• The importance of long-term genetic monitoring of reintroduced populations: inbreeding in the natterjack toad (Epidalea calamita)

      Muir, Anna P; orcid: 0000-0002-6896-6915; Phillips, Susanna; Geary, Matt; email: m.geary@chester.ac.uk; Allmark, Matthew; Bennett, Sarah; Norman, Kim; Ball, Rachel J; Peters, Catherine; University of Chester; Cheshire Wildlife Trust; Eni UK Ltd
      Genetic monitoring is an important, but frequently lacking, component of management actions to support long-term persistence in reintroduced populations. Populations that remain small, due to demographic processes and genetic diversity, are more likely to experience a second extinction event. The natterjack toad (Epidelea calamita) is legally protected in Britain and was the subject of a reintroduction programme in the 1990s. However, subsequent genetic assessment has been mostly lacking. The aim of this study was to assess the genetic diversity of two reintroduced populations of natterjack toads in order to inform conservation management. Adults were sampled and nine microsatellites amplified to assess neutral genetic variation within each site and for comparison with the source population. Inbreeding was observed at the reintroduction sites, as evidenced by high FIS values (0.43 and 0.72), low observed compared to expected heterozygosities, and significant deviation from Hardy-Weinberg equilibrium. Observed heterozygosity is currently lower in the reintroduction sites than it was in the source population at the time of the reintroductions (Red Rocks: 0.15±0.20; Talacre: 0.12±0.20; Ainsdale (source): 0.29). Evidence for a bottleneck was not found, although this is likely a result of sampling overlapping generations. No within-site population structuring was observed. Such low genetic diversity has not previously been recorded in any natterjack population. Genetic rescue, combined with pool creation, is the most viable option for safeguarding the species at these sites into the future. Our work highlights the importance of ongoing genetic monitoring, in collaboration with conservation organisations, to support conservation management.
    • Patterns in island endemic forest-dependent bird research: the Caribbean as a case-study

      Devenish-Nelson, Eleanor S.; orcid: 0000-0002-9029-4772; email: ellie.devenish@ed.ac.uk; Weidemann, Douglas; Townsend, Jason; Nelson, Howard P. (Springer Netherlands, 2019-05-04)
      Abstract: Unequal patterns in research effort can result in inaccurate assessments of species extinction risk or ineffective management. A group of notable conservation concern are tropical island endemic birds, many of which are also forest-dependent, which increases their vulnerability to extinction. Yet, island bird species have received limited research attention compared to their continental congeners, despite this taxon being globally regarded as well-studied. We used the insular Caribbean, a globally important endemism hotspot with high rates of deforestation, to explore research bias of island and regional endemic forest-dependent birds. A review of the published literature (n = 992) found no significant increase in the number of studies over the search period. Research effort was significantly higher among species with threatened status, long generation time, wide habitat breadth and low to intermediate elevational distributions. Among family groups, the Psittacidae received the highest research effort, while the Cuculidae were the most underrepresented family (30-fold higher and six-fold less than expected, respectively). We found geographic biases in effort, with Jamaica having six-fold less and Puerto Rico eight times more research than expected for their level of endemism. These patterns likely reflect individual interests and limited capacity and funding, typical of Small Island Developing States. With over 50% of species in this review having declining population trends, we recommend prioritizing research that emphasises conservation- and management-relevant data across underrepresented families and islands, by fostering greater collaboration between researchers, practitioners and the existing local amateur ornithological community.
    • Mangrove or mudflat: Prioritising fish habitat for conservation in a turbid tropical estuary

