Description of the female, nymph and larva and mitochondrial genome, and redescription of the male, of Ixodes barkeri Barker, 2019 (Acari: Ixodidae) from the short-beaked echidna, Tachyglossus aculeatus, with a consideration of the most suitable subgenus for this tick

Background Ixodes barkeri, a tick with a distinctive ventrolateral horn-like projection on palpal segment 1, was described in 2019 from two male ticks from the Wet Tropics of Far North Queensland, Australia. However, females lie at the core of the taxonomy and subgenus classification of Ixodes, hence we sought specimens of female ticks, successfully recovering females, plus nymphs and larvae. Mitochondrial genomes are also desirable additions to the descriptions of species of ticks particularly with regard to subgenus systematics. So, we sequenced the mt genomes of I. barkeri Barker, 2019 and the possible relatives of I. barkeri that were available to us (I. australiensis Neumann, 1904, I. fecialis Warburton & Nuttall, 1909, and I. woyliei Ash et al. 2017) with a view to discovering which if any of the subgenera of Ixodes would be most suitable for I. barkeri Barker, 2019. Results The female, nymph, larva, and mitochondrial genome of Ixodes barkeri Barker, 2019 are described for the first time and the male of I. barkeri is redescribed in greater detail than previously. So far, I. barkeri is known only from a monotreme, the short-beaked echidna, Tachyglossus aculeatus (Shaw, 1792), from the highland-rainforests of Far North Queensland, Australia. Conclusions Our phylogeny from entire mitochondrial genomes indicated that I. barkeri, and indeed I. woyliei Ash et al. 2017, another tick that was described recently, are best placed in the subgenus Endopalpiger Schulze, 1935.

Getting out of a mammalian egg: the egg tooth and caruncle of the echidna

In the short-beaked echidna, Tachyglossus aculeatus, after an initial period of in utero development, the egg is laid in the pouch and incubated for 10 days. During this time, fetuses develop an egg tooth and caruncle to help them hatch. However, there are only a few historical references that describe the development of the monotreme egg tooth. Using unprecedented access to echidna pre- and post-hatching tissues, the egg tooth and caruncle were assessed by micro-CT, histology and immunofluorescence, to map the changes at the morphological and molecular level. Unlike mammalian tooth germs that develop by invagination of a placode, the echidna egg tooth developed by evagination, similar to that of the first teeth in some reptiles. The egg tooth ankylosed to the premaxilla, rather than forming a mammalian thecodont attachment, with loss of the egg tooth post-hatching associated with high levels of odontoclasts, and apoptosis. The caruncle formed as a separate mineralisation from the adjacent nasal capsule, and as observed in birds and turtles, the nasal region epithelium expressed markers of cornification. Together, this highlights that the monotreme egg tooth shares many similarities with reptilian teeth, suggesting that this tooth is conserved from a common ancestor of mammals and reptiles.

The Unique Penile Morphology of the Short-Beaked Echidna, Tachyglossus aculeatus

Monotremes diverged from therian mammal ancestors approximately 184 million years ago and have a number of novel reproductive characteristics. One in particular is their penile morphology. There are differences between echidna and platypus phalluses, but both are somewhat similar in structure to the reptilian phallus. The echidna penis consists of 4 rosette glans, each of which contains a termination of the quadrifurcate urethra, but it appears that only 2 of the 4 glans become erect at any one time. Despite this, only a few historical references describe the structure of the echidna penis and none provides an explanation for the mechanisms of unilateral ejaculation. This study confirmed that the echidna penis contains many of the same overall structures and morphology as other mammalian penises and a number of features homologous with reptiles. The corpus cavernosum is well supplied with blood, extends up to the base of the glans penis and is primarily responsible for erection. However, the echidna possesses 2 distinct corpora spongiosa separated by a septum, each of which surround the urethra only distal to the initial urethral bifurcation in the glans penis. Together with the bifurcation of the main penile artery, this provides a mechanism by which blood flow could be directed to only one corpus spongiosum at a time to maintain an open urethra that supplies 2 of the 4 glans to facilitate unilateral ejaculation.

Short-beaked echidna (Tachyglossus aculeatus) home range at Fowlers Gap Arid Zone Research Station, NSW

Echidnas (Tachyglossus aculeatus) are found Australia-wide and appear to be remarkably well-adapted to the arid zone, yet nearly all echidna research has been conducted in temperate, tropical and alpine zones. This study investigated the home range and movement of echidnas in western New South Wales. Radio telemetry tracking was used to locate the echidnas daily during the study period (March-May 2018, November 2018, March-May 2019 and August 2019); the observed home range was 1.47± 1.21km2. This is over twice the reported home range of temperate environments (<0.65km2), suggesting that echidnas exhibit larger home ranges in arid zones. The home range of individual echidnas ranged from 0.02km2 to 3.56km2. Echidnas exhibited a small degree of overlap (6.6%± 19.8%) but this varied considerably between individuals (between 0 to 84.2% overlap.) Four out of the thirteen echidnas died during this study, likely due to the severe drought that occurred during the study. This study provides insight into the movement and home range of echidnas in arid zones, revealing that desert echidnas have large home ranges, probably dependent on the availability of resources.

Short-Beaked Echidna (Tachyglossus aculeatus) Home Range at Fowlers Gap Arid Zone Research Station, NSW

ABSTRACTEchidnas (Tachyglossus aculeatus) are found Australian-wide and appear to be remarkably well-adapted to arid zones, yet, nearly all echidna research has been conducted in temperate, tropical and alpine zones. This study investigated the home range and movement of echidnas in western New South Wales. Radio telemetry tracking was used to locate the echidnas daily during the study period (March-May 2019 and August 2019); the home range was 1.47± 1.21 km2. This is over twice the reported home range of temperate environments (>0.65 km2) suggesting that echidnas exhibit larger home ranges in arid zones. This study provides insight into the movement and home range of echidnas in arid zones, revealing that desert echidnas have large home ranges, dependent on the availability of resources.