Beverley Anne Holloway (25 October 1931–11 May 2023)

Beverley Anne Holloway (Figure 1) died in her sleep at her residence in Kumeu Village Rest Home, Auckland at the age of 91. She was an inspirational person as a mother, grandmother, wife and as an entomologist. Her impact on the world of entomology is felt today as a consequence of a broad career which included publishing on groups including Diplopoda, Lepidoptera, Diptera and Coleoptera, and with her research encompassing areas of systematics, evolution, natural history and behaviour, through to medical and forensic entomology. Her publications still inspire people to work on oft-forgotten groups of arthropods, particularly certain families of beetles.

Figure 1. Beverley Holloway at Mt. Albert Research Centre on 20 Feb 1991. Photograph by Andrew Thompson, provided courtesy of the New Zealand Herald Archive and the Alexander Turnbull Library, PA1-q-1058.

Beverley was born in Lower Hutt and attended Stokes Valley School and Wellington Girls’ College. From an early age she showed a love for the natural world. She especially enjoyed tramping in the hills behind Stokes Valley and in the Rimutaka Ranges. In 1952, Beverley was employed by the Dominion Museum (now the Museum of New Zealand Te Papa Tongarewa) as Assistant Entomologist, while still attending Victoria University College, Wellington, as a part-time student. Although not a foundation member of the Entomological Society of New Zealand, she attended the first conference of the Society held in Nelson in 1952. Two years later, in 1954, she was awarded a Master of Science degree with first class honours in Zoology for her study on the pill millipede, Procyliosoma tuberculatum Silvestri.

In 1955, Beverley was awarded a Fulbright Scholarship to study for her PhD in the United States of America. She moved to Cambridge, Massachusetts to study biology at Harvard University’s Museum of Comparative Zoology, where she revised the taxonomy of New Zealand stag beetles (Lucanidae) under the supervision of Professor Frank Carpenter. During her studies, she travelled to London to examine relevant type specimens in the British Museum (Natural History) and was elected to the prestigious honour societies, Phi Beta Kappa and Society of the Sigma Xi. She returned to New Zealand in 1958 and was awarded her PhD in 1959. Her experience in the USA was highly influential, and through the remainder of her life, Beverley remained in correspondence with colleagues from this time, such as P.J. Darlington, E.O. Wilson, and the chemical ecologist and naturalist Thomas Eisner. She had little time to go out and see the sights of America while doing her PhD studies, but returned to Harvard in the 1970s to visit Frank Carpenter. She dedicated her 1982 monograph on the Anthribidae to him to mark his 80th birthday.

Back at home in New Zealand, Beverley was appointed as Entomologist at the Dominion Museum in 1958, and published two significant papers on stag beetles: a taxonomic revision of the New Zealand Lucanidae, and the first phylogeny of the Lucanidae, which was to form the basis of the modern classification of the family. In 1961, Beverley hosted a visit from a Chilean entomologist, Guillermo (Willy) Kuschel, which was to prove significant for both, professionally and personally. In 1962, both Beverley and Willy accepted positions with Entomology Division in the Department of Scientific and Industrial Research (DSIR) in Nelson. Beverley and Willy were married in 1963 (Figure 2).

Figure 2. Left: Guillermo (Willy) Kuschel, Right: Beverley Holloway. Photograph taken in 1991. Photographer unknown. Reproduced with permission from the Kuschel family.

Beverley left her paid employment in 1963 to raise their three children, Gerda, Carl, and Erika, as at this time there was very little financial support for working mothers. By bearing the bulk of domestic duties, she enabled Willy to undertake the extensive collecting expeditions that allowed the rapid expansion of the New Zealand Arthropod Collection (NZAC) at this time. Family holidays doubled as collecting trips, and Beverley is remembered for always having vials of ethanol handy in her purse for those moments when an interesting insect had the misfortune to cross her path. While Willy continued his research, Beverley simultaneously maintained her interest in systematics. She significantly influenced Willy’s thoughts and ideas about taxonomic character systems and continued to publish papers on the systematics and morphology of New Zealand stag beetles and other groups of beetles.