      Marley, Guy; Deacon, Amy; Philip, Dawn; Lawrence, Andrew; University of the West Indies, University of the West Indies, University of the West Indies, University of Chester
      Mangrove habitats are typically the focus of conservation efforts in tropical estuaries because their structural complexity is thought to support greater biodiversity and nursery function than unvegetated habitats. However, evidence for this paradigm has been equivocal in turbid tropical estuaries where unvegetated mudflats are also highly productive. The present study compared the community composition, biodiversity, nursery-role and commercial fish biomass in two mangrove habitats and one mudflat habitat in the Gulf of Paria, Trinidad. A total of 12 705 fishes, comprising 63 species from 26 families, were sampled in mangrove creeks, seaward mangrove fringe and the subtidal margin of an intertidal mudflat from June 2014 to June 2015. The composition of the creek and mudflat communities were distinct, while the community of the mangrove fringe more closely resembled the mudflat than the mangrove creeks. Mean species richness (MSR), total species richness (TSR) extrapolated from species accumulation curves, and juvenile species richness (JSR) were significantly greater in the mudflat (MSR = 11.4 ± 1.0; TSR = 75 ± 14; JSR = 9.1 ± 0.8) than mangrove creeks (MSR = 9.0 ± 0.5; TSR = 49 ± 3; JSR = 6.1 ± 0.4) and the seaward mangrove fringe (MSR = 6.4 ± 0.7; TSR = 58 ± 14; JSR = 5.2 ± 0.4). Meanwhile, Shannon Weiner diversity, juvenile fish abundance and commercial fish biomass were comparable between habitats. These findings caution against the generalisation that mangroves are the most important habitat for fishes in turbid tropical estuaries. There is now a growing body of evidence that mudflats warrant consideration as important repositories of biodiversity and nursery function for juvenile fishes.
    • A review of tropical dry forest ecosystem service research in the Caribbean – gaps and policy-implications

      Nelson, Howard; Devenish-Nelson, Eleanor; Rusk, Bonnie; Geary, Matthew; Lawrence, Andrew; University of Chester;University of Edinburgh; Grenada Dove Conservation Programme
      Tropical dry forests (TDFs) are globally threatened, yet remain poorly studied. In the Caribbean, the most biodiverse of island biodiversity hotspots, TDFs have structural properties distinct from the Neotropical mainland and are important to local communities for ecosystem services. We undertook a systematic review (n = 186) of ecosystem services literature of Caribbean TDF. Only 19.89% qualified for inclusion, with the majority (56.76%) from primary literature. Research on supporting services (31.14%), particularly primary production was predominant. Most studies (70.97%) took a biophysical perspective and quantification focused on the supply of ecosystem services (43.00%), while measurement of wellbeing benefits were uncommon. Geographic coverage of all studies was patchy originating from only nine of 28 independent countries and dependent territories. Our findings highlight a lack of research, while accentuating the value of grey literature in quantifying cultural services. Of concern, are gaps in air- and water-related services and the importance of TDF to human health. To move from biophysical assessments to a broader portfolio of ecosystem services studies, research on Caribbean TDF should be collaborative and strategic. Such gaps and research biases suggest opportunities for evidence-led policy-making. These lessons are relevant for mainstreaming ecosystem services into decision-making in Small Island Developing States.
    • Current strength, temperature, and bodyscape modulate cleaning services for giant manta rays

      Murie, Calum; Spencer, Matthew; Oliver, Simon; University of Chester (Springer, 2020-04-07)
      The cleaner-client system among reef teleosts has received considerable attention in both wild and captive environments, but the spatially and taxonomically diverse associations between cleaner fish and elasmobranchs are less understood. Using remote video, we investigated interactions between giant manta rays (Mobula birostris) and cleaner wrasse at a seamount in the Philippines. Cleaning events occurred between 11:00 and 16:00 hours on a seasonal basis and were constrained by current strengths and ambient water temperatures. The frequency with which giant manta rays interacted with cleaner fish varied on an individual basis. Blue streaked cleaner wrasse (Labroides dimidiatus) and moon wrasse (Thalassoma lunare) selectively foraged on manta rays’ gills and pelvis, with L. dimidiatus also demonstrating slight preferences for the pectoral fins. Cleaners’ foraging preferences may indicate ectoparasitic infections in specific areas of a manta ray’s body. The exclusivity with which giant manta rays visited a particular cleaning station on the seamount may be a response to the quality of services that cleaners provide there. Giant mantas’ fidelity to this site may also be attributed to localised concentrations of food that are available nearby. The seamount provides habitat that appears to be important to the life history strategies of the region’s giant manta rays.
    • Alpine ibex, Capra ibex, Linnaeus 1758

      Brambillla, Alice; Bassano, Bruno; Biebach, Iris; Bollmann, Kurt; Keller, Lukas; Toïgo, Carole; von Hardenberg, Achaz; University of Zurich, Gran Paradiso National Park, Swiss Federal Research Institute, Office National de la Chasse et de la Faune Sauvage, University of Chester
      In this chapter we review the current status of knowledge about Alpine ibex Capra ibex. We cover taxonomy, systematics, distribution, habitat, genetics, life history, behaviour, parasites and disease, population ecology and conservation. We conclude examining the future challenges for research and management of this species.
    • An introduction to Phylogenetic Path Analysis

