic nuclei of the hypothalamus. We have used Nissl staining, enzyme histochemistry, immunohistochemistry for tyrosine hydroxylase, calbindin, oxytocin, neurophysin and non-phosphorylated neurofilament protein, and carbocyanine dye tracing techniques to examine the supraoptic and paraventricular nuclei and the course of the hypothalamo-neurohypophysial tract in two monotremes: the short-beaked echidna (Tachyglossus aculeatus) and the platypus (Ornithorhynchus anatinus). In both monotremes, the supraoptic nucleus consisted of loosely packed neurons, mainly in the retrochiasmatic position. In the echidna, the paraventricular nucleus was quite small, but had similar chemoarchitectural features to therians. In the platypus, the paraventricular nucleus was larger and appeared to be part of a stream of magnocellular neurons extending from the paraventricular nucleus to the retrochiasmatic supraoptic nucleus. Immunohistochemistry for non-phosphorylated neurofilament protein and carbocyanine dye tracing suggested that hypothalamo-neurohypophysial tract neurons in the echidna lie mainly in the retrochiasmatic supraoptic and lateral hypothalamic regions, but most neurophysin and oxytocin immunoreactive neurons in the echidna were found in the paraventricular, lateral hypothalamus and supraoptic nuclei and most oxytocinergic neurons in the platypus were distributed in a band from the paraventricular nucleus to the retrochiasmatic supraoptic nucleus. The small size of the supraoptic nucleus in the two monotremes might reflect functional aspects of monotreme lactation.

The cyto- and chemoarchitecture of the olfactory bulb of two monotremes (shortbeaked echidna and platypus) was studied to determine if there are any chemoarchitectural differences from therian mammals. Nissl staining in conjunction with enzyme reactivity for NADPH diaphorase, and immunoreactivity for calcium binding proteins (parvalbumin, calbindin and calretinin), neuropeptide Y, tyrosine hydroxylase and non-phosphorylated neurofilament protein (SMI-32 antibody) were applied to the echidna. Material from platypus bulb was Nissl stained, immunoreacted for calretinin, or stained for NADPH diaphorase. In contrast to eutherians, no immunoreactivity for either the SMI-32 antibody or calretinin was found in the mitral or dispersed tufted cells of the monotremes and very few parvalbumin or calbindin immunoreactive neurons were found in the bulb of the echidna. On the other hand, immunoreactivity for tyrosine hydroxylase in the echidna was similar in distribution to that seen in therians, and periglomerular and granule cells showed similar patterns of calretinin immunoreactivity to eutherians. Multipolar neuropeptide Y immunoreactive neurons were confined to the deep granule cell layer and underlying white matter of the echidna bulb and NADPH diaphorase reactivity was found in occasional granule cells, fusiform and multipolar cells of the inner plexiform and granule cell layers, as well as underlying white matter. Unlike eutherians, no NPY immunoreactive or NADPH diaphorase reactive neurons were seen in the glomerular layer. The bulb of the echidna was comparable in volume to prosimians of similar body weight, and its constituent layers were highly folded. In conclusion, the monotreme olfactory bulb does not show any significant chemoarchitectural dissimilarities from eutheria, despite differences in mitral/tufted cell distribution.

The platypus and echidna are the only extant species belonging to the clade of monotremata, the most basal mammalian lineage. The platypus is particularly well known for its mix of mammalian and reptilian characteristics and work in recent years has revealed this also extends to the genetic level. Amongst the monotreme specific features is the unique multiple sex chromosome system (5X4Y in the echidna and 5X5Y in the platypus), which forms a chain in meiosis. This raises questions about sex chromosome organisation at meiosis, including whether there has been changes in genes coding for synaptonemal complex proteins which are involved in homologous synapsis. Here we investigate the key structural components of the synaptonemal complex in platypus and echidna, synaptonemal complex proteins 1, 2 and 3 (SYCP1, SYCP2 and SYCP3). SYCP1 and SYCP2 orthologues are present, conserved and expressed in platypus testis. SYCP3 in contrast is highly diverged, but key residues required for self-association are conserved, while those required for tetramer stabilisation and DNA binding are missing. We also discovered a second SYCP3-like gene (SYCP3-like) in the same region. Comparison with the recently published Y-borne SYCP3 amino acid sequences revealed that SYCP3Y is more similar to SYCP3 in other mammals than the monotreme autosomal SYCP3. It is currently unclear if these changes in the SYCP3 gene repertoire are related to meiotic organisation of the extraordinary monotreme sex chromosome system.