In 1973, the family moved from Nelson to Auckland, accompanying the transfer of the DSIR insect collection to the Mount Albert Research Centre. In the following year, Beverley re-joined the systematics group. Instead of working on beetles, she shifted her focus to Diptera. Her research included the description of Mystacinobia zelandica Holloway, the short-tailed New Zealand bat fly (Figure 3), and erecting a new family Mystacinobidae to receive this sole member. In addition to describing the adult, larva and puparium, she presented a remarkably detailed account of the species’ lifecycle and behaviour. Her characteristically careful attention to detail extended to rearing the flies in the hot water cylinder cupboard in her home (Hunt & Morris, 2006). Beverley also conducted substantial research into medically and forensically important insects, such as providing methods for identifying maggots found on carrion (including human corpses).

Figure 3. (a) Dorsal drawing of male Mystacinobia zelandica Holloway. Artist: D. W. Helmore. Scale bars represent 1 mm. High resolution images obtained from Wikimedia Commons, CC BY 4.0. (b) Male Geodorcus helmsi collected from Sayer’s Bay, Stewart Island by Anthony Harris in 1976. Specimen deposited in Insect Collection of the Otago Museum. Photograph by On Lee Lau © Otago Museum.

Beverley’s determination to succeed meant that she excelled in circumstances where it could be perceived she was set up to fail. The taxonomy and systematics of the anthribid weevils of New Zealand had not been studied since the early work of David Sharp and Thomas Broun in the late 1800s. Although many species had been described, the bulk of these had been placed in the Palaearctic genus Anthribus Geoffrey, which at that time served as a convenient grouping containing a multitude of unrelated taxa. In 1978, when the idea of a national taxonomic monograph series was conceived, Beverley was directed to prepare a treatment on the fungus weevils (Anthribidae; Figure 4). Beverley had earlier experience on the group to draw upon, but there was still a lot to be done. In December 1982, Beverley’s work was published as one of the inaugural volumes of the Fauna of New Zealand series, which has since gone on to become one of the most prestigious outlets for taxonomic monographic research on New Zealand arthropods. Beverley’s volume is astonishing in its scope, thoroughness, and clarity, and remains the single most comprehensive work on anthribid weevils worldwide. She brought order to the taxonomy of New Zealand Anthribidae, with the description of 18 genera and 13 species, and proposing 29 new combinations and 43 synonymies. She recognised a total of 60 species in New Zealand, each of which was thoroughly described and illustrated and to which she provided exceptionally clear and innovative identification tools. This work set the standard for future Fauna of New Zealand volumes. The quality of the work becomes even more awe-inspiring when the time pressure to complete the work is considered; having been completed in four years, including a substantial period of time required for copy-editing and preparation for printing.

Figure 4. (a) Lateral view of Gynarchaeus ornatus (Sharp). Photo by Samanta Orellana. (b) Dorsal drawing of male Caliobius littoralis Holloway. Artist: D. W. Helmore. Scale bar represents 1 mm. Figure from Fauna of New Zealand Volume 3: Anthribidae. High resolution image obtained from Wikimedia Commons, CC BY 4.0.

Beverley was the leader of the Systematics Group of Entomology Division, DSIR, from 1988–1990, during which time she advocated for the inclusion of popular summaries in both English and te reo Māori in the Fauna of New Zealand series and in 1990 commissioned the carving of a pare for NZAC to recognise the 150th anniversary of the signing of Te Tiriti o Waitangi. Throughout her career, Beverley championed conditions for women at DSIR and for women in science more generally. This included serving on the Scientists’ Salary Committee.

Beverley retired in 1991, as 60 was the compulsory retirement age in the Public Service at that time. She continued her research and, in particular, she worked on an extension of the lucanid stag beetle work she began with her PhD thesis. In 2007, aged 71, she published her second Fauna of New Zealand volume, the 61st in the series (Figure 5). This monumental work could be considered the crowning jewel in her lifetime of incredible work and cemented Beverley’s reputation for being a systematist par excellence. With clear identification keys, richly detailed illustrations and insight into the biology and evolution of the New Zealand stag beetles, this volume has been an indispensable reference to the family since its publication. Any person interested in the charming and charismatic stag beetles will find untold treasures in this book.

I hope the morphological section and keys will be helpful even for those with only a general entomological knowledge and that others, who may never expect to see a stag beetle, will enjoy looking at the illustrations and be able to appreciate the amazing diversity of the unique New Zealand lucanid fauna. (Beverley Holloway, in the popular Summary of her Lucanidae Fauna of New Zealand volume, 2007).