      Gonzalez-Voyer, A.; von Hardenberg, Achaz; Estación Biológica de Doñana, Gran Paradiso National Park (Springer Verlag, 2014)
      The questions addressed by macroevolutionary biologists are often impervious to experimental approaches, and alternative methods have to be adopted. The phy- logenetic comparative approach is a very powerful one since it combines a large number of species and thus spans long periods of evolutionary change. However, there are limits to the inferences that can be drawn from the results, in part due to the limitations of the most commonly employed analytical methods. In this chapter, we show how confirmatory path analysis can be undertaken explicitly controlling for non-independence due to shared ancestry. The phylogenetic path analysis method we present allows researchers to move beyond the estimation of direct effects and analyze the relative importance of alternative causal models including direct and indirect paths of influence among variables. We begin the chapter with a general introduction to path analysis and then present a step-by-step guide to phylogenetic path analysis using the d-separation method. We also show how the known statistical problems associated with non-independence of data points due to shared ancestry become compounded in path analysis. We finish with a discussion about the potential effects of collinearity and measurement error, and a look toward possible future developments.
    • Distribution, status and recent population dynamics of Alpine ibex Capra ibex in Europe

      Brambillla, Alice; von Hardenberg, Achaz; Nelli, Luca; Bassano, Bruno; University of Zurich, University of Chester, University of Glasgow, Gran Paradiso National PArk (Wiley, 2020-04-20)
      1. Despite its recent successful and well-documented reintroduction history, a comprehensive and current update of the distribution and status of the Alpine ibex Capra ibex is lacking. As some concerns persist about its conservation, a status update appears essential for future conservation and management strategies on a large scale. 2. We provide an exhaustive update of the geographic range of the species, alongside estimates of its current abundance and population trends from 2004 to 2015. 3. We gathered census and distribution data for all the Alpine ibex colonies from management authorities and research groups that monitor them in different countries, and from the literature and publicly available reports. We produced a distribution map, reported the number of individuals observed in the most recent censuses, and estimated global, national, and local population trends using Bayesian hierarchical models. 4. Our model estimated that there were a total of 55297 Alpine ibex in the Alps in 2015 (lower 95% Credible Interval [CrI]: 51157; upper 95% CrI: 62710). The total number of individuals appears to have increased slightly over the last 10 years from the 47000-51000 estimated in previous reports. Positive population trends were observed in Switzerland and Italy, while no trend was apparent in France. For Austria, Germany, and Slovenia, there were insufficient data to estimate a trend. The slopes of the colonies’ trends were positively correlated with the year of colony foundation. 5. The geographic range of the Alpine ibex does not seem to have increased in size in recent years, although the accuracy of the spatial data varies among countries. 6. The periodic and standardised collection of census data for all colonies and a common policy of data-sharing at a European level appear essential for monitoring the global trend of this species and for planning balanced conservation and management actions.
    • Improving the practicality of using non-aversive handling methods to reduce background stress and anxiety in laboratory mice

      Gouveia, Kelly; Hurst, Jane; University of Liverpool; University of Chester
      Handling can stimulate stress and anxiety in laboratory animals that negatively impacts welfare and introduces a confounding factor in many areas of research. Picking up mice by the tail is a major source of handling stress that results in strong aversion to the handler, while mice familiarised with being picked up in a tunnel or cupped on the open hand show low stress and anxiety, and actively seek interaction with their handlers. Here we investigate the duration and frequency of handling required for effective familiarisation with these non-aversive handling methods, and test whether this is sufficient to prevent aversion and anxiety when animals then experience immobilisation and a mild procedure (subcutaneous injection). Very brief handling (2 s) was sufficient to familiarise mice with tunnel handling, even when experienced only during cage cleaning. Brief but more frequent handling was needed for familiarisation with cup handling, while pick up by tail induced strong aversion even when handling was brief and infrequent. Experience of repeated immobilisation and subcutaneous injection did not reverse the positive effects of tunnel handling. Our findings demonstrate that replacing tail with tunnel handling during routine cage cleaning and procedures provides a major refinement with little if any cost for familiarisation.
    • Combining bioacoustics and occupancy modelling for improved monitoring of rare breeding bird populations

