Rediscovery of the holotype of Petaurus notatus Peters 1859 (Petauridae; Marsupialia) and clarification of the type locality
Peter Menkhorst A B * and Tim Stranks AA
B
Abstract
A recent integrative taxonomic study concluded that the sugar glider, Petaurus breviceps (sensu lato), comprises more than one species and that Petaurus notatus Peters 1859 should be resurrected as the available name for one of the two newly circumscribed species. The type specimen of Petaurus notatus has been considered to be lost, leading to a neotype being nominated. A recent search of the type collections of the Museum für Naturkunde, Berlin, located the holotype and we provide photographs and a morphological description of the skin and partial skull. The neotype should now be set aside. The type locality of Petaurus notatus is also clarified, correcting misinterpretations of locality information provided in the original type description and recent texts.
Keywords: Blandowski Expedition, Gerard Krefft, holotype, International Code of Zoological Nomenclature, neotype, Petaurus breviceps, Petaurus notatus, type locality.
Introduction
A recent integrative taxonomic study (Cremona et al. 2021) provided convincing molecular and morphometric evidence that the sugar glider (Petaurus breviceps) (sensu lato) comprises more than one species. Cremona et al. (2021) identified Petaurus notatus Peters 1859 as the most appropriate available name for one of the two species that they split from P. breviceps. P. notatus has been regarded as a junior synonym of Petaurus breviceps for some 135 years (Thomas 1888).
McKay (1988) reported that the Petaurus notatus holotype was ‘not found’ at the Museum für Naturkunde, Berlin (MfN), in 1980, and Cremona et al. (2021) took this to mean that the holotype was missing. However, in May 2024, the P. notatus holotype was found in the type collection at MfN.
This research note has three aims relating to the resurrection of the name Petaurus notatus: (1) Cremona et al. (2021) followed McKay (1988) in assuming that the holotype had been lost; we announce its rediscovery and provide photographs and descriptions of morphological features of the skin and damaged skull and lower mandible; (2) two vague and inaccurate descriptions of the type locality appear in past and recent literature; we correct these errors with precise details of the type locality; and (3) we discuss the implications of these findings for the neotype designated by Cremona et al. (2021).
Methods
Search for the holotype
As part of a collaborative project between Museums Victoria and MfN, T. Stranks visited MfN in May 2024. During a stocktake of the mammal collection, it was determined that the holotype of P. notatus was originally registered in the old General Catalogue as ZMB_Mam_2379, Petaurus (Belideus) notatus. During a physical search of the type collection on 23 May 2024, the specimen was located, and photographs and measurements were taken.
Defining the type locality
We investigated the provenance of the holotype by reviewing relevant publications of zoologists involved in its original collection and description, namely Gerard Krefft, Wilhelm Peters and John Gould, and the literature relating to the Blandowski Expedition (1856–1857), which led to the collection of the holotype (Krefft 1866; Darragh 2009; Menkhorst 2009).
Results
Taxonomic history of Petaurus notatus
On 6 January 1859 at a general meeting of the Königlichen Preussische Akademie des Wissenschaften zu Berlin [Royal Prussian Academy of Sciences, Berlin], Dr Wilhelm Peters, Director Zoologisches Museum, formally described a new species of Petaurus, namely Petaurus (Belideus) notatus Peters 1859. In addition to the formal description (Peters 1859 p. 15), the report stated:
‘Mr. Peters reported on a new flying marsupial Petaurus (Belideus) from the southern part of New Holland. ….. The zoological museum bought this animal from Mr. Krefft, who shot it in the northern mountainous regions of Victoria in New Holland.’
Peters then sent the type specimen to John Gould in London so that it could be included and illustrated in Gould’s forthcoming ‘Mammals of Australia’ (Gould 1860; text to plate 26 issued in Part XII in November 1860 (Sauer 1982). Gould expressed reservations about whether P. notatus warranted species status (Gould 1860), but deferred to Peters’ view. However, Thomas (1888) had no such reservations and listed P. notatus as a junior synonym of P. breviceps and so it remained until the conclusions of Cremona et al. (2021) necessitated the application of the name Petaurus notatus to one of the newly circumscribed species. The holotype is registered in the General Catalogue of MfN as ZMB_Mam_2379, Petaurus (Belideus) notatus juv. The abbreviation juv presumably indicates that the animal was a juvenile when collected; unfortunately, damage to the skull precludes confirming the age class (C. Funk, MfN, pers. comm.).