Figure 5. Left: Beverley Holloway, Image from 2007 Fauna of New Zealand 61: Lucanidae. Photo taken by Birgit Rhode. Right: Dorsal drawing of male Geodorcus sororum Holloway. Artist: D.W. Helmore. High resolution image obtained from Wikimedia Commons, CC BY 4.0.

Having completed this work, Beverley ceased doing active research, proud that she had completed the projects that she had committed to. She remained keenly interested in how entomological science in New Zealand was progressing and kept an intense curiosity about developments in the field, particularly the advent of molecular data in biological systematics and the achievements of women in the New Zealand science system and the opportunities available for them.

Regrettably, shortly before Willy’s death in 2017, Beverley was struck by a car while walking and suffered a head injury. Although she quickly recovered from the physical injuries caused by the accident, her mental clarity declined irreversibly. She spent the last six years of her life in care at the Kumeu Village Rest Home.

Beverley understood that specimens are central to the realm of biological meaning. She treated insect specimens not only as items of taxonomic interest and documentation but strove to derive as much biological information from them as possible, to understand their value as living creatures. Throughout Beverley’s career, she demonstrated that often-overlooked morphological structures can play a large role in the definition and revision of large insect groups. She was fascinated by their morphology in relation to evolution and function and believed it important to highlight morphological features of significance to encourage further study into these structures. Her work on lucanid beetle wing development is of high merit. In her treatment of Anthribidae, you can find the only data known for these beetles’ use of host plants and fungi. Furthermore, in her anthribid weevil monograph she provided synoptic keys which can be more useful for identification than traditional dichotomous keys.

Beverley was an intrepid adventurer and collected insect specimens from some of the most remote and difficult-to-access offshore islands in New Zealand. Between 1952 and 1953, she visited several of the small islands in Cook Strait, including The Brothers, Stephens Island, Middle Trio Island and Sentinel Rock. In early 1955 she was part of an expedition led by the malacologist Richard K. Dell to Taukihepa/Big South Cape and other localities in the south-western area of Stewart Island where her collections make a significant contribution to the record of the insect fauna of Taukihepa prior to the catastrophic invasion of the island by rats in 1964. In 1981 she made an extended visit to Codfish Island in search of M. zelandica, collecting several other insects on the side.

Some of the species Beverley studied are on the brink of extinction. Many are range-restricted and are of high conservation value. Among the 35 beetles ranked as Nationally Critical (Leschen et al. 2012), three lucanid beetles and one anthribid weevil were listed, and three of these had been described by Beverley. There were seven beetles listed as Nationally Endangered, including two lucanids that Beverley had named. Three are classed as Nationally Vulnerable, including one lucanid.

While Beverley’s research has been instrumental towards understanding and conserving the New Zealand fauna, her work contained information of global importance. Her treatment of the New Zealand Anthribidae is unmatched: nothing like this exists in the literature for this family. With respect to lucanid beetle classification, she provided a world view on the classification that began with her seminal work in 1960s. An example is the endemic Californian genus Diphyllostoma Fall, which consists of three species that have flightless females with vestigial wings. The larvae remain unknown. These beetles were included in Lucanidae until 1972 when Beverley named Diphyllostomatidae as a separate family based on several characters. Though recent molecular data indicate that the group may be nested within Lucanidae, it may eventually be recognised as a distinct subfamily. This is the nature of classification - never permanent and subject to revision. Her views on the classification of this family, and the subfamilies, especially the limits of Aesalinae, require further study.

We couldn’t imagine a world without Beverley. She had laid down groundwork for future biologists and her work is stellar and complete. In recognition of this achievement, she was honoured by nine species patronyms (Table 1). Beverley’s other honours include a New Zealand Commemoration Medal in 1990 for services to science, and her election as a Fellow of the Entomological Society of New Zealand in 2008.

Table 1. List of Patronyms for Beverley Holloway.