      Abrahams, Carlos; Geary, Matthew; Baker Consultants Ltd; Nottingham Trent University; University of Chester
      Effective monitoring of rare and declining species is critical to enable their conservation, but can often be difficult due to detectability or survey constraints. However, developments in acoustic recorders are enabling an important new approach for improved monitoring that is especially applicable for long-term studies, and for use in difficult environments or with cryptic species. Bioacoustic data may be effectively analysed within an occupancy modelling framework, as presence/absence can be determined, and repeated survey events can be accommodated. Hence, both occupancy and detectability estimates can be produced from large, coherent datasets. However, the most effective methods for the practical detection and identification of call data are still far from established. We assessed a novel combination of automated clustering and manual verification to detect and identify heathland bird vocalizations, covering a period of six days at 44 sampling locations Occupancy (Ψ) and detectability (p ) were modelled for each species, and the best fit models provided values of: nightjar Ψ=0.684, p=0.740, Dartford warbler Ψ=0.449 p=0.196 and woodlark Ψ=0.13 p=0.996. Including environmental covariates within the occupancy models indicated that tree, wetland and heather cover were important variables, particularly influencing detectability. The protocol used here allowed robust and consistent survey data to be gathered, with limited fieldwork resourcing, allowing population estimates to be generated for the target bird species. The combination of bioacoustics and occupancy modelling can provide a valuable new monitoring approach, allowing population trends to be identified, and the effects of environmental change and site management to be assessed.
    • Mangrove and mudflat food webs are segregated across four trophic levels, yet connected by highly mobile top predators

      Marley, Guy; Lawrence, Andrew; Phillip, Dawn; Hayden, Brian; Canadian Rivers Institute
      Seascape connectivity is crucial for healthy, resilient ecosystems and fisheries. Yet, our understanding of connectivity in turbid mangrove-lined estuaries—some of the world’s most productive ecosystems—is limited to macrotidal systems, and rarely incorporates highly mobile top predators. We analysed δ13C and δ15N isotope values of 7 primary producers, 24 invertebrate taxa, 13 fishes, 4 birds and 1 reptile to reveal trophic interactions within and between a mangrove and adjacent mudflat in a microtidal system of the Gulf of Paria, Orinoco River estuary. Primary producers, invertebrates and fishes collected within the mangrove were significantly depleted in 13C and 15N compared to those collected on the mudflat. Stable isotope mixing models showed that mangrove-derived carbon was predominantly assimilated by invertebrates (78 ± 5% SE) and fishes (88 ± 11%) sampled in the mangrove. In contrast, invertebrates and fishes sampled in the mudflat derived <21% of their carbon from mangrove sources. Instead, microphytobenthos and phytoplankton underpinned the mudflat food web. Scarlet ibis Eudocimus ruber and yellow-crowned night heron Nyctanassa violacea were also highly associated with mangrove carbon sources. However, osprey Pandion haliaetus, snowy egret Egretta thula and spectacled caiman Caiman crocodilus obtained carbon from both mangrove and mudflat sources, effectively integrating the food webs. The present study demonstrates simultaneous aspects of food web segregation and connectivity, as well as the importance of surveying the entire food web across a range of tidal systems when investigating seascape connectivity.
    • A novel method to optimise the utility of underused moulted plumulaceous feather samples for genetic analysis in bird conservation.

      Peters, Catherine; Nelson, Howard, P.; Rusk, Bonnie; Muir, Anna P.; Rusk, Bonnie L.; University of Chester (Springer, 2019-10-24)
      Non-invasive sampling methods are increasingly being used in conservation research as they reduce or eliminate the stress and disturbance resulting from invasive sampling of blood or tissue. Here we present a protocol optimised for obtaining usable genetic material from moulted plumulaceous feather samples. The combination of simple alterations to a ‘user-developed’ method, comprised of increased incubation time and modification of temperature and volume of DNA elution buffer, are outlined to increase DNA yield and significantly increase DNA concentration (W = 81, p <0.01, Cohens’s d= 0.89). We also demonstrate that the use of a primerless Polymerase Chain Reaction (PCR) technique increases DNA quality and amplification success when used prior to PCR reactions targeting avian mitochondrial DNA (mtDNA). A small amplicon strategy proved effective for mtDNA amplification using PCR, targeting three overlapping 314-359bp regions of the cytochrome oxidase I barcoding region which, when combined, aligned with target-species reference sequences. We provide evidence that samples collected non-invasively in the field and kept in non-optimal conditions for DNA extraction can be used effectively to sequence a 650bp region of mtDNA for genetic analysis.
    • Preparation of Primary Rat Hepatocyte Spheroids Utilizing the Liquid-Overlay Technique.