External morphology of the type specimen
As noted by Gould (1860), the holotype is unusual in several features of its pelage colour; the dorsal surfaces of the limbs, patagia and ears are sooty black (Fig. 1) compared with the usual pearl-grey in P. breviceps (senso lato) in south-eastern Australia (Menkhorst and Knight 2011). The ventral surface is mid-grey, without evidence of a whitish fringe to the patagia. Gould emphasised and illustrated a peculiar pattern of colouring in the tail fur, namely, ‘a broad mark of light grey down the middle portion of the upper surface within the black’ (Gould 1860) (Fig. 2). He went as far as proposing the English name ‘stripe-tailed Belideus’ because of this colour pattern in the holotype. This feature is apparent in the photographs of the holotype (Fig. 1, dorsal view), but does not seem as dramatic as Gould indicated in words and art. It is possible that the atypically dark pelage colour of the holotype is due to discolouration caused by dust and grime accumulated during 165 years of storage, and that this factor has also reduced the contrasting colours in the tail compared with Gould’s description and illustration. However, Gould’s description of a broad sooty-black mark on the ‘upper edge of the flying membrane; front part of the anterior limbs and the front and hinder part of the posterior limbs’ indicates that the specimen was indeed darker than is usual for the Petaurus breviceps complex.
Dorsal (upper) and ventral (lower) views of the skin of ZMB_Mam_2379, Petaurus (Belideus) notatus juv., the holotype of Petaurus notatus. Copyright: Eran Wolff, Museum für Naturkunde Berlin.

The illustration of Petaurus notatus by J. Gould and H.G. Richter (plate 26 in Gould 1845–1863).

The skull has been extracted from the skin and is stored separately. Unfortunately, the skull and mandible are extensively damaged with only the anterior portion of each remaining, along with fragments of the brain case (Fig. 3). Fortunately, enough of the upper and lower tooth rows remain to indicate that the material does indeed belong to a small petaurid, having in the upper jaw forward of the molars, three incisors, one canine and three premolars, and in the lower jaw each partial dentary contains one incisor, no canine and four premolars (Fig. 3). The partial dentaries contain only the first two of the four molars, the upper jaw remnant contains the first two molars on the right-hand side and none on the left.
Determining the type locality
Modern statements of the type locality of P. notatus take two forms, referring either to ‘mountains of Victoria (Krefft)’ (Iredale and Troughton 1934) or to ‘Port Phillip, Victoria’ (McKay 1988; Jackson and Groves 2015; Cremona et al. 2021, 2023). However, each of these geographical statements is based on misinterpretation of the writings of the collector, Gerard Krefft.
References to ‘mountainous regions of Victoria’ come from the last sentence of Peters (1859) (quoted above). It mirrors the statement in Krefft (1866, p. 18) regarding his finding that the Indigenous people in the area around the Blandowski Expedition base camp at Mondellimin in the semi-arid north-west of Victoria were not familiar with gliding possums. This lack of local Indigenous knowledge led Krefft to speculate that petaurids were ‘restricted to the mountainous coast districts’ (Krefft 1866) and ‘prefers the mountain districts and is not found on the plains of the interior’ (Krefft 1871 text relating to plate 7). That is, Krefft was speculating that Petaurus species did not penetrate the semi-arid regions of north-western Victoria and south-western New South Wales, the geographical focus of the Blandowski Expedition. Krefft’s hypothesis has been shown to be correct (e.g. Menkhorst 1995).
References to Port Phillip being the type locality emanate from John Gould’s comment (Gould 1860) that the specimen ‘was procured by M. Gerard Krefft in the district of Victoria, generally known as Port Phillip’. McKay (1988) and subsequent authors (Jackson and Groves 2015; Cremona et al. 2021, 2023) seem to have interpreted this statement to refer to that part of Victoria proximate to Port Phillip, that is, to Melbourne. Until the mid-1850s when the Colony of Victoria was established by the British Government, the entire region south of the Murray River was known as the District of Port Phillip, part of the Colony of New South Wales. Thus, as pointed out by Goldingay et al. (2023), it is probable that Gould’s ‘district of Victoria, generally known as Port Phillip’ refers to Victoria as a whole. There is no evidence that Gould was referring to a part of Victoria centred on Port Phillip (Melbourne) or its hinterland. Cremona et al. (2021) selected a neotype from close to Melbourne partly on the basis of this misinterpretation of Gould’s statement.