Species	Author	Notes
Allograpta hollowayae	Thompson & Thompson 2010
Androporus hollowayae	Kuschel 1998	New Caledonia; paper dedicated to Beverley by Willy.
Aparua hollowayi [sic]	Forster, 1968	Now in Stanwellia
Austropsocus hollowayae	Smithers 1969
Fannia hollowayae	Dominguez & Pont 2014
Kuschelysius hollowayae	Brown & Leschen 2018
Migas hollowayi [sic]	Wilton, 1968
Omanuperla hollowayae	McLellan 1991
Perroudius hollowayae	Kuschel 1998
Plectophanes hollowayae	Forster, 1964
Pounamua hollowayi [sic]	Forster, 1956	Now in Pounamuella
Triainomyces hollowayanus	W. Rossi & A. Weir, 1998

Beverley provided an exemplary model of how to balance the dual commitments of family and systematic entomological research. Outside of her entomological work, she lived a rich and full life, with varied interests including baking, knitting and playing the guitar and the piano. Always exceptionally generous and hospitable, Beverley and Willy’s home in Tropicana Avenue, Lynfield, Auckland, became a destination for many visiting entomologists. The dining table was where entomological, linguistic, and political issues would be discussed and analysed; but in the kitchen, Beverley would turn topics towards family and personal matters. Beverley was predeceased by Willy in 2017 and is survived by her three children and four grandchildren.

Publications of Beverley Anne Holloway

Holloway, B. A. 1954. New locality records for Propolyxenus forsteri Condé (Diplopoda Polyxenidae). New Zealand Entomologist 1(4): 12-13. doi:10.1080/00779962.1954.9722725

Holloway, B. A. 1954. Notes on Lepidoptera. New Zealand Entomologist 1(4): 13-14, doi:10.1080/00779962.1954.9722726

Holloway, B. A. 1956. A new species of Rallicola (Insecta: Mallophaga) from Notornis mantelli Owen. Records of the Dominion Museum 2(3): 113-119.

Holloway, B. A. 1956. Revision of the New Zealand pill millipedes (Oniscomorpha, Sphaerotheridae). Transactions of the Royal Society of New Zealand 84(2): 431-446.

Holloway, B. A. 1960. Taxonomy and phylogeny in the Lucanidae (Insecta: Coleoptera). Records of the Dominion Museum 3(4): 321-365

Holloway, B. A. 1961. A systematic revision of New Zealand Lucanidae (Insecta: Coleoptera). Dominion Museum Bulletin No. 20, 139 pp.

Holloway, B. A. 1962. A new species of Ceratognathus Westwood (Coleoptera: Lucanidae) from New Zealand. Transactions of the Royal Society of New Zealand (zoology) 2: 69-72.

Holloway, B. A. 1962. Melanitis leda (Linn.) and other migrant butterflies in New Zealand during 1962. Records of the Dominion Museum [Wellington NZ] 4(8): 79-82.

Holloway, B. A. 1963. A new species of Lissotes Westwood from New Zealand, and a description of the female of L. oconnori Holloway (Coleoptera: Lucanidae). Transactions of the Royal Society of New Zealand Zoology 3(7): 77-80.

Holloway, B. A. 1963. Wing development and evolution of New Zealand Lucanidae (Insecta: Coleoptera). Transactions of the Royal Society of New Zealand (zoology) 3(11): 99-116.

Holloway, B. A. 1968. Relationships of Syndesus MacLeay and Sinodendron Schneider (Coleoptera-Lucanidae). New Zealand Journal of Science 11(2): 264-269.

Holloway, B. A. 1969. Further studies on generic relationships in Lucanidae (Insecta: Coleoptera) with special reference to the ocular canthus. New Zealand Journal of Science 12: 958-977.

Holloway, B. A. 1970. A new genus of New Zealand Anthribidae associated with lichens (Insecta: Coleoptera). New Zealand Journal of Science 13: 435-446.

Holloway, B. A. 1971. Entomology of the Aucklands and other islands south of New Zealand: Coleoptera: Anthribidae. Pacific Insects Monograph 27: 261-270.

Holloway, B. A. 1972. The systematic position of the genus Diphyllostoma Fall (Coleoptera: Scarabaeoidea). New Zealand Journal of Science 15: 31–38.