      Kyffin, Jonathan A; Cox, Christopher R; Leedale, Joseph; Colley, Helen E; Murdoch, Craig; Mistry, Pratibha; Webb, Steven D; Sharma, Parveen (2019-09)
      Herein, we describe a protocol for the preparation and analysis of primary isolated rat hepatocytes in a 3D cell culture format described as spheroids. The hepatocyte cells spontaneously self-aggregate into spheroids without the need for synthetic extracellular matrices or hydrogels. Primary rat hepatocytes (PRHs) are a readily available source of primary differentiated liver cells and therefore conserve many of the required liver-specific functional markers, and elicit the natural in vivo phenotype when compared with common hepatic cells lines. We describe the liquid-overlay technique which provides an ultra-low attachment surface on which PRHs can be cultured as spheroids. © 2019 The Authors. Basic Protocol 1: Preparation of agarose-coated plates Basic Protocol 2: Primary rat hepatocyte isolation procedure Basic Protocol 3: Primary rat hepatocyte spheroid culture Basic Protocol 4: Immunofluorescent analysis of PRH spheroids. [Abstract copyright: © 2019 The Authors.]
    • “You Can’t Really Hug a Tiger”: Zookeepers and Their Bonds with Animals

      Birke, Lynda; Hosey, Geoff; Melfi, Vicky (Informa UK Limited, 2019-09-20)
    • Individual, social, and environmental factors affecting salivary and fecal cortisol levels in captive pied tamarins (Saguinus bicolor)

      Wormell, Dominic; Smith, Tessa E.; Price, Eluned E.; Ahsmann, J.; Glendewar, G.; Hunt, J.; Coleman, Robert, C.; University of Chester (Wiley, 2019-08-01)
      Pied tamarins (Saguinus bicolor) are endangered New World primates, and in captivity appear to be very susceptible to stress. We measured cortisol in 214 saliva samples from 36 tamarins and in 227 fecal samples from 27 tamarins, and investigated the effects of age, sex, pregnancy, rearing history, social status, weight, group composition, and enclosure type using generalized linear mixed models. There was no effect of age on either fecal or salivary cortisol levels. Female pied tamarins in late pregnancy had higher fecal cortisol levels than those in early pregnancy, or nonpregnant females, but there was no effect of pregnancy on salivary cortisol. Females had higher salivary cortisol levels than males, but there was no effect of rearing history. However, for fecal cortisol, there was an interaction between sex and rearing history. Hand‐reared tamarins overall had higher fecal cortisol levels, but while male parent‐reared tamarins had higher levels than females who were parent‐ reared, the reverse was true for hand‐reared individuals. There was a trend towards lower fecal cortisol levels in subordinate individuals, but no effect of status on salivary cortisol. Fecal but not salivary cortisol levels declined with increasing weight. We found little effect of group composition on cortisol levels in either saliva or feces, suggesting that as long as tamarins are housed socially, the nature of the group is of less importance. However, animals in off‐show enclosures had higher salivary and fecal cortisol levels than individuals housed on‐show. We suggest that large on‐show enclosures with permanent access to off‐exhibit areas may compensate for the effects of visitor disturbance, and a larger number of tamarins of the same species housed close together may explain the higher cortisol levels found in tamarins living in off‐show accommodation, but further research is needed.
    • The importance of research applicability

      McLennan, Krista M.; University of Chester (The Humane Society Institute for Science and Policy, 2019-06-30)
      Marino & Merskin’s (2019) review contains key information about the complexity of sheep and their intelligence level, but lacks practical application. The key to making any long-term changes to sheep welfare at an industry level is by generating research that is practically relevant to the sector. The practical application of research should be considered at the design stage and in consultation with producers. Additionally, thought needs to be given to how the practical application of the research will be transferred to those people directly involved in animal care (e.g. producers, stockpersons, etc.). Focusing on the practical relevance and application of research at all stages of the process will foster changes to long-held beliefs and attitudes.
    • Conceptual and methodological issues relating to pain assessment in mammals: the development and utilisation of pain facial expression scales.