The actual type locality
This geographical confusion is unfortunate because Krefft himself was perfectly clear and concise about the place of collection of the holotype. In his seminal paper on the vertebrates collected during the Blandowski Expedition (1856–1857), Krefft (1866) wrote:
The first Belideus I captured on my return, at Mt Ida, McIvor Range, 80 miles distant from the Murray, is according to Gould, a new species and is figured by him in part XI (sic) of his Mammalia, 15, as ‘Belideus notatus’ (Krefft 1866 p. 17 under the subheading ‘14 Phalangista viverrina’).
This information was quoted in full by Wakefield (1966) and Dixon (1974) and is consistent with the information provided on the specimen label attached to the holotype, i.e. ‘Mount Igor Neuholland nordlich von Melbourne’ (Fig. 1). Mount Igor is very likely a misspelling of Mount Ida, resulting in the type locality being ‘Mt Ida north of Melbourne’.
Krefft’s wording indicates that he collected the specimen while travelling through central Victoria, during the return journey overland from Mondellimin to Melbourne, after the completion of Blandowski’s 12-month-long Lower Murray Expedition (December 1856 – December 1857). Darragh (2009, p. 40) reported that, ‘In October 1857, Blandowski was directed to recall the field party, but it was only in the middle of December that the party returned’. We presume that Krefft and the small remaining expedition team journeyed from Mondellimin to Melbourne between about mid-November and mid-December 1857 and that the glider specimen was collected during that period near Mount Ida. Mount Ida is the high point (451 m) of a narrow, north–south range of hills known as the McIvor Range that lies between the present-day towns of Heathcote and Colbinabbin in the Goldfields Region of central Victoria. It is 108 km north–north-west of Melbourne and we suggest that a representative grid reference for the type locality is −36°53′25″S, 144°41′48″E.
In the 1850s, the area around Mount Ida and nearby McIvor Creek was a rich alluvial goldfield, and home to hundreds of miners. Accommodation and supplies would have been available for travellers, making it a convenient port of call for members of the Blandowski Expedition as they headed south from the Murray River to Melbourne. The primary vegetation communities around Mount Ida are box–ironbark dry open-forest on the skeletal soils of the ranges and box–gum woodlands on the heavier and more fertile soils of the surrounding plains. Petaurus notatus is present throughout the area in both vegetation communities (Menkhorst and Gilmore 1979 as P. breviceps) and had the highest spotlighting rate of all vertebrates in a survey of the nearby Rushworth State Forest (now incorporated into Heathcote–Graytown National Park) (Myers and Dashper 1999 as P. breviceps).
Discussion
Implications for the neotype
The International Code of Zoological Nomenclature (ICZN 1999 article 75.8) states that if, after the designation of a neotype, the name-bearing type that was presumed lost is found to still exist, on publication of that discovery the rediscovered material again becomes the name-bearing type and the neotype is set aside. This paper performs that role and we hereby set aside the neotype assigned by Cremona et al. (2021) and re-instate the original holotype.
The significance of a type locality
Under ICZN (1999), any name-bearing type specimen requires a concise statement of the place from which the name-bearing type specimen was collected, known as the type locality. Although the type locality may seem of little intrinsic interest, it becomes important if subsequent taxonomic research indicates changes to the delineation of taxa, for example, the splitting of an existing species into two or more taxa. Whenever possible, the collection localities of museum specimens, and particularly type specimens, should be recorded as accurately as possible on collection databases. ICZN (1999) also stipulates that a statement of a type locality that is found to be erroneous should be corrected (article 76A.2). In the case of species that now have relictual distributions, the type locality can provide important information on historical distribution and habitat, helping counteract the insidious effects of recency bias and changing baselines in conservation planning (Menkhorst 2024).
The history of the type specimen
A remaining question is how the Mount Ida specimen came into the hands of Dr Wilhelm Peters, Director, Royal Museum Berlin, less than 2 years after it was collected. Some information is available about Krefft’s activities following his return to Melbourne in mid-December 1857. In February–March 1858, he undertook 4 weeks of temporary employment with Frederick McCoy at the National Museum of Victoria, under directions to identify and sort specimens from the [Blandowski] Expedition Collection to be kept for the Museum (Pescott 1954). In March 1858, Krefft learned of the death of his father William, and left Melbourne to travel to Germany on family business (with a stopover in England). Subsequent events indicate that he took natural history material with him, including specimens such as the Petaurus and his portfolio of illustrations from the Blandowski Expedition.