Holloway, B. A. 1976. A new bat-fly family from New Zealand (Diptera: Mystacinobiidae). New Zealand Journal of Zoology 3(4): 279-301, doi:10.1080/03014223.1976.9517919

Holloway, B. A. 1976. Pollen-feeding in hover-flies (Diptera: Syrphidae). New Zealand Journal of Zoology 3(4): 339-350, doi:10.1080/03014223.1976.9517924

Holloway, B. A. 1982. Anthribidae (Insecta: Coleoptera). Fauna of New Zealand 3, 265 pp. 2nd impression 1985 with taxonomic index (265-268) and corrigenda (269). doi:10.7931/J2/FNZ.3

Holloway, B. A. 1983. Species of ragwort seedflies imported into New Zealand (Diptera: Anthomyiidae). New Zealand Journal of Agricultural Research 26(2): 245-249, doi:10.1080/00288233.1983.10427068

Holloway, B. A. 1984. Larvae of New Zealand Fanniidae (Diptera: Calyptrata). New Zealand Journal of Zoology 11(3): 239-257, doi:10.1080/03014223.1984.10428238

Holloway, B. A. 1984. Morphology of pronotal carinae and associated surface structures in New Zealand Anthribidae (Insecta: Coleoptera). New Zealand Journal of Zoology 11(4): 445-450, doi:10.1080/03014223.1984.10428259

Smeeton, W. M.I.; Koelmeyer, T. D.; Holloway, B. A.; Singh, P. 1984. Insects associated with exposed human corpses in Auckland, New Zealand. Medicine, Science and the Law 24(3): 167-174. doi:10.1177/002580248402400304

Holloway, B. A. 1985. Comparative morphology of setiferous sex patches in male Anthribidae (Insecta: Coleoptera). New Zealand Journal of Zoology 12(2): 159-167, doi:10.1080/03014223.1985.10428275

Holloway, B. A. 1985. Immature stages of New Zealand Calliphoridae. Calliphoridae (Insecta: Diptera). Fauna of New Zealand. 8: 12-14. doi:10.7931/J2/FNZ.8

Christmas T. I.; Nicholls, D.; Holloway, B. A; Greig, D. 1987. Blister beetle dermatosis in New Zealand. The New Zealand Medical Journal, 01 Aug 1987, 100(830): 515-517. PMID: 3330591

Holloway, B. A. 1991. Identification of third-instar larvae of flystrike and carrion-associated associated blowflies in New Zealand (Diptera: Calliphoridae). New Zealand Entomologist 14(1): 24-28. doi:10.1080/00779962.1991.9722608

Holloway, B. A. 1991. Morphological characters to identify adult Lucilia sericata (Meigen, 1826) and L. cuprina (Wiedemann, 1830) (Diptera: Calliphoridae). New Zealand Journal of Zoology 18(4): 413-420. doi:10.1080/03014223.1991.10422847

Palmer, P. H.; Tingey, R. E.; Holloway, B. A. 1992. Human ophthalmomyiasis externa caused by larvae of the sheep nasal bot-fly, Oestrus ovis. The New Zealand medical journal 105(929): 84-85. PMID: 1545943.

Holloway, B. A. 1996. Two new genera of New Zealand stag beetles previously placed in Dorcus MacLeay and Lissotes Westwood (Coleoptera: Lucanidae). New Zealand Journal of Zoology 23: 61-66, doi:10.1080/03014223.1996.9518065

Holloway, B. A. 1997. Elytral surface structures as indicators of relationships in stag beetles, with special reference to the New Zealand species (Coleoptera: Lucanidae). New Zealand Journal of Zoology 24(1): 47-64, doi:10.1080/03014223.1997.9518105

Holloway, B. A. 1998. A re-evaluation of the genera of New Zealand aesaline stag beetles (Coleoptera: Lucanidae). Journal of the Royal Society of New Zealand 28(4): 641-656, doi:10.1080/03014223.1998.9517578

Holloway, B. A. 2007. Lucanidae (Insecta: Coleoptera). Fauna of New Zealand 61. 254 pp. doi:10.7931/J2/FNZ.61

Theses

Holloway, B. A. 1953. Studies on Procyliosoma tuberculatum Silv. (Diplopoda: Oniscomorpha). A Thesis Submitted to Victoria University of Wellington in partial fulfilment of the requirements for the Degree of Master of Science in Zoology.

Holloway, B. A. 1959. A Revision of the New Zealand Lucanidae (Coleoptera). Doctoral dissertation, Radcliffe College, Harvard University.

Awards

1955 – Fulbright Scholarship

1957 – Helen Marr Kirby Fellowship from the American Association of University Women 1990 – New Zealand Commemoration Medal

2008 – Elected as a Fellow of the Entomological Society of New Zealand