      McLennan, Krista M.; Miller, Amy, L.; Dalla Costa, Emanuela; Stucke, Diana; Corke, Murray J.; Broom, Donald M.; Leach, Matthew C.; University of Chester; Newcastle University; c Università degli Studi di Milano; Havelland Equine Clinic; University of Cambridge; University of Cambridge; Newcastle University (Elsevier, 2019-06-12)
      Effective management of pain is critical to the improvement of animal welfare. For this to happen, pain must be recognised and assessed in a variety of contexts. Pain is a complex phenomenon, making reliable, valid, and feasible measurement challenging. The use of facial expressions as a technique to assess pain in non-verbal human patients has been widely utilised for many years. More recently this technique has been developed for use in a number of non-human species: rodents, rabbits, ferrets, cats, sheep, pigs and horses. Facial expression scoring has been demonstrated to provide an effective means of identifying animal pain and in assessing its severity, overcoming some of the limitations of other measures for pain assessment in animals. However, there remain limitations and challenges to the use of facial expression as a welfare assessment tool which must be investigated. This paper reviews current facial expression pain scales (“Grimace Scales"), discussing the general conceptual and methodological issues faced when assessing pain, and highlighting the advantages of using facial expression scales over other pain assessment methods. We provide guidance on how facial expression scales should be developed so as to be valid and reliable, but we also provide guidance on how they should be used in clinical practice.
    • In vitro cyto-toxic assessment of heavy metals and their binary mixtures on mast cell-like, rat basophilic leukemia (RBL-2H3) cells

      Eze, Chukwuebuka T.; Michelangeli, Francesco; Otitoloju, Adebayo A.; University of Chester, University of Lagos, Nigeria, Federal University Oye-Ekiti, Nigeria (Elsevier, 2019-02-11)
      We investigated the cytotoxicity and mechanisms of cell death induced by salts of Cadmium (Cd2+), Lead (Pb2+), Arsenic (AsO4 3−) and Chromium (Cr+6) on RBL-2H3 cells (a model mast cell line). In addition, cyto-toxic effect on cell viability was assessed to reveal their nature of interaction in binary mixture. The individual cytotoxic characteristics of these metals on RBL-2H3 cell viability showed a concentration- dependent reduction of cell viability. We observed that concentration-dependent cytotoxic potency on RBL-2H3 cells of these metals range in the following order Cd2+>Cr+6>As O4 3−>Pb2+ with LC50 values of 0.11 μM, 93.58 μM, 397.9 μM and 485.3 μM respectively. Additive effects were observed with Pb2+ + Cd2+, Pb2+ + AsO4 3−, Pb2+ + Cr+6 and AsO4 3− + Cr+6. The study revealed that Pb2+, Cd2+, AsO4 3− and Cr+6 could induce significant (P<0.01) cell death by apoptosis in RBL-2H3. Highly significant necrotic cell death was observed with Pb2+ and Cr+6 (P<0.01) than Cd2+ and AsO4 3− (P<0.05). Overall, it can be deduced that several cell death executing pathways may be concomitantly activated on exposure to heavy metals and the predominance of one over others might depend on the type of heavy metal, concentration and the metabolic state of the cell. Eventually, binary mixtures of some of these metals showed less cytotoxicity than would be expected from their individual actions and may depend on the co-exposure of the metal ions and their modes of action.
    • Ida Slater: A Collection Researcher in a Male World at the Beginning of the 20th Century

      Sendino, Consuelo; Ducker, Erik; Burek, Cynthia V. (SAGE Publications, 2019-06-27)
      Ida Lilian Slater (1881-1969) was one of the first women to work as a geologist in a male world, and although her career was short, she made important contributions to the Early Palaeozoic of Wales and Scotland. Her main work was based on a collection of a group of fossil scypho-zoan polyps gathered not by her but by another significant woman, Elizabeth Anderson, widely known as Mrs. Robert Gray (1831-1924). The majority of this collection is kept at the Natural History Museum (NHM), London, and the Sedgwick Museum, Cambridge. She worked in the former one for two years describing species and comparing specimens for her monograph on British conulariids. Although her work was based not only on this group, she will be remembered by her important contribution to the conulariids through collections. The NHM collection is considered the best in the world in terms of diversity and the second best in its number of specimens, while the Sedgwick Museum has a smaller collection that is still considered the second best in diversity and number of specimens in the United Kingdom. Her work has been cited for more than 100 years and continues to be cited to this day by researchers on this group of fossils.