Details of Krefft’s voyage are sketchy but he was in London in June 1858; on 22 June, Krefft read a short paper to the Zoological Society of London on the Chestnut-crowned Babbler Pomatostomus ruficeps, on the basis of observations made during the Blandowski Expedition (Krefft 1858). While in London, Krefft met eminent British zoologists including Oldfield Thomas and Dr J. E. Gray at the British Museum and John Gould at the Zoological Society (Whitley 1959; Nancarrow 2009). Similarly, Krefft appears to have spent the second half of 1858 visiting principal German museums, universities and scientific societies, to try to gather support for his ongoing work (in addition to visiting family in Brunswick). Krefft met with representatives of the German Museums Commission and discussed arrangements for the exchange of specimens between museums in Germany and the National Museum of Victoria (Whitley 1959; Nancarrow 2009). Krefft left Germany in November 1859 and, after stays at Cape Town and Adelaide, arrived in Sydney in mid-1860. There, he was offered the position of Assistant Curator at the Australian Museum (Whitley 1959; Nancarrow 2009). By this time, Peters had long published his description of P. notatus and sent the type specimen from the Royal Museum Berlin to John Gould in London. Gould included P. notatus as a full species in part XII of ‘Mammals of Australia’ (Gould 1860; Waterhouse 1885). Part XII was published on 1 November 1860 (Sauer 1982). Thus, the Mount Ida specimen must have been sold by Krefft to the Royal Museum Berlin while he was in Berlin in 1858 (Peters having published its description in January 1859). The type specimen then dropped from sight. Clearly, John Gould returned it to the Royal Museum but a search of the collections at MfN by G. McKay in 1980 failed to locate it (McKay 1988). On this basis, subsequent authors have assumed that it had been lost or destroyed (Jackson and Groves 2015; Cremona et al. 2021).
Curiously, one of the few documented actions by Krefft while working at the National Museum in Melbourne after his return from Mondellimin and before leaving for Europe was to lodge a number of Blandowski Expedition specimens that had been given to him by Blandowski, telling the museum Director, Frederick McCoy, that ‘I was not aware at the time that Mr Blandowski had no authority to give me them’ (Pescott 1954, p. 40). Apparently Krefft did not consider that the Mount Ida petaurid specimen was similarly the property of the Victorian Museum; he stated that the genus Petaurus was ‘not represented’ in collections from the Expedition (Krefft 1866, p. 7).
All specimens from the Expedition were officially recorded in a large bound, but unpublished ‘Catalogue of all Specimens of Natural History collected by Mr Blandowski’s Party during an Expedition to the Lower Murray in 1857’ (Krefft no date a), and also in ‘List of Specimens of Natural History kept for the Museum – Mammalia collected during 1857 in different localities on the Lower Murray and Darling’ (Krefft no date b). Other details can also be found on the Consignment Lists I–XX (held in the Museums Victoria Archives). There is no mention of Petaurus (or Belideus) in any of these sources. Indeed, there are no specimens listed that were collected after the expedition members left Mondellimin in November 1857. So Krefft makes it quite clear that, in his mind, the Petaurus was not officially collected during the expedition. Krefft’s inclusion of a statement about the lack of specimens of Belideus collected during the expedition is not at odds with this stance. Krefft may have thought that his official expedition collecting duties were over once he left Mondellimin, and perhaps felt free to make his own personal collections as he travelled on his own time back to Melbourne, with no obligation to lodge these at the museum in Melbourne.
This investigation has highlighted the value of searching beyond the contents of museum specimen registers, especially recently digitised registers, when looking for information pertaining to historical specimens. Often, digging a little deeper into the wealth of available published material and unpublished archives, reveals useful information that had been considered less important when modern digitised specimen registers were created. Such information can be important both for its relevance to the taxonomy and nomenclature of the taxon and because it can have significant implications for conservation planning by providing refined locality information (e.g. Menkhorst 2024).
Data availability
This paper is a review of historical literature and museum specimens. It does not present data as such.
Declaration of funding
Tim Stranks received funding support from Museums Victoria for his visit to MfN.
Acknowledgements
We thank Eva Diestelhorst for translating Peters (1859) from German. Christiane Funk (Collections Manager recent mammal collection) and Eran Wolff (photographer) at MfN kindly provided the photographs used in Figs 1 and 3. Museums Victoria created the opportunity for T. Stranks to travel to Berlin to work with staff of the MfN on its holdings of specimens from Australia. Ross Goldingay and an anonymous reviewer provided helpful comments on a draft.
References
Cremona, T., Baker, A. M., Cooper, S. J. B., Montague-Drake, R., Stobo-Wilson, A. M., and Carthew, S. M. (2021). Integrative taxonomic investigation of Petaurus breviceps (Marsupialia: Petauridae) reveals three distinct species. Zoological Journal of the Linnean Society 191, 503-527.
| Crossref | Google Scholar |
Darragh, T. A. (2009). William Blandowski: a frustrated life. Proceedings of the Royal Society of Victoria 121(1), 11-60.
| Crossref | Google Scholar |
Goldingay, R. L., Jackson, S. M., Winter, J. W., Harley, D. K. P., Bilney, R. J., Quin, D. G., Smith, G. C., Taylor, B. D., and Kavanagh, R. P. (2023). What’s in a name? Selection of common names among new and revised species of Australian mammals, and the case of the sugar glider. Australian Mammalogy 46(1), AM23017.
| Crossref | Google Scholar |
Iredale, T., and Troughton, E. L. G. (1934). A check-list of the mammals recorded from Australia. Australian Museum Memoir 6, 1-122.
| Crossref | Google Scholar |
Krefft, G. (no date a). ‘List of Specimens Kept for the Museum.’ (Museum Victoria) Available at Catalogue – Mammalia Collected from Murray and Darling Expedition – National Museum of Victoria – 1857
Krefft, G. (no date b). ‘Blandowski Catalogue of all specimens of natural history collected by Mr Blandowski’s party during an expedition to the lower Murray in 1857.’ (Museum Victoria) Available at Catalogue of all specimens of natural history collected by Mr Blandowski’s party during an expedition to the Lower Murray in 1857 - Biodiversity Heritage Library (biodiversitylibrary.org)
Krefft, G. (1858). A few remarks on the habit and economy of the brown-capped pomatorhinus (P. ruficeps, Hartlaub). Proceedings of the Zoological Society London 26, 352-353.
| Crossref | Google Scholar |
Krefft, G. (1866). On the vertebrated animals of the Lower Murray and Darling, their habits, economy and geographical distribution. Transactions of the Philosophical Society of New South Wales, 1862-1865 1-33.
| Crossref | Google Scholar |
Menkhorst, P. W. (2009). Blandowski’s mammals: Clues to a lost world. Proceedings of the Royal Society of Victoria 121(1), 61-89.
| Crossref | Google Scholar |
Menkhorst, P. (2024). Ecological history can inform current conservation actions: lessons from an Australian rodent. Pacific Conservation Biology 30, PC23049.
| Crossref | Google Scholar |
Menkhorst, P. W., and Gilmore, A. M. (1979). Mammals and reptiles of North Central Victoria. Memoirs of the National Museum of Victoria 40, 1-33.
| Crossref | Google Scholar |
Myers, S. D., and Dashper, S. G. (1999). A survey of the vertebrate fauna of the Rushworth State Forest. The Victorian Naturalist 116, 131-141.
| Google Scholar |
Nancarrow, J. (2009). Gerard Krefft: a singular man. Proceedings of the Royal Society of Victoria 121(1), 146-154.
| Crossref | Google Scholar |
Peters, W. (1859 [In German]). Über ein neues Flugbeutelthier, Petaurus (Belideus), aus dem südlichen Theile von Neuholland. Monatsberichte der Königlichen Preussischen Akademie der Wissenschaften zu Berlin for 1859 1859, 14-15.
| Google Scholar |
Wakefield, N. A. (1966). Mammals of the Blandowski Expedition to north-western Victoria. Proceedings of the Royal Society of Victoria 79, 371-391.
| Google Scholar |
Whitley, G. P. (1959). The life and work of Gerard Krefft (1830-1881). Proceedings of the Royal Zoological Society of New South Wales 1958-1959 21-34.
| Google Scholar |